Conservation Biology of Lycaenidae (Butterflies) - IUCN

The IUCN Species Survival Commission 

Conservation Biology of 

Lycaenidae 

(Butterflies) 

Edited by T.R. New 

Occasional Paper of the IUCN Species Survival Commission No.8 

IUCN 

The World Conservation Union

Conservation Biology of Lycaenidae was made possible through the generous support of: 

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DEJA, Inc. 

IUCN/SSC Peter Scott Action Plan Fund (Sultanate of Oman) 

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World Wide Fund For Nature 

© 1993 International Union for Conservation of Nature and Natural Resources. 

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ISBN 2-8317-0159-7 

Published by IUCN, Gland, Switzerland. 

Camera-ready copy by The Nature Conservation Bureau Limited, 

36 Kingfisher Court, Hambridge Road, Newbury RG14 5SJ, U.K. 

Printed by Information Press, Oxford, U.K. 

Cover Photo: Karner Blue, Lycaeides melissa samuelis (Photo: Richard A. Arnold).

The IUCN Species Survival Commission 

Conservation Biology of 


(Butterflies) 

Edited by T.R. New

Preface iv 

Acknowledgment v 

List of Contributors 

PART 1. INTRODCTION 

Introduction to the Biology and Conservation of the 


T.R. NEW 1 

PART 2. REGIONAL ASSESSMENTS 

Introductory Comment 22 

Conservation biology of Lycaenidae: a European overview 

M.L. MUNGUIRA, J. MARTIN & E. BALLETTO 23 

Overview of problems in Japan 

T. HlROWATARI 35 

Conservation of North American lycaenids- an overview 

J. HALL CUSHMAN & D.D. MURPHY 37 

Neotropical Lycaenidae: an overview 

K.S. BROWN, JR. 45 

Threatened Lycaenidae of South Africa 

M.J. SAMWAYS 62 

Australian Lycaenidae: conservation concerns 

T.R. NEW 70 

PART 3. ACCOUNTS OF PARTICULAR TAXA OR 

COMMUNITIES 

Introductory comment 77 

Europe 

The mariposa del Puerto del Lobo, Agriades zullichi 

M.L. MUNGUIRA & J. MARTIN 78 

The Large Copper, Lycaena dispar 

E. DUFFEY 81 

The Adonis Blue, Lysandra bellargus 83 

Large Blues, Maculinea spp 85 

Polyommatus humedasae 

E. BALLETTO 88 

Polyommatus galloi 

E. BALLETTO 90 

The Sierra Nevada Blue, Polyommatus golgus 


Tomares ballus 

H.A. DESCIMON 95 

The Silver-studded Blue, Plebejus argus 

CD. THOMAS 97 

The Zephyr Blue, Plebejus pylaon 


Contents 

vi 

ii 

The Pannonian Zephyr Blue, Plebejus sephirus 

kovacsi 

Z. BÁLINT 103 

The threatened lycaenids of the Carpathian Basin, 

east-central Europe 

Z. BÁLINT 105 

Aricia macedonica isskeutzi 105 

Aricia eumedon 105 

Cupido osiris 106 

Jolana iolas 106 

Maculinea alcon 106 

Maculinea nausithous 107 

Maculinea sevastos limitanea 107 

Plebejus (Lycaeides) idas 107 

Polyommatus (Agrodiaetus) admetus 108 

Polyommatus (Agrodiaetus) damon 108 

Polyommatus (Vacciniina) optilete 108 

Polyommatus eroides 109 

Polyommatus dorylas magna 109 

Lycaena helle 109 

Lycaena tityrus argentifex 110 

Pseudophilotes bavius hungaricus 110 

Tomares nogelii dobrogensis 110 

Japan 

Coreana raphaelis 

T.HIROWATARI 112 

Shijimiaeoides divinus 

T. T.HIROWATARI 114 

North America 

Lange's Metalmark, Apodemia mormo langei 

J.A. POWELL & M.W. PARKER 116 

The Hermes Copper, Lycaena hermes 

D.K. FAULKNER & J.W. BROWN 120 

Thome'sHairstreak, Mitoura thornei 

J.W. BROWN 122 

Sweadner's Hairstreak, Mitoura gryneus sweadneri 

T.C. EMMEL 124 

Bartram' s Hairstreak, Strymon acis bartrami 

T.C. EMMEL & M.C. MINNO 126 

The Avalon Hairstreak, Strymon avalona 

T.C. EMMEL & J.F. EMMEL 128 

TheAtala, Eumaeus atala florida 

T.C. EMMEL & M.C. MINNO 129 

Smith's Blue, Euphilotes enoptes smithi 


The El Segundo Blue, Euphilotes bernardino allyni 

R.H.T. MATTONI 133

The Palos Verdes Blue, Glaucopsyche lygdamus 

palosverdesensis 

R.H.T. MATTONI 135 

The Xerces Blue, Glaucopsyche xerces 


The Mission Blue, Plebejus icarioides missionensis 

J.H. CUSHMAN 139 

The San Bruno Elfin, Incisalia mossii bayensis 

S. B. WEISS 141 

The Lotis Blue, Lycaenides idas lotis 

R.A. ARNOLD 143 

Additional taxa of concern in southern California 

R.H.T. MATTONI 145 

Philotes sonorensis extinctis 145 

Satyrium auretorum fumosum 145 

Plebejus saepiolus ssp 145 

South America 

Selected Neotropical species 


Styx infernalis 146 

Nirodia belphegor 146 

Joiceya praeclarus 147 

Areas spp 148 

Arawacus aethesa 149 

Cyanophrys bertha 149 

Neotropical Lycaenidae endemic to high elevations in 

south eastern Brazil 


Riodininae: Amazonian genera with most species very 

rare or local 


iii 

Theclinae endemic to the Cerrado vegetation (central 

Brazil) 


Neotropical Riodininae endemic to the Chocó region 

of western Colombia 

C.J. CALLAGHAN 153 

South Africa 

Aloeides dentatis dentatis and A. d. maseruna 

S.F. HENNING, G.A. HENNING & M.J. SAMWAYS.........154 

Erikssonia acraeina 


Alaena margaritacea 


Orachrysops (Lepidochrysops) ariadne 


Australia 

Hypochrysops spp. 

D.P.A. SANDS....................................................................160 

Illidge's Ant- Blue, Acrodipsas illidgei 

P.R. SAMSON..............................................................................166 

The Eltham Copper, Paralucia pyrodiscus lucida 

T.R. NEW...............................................................................166 

The Bathurst Copper, Paralucia spinifera 

E.M. DEXTER & R.L. KITCHING..........................................168 

The Australian Hairstreak, Pseudalmenus chlorinda 

G.B. PRINCE................................................................................171 

APPENDIX 1 

IUCN Red Data Book Categories 173

Butterflies have long been accorded 'special' status by many 

people who do not like insects. Ever since the ancient Greeks 

employed the same word (psyche) for 'soul' and 'butterfly', an 

aura of spiritual or aesthetic appreciation has enhanced their 

generally charismatic popularity, so that people to whom 'the 

only good insect is a dead insect' accept readily that butterflies 

merit conservation (New 1991). Perhaps the most widely 

appreciated butterflies are the swallowtails, birdwings and their 

allies (Papilionidae), which have recently received substantial 

conservation impetus through production of a global survey 

(Collins and Morris 1985) and a Swallowtail Action Plan (New 

and Collins 1991) based on this, and produced under the 

auspices of the IUCN Species Survival Commission's 

Lepidoptera Specialist Group. 

Many other butterflies are much less well known than the 

swallowtails, and such a comprehensive appraisal of them 

would be difficult or impossible to achieve. But, especially in 

north temperate regions of the world, very substantial 

conservation effort has been directed to members of the largest 

family of butterflies, the Lycaenidae – the blues, coppers, 

hairstreaks, metalmarks and related forms. In 1989, I suggested 

that the Lepidoptera Specialist Group should seek to complement 

the swallowtail studies by an appraisal of the Lycaenidae, to 

'round out' the emerging picture of butterfly conservation by 

gathering together some of the data on this family. Lycaenidae 

exemplify a wide spectrum of concerns: populations are often 

extremely localised with colonies occupying a few hectares or 

less; many are associated with early successional stages of 

vegetation in grasslands or herb associations; and many 

participate in subtle ecological associations with ants or (more 

rarely) Homoptera. 

Preface 

iv 

A number of species or subspecies of the Lycaenidae have 

been the targets of major conservation campaigns which have 

been vitally important in raising public awareness of insect 

conservation in areas where swallowtails are scarce, and it is no 

exaggeration to claim that they have been the most important 

butterfly family in fostering conservation concern in temperate 

regions. 

This is not a Red Data Book but, rather, an introduction to 

the conservation biology of the Lycaenidae. It emphasises the 

very different knowledge base available for lycaenids compared 

with the swallowtails, and should be a salutary warning against 

feelings of complacency that butterflies are well known! 

It draws on the expertise of many experienced practitioners 

of butterfly conservation, and on the large literature of lycaenid 

biology. The account consists of three sections. The first is a 

brief general introduction to the Lycaenidae and their place in 

butterfly conservation. The second is a series of regional 

overviews of lycaenids for several parts of the world where 

interest and knowledge has been sufficient to prepare such an 

essay; and the third is a series of selected case-histories or 

species accounts which range from the well known to the novel. 

This section is not in any sense encyclopedic, but provides a 

tentative basis for direction and for future synthesis and 

development of more general conservation strategies. 

References 

COLLINS, N.M. and MORRIS, M.G. 1985. Threatened Swallowtail Butterflies 

of the World. The IUCN Red Data Book, IUCN, Gland and Cambridge. 

NEW,T.R. 1991. Butterfly Conservation. Oxford University Press, Melbourne. 

NEW, T.R. and COLLINS, N.M. 1991. Swallowtail Butterflies. An Action 

Plan for their Conservation. IUCN, Gland.

The task of preparing this volume has been a pleasant one, 

leading me to appreciate very deeply the spirit of cooperation 

engendered by a common concern for the well being of 

butterflies. Most authors approached to contribute to the 

compilation agreed promptly and willingly, and I am very 

grateful for their participation and encouragement. Other people 

responded with constructive advice and suggestions in response 

to my queries, and most members of the Lepidoptera Specialist 

Group in 1989 and 1990 suggested possible species or candidate 

authors. Dr. Simon N. Stuart and his dynamic team, including 

Acknowledgements 

V 

Dr. Mariano Gimenez-Dixon and Ms Linette Humphrey, in the 

SSC Office in Gland, have been continually encouraging and 

supportive, and their rapid responses to my numerous questions 

have been appreciated greatly. Editorial advice from Dr. 

Alexandra Hails has also been welcomed. 

Transformation of a miscellany of scripts into a relatively 

harmonic and consistent typescript was accomplished patiently 

by Mrs. Tracey Carpenter at La Trobe University; she also drew 

a number of the figures. The photographers are acknowledged 

individually in the photograph legends.

Dr. R.A. Arnold 

Entomological Consulting Services Limited, 

104 Mountain View Court, 

Pleasant Hill, 

California 94523, 

U.S.A. 

Dr. Z. Bálint 

Zoological Department, 

Hungarian Natural History Museum, 

Baross utca 13, 

Budapest, H-1008, 

Hungary 

Prof. E. Balletto 

Dipàrtimento di Biologia Animale, 

Università di Torino, 

V. Accademia Albertina 17, 

Torino, 

Italy -10123 

Dr. J.W. Brown 

Entomology Department, 

San Diego Natural History Museum, 

P.O. Box 1390, 

San Diego, 


U.S.A. 

Dr. K.S. Brown, Jr. 

Departmento de Zoologia, 

Instituto de Biologia, 

Universidade Estadual de Campinas, 

C.P. 6109. Campinas, 

Sao Paulo, 13. 081. 

Brazil 

Dr. C.J. Callaghan 

Louis Berger International Inc., 

100 Halsted Street, 

P.O. Box 270, 

East Orange, NJ 07019, 

U.S.A. 

Dr. J. Hall Cushman 

Center for Conservation Biology, 

Department of Biological Sciences, 

Stanford University, 

Stanford, California 94305, 

U.S.A. 

List of contributors 

vi 

Dr. H.A. Descimon 

Laboratoire de Systématique évolutive, 

Université de Provence, 

3 place Victor Hugo, 

13331 Marseille Cedex 3, 

France 

Ms E.M. Dexter 

School of Australian Environmental Studies, 

Griffith University, 

Nathan, 

Queensland 4111, 

Australia 

Dr. E. Duffey 

Cergne House, 

Church Street, 

Wadenhoe, 

Peterborough PE8 5ST, 

U.K. 

Dr. J.F. Emmel 

26500 Rim Road, 

Hemet, 


U.S.A. 

Dr. T.C. Emmel 

Department of Zoology, 

University of Florida, 

Gainesville, 

Florida 32611, 

U.S.A. 

Dr. D.K. Faulkner 

Entomology Department, 

San Diego Natural History Museum, 

P.O. Box 1390, 

San Diego, 


U.S.A. 

Mr. G.A. Henning 

17 Sonderend Street, 

Helderkruin 1724, 

South Africa 

Mr. S.F. Henning 

5 Alexander Street, 

Florida 1709, 

South Africa

Dr. T. Hirowatari 

Entomological Laboratory, 

College of Agriculture, 

University of Osaka Prefecture, 

Sakai, 

Osaka, 591 

Japan 

Prof. R.L. Kitching 

School of Australian Environmental Studies, 

Griffith University, 

Nathan, 

Queensland 4111, 

Australia 

Dr. J. Martin 

Departamento de Biologia (Zoologia), 

Facultad de Ciencias, 

Universidad Autonoma de Madrid, 

Madrid, 

Spain 

Dr. R.H.T. Mattoni 

9620 Heather Road, 

Beverly Hills, 


U.S.A. 

Dr. M.C. Minno 


University of Florida, 

Gainesville, 

Florida 32611, 

U.S.A. 

Dr. M.L. Munguira 

Departamento de Biologia (Zoologia), 

Facultad de Ciencias, 

Universidad Autonoma de Madrid, 

Madrid, 

Spain 

Dr. D.D. Murphy 



Stanford, 


U.S.A. 

Dr. T.R. New 


La Trobe University, 

Bundoora, 

Victoria 3083, 

Australia 

vii 

Dr. M.W. Parker 

U.S. Fish & Wildlife Service, 

San Francisco Bay National Wildlife Refuge Complex, 

Newark, 


U.S.A. 

Dr. J.A. Powell 

Department of Entomological Sciences, 

201 Wellman Hall, 

University of California, 

Berkeley, 

California 94720-0001, 

U.S.A. 

Dr. G.B. Prince 

Department of Lands, Parks and Wildlife, 

Mrs Macquarie's Road, 

Hobart, Tasmania 7001, 

Australia 

Dr. P.R. Samson 

Bureau of Sugar Experiment Stations, 

P.O. Box 651, 

Bundaberg, 

Qld 4650, 

Australia 

Prof. M.J. Samways 

Department of Zoology and Entomology, 

University of Natal, 

Pietermaritzburg 3200, 

South Africa 

Dr. D.P.A. Sands 

Division of Entomology, 

CSIRO, 

Meiers Road, 

Indooroopilly, 

Qld 4068, 

Australia 

Dr. CD. Thomas 

School of Biological Sciences, 

University of Birmingham, 

Edgbaston, 

Birmingham B15 2TT, 

U.K. 

Dr. Stuart B. Weiss 


Department of Biological Sciences, 


Stanford, California 94305, 

U.S.A.

PART 1. INTRODUCTION 

Introduction to the biology and conservation of the Lycaenidae 

Introduction 

T.R. NEW 

Department of Zoology, La Trobe University, Bundoora, Victoria 3083, Australia 

The Lycaenidae, the 'blues', 'coppers', 'hairstreaks','metalmarks' 

and related butterflies, are the most diverse family of 

Papilionoidea and comprise between 30 and 40% of all butterfly 

species. They are mostly rather small. The world's smallest 

butterfly may be the lycaenid Micropsyche ariana Mattoni 

from Afghanistan with a wingspan of only about 7mm (although 

some individuals of Brephidium exilis (Boisduval) can have as 

small a wingspan as 6mm). A few Lycaenidae are relatively 

large: Liphyra brassolis Westwood has a wingspan of 8–9cm, 

and is the largest known species. The family occurs in all 

major biogeographical regions in temperate and tropical zones. 

As with other Lepidoptera, the life cycle comprises egg, 

larva (caterpillar) passing through several instars, pupa and 

adult, with the cycle occupying several weeks to a year. 

Particularly in temperate regions, there may be a well-defined 

phenological break during winter which is passed in an inactive 

stage. The early stages of many taxa have been described, and 

a consideration of lycaenid conservation biology must include 

the biology of all of these life forms, from oviposition site 

selection by reproductive females to larval life and adult biology. 

In general, far more distributional and biological information is 

available on adults, which tend to be conspicuous and actively 

sought by collectors and photographers, than on the relatively 

inconspicuous and cryptic immature stages. 

Many species have very precise environmental requirements, 

but the family occurs in many major biomes and vegetation 

associations from climax forests to scrublands, grasslands, 

wetlands and semi-arid desert communities, many of which 

could be viewed as early seres in terrestrial successions. Some 

lycaenids have considerable potential for use as indicator species 

as their incidence and abundance reflects rather small degrees 

of habitat change. 

The larvae of some taxa feed on flowerbuds, flowers and 

fruits (Downey 1962), and thus may exert stronger selective 

pressures on their foodplants than many foliage feeders 

(Breedlove and Ehrlich 1968). Collectively, a very broad range 

of foods are utilised and many lycaenids have departed from the 

normal lepidopteran dependence on angiosperm plants to feed 

1 

on lower plants or animal material. The extent of aphytophagy, 

which includes predacious and mutualistic relationships with 

ants and various Homoptera, is sometimes both pronounced 

and obligatory (Cottrell 1984), so that lycaenids, as a group, 

participate in a wider range of ecological interactions than 

perhaps any other Lepidoptera. 

This ecological breadth has been the basis for designation of 

'biological groups' in the family (Hinton 1951; Henning 1983). 

Together with the relatively comprehensive knowledge of the 

systematics and distribution of many temperate region taxa 

through longterm collector accumulation, this ecological breadth 

renders the family of very considerable value in conservation 

studies. Several species have been the targets of detailed practical 

measures related to their conservation in recent years, and many 

of these case histories are summarised in the third section of this 

volume. 

This introductory chapter enlarges on some of the topics 

noted above, to provide a general background to the regional 

and species accounts which follow. 

Taxonomy 

In this volume the Lycaenidae is taken to include the Riodininae 

(= Nemeobiinae) and the Styginae, both of which have been 

given family status by some researchers. 

Early classifications grouped the Riodinidae and Lycaenidae 

s. rest, as the superfamily Lycaenoidea. Clench (1955) divided 

the Lycaenoidea into three families: Lycaenidae s. str., Liptenidae 

and Liphyridae, to which Shirozu and Yamamato (1957) added 

the Curetidae. In contrast, Ehrlich (1958) considered the 

Lycaenoidea to be a single family with the major subfamiliar 

groupings of Riodininae, Styginae and Lycaeninae – the latter 

including the four families noted in the last sentence. 

While the higher classification of the two 'problem' groups 

has proved to be controversial, the scheme proposed by Eliot 

(1973) (Figure la) has, with some modification, received 

strong support. As Eliot (1973) noted, no satisfactory 

classification for the whole of the Lycaenidae had been produced 

until then, despite notable attempts by Clench (1955) and

Figure 1. Classification of the Lycaenidae into major divisions: (a) after Eliot (1973); (b) after Ackery (1984). 

Stempffer (1957, 1967). Other recent authorities (such as 

Harvey 1987), while maintaining the Riodinidae as distinct, 

have allied it with the Nymphalidae. However, yet others have 

retained the Riodinidae and Stygidae in a somewhat broader 

concept of the family (Ackery 1984 and Figure lb). Thus the 

more recent classifications recognise eight (Eliot) or ten (Ackery) 

subfamilies (Figure 1). Eliot's (1973) more extensive 'tribal' 

divisions (Figure 2) have been more generally adopted as a 

working scheme by many researchers: while some of the formal 

divisions remain uncertain, the relationships implied appear to 

be valid. 

There is a very wide range of chromosome numbers in the 

family, but the modal number appears to be n = 24 (Robinson 

1971). 

Diversity and distribution 

The trio of Theclinae (hairstreaks), Riodininae (metalmarks) 

and Polyommatinae (blues) together comprise about 90% of the 

family. Some other subfamilies are small: Styginae, for example, 

(if recognised as distinct from Riodininae) includes only Styx 

infernalis Staudinger, from Peru. Theclinae, with well over 

2 

2000 species, is the most diverse section of the Lycaenidae. 

The number of species of Lycaenidae can only be estimated. 

In a survey Robbins (1982) produced figures of 6000–6900, an 

estimate which placed Lycaenidae clearly ahead of the next 

most diverse family, Nymphalidae. Shields (1989) noted totals 

of 4089 Lycaenidae s.str. and 1366 'Riodinidae' for described 

taxa only. Bridges (1988) listed 16,475 species-group names 

and more than 4000 taxonomic publications for these groups. 

Lycaenidae are most diverse in the tropics, especially the 

neotropics and southeast Asia, followed by Africa, and most 

species occur in tropical rainforest regions. For the neotropics, 

with 2650 ±350 spp., Riodininae and Theclinae are codominant, 

with only a few Polyommatinae, a single lycaenine, and very 

few others. The nearctic fauna, surprisingly, is not diverse, with 

only about half the number of species which occur in the 

Palaearctic. But, as any regional synopsis indicates, endemism 

at levels of both major geographical region and local community 

tends to be high. 

Very few Lycaenidae are at all widely distributed. Exceptions 

include Lampides boeticus (L.) which extends from Europe to 

Australia and Hawaii, and Celastrina argiolus (L), which Eliot 

and Kawazoe (1983) regard as 'one of the world's commonest 

and most widespread butterflies'. The latter species occurs 

throughout most of the Palaearctic, Oriental and Nearctic

Figure 2. Classification of the Lycaenidae (after Eliot 1973). 

3

egions, and has formed numerous putative subspecies over this 

broad range. The American subspecies have sometimes been 

regarded as separate species, C. ladon (Cramer) (Clench and 

Miller 1980). 

Most lycaenids are not particularly vagile and some cannot 

cross even small spaces between habitat patches. Although a 

few are occasional migrants from Europe to Britain (Lampides 

boeticus, Everes argiades (Pallas), Cyaniris semiargus 

(Rottemburg)), these are apparently rather exceptional. Small 

Lycaenidae have apparently crossed oceanic barriers on 

occasion, but this process may have played only a minor role in 

their evolution: Vaga Zimmerman (in the Bonins and Hawaii) 

and Hypojamides Riley (in Tahiti) are putative examples. L. 

boeticus probably reached New Zealand from Australia (Gibbs 

1980). A few lycaenids have been deliberately introduced to 

areas where they would not naturally occur. For example both 

Strymon bazochi Godart and Tmolus echion (L.) were introduced 

into Hawaii from Mexico early this century as potential agents 

for the control of lantana (Riotte and Uchida 1979). 

Poor representation of many lycaenid groups in North 

America led Eliot (1973) to suggest that the fauna is derived 

from the Old World. Tentative explanations of present 

distribution patterns, sometimes involving a Gondwanan origin 

for the group with dichotomy into Riodininae and other tribes 

at the time of South American separation (Eliot 1973), are not 

wholly convincing. Anomalies occur, perhaps related to 

dispersal: 'coppers' (Lycaena F.) occur in New Zealand but not 

in Australia or southeast Asia, and their geographically nearest 

congenors are in the Himalaya. Other distribution patterns are 

perhaps easier to explain: Udara (Vaga) blackbumi (Tuely) is 

one of only two endemic butterflies in Hawaii and it is thought 

that its ancestors may have progressively 'island-hopped' to 

this location. Members of another genus of small lycaenids, 

Zizula Chapman, may also have crossed substantial water 

barriers in the past; such dispersal by wind may be more 

frequent than commonly supposed. In Panama, Robbins and 

Small (1981) reported 128 species of hairstreaks being blown 

by the seasonal trade winds. More than 80% of these were 

blown through habitats where they do not normally occur. 

In general, though, lycaenid faunas on remote islands are 

small, and climatic barriers (e.g. desert) may also counter 

dispersal on larger land masses. Colonisation of new habitats is 

clearly facilitated by the presence of larval foodplants and 

suitable ants, but there are few quantitative data on natural 

colonisation of islands or other 'new' habitats. However, on the 

Krakatau Islands (Indonesia), 24 species have been recorded 

since the sterilising volcanic eruptions of 1883: none were 

found in 1908, seven from 1919–1921, eight by 1928–1934, 

and 23 were present in the 1980s (New et al. 1988). Very few 

of these are habitual deep forest dwellers, and perhaps 15–18 of 

the species currently present there depend on early successional 

Ipomoea pes-caprae associations on accreting beach 

environments. Preservation of these rather transient associations 

on the islands appears to be a key theme to facilitate lycaenid 

colonisation from Java and/or Sumatra until more diverse 

vegetation is present. The eight species which have colonised 

4 

Anak Krakatau, last sterilised by volcanic activity in 1952, are 

all associated with such 'strand-line' vegetation. Natural 

opportunities to detect colonisation patterns of Lycaenidae on 

this scale are indeed rare. 

Life histories and biology 

Many species of Lycaenidae have very precise environmental 

needs, and a number of recent studies (especially in temperate 

regions) show that they may have considerable value as 

environmental indicators and, thus, an enhanced role in 

conservation studies. At this stage, though, the biology of most 

tropical taxa is almost entirely unknown, and a tendency to 

focus on the better understood temperate region forms is 

inevitable. 

Myrmecophily 

The life histories of slightly over 900 species (Downey 1962 

noted 838 species) have been documented to varying extents. A 

remarkable feature of many of these is dependence on ants: 245 

of the species noted by Downey had myrmecophilous larvae, 

and this dependency has attracted much attention. A tentative 

classification of different degrees of myrmecophily was 

proposed by Fiedler (1991a, 1991b). 

The possible advantages of myrmecophily have been 

addressed by, inter al., Henning (1983), Pierce (1984) and De 

Vries (1991 a), and a broader overview of aphytophagy is given 

by Cottrell (1984). In general, the ants tend to protect the 

caterpillars from natural enemies (Pierce and Mead 1981; 

Pierce and Easteal 1986) and gain additional food from caterpillar 

secretions (Fiedler and Maschwitz 1988). Some of these 

mutualisms are very complex. In Panama, the ant Ectatomma 

ruidum protects caterpillars of Thisbe irenea (Stoll) (Riodininae) 

from attacks by predatory wasps, but not from tachinid fly 

parasitoids, for example (De Vries 1991b). 

The caterpillars of T. irenea have an elaborate 'calling' 

system, and attract ants by producing sounds (De Vries 1990). 

Similar noises were recorded in other species, all of which 

depended on associations with ants to enhance their well being. 

The noises mimic vibrations used by the ants in their own 

communications. Stridulation is also well known in the pupae 

of many lycaenids, and one function of pupal sound production 

might also be to attract ants (Downey 1966). 

Females of Hypochrysops ignitus (Leach) are attracted to 

colonies of ant-tended Membracidae (Sands 1986). Those of 

Allotinus major Felder & Felder lay eggs near or on membracids 

(probably a species of Terentius), using brooding adult female 

membracids as oviposition cues, and the caterpillars eat the 

membracid nymphs (Kitching 1987). Younger caterpillars sit 

on the membracids to feed and are thus exposed – unlike larvae 

of Feniseca Grote and Taraka Doherty which form silken 

retreats amongst their prey (Edwards 1886; Iwase 1953). Both 

the membracid and the caterpillars of A. major are tended by

workers of the same ant, Anoplolepis longipes (Jerdon) (Kitching 

1987). 

In many cases, relationships between myrmecophily or 

aphytophagy and oviposition patterns are not as clear as in 

Allotinus Felder & Felder. Laying eggs in clusters ('clustering') 

in many Australian Lycaenidae is strongly associated with 

obligate myrmecophily (Kitching 1981), but this is not the case 

for neotropical Riodininae, in which myrmecophilous species 

lay eggs singly (Callaghan 1986). In the latter group, clustering 

is associated with gregarious behaviour. Myrmecophilous 

riodinine larvae are solitary, and gregarious larvae are not 

myrmecophilous. The latter may be aposematic and conspicuous, 

so that their protection against many predators is a function of 

their distastefulness. Many myrmecophilous larvae, in contrast, 

are rather cryptic, and accompanying ants may help to prevent 

them being attacked by predators and parasitoids. 

Such relationships between caterpillars and ants are thus of 

central importance in considering the evolution and biology of 

Lycaenidae and have attracted much attention. 

Myrmecophily and evolution in lycaenids 

Symbiosis with ants may have been an early development in the 

evolution of Lycaenidae (Eliot 1973), and both Hinton (1951) 

and Malicky (1969) suggested that ancestral lycaenids were 

myrmecophilous. Contrary to Pierce's (1987) conclusion that 

the distribution of myrmecophily in Lycaenidae does not reflect 

phylogeny, Fiedler (1991a, 1991b) believed that there was a 

strongly phylogenetic relationship present. 

However, there is some possible confusion over roles of 

myrmecophily in lycaenoid evolution as their influences on 

Riodinines and the other taxa may be markedly different (De 

Vries 1991a). Not only are they the most common basis for 

suggesting ecological groupings in the family (Henning 1983), 

but the evolution of lycaenid diversity itself may also be 

involved. Pierce (1984) suggested that lycaenid diversity may 

reflect speciation in relation to other butterfly families, and that 

this could be influenced by larvae/ant associations in two 

important ways: 

1. Female lycaenids may adopt ants as oviposition cues 

(Fiedler and Maschwitz 1989, on Anthene emolus (Godart)) so 

that the presence of ants on a novel foodplant may induce a 

rapid host switch. Although few such 'oviposition mistakes' 

(Pierce 1984) may actually lead to range extensions, it may be 

more important for a given lycaenid to retain a particular ant 

association than a particular foodplant, and an increase in the 

number of ovipositions on different foodplants may increase 

the number of opportunities for subsequent speciation. 

Essentially, novel foodplant choices may be made by female 

lycaenids to an unusually high degree because they select for 

ants as well as for chemically and physically suitable foodplants. 

A 'new' hostplant may occupy a different ecological range 

from those utilised earlier, and population isolates could thus be 

formed. 

2. The general non-vagility of many lycaenids results in 

their occurrence in small, semi-isolated populations with rather 

5 

little regular genetic interchange between them. Pierce (1983) 

showed that a deme of the Australian Jalmenus evagoras 

(Donovan) may be restricted to a single Acacia tree, where 

males aggregate and compete for emerging females so that 

variability in male reproductive success effectively reduces 

population size further. Such patchy distributions (also noted in 

the North American Glaucopsyche lygdamus Doubleday: Pierce 

1984) occur in spite of apparent continuous foodplant availability 

and it is quite possible that they result from selection of 

foodplant areas which are high in nitrogen, as well as having the 

required ants. Many myrmecophilous lycaenid larvae actively 

prefer nitrogen-rich foodplants and plant parts such as seed 

pods and flowers. This may be explained in part by the need to 

supply ants with amino acids as a 'nutrient reward' for tending 

the larvae (Pierce 1984). 

Larval feeding 

The overall importance of plant-feeding to caterpillars of 

Lycaenidae differs substantially between different subfamilies, 

and those of some groups rarely take plant food. As far as is 

known, all species of Poritiinae and Lycaeninae are normally 

phytophagous. Some Curetinae are phytophagous. Lipteninae 

are also plant feeders, but are highly unusual amongst butterflies 

in that larval food usually consists of algae, fungi or lichens (see 

Cottrell 1984, for summary). Most genera of the two largest 

subfamilies, Theclinae and Polyommatinae, appear to be 

phytophagous or opportunistically carnivorous with varying 

degrees of dependence on prey. Maculinea van Eecke and 

Lepidochrysops Hedicke larvae are phytophagous when young, 

but the late instars are obligate predators of ant larvae. Other 

aphytophagous genera are noted in Table 1. Both Liphyrinae 

and Miletinae appear to be entirely aphytophagous, and the 

unlisted genera in Table 1 reflect ignorance of their larval 

biology, rather than known phytophagy. Liphyra Westwood 

and Euliphyra Holland larvae are probably specific feeders on 

early stages of tree ants (Oecophylla spp): their larvae are 

flattened and have a heavily armoured cuticle which enables 

them to withstand ant attacks. The pupa of Liphyra remains 

inside the last larval skin, which thereby functions as a puparium. 

Aslauga larvae are predators of Homoptera, at least as late 

instars. Eggs of Miletinae are typically laid near colonies of 

Homoptera, including aphids, coccids and membracids and 

some, at least, are found on a wide range of different hostplants. 

Although the larvae are predominantly predators, some younger 

instars may also feed on honeydew or other insect secretions 

such as aphid cornicle secretions. 

Selection of foodplant species by phytophagous species, 

and their effects on foodplants, are difficult to study. Flower 

predation of a range of perennial herbaceous legumes by 

Glaucopsyche lygdamus in Colorado differed substantially 

between species (Breedlove and Ehrlich 1972), with either 

Lupinus or Theropsis being by far the most heavily attacked 

plant at each of a series of sites. On both plant genera, flowerfeeding 

can markedly reduce seed-set (Breedlove and Ehrlich 

1968,1972). Whereas G. lygdamus females select inflorescences

Table 1. Lycaenidae with larvae which do not feed on plants (after Cottrell 1984, based on Eliot 1973). 

Subfamily 

Lipteninae 

Poritiinae 

Liphyrinae 

Milelinae 

Curetinae 

Theclinae 

Lycaeninae 

Polyommatinae 

Number of 

Tribes Genera 

2 

1 

1 

4 

1 

19 

1 

4 

46 

7 

5 

12 

1 

241 

18 

149 

Aphytophagous genera* 

(none) 

(none) 

(3) Liphyra, Euliphyra, Aslauga 

(8) Miletus, Allotinus, Megalopalpus, Taraka, Spalgis, Feniscea, 

Lachnocnema, Theslor 

(none) 

(7) Acrodipsas, Shirozua, Zesius, Spindasis, Oxychaela, Trimenia, 

Argyrocupha 

(none) 

(4) Triclena, Niphanda, Maculinea, Lepidochrysops 

• Members of some genera are partially phytophagous, some only in the early instars (e.g. Maculinea, Lepidochrysops). 

of less hairy Lupinus species on which to lay, the reverse trend 

is true of Plebejus icarioides Boisduval, which oviposits on the 

most hairy (non-floral parts) of Lupinus by preference (Downey 

1962). Some caution is needed in extrapolating from laboratory 

feeding trials to the field - thus, captive larvae of P. icarioides 

will feed on any species of Lupinus proffered (Downey and 

Fuller 1961) but wild populations normally utilise only a few of 

the species available in their habitat. There may also be a 

pronounced seasonal variation in foodplant quality: P. acmon 

(Westwood) and related species utilise some foodplants only at 

particular times of the year (Goodpasture 1974). J. evagoras 

females preferred to oviposit on potted Acacia which had been 

given nitrogenous fertiliser rather than on similar plants given 

water alone (Baylis and Pierce 1991). 

Comparatively few species of Lycaenidae seem to be 

markedly polyphagous and many clearly have very restricted 

ranges of foodplants. Broad taxonomic ranges of foodplants – 

such as the 30 or so species (representing the families 

Selaginaceae, Lamiaceae, Verbenaceae, Fabaceae and 

Geraniaceae) recorded for Lepidochrysops in Africa (Cottrell 

1984) – are commonly recorded only for taxa which are later 

ant-feeders. However, there is evidence of more local host 

restriction for Lepidochrysops, so that in any one area only a 

single plant genus is used for oviposition (Cottrell 1984). In 

general support of this relationship between poly phagy and ant 

attendance, a broad foodplant range for Hypochrysops miskini 

(Waterhouse) in Queensland led Valentine and Johnson (1989) 

to suggest that rainforest lycaenids may be polyphagous because 

a narrow choice of plants may be restrictive and cancel out 

advantages of ant-attendance. In Queensland, rainforest species 

confined to single host plant species tend not to be ant-attended. 

Rarely, different generations of the same species may utilise 

different foodplants: the first (spring) generation of Celastrina 

argiolus (L.) in Europe feeds on holly, while the second 

(summer) generation larvae eat ivy. This species also shows 

6 

marked seasonal dimorphism, but it is not clear if this is foodrelated. 

Nitrogen-rich plant feeding is a characteristic of many 

lycaenid taxa. Pierce (1985) has noted that caterpillars of 

lycaenids from Australia, South Africa and North America 

have been recorded feeding on most known non-leguminous, 

nitrogen-fixing plant families. Several species of Cycadaceae 

are included, and most Lipteninae are probably specialised 

lichen-feeders. There are, of course, exceptions: the nonmyrmecophilous 

riodinine Sarota gyas (Cramer) in Panama 

and Costa Rica feeds exclusively on epiphyUs (De Vries 1988), 

and those epiphylls tested were not nitrogen-fixers. Females of 

this species lay eggs on taxonomically disparate hostplants with 

old leaves covered by epiphylls. In many other lycaenids, 

though, ovipositions on reproductive structures of larval 

foodplants may reflect selection for high nitrogen levels in the 

future larval food. 

Feeding on floral structures may itself engender polyphagy: 

Robbins and Aiello (1982) noted that the thecline Strymon 

melinus (Hübner) is one of the most polyphagous butterfly 

species known, and has been recorded feeding on flowers of 46 

genera in 21 families of plants. Members of the genus Callophrys 

Billberg also feed on plants of some 11 families (Pratt and 

Ballmer 1991). Polyphagy of this sort might be an important 

ecological strategy for the lycaenids in exploiting and adjusting 

to changing environmental conditions and could be a correlate 

of r-selection. Flower-feeding could, perhaps, allow larval 

access to plants when the foliage is not available due to their 

chemical defences (Robbins and Aiello 1982), and it is likely 

that some lycaenids in the tropics have life cycles which are 

well adapted to exploit peak flowering seasons of putative 

foodplants. Robbins and Aiello cite data (Croat 1978) that peak 

flowering on Barro Colorado Island (Panama) occurs at the end 

of the dry season, a period corresponding with a marked 

increase in abundance of Lycaenidae but a decrease of many

other butterflies. Such seasonal apparency may be an important 

feature facilitating the use of tropical lycaenids as indicators of 

habitat quality. It may also facilitate ecologically important 

dispersal, as the butterflies are abundant at the time of strong 

sustained dry season trade winds (Robbins and Small 1981). 

Samples were skewed heavily toward females, and many of the 

128 species were being blown through habitats where they do 

not occur normally. 

Most phytophagous lycaenid larvae are external feeders, 

either eating whole plant tissues or organs, or skeletonising 

foliage. However, Callophyrysxami (Reakirt) in Mexico feeds 

on fleshy-leaved Crassulaceae and, on Echeveria gibbiflora, 

caterpillars may burrow completely into the leaves and feed on 

the fleshy internal pulp. Wastes are expelled through the entrance 

hole (Ziegler and Escalante 1964). Larvae of some other 

Callophrys (such as C. viridis (Edwards)) feed on flowers of 

Eriopsis and, as with other larvae with similar habits, they 

resemble their foodplant flowers in coloration (Brown and 

Opler 1967). Variation in larval coloration occurs in other taxa, 

such as Zizina labradus (Godart) in Australia (Sibatani 1984). 

Although it is clear that very considerable trophic specialisation 

occurs in many lycaenids – both in phytophagous and 

aphytophagous species where intricate and obligate relationships 

with other animals are common – little is known of the biology 

of most species of the family. The precise resource needs of many 

specialised species cannot yet be appreciated fully, but they are 

likely to render the species very susceptible to habitat change and 

consequent changes in resource supply. 

Adult feeding 

Adult lycaenids (of Theclinae, Lycaeninae and Polyommatinae) 

commonly seek nectar from flowers, although some Theclinae 

may depend more on aphid honeydew. In contrast, Lipteninae 

and Miletinae are not known to visit flowers, and depend on 

secretions from extrafloral nectaries or on homopteran 

honeydew. Adult Liphyra lack a proboscis and thus cannot 

feed. A few taxa may feed on secretions from the dorsal nectary 

organ of other ant-tended lycaenid larvae. 

Adult sexual dimorphism 

This is widespread in the family. Commonly, males are brighter 

(either the females are browner or have wider, dark borders to 

the wings). Indeed, in some taxa it is difficult to associate the 

sexes correctly because of relatively extreme dimorphism. 

Males may have modified scales or sex-brands on either surface 

of the forewing or on the upperside of the hindwing. These may 

be associated with hair-tufts but, more commonly, distinct sex 

brands are absent and androconia are scattered irregularly over 

the upper surface of the wings. 

Adult behaviour 

Males of some species are regular hill-toppers: some species of 

the 'Lycaenopsis-group', for example, are rarely seen elsewhere, 

7 

and both sexes of some species of Monodontides Toxopeus 

appear to be hill-top residents (Eliot and Kawazoe 1983). Hilltopping 

may be common in riodinines (Shields 1967). 

Protandry may occur regularly in some lycaenids. Males of 

Polyommatus icarus (Rottemburg) not only emerge before the 

females but also fly more strongly. Males of this genus interact 

strongly with other males, and it seems that the bright male 

colours may indeed be 'directed' at other males. Lundgren 

(1977) made the intriguing suggestion that the high frequency 

of the inter-male encounters may aid in regulating dispersal and 

population density, and this may occur also between groups of 

sympatric species with similar appearance. 

Displays and pheromones both appear to be involved in 

ensuring conspecific matings in lycaenids living in the same 

area. The female anal hair-tuft of Nordmannia Tutt may play a 

role in courtship (Nakamura 1976). Multiple copulations can 

occur (Fujii, 1989, on Shirozua janasi (Janson)), and it is 

possible that copulatory mate guarding occurs in this thecline. 

The time of day for mating differs between species: S. janasi 

copulates during the day whereas two species of Japonica Tutt 

mate at dusk (Fujii 1989). There are occasional records of 

cross-taxon matings, sometimes even between subfamilies 

(Shapiro 1985), or even families (Shapiro 1982; Johnson 1984), 

usually between phenotypically similar butterflies. 

'Perching' behaviour appears to be an important isolating 

mechanism in Riodininae (Callaghan 1979,1983). Neotropical 

riodinines appear to have developed a range of different perching 

strategies which help to maintain specific isolation within the 

habitat. Habitat complexity is an important facet of this aspect 

of these forest butterflies, as the spacing of perching sites and 

positions of perching differ considerably between genera – by 

topographic features, by light/shadow regimes and by time of 

day. Following earlier accounts by Scott (1968) and Shields 

(1967), Callaghan suggested that perching (and hill-topping) 

species tend to have low population densities, and 'rendezvous' 

localities help such rarer species to find mates. Perching for 

short periods (an average of 2.4h in 10 riodinine genera) 

minimises exposure to predators. Different perching strategies 

are enhanced by ethological strategies such as displays. When 

males are scarce females may actively search out perching sites 

and await them if they are ready to mate. Ethological isolating 

mechanisms of this sort may be more important in taxa which 

are not strict deep forest dwellers. 

Perching behaviour of different species of Cupressaceaefeeding 

Callophrys is distinctly non-random (Johnson and 

Borgo 1976). The insects apparently orientate to the position of 

the sun, and taller trees are most often selected for perching on. 

Perching posture in lycaenids, as in other butterflies, may have 

an important role in thermoregulation (Clench 1966). 

Lycaenidae which are normally 'perchers' rather than 

'patrollers' may at times undergo sustained flights and the 

down-valley flights of four such species of Theclini in Colorado 

and Arizona (Scott 1973) appeared to be a food-seeking 

behaviour. This may also be so for Satyrium Scudder in California 

(MacNeill 1967). Some species fly only at particular times of 

the day and, within a genus, species co-occurring at the same

locality may not overlap in their flight periodicity (see Sibatani 

1992, on Favonius Sibatani & Ito). 

Some very small blues are extremely weak fliers, as suggested 

earlier. Some have elongated narrow wings which, at least in 

Zizula hylax (F.), enable the butterflies to crawl into the corollas 

of long-tubed flowers and obtain nectar (Ehrlich and Ehrlich 

1972). 

Defence against predators 

It has been suggested that a few adult Lycaenidae participate in 

mimicry complexes, either with other blues or other kinds of 

Lepidoptera. I have netted specimens of Udara Toxopeus in 

stream beds in New Guinea in the belief that they were small 

members of one of the many species of Delias Hübner (Pieridae) 

in the same habitat; Plebejus icarioides may be a model for the 

noctuid moth Caenurgina caerulea Grote in North America 

(Downey 1965); and male Lycaena heteronea Boisduval may 

be a mimic of G. lygdamus and other sympatric blues (Austin 

1972). The latter feed on plants containing alkaloids (Lupinus) 

or selenium (Astragalus), while the foodplants of the Lycaena 

(Eriogonum) apparently do not. However, experimental 

investigations of this scenario do not appear to have been made. 

In Sierra Leone, Pseudaletis leonis (Staudinger) may be a 

mimic of a similarly-patterned diurnal arctiid moth, or of a 

larger danaine (Owen 1991). 

Other putative defences against predators include the 

chemical defence of Eumaeus atala (Poey). This species 

sequesters a deterrent defensive chemical (cycasin) from larval 

cycad foodplants, and this clearly renders the adults unpalatable 

to birds (Bowers and Larin 1989). Cycasin also deters attack by 

Camponotus ants. Adult E. atala are aposematic, and the genus 

is mimicked by several other Lepidoptera (including species of 

Castniidae and Noctuidae) in South America. Further examples 

of defence can be found at all growth stages: the 'false heads', 

including hindwing tails of many adults (Robbins 1980,1981); 

the 'monkey head' pupae of Spalgis Moore (Hinton 1974); 

sound production by pupae of some taxa; unusual hairiness or 

thickened cuticle of some caterpillars; and the female anal hairtuft 

of some species of Nordmannia which is used to cover eggs 

with long scales at the time of oviposition. 

Even mode of hatching from the egg may be correlated with 

protection from natural enemy attack: caterpillars of Maculinea 

alcon (Denis & Schiffermüller) and M. rebeli (Hirschke) hatch 

through the base of the eggshell and emerge on the opposite side 

of the leaf (Thomas et al. 1991). The 'normal' exposed eggshells 

are unusually thick, perhaps to counter enemy attack. 

Particularly in the tropics, lycaenid biology is almost 

unstudied at other than the most superficial levels. Most 

information, including that which has led to tentative 

generalisations for the family, has been derived from temperate 

region species. Even for the best-studied lycaenid faunas, there 

are many gaps in fundamental biological knowledge. 

8 

Endangering processes 

In their review of threatened swallowtail butterflies, Collins 

and Morris (1985) enumerate a number of processes which may 

lead to the decline of these butterflies, and it is instructive to 

assess the effects of these processes on the ecologically very 

different Lycaenidae. 

Collecting and trade 

The effects of collecting are controversial and difficult to 

evaluate fully. Many lycaenids occur in small closed populations 

and because of this may be much more vulnerable to localised 

collector pressure than many other butterflies. There is at least 

some suggestion, for example, that the demise of the Large 

Copper, Lycaena dispar (Haworth), in Britain in the midnineteenth 

century was in part due to increased collector pressure 

on populations which had been rendered vulnerable by habitat 

destruction. In most cases collecting is probably the subsidiary 

rather than the prime cause of decline or extinction. Perhaps, 

especially in northern temperate regions, there is reason to 

suggest that this syndrome could occur for many localised taxa 

sought by collectors who visit the same colony year by year. 

Even specialist collecting to help monitoring of restricted 

colonies for conservation assessment may cause direct damage 

to populations: Murphy (1989) has recently pointed out that 

conventional mark-recapture studies used to estimate population 

sizes of taxa of conservation interest may cause inadvertent 

damage, either by mutilation through handling, or by inducing 

changes in individual behaviour. Small delicate butterflies, 

such as most lycaenids, may be particularly vulnerable to such 

effects. 

Commercial collecting is of considerably less importance 

for Lycaenidae than for large showy butterflies. Rare species 

from unusual localities will always find a market amongst 

wealthy collectors and museums, but the 'ornament' and 

'souvenir' trades in Lycaenidae are very low. In Malaysia, for 

example, the vast bulk of this trade is in Papilionidae and 

Nymphalidae: in stores in Kuala Lumpur in 1988,I noticed only 

very few Lycaenidae (all relatively large species of Arhopala 

Boisduval and related genera). In general, for this trade, 'small' 

is distinctly not 'beautiful' or desirable. 

Similarly, Lycaenidae are not particularly desired as exhibits 

in butterfly houses, and the difficulty of rearing many of the 

species may also be a deterrent in this context. Collins (1987a) 

lists only eight species of Lycaenidae (of 223 butterfly species 

in total) on show in a sample of 18 butterfly houses in Britain 

in 1986 (Table 2). Seven of these were obtained as pupae, but 

the Malaysian Spindasis syama (Horsfield) were apparently 

obtained as adults by the one exhibitor. 

A total collecting ban is difficult to enforce, especially on 

unreserved public land – indeed, it is impossible without 

wardenship or other constant (and probably expensive) security 

measures. Even responsible, private collectors obeying voluntary 

restrictive quota codes may cause harm if they are in sufficient

numbers. As Collins and Morris (1985) commented for 

Papilionidae 'there is a danger that collectors may be unable to 

recognise when they are depleting butterfly stocks below the 

threshold of recovery, particularly when they only visit the 

breeding area for short periods of time'. Although no Lycaenidae 

are currently listed in CITES Appendices, a number of rare or 

local species have received local legislative protection (if not 

more tangible conservation) through bans on collecting. In 

addition, over 40 species are mentioned in various countrybased 

European legislation (Collins 1987b and Table 3) and 28 

taxa are listed in the United States Federal Register of Endangered 

and Threatened Wildlife. The most prolific, and probably the 

least discriminating legislation is in India, where some 160 

species are listed under the Wildlife Protection Act (Table 4). 

The Code of Conservation Responsibility adopted by 

commercial entomologists in Britain in 1974, restricts trade in 

a number of species (including Maculinea arion (L.), then still 

extant in Britain) to specimens already in 'circulation', so that 

the only legal way that a collector can purchase examples of 

these species is from the limited pool already in collections. 

Seventy-nine lycaenid taxa arc included in the IUCN Red 

List of Threatened Animals (1990) (Table 5). Many of these are 

also cited in various country-based legislations. 

Habitat alteration and destruction 

This is the prime threat to all lycaenid species with limited 

distributions and low vagility, and has already been the agent of 

major declines of many of these – as the examples discussed 

later in this volume attest. Lycaenid taxa are particularly 

susceptible to certain types of habitat alteration including: 

changes in forestry practices in tropical and temperate regions; 

conversion of shrublands to pasture and agricultural lands; 

wetland drainage; heathland succession; grassland management 

practices; the effects of grazing animals such as rabbits; and 

expanding urban, industrial and recreational land use. 

In common with all other animals and plants, levels of 

concern therefore range from large-scale destruction of tropical 

rainforests whose biota have scarcely been documented (and in 

many cases never now can be), to small, local habitat changes 

Table 2. Lycaenidae flown in butterfly houses in Britain (Collins 1987a). 

Species 

Eumaeus alula 

I ampules boeticus 

Lycaena helle 

L. phlaeas 

Narathura centaurus 

Polyommatus icarus 

Spindasis syama 

Telicada nyseus 

Native range 

Caribbean 

(widespread) 

Europe, Asia 

Europe, Asia 

Malaysia 

Europe 

Asia 

East Asia 

9 

in 'ordinary' vegetation such as grassland or heathland. Such 

changes have, of course, occurred in many parts of the world, 

and in many major areas their effects have been both unheralded 

and undocumented so that we can merely infer, from the present 

status of taxa and knowledge of their biology as this accumulates, 

the magnitude of their effects. Only rarely outside northern 

temperate regions has conservation awareness for Lycaenidae 

progressed to the stage where concern can be shown in any 

practical manner. There, it is sometimes abundantly clear that 

decline of species or assemblages, and the initiation of concern 

for their conservation, has been engendered by particular 

localised human activities – sometimes as 'one-off destructive 

events, sometimes more broadly. Many of the former apply to 

remnant populations which represent formerly much more 

widespread taxa which have become progressively vulnerable 

over a longer time. In other parts of the world, very many 

species have been recorded only from single or highly disjunct 

localities, and even sound demonstration of their abundance or 

dependence on particular habitats is difficult or impossible at 

this time. In short, status evaluation is difficult or impossible, 

and the option of habitat protection, in the interest of decreasing 

or eliminating perceived threats, is the most urgent option. 

In contrast to most Papilionidae, grassland and other open 

vegetation types are vitally important habitats to many 

Lycaenidae. In Europe, calcareous grassland is a particularly 

important lycaenid habitat which has undergone large scale and 

sometimes dramatic changes. The extinction of Cyaniris 

semiargus (Rottemburg) in Britain as long ago as 1877 has been 

attributed to changes in grassland management (Heath 1981). 

While 'traditional' methods of land and vegetation 

maintenance, such as coppicing of forests, may foster the wellbeing 

of some species, intensification of agricultural practices 

has caused concern for some. Wetlands are particularly 

vulnerable to such changes and a number of European wetland 

Lycaenidae are under threat. Some species of Maculinea and 

Lycaena restricted to this habitat are particularly endangered. 

Drainage of the fens in England last century was a prime cause 

of decline of Lycaena dispar. 

Urbanisation has caused concern for lycaenids in places as 

far apart as Los Angeles, California and Melbourne, Australia. 

Origin of material 

U.S.A.: Florida 

Sri Lanka, Malaysia 

France 

France 

Malaysia 

U.K 

Malaysia 

Sri Lanka, Malaysia

Table 3. Lycaenidae protected by legislation in Europe (from Collins 1987b). 

(i) Areas which protect Lycaenidae as part of 'all butterflies' or a similar ordinance, sometimes with exceptions for particular Pieridae stated. Date of legislation 

given in parentheses: 

Austria: Niederosterreich (1978), Salzburg (1980), Steiermark (1936), Tyrol (1975), Oberosterreich (1982), Vienna (1985), Vorarlberg (1979). 

Germany: (Law of German Democratic Republic 1984*). 

Luxembourg: (1986). 

(ii) Particular taxa 

Taxon 

Agrodiaetus admelus 

A. damon 

A, riparlii 

Aricia agestis 

A. crassipuncta 

A. taberdiana 

Callophrys mystaphia 

C. suaveola 

Cupido osiris 

Cyaniris Helena 

Eumedonia damon 

Freyeria trochylus 

lolana iotas 

Kretania eurypilus 

K. psylorita 

Lycaeides argyrognomon 

L. idas 

L. helle 

Lysandra bellargus 

L. caucasica 

Maculinea alcon 

M. arion 

M. nausithous 

M. rebeli 

Country 

Greece 

Hungary 

Greece 

Hungary 

Germany (FR) 

Germany (FR) 

Germany (FR) 

Germany (FR) 

Hungary 

Greece 

Hungary 

Greece 

Hungary 

Germany (FR) 

Germany (FR) 

Greece 

Belgium: Wallone Region 

Belgium: Flemish Region 


Finland 

France (females only) 

Germany (FR) 

Hungary 

Netherlands 


France (females only) 

Germany (FR) 

Hungary 


France (L. b. coelestis, females only) 

Germany (FR) 


Germany (FR) 



Germany (FR) 

United Kingdom 

Germany (FR) 

Hungary 


Germany (FR) 

10 

Date of legislation 

(1980) 

(1982) 

(1980) 

(1982) 

(1986)* 

(1986) 

(1986) 

(1986) 

(1982) 

(1980) 

(1982) 

(1980) 

(1982) 

(1986) 

(1986) 

(1980) 

(1985) 

(1980) 

(1985) 

(1976) 

(1979) 

(1986) 

(1982) 

(1973) 

(1985) 

(1979) 

(1986) 

(1982) 

(1985) 

(1979) 

(1986) 

(1980) 

(1986) 

(1980) 

(1985) 

(1986) 

(1981) 

(1986) 

(1982) 

(1985) 

(1986)

Table 3 (cont.). Lycaenidae protected by legislation in Europe (from Collins 1987b). 

Taxon 

M. teleius 

Neolycaena coelestina 

Nordmannia acaciae 

N. armenia 

N. marcidus 

N. sassanides 

Paleochrysophanus hippothoe 

Plebejus pylaon 

Polyommatus eros 

Pseudophilotes bavius 

Strymonidia pruni 

S. myrrhina 

Thersamonia thetis 

Tomares callimachus 

T. romanovi 

Turanana panagea 

Vacciniina optilate 

Zizeeria knysna 

Country 


France ( M. t. burdigalensis, females only) 

Germany (FR) 


Germany (FR) 

Germany (FR) 

Germany (FR) 

Hungary 

Hungary 

Greece 

Germany (FR) 

Greece 

Germany (FR) 

Greece 

Germany (FR) 

Germany (FR) 

Greece 

Germany (FR) 

Greece ( Z. k . cassandra) 

• FR = Federal Republic: it is not yet clear how the former laws for FRG and DDR will operate for a unified Germany. 

Table 4. Lycaenidae listed in The Indian Wildlife (Protection) Act, 1972. 

Note: the scientific names are given as spelled in the Act – a number of spelling errors are present in the listing. 

1. Schedule 1. Part IV (Collection and trade, including gift, prohibited) Liphyra brassolis 

Allotinus drumila 

A. fabious penormis 

Amblopala avidiena 

Amblypodia ace arala 

A. alea constanceae 

A. ammonariel 

A. arvina ardea 

A. asopia 

A. comica 

A. opalina 

A. zeta 

Biduanda melisa cyana 

Callophyrs leechii 

Castalius rosimon alarbus 

Charana cepheis 

Chlioria othona 

Deudoryx epijarbas amatius 

Everes moorei 

Gerydus biggsii 

G. symethus diopeithes 

Heliophorus hybrida 

Horaga albimacula 

Jamides ferrari 

Darkie, crenulate/great 

Angled darkie 

Hairstreak, Chinese 

Leaf blue 

Rosy oakblue 

Malayan bush blue 

Purple brown tailless oakblue 

Plain tailless oakblue 

Comic oakblue 

Opal oakblue 

Andaman tailless oakblue 

Blue posy 

Hairstreak, ferruginous 

Pierrot, common 

Mandarin blue, Cachar 

Tit, orchid 

Cornelian, scarce 

Cupid, Moore's 

Bigg's brownie 

Great Brownie 

Sapphires 

Onyxes 

Caeruleans 

11 

Listeria dudgenni 

Logania watsoniana subfasciata 

Lycaenopsis binghami 

L. haraldus ananga 

L. puspa prominent 

L. quadriplaga dohertyi 

Nacaduba noreia hampsoni 

Polymatus orbitulus leela 

Pratapa icetas mishmia 

Simiskina phalena harterti 

Sinthusa virgo 

Spindasis elwesi 

S. rukmini 

Strymon mackwoodi 

Tajuria ister 

T. luculentus nela 

T. yajna yajna 

Thecla ataxua zulla 

T. bieti mendera 

T. letha 

T. paona 

T. pavo 

Virachala smilis 

Date of legislation 

(1980) 

(1979) 

(1986) 

(1985) 

(1986) 

(1986) 

(1986) 

(1982) 

(1982) 

(1980) 

(1986) 

(1980) 

(1986) 

(1986) 

(1986) 

(1986) 

(1980) 

(1986) 

(1980) 

Butterfly, moth 

Lister's hairstreak 

Mottle, Watson's 

Hedge blue 

Hedge blue, Felder's 

Common hedge blue 

Naga hedge blue 

Limeblue, white-tipped 

Greenish mountain blue 

Royal, dark blue 

Brilliant, broadbanded 

Spark, pale 

Silverline, Elwe's 

Silverline, khaki 

Hairstreak, Mackwood's 

Royal, uncertain 

Royal, Chinese 

Royal, chestnut and black 

Wonderful hairstreak 

Indian purple hairstreak 

Watson's hairstreak 

Paona hairstreak 

Peacock hairstreak 

Guava blues 

Continued...

Table 4 (cont.). Lycaenidae listed in The Indian Wildlife (Protection) Act, 1972. 

2. Schedule II. Part II. ('Special game': 

Allotinus subviolaceous manychus 

Amblypodia abetrans 

A. aenea 

A. agaba aurelia 

A. agrata 

A. alesia 

A. apidanus ahamus 

A. areste areste 

A. bazaloides 

A. camdeo 

A. ellisi 

A. fulla ignara 

A. ganesa watsoni 

A. paragenesa zephpreeta 

A. paralea 

A. silhetensis 

A. suffusa stiffusu 

A. yendava 

Apharitis lilacinus 

Araotes lapithis 

Artipe eryx 

Bindahara phocides 

Bolhrinia chennellia 

Castraleus roxus manluena 

Catapoecilma delicalum 

C. elegans myoslina 

Charana jalindra 

Cheritrella truncipennis 

Chliaria kina 

Deudoryx hypargyria gaetulia 

Enchrysops enejus 

Everes kala 

Helipphorus androcles moorei 

Horaga onyx 

H. viola 

Hypolycaena nilgirica 

H. ihecloides nicobarica 

lraota rochana boswelliana 

Jamides alectokandulana 

J. exleodus para 

J. coeruleus 

J. kankena 

Lampides boeticus 

Lilacea albocaerulea 

L. atroguttata 

L. lilacea 

L. melaena 

L. minima 

Logania massalia 

Lycaenesthes lycaenina 

Mahathala ameria 

M. atkinsoni 

Magisba malaya presbyter 

Nacaduba aluta coelestis 

N. ancyra aberrans 

N. dubiosa fulva 

N. helicon 

N. herus major 

N. pactolus 

Neucheritra febronia 

licence needed to collect or trade.) Niphanda cymbia 

Orthomiella ponlis 

Pithecops fulgens 

Polymmatus devanica devanica 

P. metallica metallica 

P. orbitulus jaloka 

P. younghusbandi 

Poritia erycinoides elisei 

P. hewitsoni 

P. plusrata geta 

Pralapa bhetes 

P. blanka 

P. deva 

P. icetas 

Rapala buxaria 

R. chandrana chandrana 

R. nasaka 

R. refulgens 

R. rubida 

R. scintilla 

R. sphinx sphinx 

R. varuna 

Spindasis elima elima 

S. lohita 

S. nipalicus 

Suasa lisidus 

Surendra todara 

Tajuria albiplaga 

T. cippus cippus 

T. culta 

T. diaeus 

T. illurgioides 

T. illurgis 

T. jangala andamanica 

T. melastigma 

T. sebonga 

T. thyia 

T. yajna istroides 

T. callinara 

Tarucus dharta 

Thaduka multicaudata kanara 

Thecla ataxus ataxus 

T. bitel 

T. icana 

T. jakamensis 

T. kabreea 

T. khasia 

T. kirbariensis 

T. suroia 

T. syla assamica 

T. vittata 

T. ziha 

T. zoa 

Una usta 

Yasoda tripunctata 

12 

3. Schedule IV. ('Small game': 

collect.) 

Tarucus ananda 

small game hunting licence needed to

Table 5. Lycaenidae included on the 1990 IUCN Red List of Threatened Animals. 

Taxon 

Alaena margaritacea 

Aloeides caledoni 

A. dentatis 

A. egerides 

A. lutescens 

Argyrocupha malagrida malagrida 

A. m. paarlensis 

Callophrys mossii bayensis 

Capys penningtoni 

Chrysoritis cotrelli 

C. oreas 

C. zeuxo 

Cyclyrius mandersi 

Deloneura immaculata 

D. millari millari 

Deudorix penningtoni 

D. vansoni 

Durbania limbata 

Erikssonia acraeina 

Eumaeus atala florida 

Everes comyntax texanus 

Glaucopsyche lygdamus palosverdesensis 

G. xerces 

Hemiargus thomasi bethune-bakeri 

Icaricia icarioides missionensis 

I. i. moroensis 

I. i. pheres 

Lepidochrysops ariadne 

L. bacchus 

L. hypolia 

L. loewensteini 

L. lotana 

L. methymna dicksoni 

L. titei 

Lycaeides argyrognomon lotis 

L. melissa samuelis 

Lycaena dispar 

L. dorcas claytoni 

L. hermes 

Maculinea alcon 

M. arion 

M. arionides 

Status 

V 

V 

R 

V 

v 

V 

V 

E 

E 

I 

I 

I 

I 

Ex? 

V 

V 

V 

V 

R 

V 

Ex 

Ex? 

Ex 

I 

E 

I 

I 

E 

V 

Ex? 

V 

E 

E 

V 

E 

I 

E 

I 

I 

V 

V 

V 

13 

Country 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

U.S.A. 

South Africa 

South Africa 

South Africa 

South Africa 

Mauritius 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

U.S.A. 

(U.S.A.) 

U.S.A. 

(U.S.A.) 

U.S.A. 

U.S.A. 

U.S.A. 

U.S.A. 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

U.S.A. 

U.S.A. 

northern Europe 

U.S.A. 

Mexico, U.S.A. 

Europe, USSR 

Europe, USSR 

China, Japan, USSR 

Continued...

Table 5. (cont.) Lycaenidae included on the 1990 IUCN Red List of Threatened Animals. 

Taxon 

M. nausithous 

M. rebeli 

M. teleius 

M. t. burdigalensis 

Notarthrinus binghami 

Oreolyce dohertyi 

Ornipholidotos peucetia penningtoni 

Oxychaeta dicksoni 

Panchala ganesa loomisi 

Philotiella speciosa bohartorum 

Plebejus emigdionis 

P. icarioides missionensis 

Plebicula golgus 

Poecilmitis adonis 

P. aureus 

P. endymion 

P. lyncurium 

P. nigricans 

P. rileyi 

Pseudalmenus chlorinda chlorinda 

P. c. conara 

Pseudiolaus lulua 

Shijimiaeoides battoides allyni 

S. b. comstocki 

S. enoptes smithi 

S. lanstoni 

S. rita mattoni 

Spindasis collinsi 

Strymon acis bartrami 

S. avalona 

Thestor dicksoni dicksoni 

T. kaplani 

T. tempe 

Trimenia wallegrenii 

Uranothauma usambarae 

(Categories defined in Appendix 1). 

Status 

E 

V 

E 

E 

R 

R 

I 

E 

E 

I 

I 

E 

E 

V 

I 

V 

V 

V 

V 

I 

I 

V 

E 

I 

E 

I 

I 

V 

I 

K 

V 

V 

V 

V 

V 

14 

Country 

Europe, USSR 

south & central Europe 

Europe, northern Asia 

France 

India 

India 

Mozambique, S. Africa 

South Africa 

Japan 

U.S.A. 

U.S.A. 

U.S.A. 

Spain 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

South Africa 

Australia: Tasmania 

Australia: Tasmania 

South Africa 

U.S.A. 

U.S.A. 

U.S.A. 

U.S.A. 

U.S.A. 

Tanzania 

U.S.A. 

U.S.A. 

South Africa 

South Africa 

South Africa 

South Africa 

Tanzania

Increasing human recreational activities constitute another 

serious threat to many habitats all over the world and the 

following list, while far from exhaustive, gives some idea of the 

range of habitats involved: 

• coastal sand dunes in California are threatened by off-road 

vehicles and trampling; 

• alpine heathlands and meadows in Europe and southeastern 

Australia are threatened by the construction of ski-lifts, 

runs, access roads, car parks and resort accommodation and 

facilities; 

• Pacific islands habitats are threatened by the proliferation of 

golf courses and by the exotic vegetation often introduced; 

• mangrove swamps in eastern Australia are threatened by 

coastal resort development. 

In many cases these recreational activities involve the 

degradation of particularly sensitive habitats which tend to 

support isolated, relict and often taxonomically discrete 

populations of lycaenids and other insects. 

Examples could be multiplied several-fold, but the principle 

is well established, and the vital importance of suitable habitat 

and resources for conserving small animal and plant populations 

should not need further emphasis. 

Pollution 

The effects of chemical pollution on lycaenids are difficult to 

assess, but a number of declines of particular species in Europe 

have been attributed in part to atmospheric pollution, including 

acid rain. Such pollution is likely to affect the well being of 

sensitive foodplants and a wide spectrum of invertebrates 

associated with vegetation. Likewise, pesticide drift may cause 

occasional hazard, both in agricultural and forest environments. 

Exotic introductions 

No significant information is available on the deleterious effects 

of exotic taxa on native Lycaenidae in many parts of the world. 

The introduction of Dutch Elm Disease into Britain, with 

consequent large scale demise of Ulmus trees, led to a reduction 

in the numbers of the Whiteletter Hairstreak, Strymonidia walbum 

(Knoch). Similarly, the introduction of myxomatosis to 

Britain in the 1950s resulted in a drastic reduction of the 

intensity of rabbit grazing on chalk grasslands, with a resultant 

reduction in the numbers of several butterfly species, such as 

Lysandra bellargus (Rottemburg) and Maculinea arion. In 

New Zealand, Gibbs (1980) noted the trend towards decline of 

Zizina oxleyi (Felder & Felder) in parts of the South Island 

through hybridisation with the invasive Australian Z. labradus 

(both are sometimes treated as subspecies of Z. otis (F.) (Figure 

3)). Claims that hybridisation with the introduced Strymon 

melinus Hübner could threaten the Avalon Hairstreak, S. avalona 

(Wright), on Santa Catalina Island, California, need further 

investigation (Wells et al. 1983). 

In a rather different interaction between exotic and native 

species, Brown (1990) reported that the widespread Leptotes 

marina (Reakirt) had adapted well to urbanisation in North 

15 

America with its range expansion largely due to a 'switch' to a 

South African larval foodplant (Plumbago auriculata), which 

is used widely in freeway landscaping and as an ornamental. 

Furthermore, larvae associate closely with the introduced 

Argentine ant, and the prime nectar source for adults is a 

Brazilian tree. Leptotes thus has benefited from the presence of 

several different exotic species. 

Conservation of Lycaenidae 

The major sequence of needs in order to effect conservation 

programmes for any form of terrestrial wildlife is as follows: 

i) Documentation and education, to increase awareness at all 

levels and as a mode of communication between informed 

scientists and those who make practical decisions over 

priorities for land use; 

ii) Detection of habitats supporting either critical faunas or 

single notable or vulnerable species which merit protection 

and the promotion of their continued protection in existing 

National Parks and other reserves; 

iii) Investigation of the limits and/or wisdom of legislative 

protection for particular taxa or habitats, as an interim 

measure whilst additional documentation is obtained, and 

iv) Autecological studies of selected taxa as a basis for 

formulating sound management plans, a step which can 

come only from a basis of substantial research rather than 

haphazard extrapolation and which is, therefore, costly. 

v) Investigation of techniques for captive rearing, in case of 

need for ex situ conservation, or translocation. This should 

not be seen as a replacement option for in situ conservation. 

The information contained in this volume has hitherto been 

scattered through a wide range of reports (of varying degrees of 

formality and distribution) and scientific papers. In dealing 

with such a diverse group of insects, this book cannot be as 

definitive as 'Threatened Swallowtail Butterflies' (Collins and 

Morris 1985), but the examples given reflect a growing number 

of detailed studies on lycaenids and concern over their 

conservation. It represents a useful starting point for the 

development of conservation programmes for the Lycaenidae 

and should help to focus attention on groups or geographical 

regions in need of urgent attention. 

Public awareness 

Most conservation projects to date have been species orientated 

and many of these have done much to improve public awareness 

of the threats to lycaenid species. The potential for 

reintroductions is demonstrated well by the recent project 

involving the successful liberation of the Large Blue, Maculinea 

arion L. in Britain. This case is discussed in detail by Elmes and 

Thomas (1992) and by New ('Large Blues', this volume). 

Attempts to introduce the continental subspecies of the Large 

Copper, Lycaena dispar, into Britain, are of long standing 

(Duffey 1977 and this volume). Both projects highlight the

Figure 3. Distribution of Zizina oxyleyi (native) and Z. labradus (exotic) in 

New Zealand, and their hybrid zone (after Gibbs 1980). 

need for substantial amounts of ecological information for such 

management initiatives and they have done much to improve 

public awareness of butterfly conservation issues. 

There is no doubt that particular 'charismatic' butterflies 

amongst the Lycaenidae can do much to increase public 

awareness of conservation at a level which is otherwise difficult 

to realise, as witness the Large Blue campaign in Britain, the 

Mission Blue (Plebejus icarioides missionensis Hovanitz) in 

California and the more recent Eltham Copper (Paralucia 

pyrodiscus lucida Crosby) campaign in Australia. 

However, such species-orientated conservation is essentially 

confined to well-developed countries. It could not be achieved 

practically in much of the rest of the world, where conservation 

of any sort is considered a luxury and where there are few 

skilled entomological or conservationist practitioners and little 

if any local funding or sympathy for insect well-being. 

Any opportunity for promotion of significant species as 

' local emblems' is worth pursuing. For example the appearance 

of butterflies on stamps might help to gain public sympathy: 

lycaenids have been depicted on postage stamps of nearly 70 

countries (Coles et al. 1991 and Table 6) and these have 

included both less developed and developed countries. 

16 

Identification of threatened species and critical 

faunas 

The identification of these is clearly an immediate need in 

lycaenid conservation. The information contained in this volume 

indicates that while the former is the main focus in the developed 

world it is the latter which may be the most relevant path to 

follow elsewhere. The largest group of oriental Polyommatini 

(the 'Lycaenopsis group') was appraised by Eliot and Kawazoe 

(1983), and additional comprehensive studies of this sort would 

be of considerable value in setting conservation priorities on a 

faunistic basis. That study indicated a number of regions with 

high levels of diversity and/or endemism which could thus be 

considered as critical faunas (see Collins and Morris 1985 on 

Papilionidae, Ackery and Vane-Wright 1984 on Nymphalidae: 

Danainae). 

Other examples of critical faunas can be cited: the Philippines 

support more species of the Polyommatini than any other area 

of comparable size and some are endemic; Sulawesi is less rich 

than the Philippines but about half the species there are relict 

endemics or have only a narrow range elsewhere; most taxa in 

the Papuan subregion are endemic; centres of diversity (?refugia) 

for Riodininae occur in the neotropics (see taxa accounts for 

South America, this volume) and of other taxa in various parts 

of the world. Many of these areas are undergoing substantial, 

rapid and irreversible changes at present, and the prime need is 

to increase reservation of ecologically representative areas as a 

prelude to more detailed studies of their needs. Unless additonal 

reserves are forthcoming the rich diversity of Riodininae in the 

neotropics (see essays by Brown and Callaghan, this volume) 

and the substantial numbers of unusual Polyommatinae in 

southeast Asia will be seriously reduced. 

The emphasis on species-orientated lycaenid conservation 

is likely to occupy entomologists in temperate regions for some 

time to come. This has two important results. Firstly, detailed 

studies of particular taxa, which would be impossible in a 

broader context, may give a good basis for their proper 

management. Various projects on the reintroduction of species 

and successful management of threatened species, which are 

documented in the taxa accounts, attest to the depth of ecological 

knowledge necessary for the successful management of habitat 

for a particular species. In fact, for most Lycaenidae, preservation 

of suitable habitat is generally not in itself sufficient to ensure 

their long-term well-being: intricate management plans to 

conserve complex tripartite associations involving butterfly, 

ant and plant and to control seral succession by ensuring that 

some rapidly developing early stages may be continually 

available, are integral facets of successful conservation. 

Secondly, particular critical habitats may be reserved for the 

species and give benefit also to less obvious rare biota. However, 

few such studies result in reservation of large or well-buffered 

habitats. Systems akin to the British SSSI (Sites of Special 

Scientific Interest, history documented by Moore 1987) are not 

yet widespread elsewhere but merit serious attention in other 

parts of the world to ensure the adequate reservation of wellbuffered 

habitats.

Table 6. List of Lycaenidae which have been depicted on postage stamps up to October 1991. (Data from Coles et al. 1991; identifications not checked 

further, authors' names omitted). 

Species 

Aethiopana honorius 

Citrinophila erastus 

Hewitsonia boisduvali 

Mimacraea marshalli 

Pentila abraxas 

Telipna acraea 

Ogyris amaryllis 

Aphnaeus questiauxi 

Aphniolaus pallene 

Atlides polybe 

Axiocerses amanga 

A. harpax 

A. styx 

Bindahara phocides 

Callophrys crethona 

Chrysozephyrus mushaellus 

Deudorix epijarbas 

Epamera handmani 

E. sidus 

Eumaeus alula 

Evenus coronata 

E. dindymus 

E. regalis 

Hypokopelates otraeda 

Hypolycaena antifaunus 

Japonica lutea 

Lipaphnaeus leonina 

Loxura atymnus 

Myrina silenus 

Narathura centaurus 

Panthiades bathildis 

Pratapa ctesia 

Pseudalmenus chlorinda 

Pseudolycaena marsyas 

Rapala arata 

Ritra aurea 

Scoptes alphaeus 

Spindasis modesla 

S. natalensis 

S. nyassae 

Strymon maesites 

S. martialis 

S. melinus 

S. rufofusca 

S. simaethis 

Stugeta marmorea 

Tanuetheira timon 

Thecla betulae 

Agrodiaetus amanda 

A. damon 

Country, year of issue 

Ghana 1990 

Ghana 1990 

Uganda 1989 

Uganda 1989 

Ghana 1990 

Ghana 1990 

Australia 1981 

Zambia 1980 

Uganda 1989 

Grenada 1975 

Uganda 1989 

Mozambique 1953, Togo 1990 

Tanzania 1973 

Palau 1990 

Jamaica 1978 

China 1963 

Samoa 1986 

Malawi 1966 

Kenya 1988 

Cuba 1974 

Ecuador 1970, Nicaragua 1986 

Grenadines of St. Vincent 1975 

Belize 1990, Brazil 1979, Nicaragua 

1967 

Mali 1964 

Togo 1990 

Laos 1986 

Mali 1964 

China 1963 

Angola 1982, Mauritania 1966, Togo 

1990. Sierra Leone 1979, Tanzania 1988 

Malaysia 1970 

Belize 1974 

Laos 1986 

Australia 1981 

Grenada 1990, Grenadines of Grenada 

1985, St. Vincent 1978 

Korea (North) 1977 

Nicaragua 1986 

South Africa 1977 

Zambia 1980 

Swaziland 1987 

Malawi 1984 

Dominica 1988, Turks & Caicos Islands 

1982, Grenadines of St. Vincent 1989 

Cayman Islands 1988 

Honduras 1991 

Grenadines of Grenada 1985 

Grenada 1989, Grenadines of Grenada 

1985 

Sierra Leone 1987 

Congo 1971, Sierra Leone 1987 

Bulgaria 1990 

Finland 1990 

Mongolia 1963 

17 

Species 

Aricia agestis 

Brephidium exilis 

Catochrysops taitensis 

Celastrina argiolus 

Cupidopsis jobates 

Danis danis 

Glaucopsyche melanops 

Hemiargus ammon 

H. hanno 

H. thomasi 

Jamides bochus 

J. cephion 

Lampides boelicus 

Leptotes cassius 

Luthrodes cleotas 

Lycaeides idas 

Lysandra albicans 

L. beltargus 

L. coridon 

Maculinea arion 

Meleageria daphnis 

Polyommatus icarus 

Tarucus balkanicus 

Uranothauma crawshayi 

Heliphorus epicles 

Heodes solskyi 

H. virgaureae 


L. helle 

L. salustius 

Palaeochrysophanus hippothoe 

Thersamonia phoebus 

T. thersamon 

Abisara talantus 

A ncyluris formosissima 

A. jurgenseni 

Chorinea faunus 

Dodona adonira 

Helicopis cupido 

Melanis pixe 

Nymphidium mantus 

Nymula orestes 

Rhetus thia 

R. sp. 

Stalachtis calliope 

S. phlegia 

Country, year of issue 

Cyprus 1983 

Cayman Islands 1990, Turks & Caicos 

Islands 1990 

Samoa 1986 

Libya 1981 

Togo 1990 

Netherlands New Guinea 1960 

Cyprus 1983, Libya 1981 

Cayman Islands 1988, Cuba 1991 

Barbados 1983, British Virgin Islands 

1978 

Turks & Caicos Islands 1990 

Tonga 1989 

Solomon Islands 1980 

Ascension 1987, Fiji 1985, Vanuatu 

1991 

St. Vincent 1989 

Vanuatu 1983, Wallis & Futuna Islands 

1987 

Canada 1988 

Libya 1981 

Hungary 1959 

Switzerland 1952 

Gibraltar 1977, Great Britain 1981, 

Hungary 1969, Poland 1967 

Bulgaria 1962, Hungary 1966, Rumania 

1969 

Guernsey 1981, Ireland 1 985, Malta 

1986 

Turkey 1958 

Malawi 1973 

Hong Kong 1979, Macau 1985 

China 1963 

Finland 1990, Hungary 1959, Mongolia 

1977, Rumania 1960 

Germany (F.R.) 1991 

Germany (F.R.) 1991 

New Zealand 1970 

Hungary 1974, Nicaragua 1986 

Ifni 1963 

Hungary 1969 

Sierra Leone 1987 

Guinea 1973, Hungary 1984 


Guyana 1989 

China 1963 

Guyana 1983, Surinam 1971 


Guyana 1983 

Grenada 1975 


Honduras 1991 

Guyana 1989, Surinam 1971 

Surinam 1971

Managing lycaenid populations 

Various management plans for particular Lycaenidae have 

been produced, and these have several elements in common: 

habitat security; the need for management based on sound 

biological and ecological understanding of the target species; 

and the need for an understanding of the endangering processes. 

Increasing public awareness of the particular lycaenid is also 

commonly seen as important. The processes needed are well 

exemplified in a 'flow-chart' produced by Arnold (1983a) for 

Smith's Blue (Euphilotesenoptessmithi(Mattoni)) in California 

and augmented in his Recovery Plans for the Lotis Blue 

(Lycaeides idas lotis (Lintner) and other species (USFWS 

1984, 1985), and this has been used as a basis for the scheme 

outlined in Table 7. It is clear from this that the basic information 

necessary to formulate sound management for any ecologically 

sensitive species cannot be gathered instantly. All too often the 

time available is insufficient once a decision has been taken to 

develop a habitat containing a threatened species or population. 

The approach of 'Population Viability Analysis' (sometimes, 

'Population Vulnerability Analysis'), PVA, is receiving 

substantial attention in assessing conservation needs of vertebrate 

animals. In general, the detailed demographic and reproductive 

data needed for such predictive modelling are not available for 

extending this approach to invertebrates. However, a North 

American lycaenid, the Karner Blue (Lycaeidesmelissa samuelis 

Nabokov) has recently (April 1992) been the focus of the first 

specialist workshop held to appraise an insect in this context. 

This species lives in fire-successional vegetation, and the 

metapopulations are associated with local extinctions and 

recolonisation or habitat shifts linked with climate and fire 

regimes (Cushman and Murphy, this volume). The outcome 

from this workshop may mark a substantial augmentation to 

current methods of formulating management programmes for 

insects of conservation concern. 

Ex situ conservation 

Table 7. A pro-forma scheme for species-orientated conservation of Lycaenidae (after Arnold 1983a). 

1. Preserve, protect and manage known existing habitat to provide conditions needed by the species. 

(a) 

(b) 

(c) 

(d) 

Preserve: prevent further degradation, development or environmental modification. 

Steps (some or all): 

(i) Cooperative agreements with landowners/managers. 

(ii) Memoranda or undertakings. 

(iii) Conservation easements. 

(iv) Site acquisition (purchase/donation): private land. 

(v) Site reservation: public land. 

Maintain larval and adult resources at known habitat(s). 

Steps (some or all): 

(i) Minimise use of toxic substances: herbicides, pesticides. 

(ii) Minimise uncontrolled intrusion by humans: trampling, off-road vehicles, etc. 

(iii) Minimise intrusion by domestic stock: cattle, horses, sheep, etc. 

(iv) Minimise exotic vegetation planting. 

(v) Minimise removal of native vegetation, unless controlled. 

The controlled harvesting and ranching of Papilionidae to 

reduce pressure on natural populations and supply collector 

needs has proved to be an important conservation avenue for 

this group. It probably will never be achievable (or, indeed, 

needed) for lycaenids on any large scale, despite their uniqueness 

and vulnerability and their geographical overlap with areas 

where Papilionidae are also vulnerable. However, rearing of 

particular rare species in captivity does have some potential for 

augmenting field populations and for effecting translocations. 

The Atala Hairstreak, Eumaeus atala, has proved to be relatively 

easy to rear in captivity, and this practice is pursued by several 

butterfly houses in Britain using stock from Florida (Collins 

1987a). Some success has been obtained with attempts to start 

new colonies of E. atala in the wild, and although the species 

continues to be regarded as vulnerable to habitat loss it has 

become more widespread in recent years (Lenczewski 1980). 

Some Japanese Theclini may be reared relatively easily from 

eggs collected in the field, and eggs of species which are 

otherwise very hard to collect have been found (Kuzuya 1959). 

The larval stage lasts for 3–4 weeks, as does the pupal period, 

and the only major problem in rearing was a tendency for older 

larvae to be cannibalistic. 

Unlike most other butterflies, captive rearing of lycaenids 

may need to incorporate provision for ants, often of particular 

species for any given lycaenid, as well as foodplants. 

Propose critical habitat (U.S.). 

If necessary, clarify taxonomic status of target lycaenid in habitat and, where known, outlier or other populations. 

18 

Continued...

Table 7 (cont.). A pro-forma scheme for species-orientated conservation of Lycaenidae (after Arnold 1983a). 

2. Manage and enhance lycaenid population(s) by habitat maintenance and quality improvement, and reducing effects of limiting factors. 

(a) Investigate and initiate habitat improvement methods as appropriate. 

Examples: 

(i) Remove or control exotic weedy or noxious plants. 

(ii) Promote natural establishment of foodplants and other natural vegetation – if necessary, propagate and transplant, 

(iii) Promote particular grazing (grassland) or coppicing (woodland) regimes. 

(b) Determine physical and climatic regimes/factors needed by species and relate to local habitat enhancement in overall site. 

(c) Investigate ecology of the lycaenid species 

References 

(i) Life history and phenology; dependence on particular plant species/stages/organs. 

(ii) Dependence on ants or other animals, and their role in protection from predators and parasites. 

(iii) Population status; size, movement, degree of isolation, sex ratio, etc. 

(iv) Adult behaviour: mating, oviposition cues and sites, activity rhythms, 

(v) Determine predators, parasitoids and other factors which cause mortality or limit population growth. 

(vi) Investigate possibility of captive rearing from local population enhancement or range extension. 

(d) Investigate ecology of tending ant species and (if needed) homoptcrous prey. 

(e) Investigate ecology of food plant species 

(i) Life history and recruitment processes. 

(ii) Mortality and debilitatory factors, including other consumer species. 

(iii) Limiting factors – edaphic conditions, slope, exposure, etc. 

(iv) If needed, horticultural studies to determine propagation techniques for transplantation/augmentation of food plant stocks. 

3. Evaluate above, and incorporate into development of long-term management plan. Computer modelling may assist in making management 

decisions. 

4. Monitor lycaenid populations to determine their status and to evaluate success of management. 

(a) Determine sites to be surveyed, if choice available and/or logistics limited. 

(b) Develop methodology to estimate population numbers, distribution and trends in abundance. 

Examples: 

(i) Counts of larvae when feeding at night. 

(ii) Counts of adults by transect patrols, mark-recapture, etc. 

5. Throughout all above, increasing public awareness of the species by education/information programmes. 

Examples: 

(i) Information signs at key sites (unless risk of inducing unwanted intrusion, collecting, etc.) 

(ii) Interpretive tours, if secure sites permit without causing additional problems. 

(iii) Audio and visual programmes, publications. 

– TV/radio interviews and information on the species and its management. 

– Conservation education programmes for schools and community groups. 

– 'Popular style' newspaper articles. 

– Continuing liaison with all interested parties. 

6. Enforce available regulations and laws to protect species. Determine whether additional legal steps needed, and promote these if necessary. 

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19 

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20 

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Callophrys (Mitoura) (Lycaenidae). J. Lepid. Soc. 30: 169–183. 

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21 

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PART 2. REGIONAL ASSESSMENTS 

This section is important both for what it contains and for what 

it does not contain. It demonstrates that most of our knowledge 

of the status of particular Lycaenidae is derived from temperate 

regions, particularly in the northern hemisphere, and that the 

fauna of much of the Old World tropics (in particular) cannot 

yet be evaluated in this way. Much of the tropics supports a high 

diversity of butterflies, but few resident lepidopterists with the 

facilities for undertaking biological studies: much of what can 

be inferred is based less on ecological surveys and a detailed 

knowledge of biology and more on old collections and synoptic 

works. For example, books such as those by Corbet and 

Pendlebury (1978, Malaysia) and Seki et al. (1991, Borneo) 

give invaluable appraisals of those magnificent local faunas. 

The introduction to the latter includes passionate comment on 

conservation of butterflies in Borneo, and the text, photographs 

and keys facilitate identification of 379 species and subspecies 

– but the status of many of these is largely unknown, and the 

knowledge of their distribution often fragmentary. Many 

lycaenids there, as elsewhere, are indeed both rare and restricted 

– perhaps threatened by habitat alteration or destruction — but 

Introductory comment 

22 

it is not possible to give a sound biological overview of these 

unique faunas, except in such very general terms. 

There is abundant need to document lycaenid diversity in 

many of the nominal 'protected areas' in the tropics, and to 

ensure that the widest possible range of habitats is conserved, 

especially for the many specialist forest-frequenting taxa. 

Though few specific details are available, there seems little 

doubt that continuing forest destruction throughout the African 

and Asian tropics is adversely affecting these insects. 

References 

CORBET, A.S. and PENDLEBURY, H.M. 1978. The Butterflies of the Malay 

Peninsula. 3rd ed., revised by Eliot, J.N., Malayan Nature Society, Kuala 

Lumpur. 

SEKI, Y., TAKANAMI, Y. and OTSUKA, K. 1991. Butterflies of Borneo 

2(1). Lycaenidae. Tobishima Corporation, Tokyo.

Conservation biology of Lycaenidae: A European overview 

Miguel L. MUNGUIRA 1 , José MARTIN 1 and Emilio BALLETTO 2 

1 Departamento de Biología, Universidad Autónoma de Madrid, Cantoblanco, 28049 Madrid, Spain 

2 Dipartimento di Biologia Animate, Università di Torino, Via Accademia Albertina 17, 10123 Torino, Italy 

Biology of European Lycaenidae 

The biology of European lycaenids is generally well known as 

a result of both general and specific studies. A review of central 

European species was made by Malicky (1969a) who dealt with 

larval foodplants, phenology and overwintering stages. 

Relationships with ants were also reviewed by Malicky (1969b) 

and Fiedler (1990a, 1991). Books on regional faunas including 

detailed ecological information on a species-by-species basis 

are available for Italy (Verity 1943), the Netherlands (Tax 

1989), British Isles (Ford 1970; Emmet and Heath 1989) and 

Switzerland (SBN 1987). Specific autecological studies cover 

a wide range of species from almost every major group within 

the European fauna. Some of these studies are listed in Table 1, 

together with the geographic areas in which they were 

undertaken. Several studies in press or short notices have not 

been listed and a wider range of species is covered by these. 

Thus the information available covers roughly 50% of the 

European lycaenids. The Table clearly shows wider coverage 

of northern species and of the countries where the present 

authors are based (Italy and Spain), but nevertheless it gives a 

general idea of our knowledge of European lycaenids. 

While most of the northern species are well known as far as 

their biology is concerned, information on most Mediterranean 

species is not available, even on such basic topics as foodplants 

or habitat preferences. Greece, with its 12 endemic species, is 

the area where basic studies are most urgently needed. High 

altitude species are also under-represented in ecological studies 

due to the difficulties of reaching their habitats for long-term 

studies, but their conservation seems to pose fewer problems 

than that of lowland species. 

Synecological studies are generally less abundant and only 

refer to some geographic areas such as southern France (Cléu 

1950; Bigot 1952, 1956; Dufay 1961, 1965–66), Hungary 

(Uherkovic 1972,1975,1976), Czechoslovakia (Kralichek and 

Povolny 1978), Italy (Balletto et al. 1977, 1982a–e, 1985, 

1988, 1989), Switzerland (Erhardt 1985) and Germany 

(Kratochwil 1989a, 1989b, 1989c), or to some particular 

ecosystems (Fouassin 1961; Janmoulle 1965; Cléu 1957; 

Mikkola and Spitzer 1983; see also Ehrlich 1984). 

23 

Larval foodplants 

Most of the European lycaenids are polyommatines (Kudrna 

1986). The data on lycaenid larval foodplants in Table 2 clearly 

illustrate the fact that polyommatines have radiated in our area 

more than the other groups, using wider niches and evolving to 

produce a wider range of endemic species and peculiar ecotypes. 

Life cycles 

The most common strategy adopted by the European lycaenids 

to endure adverse seasons is larval quiescence. Most 

polyommatines hide away and spend the winter season (in 

northern Europe) or the summer and winter (in the Mediterranean 

areas) as second or third instar larvae. When the egg or pupa is 

the overwintering stage, a diapause normally takes place (e.g. 

Iolana iolas (Ochsenheimer), Tomares ballus (F): Munguira 

1989, Jordano et al. 1990). Theclines typically overwinter as 

eggs, lycaenines and polyommatines as larvae. In the last case, 

however, there is considerable variation, again supporting the 

idea of a wider radiation of the group in our area. Table 3 

summarises the data on overwintering stages for Spanish 

lycaenids (a sample covering 70% of European species). 

There is one generation per year for most species. This 

seems to be a clear adaptation to temperate climates which have 

a favourable summer and a cold winter in which insect life 

comes through a dormant period. All the theclines follow this 

pattern, but again the polyommatines show some variation, 

having species with two generations (e.g. Polyommatus [or 

Lysandra] bellargus (Rottemburg), Lycaeides idas (L.)) or as 

many generations as weather allows (e.g. Tarucus theophrastus 

(F.), Aricia cramera Eschscholtz, Polyommatus icarus 

(Rottemburg)). It is possible that species with two generations 

are not 'fixed' to this condition, but constrained by food 

availability or unfavourable weather conditions. Thus, many 

species with a low number of generations may increase the 

number when resource limits change. An example of this is 

Polyommatus icarus with one generation in northern Britain, 

two in southern Britain (Emmet and Heath 1989) and four to 

five in Spain (Martin 1982).

Table 1. European Lycaenid species which have been the subject of thorough autecological studies, and the areas where these studies were made. 

Nomenclature here, and in the text follows Kudrna (1986). 

Species 

Thecla betulae 

Satyrium pruni 

S. ilicis 

Quercusia quercus 

Callophrys rubi 

Tomares ballus 

Lycaena virgaureae 

L. phlaeas 

L. dispar 

Lampides boeticus 

Leptotes pirithous 

Glaucopsyche alexis 

G. melanops 

Maculinea rebeli 

M. alcon 

M. arion 

M. nausithous 

M. teleius 

Cupido minimus 

C. lorquinii 

Iolana iolas 

Cyaniris semiargus 

Plebejus argus 

P. pylaon 

Lycaeides idas 

Aricia morronensis 

A. agestis 

A. artaxerxes 

A. cramera 

A. nicias 

A. eumedon 

Polyommatus golgus 

P. dorylas 

P. nivescens 

P. thersites 

P. bellargus 

P. albicans 

P. hispana 

P. humedasae 

P. galloi 

P. icarus 

Agriades glandon 

A. zullichi 

Area 

U.K. 

U.K. 

Italy 

U.K. 

Germany 

Spain 

Sweden 

U.K. 

U.K. 

Spain 

Spain 

Spain 

Spain 

France 

Spain 

France 

Spain 

U.K. 

France 

France 

U.K. 

Spain 

Spain 

Spain 

U.K. 

Germany 

Spain 

Hungary 

Austria 

Sweden 

Belgium 

Spain 

U.K. 

U.K. 

Denmark 

Italy 

Sweden 

Spain 

Spain 

Spain 

Spain 

France 

U.K. 

France 

Italy 

Italy 

Spain 

U.K. 

Spain 

Spain 

24 

Reference 

Thomas 1974 

Thomas 1974 

Fiori 1957 

Thomas 1975 

Fiedler 1990b 

Jordano et al. 1990 

Douwes 1975 

Dempster 1971; Emmet and Heath 1989 

Duffey 1968 

Martin 1984 

Martin 1984 

Martin 1981 

Martin 1981 

Thomas et al. 1989 

Munguira 1989 

Thomas et al. 1989 

Munguira 1989 

Thomas 1980 

Thomas 1984a 

Thomas 1984a 

Morton 1985 

Munguira 1989 

Munguira 1989 

Rodriguez 1991 

Thomas 1985; Ravenscroft 1990 

Weidemann 1986 

Munguira 1989 

Bálint and Kertész 1990 

Malicky 1961 

Pellmyr 1983 

Leestmans 1984 

Munguira and Martin 1988 

Jarvis 1958, 1959; Hoegh-Guldberg and Jarvis 1970 

Hoegh-Guldberg and Jarvis 1970 

Balletto et al. 1981 

Wiklund 1977 

Munguira et al. 1988 




Chapman 1914 

Davis et al. 1958; Thomas 1983 

Sourès 197;, Nel 1978 

Schurian 1980 

Manino et al. 1987 

Balletto and Toso 1979 

Martin 1984 

Dennis 1984 

Munguira 1989 

Munguira 1989

Table 2. Larval foodplants in the European Lycaenidae. 

Subfamily 

Polyommatinae 

Theclinae 

Lycaeninae 

Theclinae 

Lycaeninae 

Polyommatinae 

TOTAL 

Taxa 

Most feed mainly on legumes 

Some genera have shifted to other food plants, e.g. 

Tarucus Moore, Celastrina Tutt 

Some Maculinea Ecke; Pseudophilotes Beuret 

Maculinea arion (L.) 

Freyeria Courvoisier 

Scolitantides Hubner 

Plebejus argus (L.) 

Vacciniina Tutt 

Aricia Reichenbach 

Agriades Hubner 

Cyaniris Dalman 

Most species feed mainly on trees and shrubs 

Some are adapted to legumes, e.g. 

Tomares ballus (F.), Callophrys rubi (L.) 

All species are restricted to: 

Egg 

9 (75%) 

1 (17%) 

4 ( 8%) 

14 (20%) 

Food Plants 

Fabaceae 

Rhamnaceae, Aquifoliaceae, Cornaceae, Araliaceae 

Gentianaceae, Rosaceae, Lamiaceae 

Boraginaceae 

Crassulaceae 

Fabaceae, Ericaceae, Cistaceae 

Ericaceae 

Geraniaceae, Cistaceae, 

Primulaceae 

Fabaceae, Plumbaginaceae 

Fagaceae, Rhamnaceae, Oleaceae, Ulmaceae, Ericaceae 

Fabaceae 

Polygonaceae (Rumex and Polygonum spp.) 

Table 3. Frequencies and percentages of species overwintering at different stages in the three subfamilies of Spanish Lycaenidae. Data taken from Martin 

(1982). 

European habitats of importance for 

lycaenids 

Although conservation practice involves a battle for every 

piece of land containing wild animals or plants, some ecosystems 

in Europe are particularly important because they have become 

rare or support valuable species or communities (Blab and 

Kudrna 1982; Kudrna 1986; Balletto and Casale 1991). The 

following are some of the habitats of special importance for 

lycaenid conservation. 

25 

Larvae 

_ 

5 (83%) 

40 (78%) 

45 (65%) 

Woodlands 

Pupae 

3 (25%) 

– 

7(14%) 

10(15%) 

Apart from regions of the extreme north of Europe, woodlands 

represent the only stable ecosystems from sea level up to the 

high altitude vegetational stages. Nevertheless they have been 

subject to the most severe human pressures, and very few 

remain in their natural condition. 

In the case of lycaenids, woodlands are generally used only 

by theclines which are mostly encountered on the borders of 

natural or artificial clearings. Whether this is the natural 

condition, or if theclines (as inhabitants of the canopy) can 

rarely be observed in closed woods, remains for the moment a 

matter of speculation.

Woodland damage has been particularly severe in central 

Europe (e.g. Germany), but the wide distribution of most 

theclines makes threats to whole individual species rather 

unlikely. Only some species such as Satyrium pruni (L.), S. 

acaciae (F.), Thecla betulae (L.), or Callophrys avis Chapman 

are locally rare or have declined in the last few decades (Heath 

1981; Heath et al. 1984). 

Screes 

Rocky areas with sparse vegetation usually have a very interesting 

flora and fauna in Mediterranean countries. Limestone outcrops 

and schists are particularly interesting habitats for some rare 

endemics such as Aricia morronensis Ribbe. The most interesting 

areas are high altitude habitats where vegetation is sparse and the 

climate makes living conditions particularly severe for butterflies: 

here only a few well adapted species can survive. Some of the 

species living on these areas are among the rarest European 

endemics: Agriades zullichi Hemming and Polyommatus golgus 

(Hübner). Turanana panagaea (Herrich-Schaeffer), a rather 

widespread xerophilous species in Turkey, becomes extremely 

scarce and localised in Europe and although no appropriate 

habitat study has been made, it is possible that it may qualify as 

a member of this group on the western side of the Aegean. 

Grasslands 

Ecologically, grasslands can be subdivided into two major 

types on the basis of their being climacic or seral. The former 

occur typically in the far north of Europe and at high elevations 

of mountain ranges. The latter are found over a wider range of 

altitudes and generally represent the direct or indirect result of 

human activities. Within these grasslands some areas defined as 

'seminatural' have conserved a very rich soil and diverse 

vegetation and are by far the most important habitats for 

lycaenid conservation. 

Grassland management is changing dramatically throughout 

Europe, and the effects of this upon butterflies are just beginning 

to be understood. Erhardt and Thomas (1991) and Balletto et al. 

(1989) have dealt with the effect of grassland management on 

butterfly populations, stating that traditional land uses are the 

best way to preserve the existing butterfly diversity and are 

essential for the survival of several endangered species. 

As far as butterfly conservation is concerned several types 

of grasslands can be considered although these categories are 

very heterogeneous from the vegetational point of view. 

Dry grasslands and steppes are only found in Europe in some 

Mediterranean countries and in central east Europe. They often 

develop on stony, poor soils from the Mediterranean to the 

montane vegetation stage and are generally characterised by a 

comparatively low turf, or in some cases by the presence of one 

or more grass species of the genus Stipa from which some of 

them derive their name. 

The xeric character of this type of vegetation is reflected in 

the structure of their butterfly communities which are dominated 

26 

by grass feeders (Satyrinae and Hesperiidae). Some more or 

less restricted endemics and other interesting lycaenids occur in 

these habitats, the most notable of these species being 

undoubtedly Plebejus pylaon Fischer von Waldheim. In 

mountain areas these grasslands host many Polyommatus species 

of the subgenus Agrodiaetus, some of which are extremely 

local. 

The conservation of these habitats is particularly important 

because traditionally they have been considered useless, and 

steppe landscapes are generally unattractive for the public and 

therefore given low priority for conservation. At low elevations, 

however, they have been used for sheep or goat grazing, mainly 

during the winter, or locally (e.g. Alps) as vineyards. 

Mesophilous grasslands represent the lusher, more humid 

counterpart of dry grasslands. They generally develop on deeper, 

richer soils and are very rich in lycaenid butterflies. The rare 

species present in such areas include Lycaena hippothoe (L.), 

Maculinea rebeli (Hirschke), M. arion, Aricia eumedon, 

Pseudophilotes bavius (Eversmann), Scolitantides orion (Pallas) 

and Lycaenides argyrognomon (Bergsträsser). 

Mountain meadows are maintained by extensive cattle or 

sheep grazing, or locally by seasonal mowing. As a consequence 

of the milk surplus in the EEC this management is changing to 

reafforestation for paper production, particularly in southern 

countries. On far too many occasions this management practice 

leads to plantations with alien tree species and to the total 

destruction of the original habitat. 

It is important to point out that the greatest diversity of 

European lycaenids occurs in this kind of habitat, due to the 

ecotone nature of most meadows in which patches of trees, 

shrubs and grasslands occur in the same places. The beautiful, 

flowered meadows blooming with butterflies are disappearing 

very quickly in Europe and only some mountain areas such as 

the Alps or the Pyrenees act as refuge areas. The role of 

extensive grazing is crucial to conserve these habitats, and 

subsidies will probably soon be needed to support some of the 

less productive farming practices. Other aggressive uses related 

to skiing and the expansion of mountain tourism are deleterious 

for the conservation of these habitats. 

High altitude grasslands are only present above the natural 

tree line, either at the highest elevations on the major mountains 

or in the extreme north of the continent. Their climacic character 

makes management less necessary and thus their conservation 

easier. An important number of the European rare lycaenids 

live in these habitats: all the Agriades species (including the 

extremely rare A. zullichi which is found in a habitat transitional 

with high altitude screes); Albulina orbitulus (Prunner), Cyaniris 

helena (Staudinger); and some subspecies of Aricia morronensis, 

Polyommatus eros (Ochsenheimer), P. eroides Firvaldszky, 

and P. golgus. 

Threats to this type of environment are comparatively 

limited and are mainly represented by the spreading of ski 

resorts and mountain tourism, or to extensive overgrazing in 

southwest Europe.

Wet meadows and grasslands are azonal, like screes, in the 

sense that they can develop almost at any altitude where 

appropriate soil conditions are met (in this case a high humidity). 

Particularly at low elevations they are among the most 

endangered habitats in Europe. Species living on these grasslands 

are threatened throughout their range and examples of these are 

some of the most endangered species of Maculinea: M. alcon 

(Denis and Schiffermüller); M. teleius (Bergsträsser); and M. 

nausithous (Bergsträsser). Threats to this type of habitat are 

often related to land drainage and include changes in land 

management as a consequence of the very productive nature of 

these grasslands. 

Shrublands 

Shrublands normally grow as a consequence of the action of 

disturbing ecological factors such as strong winds, short 

vegetation times, human management or recurrent fires. They 

are interesting because some rare species are almost restricted 

to these formations. 

True heathlands dominated by Erica species are the habitat of 

Plebejus argus, a species that has declined rapidly in England 

but which does not seem to have similar conservation problems 

in other countries. 

Mediterranean shrublands (garigues or chaparral formations) 

also support interesting rare species such as Iolana iolas, a 

circum-Mediterranean species living on chaparral. Another 

peculiar type of Mediterranean shrubland is represented by 

heathlands (with Erica arborea and E. scoparia) found on the 

highest, windswept elevations of Corsica, Sardinia and Elba 

(France and Italy, Balletto et al. 1989). This is the exclusive 

habitat of the endemic Lycaeides Corsica (Tutt). 

Kretania psylorita (Freyer) is a rare endemic living in 

Cretan shrublands dominated by an Astragalus species (Leigheb 

et al. 1990). Similar to these are some species living on biotopes 

dominated by Thymus species (called locally in Spain 

'tomillares') where some endemics (Cupido lorquinii Herrich- 

Schaeffer) or rare species (Pseudophilotes bavius, P. 

abencerragus (Pierret) and Scolitantides orion (Pallas)) live. 

Subalpine-type shrublands are found on mountains above the 

tree line. They are characterised by the presence of 

Rhododendron and Vaccinium, often in association with dwarf 

pine trees. Their butterflies are mainly boreal-alpine elements 

which include, in wet areas, Aricia nicias Meigen, and Vacciniina 

optilete (Knoch) among the lycaenids. 

Wetlands 

These are probably the most endangered among European 

habitats as a consequence of drainage either to control mosquitoes 

or to transform the habitat into agricultural land (particularly 

rice fields). Wetland drainage has affected huge areas, both in 

central Europe (see Kudrna 1986 for the case of Bohemia in the 

27 

16th century) and in the south of the continent (Italy in the 19th 

century). Typical lycaenids living on wetlands are Lycaena 

dispar (Haworth), L. helle (Denis and Schiffermüller) and at 

higher altitudes or latitudes, Vaciniina optilete. These three 

species are listed as threatened by Heath (1981) and L. dispar 

is listed in the Berne Convention as being one of the most 

endangered European lycaenids. 

Causes of decline and extinction of 

European lycaenids 

This subject has been considered by several authors when 

dealing with the broader topics of European butterflies (Heath 

1981, Thomas 1984b, Kudrna 1986) and insects of the 

Mediterranean Basin (Balletto and Casale 1991). A considerable 

amount of information and a number of opinions are also 

available for individual countries (Blab and Kudrna 1982; 

Viedma 1984; Balletto and Kudrna 1985; SBN 1987; Gonseth 

1987; Swaay 1990; Bàlint 1991; Kulfan and Kulfan 1991). 

Although no single review has dealt with the subject solely with 

reference to lycaenids, many papers pointing out causes of 

decline of individual species are now available (see Table 1 for 

references). An overview of the topic is provided in Table 4, 

where information on butterfly conservation at a European 

level is summarised. 

Habitat alteration or destruction 

All three authors referred to in Table 4, as well as various 

national reports, identify this factor as the most important one 

for butterfly decline throughout Europe. A change in habitat 

quality is the cause of all extinctions documented to have taken 

place. This applies also to the lycaenids Lycaena dispar and 

Maculinea arion in the United Kingdom (Duffey 1968; Thomas 

1980) and Lycaena hippothoe (L.), M. arion, M. nausithous and 

M. teleius in The Netherlands (Heath 1981). 

Some examples of documented extinctions in Europe are 

given in Table 5. Wetland and grassland destruction or alteration 

are the main causes of recent extinctions. 

Butterfly declines are reaching an alarming scale in most 

central and north European countries where a high proportion 

of the fauna is experiencing dramatic range reductions (Heath 

et al. 1984). The range reductions of the following lycaenid 

species have been caused by habitat changes: Plebejus argus 

(United Kingdom, Ravenscroft 1990); Polyommatus bellargus 

(United Kingdom, Thomas 1983); Satyrium pruni (United 

Kingdom, Thomas 1974);LycaenadisparandL.helle(Germany, 

Kudrna 1986); and Polyommatus exuberans Verity and L. 

dispar (Italy, Balletto et al. 1982a–e, Balletto in press). 

Habitat alterations can be quite subtle: for example, a slight 

change of growth in grass height on British Maculinea arion 

sites is enough to make the habitat unsuitable for the butterfly 

host ant (Thomas 1989). This change produced by grazing 

relaxation was sufficient to cause the disappearance of the

Table 4. Causes of decline or extinction of European butterflies in three different reviews of the topic. Similar causes (abbreviated) are listed on the 

same line. 

Heath 1981 

• Habitat destruction 

wetland drainage, changes in 

grassland management, 

forestry practices 

• Air pollution 

• Pesticides 

• Climatic change 

• Urbanisation 

• Tourism 

• Collecting 

• Commerce 

Thomas 1984b 

• Habitat changes 

wetlands, woodlands, 

agricultural habitats 

• Forestry management 


• Insecticides 

• Weather & climate 

• Butterfly collectors 

• Isolation and area 

Kudrna 1986 

• Wetland drainage 

• Grassland management 

• Afforestation 


• Weed & pest control 

• Urbanisation 

• Tourism & transport 

• Overcollecting 

• Earthworks 

Table 5. Extinctions of European lycaenids documented to have taken place in individual countries with an indication of the total number of extinct 

butterfly species in each case. 

Country 

United Kingdom 

Netherlands 

Denmark 

Luxembourg 

Extinct 

lycaenids 

3 

5 

1 

3 

Species 

butterfly from formerly suitable habitats. Extensive grazing is 

declining rapidly over almost all of Europe due to intensification 

of agricultural practices and abandonment of former grazing 

areas. Neither grazing relaxation nor the decline in extensive 

grazing is good news for butterflies because these insects have 

evolved in Europe over thousands of years together with man, 

and have probably benefited from the patchy structure of 

agrobiosystems in which grasslands, hedges and woodlands 

occur together. The apparent phenological synchronization 

observed in the Dolomites between butterfly cycles and 

traditional mowing cycles (Balletto et al. 1988) may be another 

example of such a process. 

Air pollution 

Almost every author identifies air pollution as a cause of 

butterfly decline, but this is not supported by detailed evidence 

in any case (Thomas 1984b). The effect of pollution on insects 

is better understood in soil or aquatic species, although it seems 

obvious that it may have a real effect upon terrestrial insects. A 

dispar, arion, semiargus 

arion, teleius, nausithous, hippothoe, semiargus 

dispar 

argiades, idas, dorylas 

28 

Total extinct 

butterflies 

high diversity and population number have been recorded on 

road verges in the United Kingdom (Munguira and Thomas 

1992), a habitat where pollution caused by car exhausts is 

expected to be particularly severe. This probably suggests that, 

at least for some species, direct effects of pollution are negligible, 

whereas indirect effects (for example, of acid rain on forests) 

may prove to be more important for butterfly populations. 

Chemicals (pesticides, herbicides, fertilizers) 

A negative effect on insect diversity is often attributed to any 

chemicals sprayed over natural or semi-natural habitats. The 

effect of pesticides has only been studied on common species, 

but it is also evident that no rare or endangered species can 

survive on heavily sprayed localities. Some indirect evidence 

for this can be found in the literature. In the Padano-venetian 

plains, Balletto et al. (1982e) found considerable differences 

between butterfly communities in heavily sprayed and 

traditionally managed rice fields. Whereas on average five 

butterfly species were observed on each of the relatively natural 

5 

15 

2 

8

habitats which supported Lycaena dispar, only one or two 

species could be found in the more heavily sprayed fields and 

here L, dispar was totally absent. Other unpublished observations 

give further evidence of this effect. The conclusion is that 

chemicals can reduce butterfly diversity and that they may be 

implicated in the disappearance of L. dispar from vast areas in 

the region. 

Erhardt (1985) showed a negative effect of grassland 

fertilization: whereas six lycaenids among 30 Lepidoptera were 

more abundant in an unfertilized meadow than in unfertilized 

areas only one lycaenid out of two Lepidoptera was more 

abundant in fertilized meadows. Butterfly diversity was also 

drastically reduced in fertilized compared with unfertilized 

meadows. Balletto et al. (1988), however, failed to demonstrate 

the same effect on six fertilized and three unfertilized plots in 

the Dolomites. 

Climatic change 

Any relationship between climate and butterfly fluctuations is 

also hard to establish. In the study of this process, authors have 

referred to short-term climatic fluctuations, which are the only 

ones we can analyse. Even this needs a database maintained 

over several years, and only the British Butterfly Monitoring 

Scheme (BMS) is now available for such studies. Pollard 

(1988) stated that some climatic parameters such as summer 

temperatures are correlated with high butterfly numbers. Climate 

can also represent an important factor when fluctuations in 

numbers occur in populations previously isolated by other 

means such as habitat destruction. The synergetic effect of both 

factors can certainly endanger populations that are already 

declining. For example, a coincidence of habitat damage and 

unfavourable weather accelerated the extinction of Maculinea 

arion in Great Britain (Thomas 1989). Long-term climatic 

changes normally represent a natural process, but it is now 

uncertain if the short-term fluctuations derived from the 

greenhouse effect will have any influence on butterfly 

populations. This certainly adds another factor probably having 

some effect on butterfly abundance or survival, particularly 

when considering rare species. 

Tourism and urbanisation 

Ever-expanding tourist facilities and advancing urbanisation 

are obviously among the factors that threaten many butterfly 

populations. Nevertheless their effects are far more restricted 

geographically than habitat destruction or alteration due to 

changes in land management practices. Tourism is particularly 

aggressive in some areas, such as the Mediterranean coast, the 

Alps or some parts of the Pyrenees, where huge areas have 

literally been covered by urbanisation or ski courses. 

There is no particular lycaenid restricted to coastal areas 

around the Mediterranean, but the effect of tourism clearly 

makes the species' habitats smaller, acting together with other 

more extensive impacts. In northwestern Italy Glaucopsyche 

melanops (Boisduval) and Satyrium esculi (Hübner) are 

29 

threatened by tourist resorts which are now spreading inland 

from the sea borders (already totally covered in tarmac and 

concrete). Some previously abundant populations of Lycaena 

thesarmon (Esper) have become extinct as a consequence of the 

spreading urbanisation around Rome. 

High mountain habitats are particularly susceptible to the 

impact of expanding tourist facilities because they host scarce 

and ecologically specialised forms. One example is represented 

by Vacciniina optilete in the Alps, which is declining in many 

parts of its former range (Balletto in press). Another example is 

represented by Agriades zullichi and Polyommatus golgus, 

very rare endemic lycaenids in Sierra Nevada (southern Spain). 

Their range is already restricted by a road and a ski resort 

development, but they may now disappear from one of their 

localities if the planned redevelopments really take place. 

Collecting and commerce 

Again, every review on the causes of butterfly decline and 

extinction deals with this topic, but appropriate studies on the 

effects of collectors on butterfly populations remain wanting. 

The appeal of this topic probably has something to do with 

ethics and with the fact that treating some sophisticated and 

non-renewable products of nature as items of commerce is now 

unacceptable; neither does it seem correct to kill animals that 

other agencies are striving to conserve. Some collecting is still 

necessary in areas where our faunistic knowledge is poor. 

Sometimes forbidding collection does little if any good for 

butterflies (Kudrna 1986). Results obtained from the 

enforcement of bans on butterfly collection in some cases have 

shown this to be an unsuccessful management practice: in 

Germany a ban on the collection of four butterfly species passed 

in 1936 has not prevented the dramatic decline of these species 

in the course of the last 55 years (Kudrna 1989). 

In large populations the number of butterflies a collector can 

take is really negligible, not reaching 10% of the total daily 

population estimates, while small populations are normally of 

little interest to commercial collectors. To destroy one of these 

small populations by collection it is necessary to kill almost all 

the butterflies seen during the flight period: this represents a 

highly time-consuming job with slight rewards for the collector 

(Munguira et al. in press). 

Legislation to protect species and habitats 

In the last 30 years some European countries have passed laws 

protecting butterfly species. Heath (1981) reviewed this topic 

gathering data from 25 European countries, 13 (52%) of which 

had some legislation on the matter while only three countries 

(12%) included lycaenids among the protected species. In this 

review protected lycaenids were Maculinea alcon, M. alcon, M. 

teleius, Lycaena dispar, L. helle and Polyommatus bellargus. 

Most of this legislation has been ineffective because it was based 

on the species themselves and paid little attention to habitats.

At the European level, the Council of Europe, an international 

organisation based in Strasbourg (France) that groups together 

almost all European countries, has endeavoured in recent years 

to provide collective tools for animal and plant conservation. 

This organisation sponsored the publication of Heath's 

Threatened butterflies in Europe (1981) and Collins and Wells' 

Invertebrates in need of Special Protection in Europe (1987). 

Partially as a result of these two reviews some invertebrates 

were included in the appendices to the Berne Convention 

(Convention on the Conservation of European Wildlife and 

Natural Habitats) in 1988. The Convention has been signed by 

18 States and is important because it takes into account the need 

for habitat protection together with species protection. Appendix 

II of the Convention protects 24 lepidopteran species, five of 

which are lycaenids (Lycaena dispar, Maculinea arion, M. 

teleius, M. nausithous and Polyommatus golgus). 

Another potentially useful international convention yet to 

be considered in this framework is the International Convention 

for the Conservation of Wetlands (Ramsar), 1971. Devised to 

ensure protection of all European important wetlands, Ramsar 

legislation is, in principle, an ideal instrument for the 

conservation of all wet meadows and wetlands including any 

threatened lycaenid species. The problem with this convention 

is that it was aimed at the conservation of birds and has been 

opened only in recent times to include other vertebrates 

(amphibians, reptiles). The possibility that butterflies may be 

also considered in the selection of Ramsar sites is unfortunately 

nowhere in sight, but insect conservationists should be aware 

that such a possibility exists for the future. 

Species used as emblems, highlights of the 

fauna 

Some individual species, generally characterised by a 

particularly striking appearance, have often been used to attract 

the attention of the general public toward conservation issues. 

These so-called 'panoramic species' have proved useful to 

promote the conservation of many animal groups. 

Although the importance of lycaenids from this point of 

view is not as clear as with papilionids (e.g. Parnasius apollo 

(L.), Papilio hospiton Géné) some species have been 

instrumental in arousing considerable attention from the public 

towards butterfly conservation. The best example for this is 

Maculinea arion in Great Britain. During the 1970s the dramatic 

decline of this species in its few remaining biotopes was closely 

followed by many amateur lepidopterists and the topic appeared 

in several local and national press releases. A fund was 

established to support the preservation of the species and 

sponsor scientific research on its habitat requirements. 

Unfortunately all this interest came too late to save the species 

which became extinct in 1979 (Thomas 1989). However, this 

process helped in advancing the knowledge of the biology of 

the species to such an extent that it was successfully 

reintroduced to some sites a few years later. Since then, the 

30 

peculiarities of the biology of the 'large blue' and of the 

Maculinea species in general have fascinated the public and 

they are now certainly emblems of European efforts to preserve 

butterflies. 

The reasons for the success of Maculinea species in this 

respect have something to do with their relatively large size and 

beauty among lycaenids, their obligate dependence on Myrmica 

ants and the fact that they are very sensitive to subtle changes 

in habitat quality (a perfect illustration of the reasons for 

butterfly decline in Europe). Maculinea species are subjects of 

conservation studies throughout Europe from Spain to Hungary 

and are being reintroduced in several countries where they had 

previously become extinct. They are also currently included in 

every European list of endangered Lepidoptera. 

Another emblematic lycaenid, from the conservation point 

of view, is Lycaena dispar, the first butterfly whose extinction 

was documented in the United Kingdom as early as 1851. The 

habitat of this species consists of marshes, fens, wet meadows 

and oxbow lakes. Drainage of such biotopes has been the main 

cause of its decline throughout its range. It is protected by 

national laws in the following seven countries: United Kingdom, 

France, Netherlands, Hungary, Germany, Finland and Belgium; 

and also by all the Contracting Parties of the Berne Convention 

(Collins 1987). Apart from the United Kingdom it has also 

become extinct in Denmark and endangered in all the rest of its 

European range. Lycaena dispar is emblematic because it is 

sensitive to a very common practice in Europe, wetland drainage, 

and because it is conveniently large in size and colourful like the 

Maculinea species. It is the only lycaenid included by Kudrna 

(1986) in his short list of endangered European butterflies. 

Species of economic importance as pests 

There are no real pests among European lycaenids. Only some 

very common species can live on Mediterranean legume crops 

and could be considered as potential enemies of cultivated 

plants, but their presence has never caused serious problems. 

Lampides boeticus (L.) is occasionally listed among pests of 

peas, and together with Polyommatus icarus and Leptotes 

pirithous (L.) can also be a potential pest of alfalfa crops 

(Martin 1984). 

Red Data List of European lycaenids 

Any Red Data List is subjective and normally tends to be 

influenced by the authors' personal experience. Even the Red 

Data List concept is controversial and many authors have 

expressed opinions against it (Diamond 1988). Some authors 

have suggested that a list of endangered habitats may be far 

more useful than species orientated red data lists (Balletto and 

Casale 1991), and Kudrna (1986) has proposed that the latter 

should only include species which are in need of emergency

action. In our opinion, policies concentrating on habitat 

conservation should be given priority over species-centred 

schemes although it must be remembered that it is sometimes 

easier for policy makers to focus on the protection of a species 

rather than the more complicated process of habitat protection. 

In general, we consider that our section on important habitats 

for lycaenids is more relevant than the present paragraph on 

threatened lycaenids. 

In an attempt to make our list as impartial as possible we 

have pooled data from five European-level compilations of 

threatened species (Heath 1981; Kudrna 1986; Collins and 

Wells 1987; the Berne Convention (Appendix II, 1988) and an 

unpublished list drafted by Van der Made during the Wageningen 

Symposium on 'Status of Butterflies in Europe' in 1989). For 

every species appearing on a list we have given a score depending 

on whether it was listed as endangered (3), vulnerable (2), or 

rare (1). Species appearing as indeterminate or other categories 

were not considered, and the Berne Convention species 

(Appendix II) have all been scored as 3. In this way the highest 

possible score for any species is 15 (if 'endangered' on all five 

lists) and the lowest is 1. 

With a few exceptions that will be dealt with later the results 

shown in Table 6 are surprisingly consistent for the high 

ranking categories ('endangered' and 'vulnerable' in our list). 

All the species listed as 'endangered' have been accorded this 

status in at least one of the lists referred to above. Species in the 

'rare' category are a more mixed group that can be interpreted 

as species on which different authors do not agree, or that are 

localised but common. 

Such a list, however, is far from perfect, mainly because for 

the north and centre of Europe there is a long tradition in 

conservation studies, whereas in the south of the continent 

comparable research has long been neglected. Polyommatus 

coelestinus (Eversmann), for example, is categorised only as 

'rare' because it is part of a complex which may be abundant in 

Anatolia and the Caucasus, but is certainly more than rare in 

other parts of Europe (e.g. Peloponnisos). There is a similar 

situation with Turanana panagea. 

Another problem clearly under-ranking some species has to 

do with taxonomy (Daugherty et al. 1990). A number of 

Polyommatus of the subgenus Agrodiaetus, for instance, have 

been recognised as separate species only recently. Accordingly 

they were not included in older lists. Another case is Agriades 

zullichi which was considered a subspecies of the comparatively 

common A. glandon (Prunner), a fact that previously obscured 

the true endangered status of what is now considered a distinct 

species. As a consequence, we think that Agriades zullichi, 

Polyommatus exuberans and P. humedasae (Toso and Balletto) 

should be listed as 'endangered' and Turanana panagea, 

Pseudophilotes barbagiae Prins & Poorten, Kretania psylorita, 

Polyommatus galloi Balletto & Toso, P. aroaniensis Brown 

and P. coelestinus as 'vulnerable'. 

31 

Priorities in the conservation of European 

lycaenids 

Some conclusions can be drawn from all that has been discussed 

in the preceding sections. Most butterfly conservationists agree 

that some general considerations are to be kept in mind if any 

conservation measure is intended to be effective. Therefore, the 

duty of ecologists and butterfly conservationists is to make 

these available in an understandable way to the people who 

make decisions. 

The main considerations which emerge from our appraisal 

are: 

(1) The fact that many lycaenid species are declining is 

supported by evidence derived from most European countries. 

Habitat destruction and changes in land use are apparently the 

two most important factors responsible for species decline. As 

a consequence, such alterations should be stopped in the areas 

where endangered species have their habitat. 

(2) Vulnerable species suffer the same threatening factors 

on a more local scale, but any action liable to pose a threat for 

them should be carefully considered, and the status of the 

species over its distributional range taken into account. 

(3) Wetlands and wet meadows are among the most 

vulnerable habitats in Europe. Accordingly, wetland lycaenids 

are unanimously regarded as the most endangered by butterfly 

specialists. A European wetland register is urgently needed and 

legislation passed in order to protect all European wetlands 

known to include populations of threatened species. As we have 

already said, the most suitable international framework for such 

an action is Ramsar which might be persuaded, at a future date, 

to incorporate butterfly data into the listing criteria. 

(4) Traditional land practices have been compatible for 

centuries with the survival of species living on those sites and 

their continuation may occasionally prove necessary. They 

should be maintained wherever they are vital for the survival of 

declining species, and enhanced in those places where they 

gave way to more aggressive practices. Nevertheless, this fact 

cannot be generalised and some species will certainly do better 

in a totally natural habitat. 

The policy of continuing with traditional practices may 

sometimes need subsidies, but may be enough to keep high 

standards of living in many areas as suggested by the IUCN 

conservation policies, and by several conservation specialists 

(Thibodeau and Field 1984). Abandonment of former extensive 

agricultural areas should be monitored, because this may 

eventually lead to changes in habitat as a result of natural 

succession and the resultant loss of many species adapted to 

traditional agrobiosystems. 

(5) Nature reserves should only be declared for those 

habitats and species whose conservation cannot be assured by 

current land uses. Special care should be taken to satisfy the 

ecological requirements of protected species to ensure their 

survival. This will mean in many cases that management will 

have to oppose the natural succession of vegetation. Conflicts 

with the requirements of other endangered or rare species

Table 6. A Red Data List of European Lycaenidae, obtained by summing 

scores drawn from other previous lists (see text). 

ENDANGERED (score 15–8) 


Maculinea teleius 

M. nausithous 

M. alcon 

Polyommatus golgus 

Maculinea arion 

VULNERABLE (score 7–3) 

Vacciniina optilete 

Lycaena helle 

Maculinea rebeli 

Cupido lorquinii 

Pseudophilotes bavius 

Agriades pyrenaicus 

Polyommatus humedasae 

P. exuberans 

Callophrys avis 

RARE (score 2–1) 

Plebejus pylaon 

Polyommatus eroides 

P. ainsae 

P. galloi 

P. violetae 

Aricia morronensis 

A. eumedon 

Pseudophilotes baton 

P. barbagiae 

Agriades zullichi 

Kretania psylorita 

Turanana panagea 

Cupido carswelli 

Iolana iolas 

Scolitantides orion 

Kretania euripilus 

Polyommatus aroaniensis 

P. coelestinus 

Thesarmonia thetis 

(15) 

(14) 

(14) 

(9) 

(9) 

(8) 

(6) 

(6) 

(4) 

(4) 

(4) 

(3) 

(4) 

(3) 

(3) 

(2) 

(2) 

(2) 

(2) 

(2) 

(2) 

(2) 

(2) 

(2) 

(2) 

(2) 

(2) 

(1) 

(1) 

(1) 

(1) 

(1) 

(1) 

(1) 

should also be considered in order to make management decisions 

which ensure the conservation of all target species. 

(6) Natural habitats where rare or endangered species live 

on climacic plant communities should also be preserved from 

any possible alteration by the creation of Natural or National 

32 

Parks. Priority should be given to those places where extremely 

localised lycaenids are present, such as the high elevations of 

Sierra Nevada (Spain), or the Idhi Mountain (Crete). Climacic 

forests or shrublands must also be preserved when listed species 

live on them, but special effort should be made to ensure that the 

ecological requirements of target species are really those of the 

climax and do not depart from it in any way, however subtle. 

(7) Finally, more research is still needed, even for the most 

well known among the European lycaenids. We suggest three 

topics of special relevance for lycaenid conservation: 

• the creation of a European database of all the areas relevant 

for lycaenid conservation with special attention to the needs 

of endangered or rare species; 

• the monitoring and mapping of each endangered or 

vulnerable species both on a national and European scale; 

• the study of the ecological requirements of every target 

species so that sound recommendations can be made to 

ensure their long-term survival. 

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THOMAS, J.A., ELMES, G.W., WARDLAW, J.C. and WOYCIECHOWSKY, 

M. 1989. Host specificity among Maculinea butterflies in Myrmica ant 

nests. Oecologia 79: 452–457. 

UHERKOVICH, A. 1972. Adatok a Dravasik nagylepkefaunajanak 

(Makrolepidoptera) ismeeretéhez. Vas. megvei Muz. Ert. 5–6: 115–145. 

UHERKOVICH, A. 1975. Adatok Buranya nagylepkefaunajanak ismeeretéhez. 

iv. A Villanyi-hegiség nappali lepkëi. Janus Pannonius Muz. Evk. 17–18: 

33–42. 

UHERKOVICH, A. 1976. Adatok a Dél-Dunanuntul nagylepkefaunajanak 

(Makrolepidoptera). Folia ent. Hung. 29: 119–137. 

VERITY, R. 1943. Le Farfalle diurne d'Italia. Vol. 2. Marzocco, Firenze. 

VIEDMA, M.G. 1984. Consideraciones acerca de la conservaación de 

especies de insectos. Real Acad. Ciencias, Madrid. 

WEIDEMANN, H.-J. 1986. Tagfalter. Band I. Entwicklung, Lebensweise. 

Neumann-Neudamm GmbH and Co. KG, Melsungen. 

WIKLUND, C. 1977. Observationer över ägglaggning födosöl och vila hos 

donzels blavinge, Aricia nicias scandicus Wahlgr. (Lep., Lycaenidae). 

Entomol. Tidskr. 98: 1–4.

Overview of problems in Japan 

Toshiya HIROWATARI 

Entomological Laboratory, College of Agriculture, University of Osaka Prefecture, Sakai, Osaka, 591 Japan 

In Japan, there are about 240 resident butterfly species which 

comprise palaearctic and oriental faunal elements. Fortunately, 

none of them has been rendered completely extinct but many 

local butterfly colonies seem to have been totally eradicated. 

The first legislation to protect butterflies as 'Tennen 

Kinenbutu' or 'Natural monuments' was promulgated by the 

national government in 1932 for Panchala ganesa (Lycaenidae, 

Arhopalini) in Nara City. Until now, a total of 37 species have 

been designated as 'Tennen Kinenbutu' by the national and 

local governments. However, in some cases, legislation for the 

prohibition of collecting without any effective measures for 

conservation seems to have been ineffective, especially when a 

taxon is designated as a protected species rather than as a local 

population with a definable habitat or biotype. In fact, the Nara 

population of P. ganesa seems to have become extinct without 

any precise records because collectors lost interest in studying 

protected species. Another case is that of Shijimia moorei 

(Leech) (Lycaenidae, Polyommatini). This species had been 

known to occur in east Asia in places such as China and Taiwan. 

It was not until 1973 that this species was discovered in Kyushu, 

Table 1. Lycaenidae from Japan listed by Hama et al. (1989). 

Species 

Artopoeles pryeri Murray 

Coreana raphaelis Oberthur 

Niphandra fusca Bremer 

Shijimiaeoides divinus Fixcen 

Tongeia fisheri Eversmann 

Lycaeides subsolana Eversmann 

Causes of decline 

Urbanisation 

Urbanisation; deforestation 

Urbanisation 

Road construction 

Larch forestation 

Habitat degradation 

Orchard and golf course construction 

Agriculture and spraying 

Factory construction 

Flood: foodplant extinction 

Urbanisation 

Succession 

Flood control works 

35 

Japan: its distribution is extremely local, feeding on Lysionotus 

pauceflorus (Gesneriaceae) which usually grows on Quercus 

trees (Fagaceae) in humid evergreen forest. Just after that, in 

1975, this species was designated as 'Tennen Kinenbutu' by the 

national government. In this case, the original colonies of 

Kyushu seem to have been conserved. However, there have 

been few additional records from other areas (except one from 

Nara, Honshu) because collectors do not publish records of 

protected species even if these are caught. It is believed that 

some populations of S. moorei other than that in Kyushu 

become extinct without any definite records of this. 

Apart from Shijimia moorei, the decline of Japanese 

butterflies is attributable to alteration in land management 

practices and its effect on butterfly habitat. In Japan, most of the 

victims, such as Shijimiaeoides divinus Fixcen (Lycaenidae, 

Polyommatini), Coreana raphaelis Oberthur (Lycaenidae, 

Theclini) and FabriciananerippeC & R. Felder(Nymphalidae), 

depend on habitats such as coppice or grassland which have 

been maintained by traditional agricultural practices such as 

slash and burn, periodical coppicing for fuels and charcoal 

Locality 

Setagaya, Tokyo 

Kawanishi, Yamagata 

Kiso, Nagano; Shiga 

Kiso, Nagano; Shiga 

Kiso, Nagano 

Aomori 

Aomori 

Azumino, Nagano 

Komagano, Nagano; Matsumato, Nagano 

Matsumato, Nagano 

Azumino, Nagano 

Minami-azumi, Nagano

production, grass-cutting, animal husbandry, and so on (Sibatani 

1989; Ishii 1990). 

In Japan, few projects on butterfly conservation are supported 

by governmental funding, but awareness is increasing in many 

local research groups. Primary interest on the decline of 

butterflies and campaigns for their conservation have mainly 

highlighted rather large and beautiful butterflies, i.e. the 

'Luehdorfia Butterflies', Luehdorfia japonica Leech and L. 

puziloi Ehrschoff (Papilionidae) and the 'Great Purple' Sasakia 

charonda (Nymphalidae). However, urgent management is 

required for some lycaenid species which have survived in 

harmony with the traditional land management that is now 

being superceded. 

In 1989, the Lepidopterological Society of Japan (LSJ) (a 

group with nearly 1500 members) published the first volume of 

'Decline and Conservation of butterflies in Japan' (edited by 

Hama, Ishii and Sibatani 1989) which was the first systematic 

approach to the problem of butterfly conservation in Japan. Six 

species of Lycaenidae were included in that book (Table 1). 

Soon after publication, the LSJ held its first seminar on 

'Conservation of Butterflies' in June 1990. We found that there 

36 

are many more local extinctions of butterflies than we expected, 

and fewer conservation projects supported by adequate budgets. 

Attempts to find adequate funds for protecting butterflies or 

their habitats and for monitoring and overseeing activities 

continue. In early 1990, a group of the LSJ was granted funds 

of JPY 3,450,000 from the Nippon Life Insurance Foundation 

for monitoring the latest states of distribution and changes in 

population size of Japanese butterflies as environmental 

indicators for quality of human life. 

References 

HAMA, E., ISHII, M. and SIBATANI, A. (Eds) 1989. Decline and conservation 

of butterflies in Japan. I. Lepidopterological Society of Japan, Osaka. 

ISHII, M. 1990. What has led to the butterflies declining? Shizen Hogo (The 

conservation of Nature) 337: 14–15. (In Japanese). 

SIBATANI, A. 1989. Decline and conservation of butterflies in Japan. In: 

Hama, E., Ishii, M. and Sibatani, A. (Eds) Decline and Conservation of 

Butterflies in Japan. I Lepidopterological Society of Japan, Osaka, pp. 

16–22.

Conservation of North American lycaenids – an overview 

J. HALL CUSHMAN and Dennis D. MURPHY 

Center for Conservation Biology, Department of Biological Sciences, Stanford University, Stanford, California 94305-5020, 

U.S.A. 

Introduction 

Two distinct patterns emerge when one examines the United 

States' Endangered Species List. First, although butterflies 

probably constitute less than 1 % of global insect species richness, 

they are disproportionately represented on the list: 53% (14 of 

26) of the insects currently afforded federal protection are 

butterflies. Second, members of the Lycaenidae (including the 

blues, coppers, hairstreaks and metalmarks) are 

disproportionately represented: the family comprises only 21% 

of the species-level butterfly fauna of North America (Scott 

1986), but constitutes 50% of the listed butterfly taxa: see 

Table 1, (Federal Register 1991a). 

This over-representation of lycaenids also extends to the list 

of candidate species awaiting protection, including some taxa 

at risk of imminent extinction (see Table 2). For the U.S. as a 

whole, lycaenids comprise 37% of all butterflies that are 

candidates for listing as endangered species, and this number 

rises to 49% if skippers (a group currently treated as a superfamily 

distinct from the 'true butterflies') are excluded (Federal Register 

1991b). In California, lycaenids are substantially overrepresented 

among those taxa listed as candidates for federal 

protection or known to be at particular risk, making up 10 of 

those 20 taxa, whereas the state contains only a third of the 

nation's lycaenid fauna (Murphy 1987a). In addition, the 

candidates list is dominated by taxa from California and/or 

Nevada, making up 71% of the candidates (20 of 28; Table 2). 

They include a subspecies of Plebejus saepiolus (Boisduval) 

(soon to be described) that is probably already extinct. Another 

undescribed subspecies of hairstreak, Incisalia mossii (Edwards), 

has been pushed towards extinction by overzealous collectors 

who have removed hostplants and larvae. Also included is a 

copper, Lycaena hermes (Edwards), that may be differentiated 

sufficiently from all known relatives to warrant its recognition 

as a monotypic genus. It has been extirpated in significant 

portions of its historical range on both sides of the California- 

Mexico border. 

There are two opposing ways of viewing the lycaenid 

dominance of endangered and threatened species, and the list of 

candidates for this status. The first interpretation of this pattern 

37 

is that it results from the biased study of U.S. butterfly families. 

It may be that non-lycaenid butterflies are just as endangered as 

lycaenids, but the latter have received more attention from 

biologists, and thus more is known about their imperilled state. 

While it is difficult to evaluate this contention, there are no 

obvious reasons why lycaenids should have received more 

attention than other families, given that they are small in size 

and not nearly as showy. We favour a second interpretation, 

which is that the patterns reflect ecological differences among 

the butterfly families. 

Here, we first provide a brief overview of the taxonomic and 

geographic distributions of North American lycaenids. We then 

discuss five interrelated characteristics of lycaenids that we 

suspect are responsible for, or at least contribute to, the group's 

extreme susceptibility to endangerment and extinction. We 

conclude by summarising the ongoing efforts to conserve North 

American lycaenids. Throughout the chapter, we focus primarily 

on lycaenids and conservation programmes in the U.S. In part 

this is a reflection of our experience with lycaenids in this 

region. However, our restricted emphasis also occurs because 

insect conservation in Canada and Mexico is far less developed 

(see review by Opler 1991). Particularly in Mexico, a great 

number of insect taxa are at risk, but specific details are lacking. 

Taxonomic and geographic distributions 

Lycaenids are somewhat under-represented in North America 

and many of them just barely enter the United States from 

Mexico. The subfamily Riodininae (the metalmarks) is especially 

under-represented when compared to the equatorial latitudes in 

the New World, with just 20 species (14%) in North America. 

By comparison, riodinines make up about two-thirds of the 

local lycaenid fauna in equatorial lowland communities of 

South America. Indeed, the genera that include all but three of 

the North American species – Apodemia Felder & Felder, 

Calephelis Grote & Robinson, and Emesis F. – reach much 

greater species richness to the south of the United States. 

The subfamily Lycaeninae (the harvesters, hairstreaks, 

coppers, and blues) is represented by more than 120 species,

Table 1. Lycaenid taxa that are listed as endangered by the U.S. government as of 15 July 1991 (Federal Register 1991a). CA = California stale. 

Species 

Apodemia mormo langei 

(Lange's Metalmark) 

Glaucopsyche lygdamus palosverdesensis 

(Palos Verdes Blue) 

Euphilotes baltoides allyni 

(El Segundo Blue) 

Euphilotes enoptes smithi 

(Smith's Blue) 

Icaricia (=Plebejus) icarioides missionensis 

(Mission Blue) 

Lycaeides argyrognomon (=idas) lotis 

(Lotis Blue) 

Incisalia (=Callophrys) mossii bayensis 

(San Bruno Elfin) 

Range 

including some endemic species groups. Feniseca Grote is an 

endemic, monotypic genus of the small tribe of harvesters 

(Miletini) whose predaceous larvae feed on homopterans. 

Among the hairstreaks (Theclini) are a number of largely 

neotropical genera that are represented by one or just several 

species, including Eumaeus Hübner, Atlides Hübner, 

Chlorostrymon Clench, Tmolus Hübner, Calycopis Scudder, 

Cyanophrys Clench, Strymon Hübner, Erora Scudder and 

others. The genus Callophrys Billberg (sensu stricto) is holarctic. 

Genera that are largely restricted to North America (and mainly 

distributed in the west) include the species-rich Satyrium Scudder, 

Mitoura Scudder, and Incisalia Scudder (lumped with Callophrys 

by many), and the striking monotypic Sandia Clench & Ehrlich. 

Representatives of the coppers (Lycaenini) are holarctic in 

distribution, but many representatives are endemic to North 

America. Two species are related to Old World species, Lycaena 

cupreus (Edwards) and L. phlaeas (L.), the latter being found 

also in Asia and Africa. Three other species are represented in 

eastern North America. The remaining eight species, with the 

exception of the aforementioned highly restricted Lycaena 

hermes, are distributed across the intermountain west. 

Like the hairstreaks, the blues (Polyommatini) include a 

number of genera that are more diverse in the neotropics, just 

barely reaching into the U.S. (i.e. Hemiargus Hübner and 

Leptotes Scudder), and genera that share a similar distribution, 

but are also represented in Africa (i.e. Brephidium Scudder and 

Zizula Chapman). Celastrina Tutt and Everes Hübner are 

holarctic in distribution as functionally are Glaucopsyche 

Scudder and Plebejus Kluk (includingAgriades Hübner, Icaricia 

Nabokov, Lycaeides Hübner, and Plebulina Nabokov), genera 

that show high affinity with many European genera (i.e. 

Maculinea van Ecke, Aricia Reichenbach, Polyommatus Kluk, 

Plebicula Higgins, Lysandra Hemming, and Agrodiaetus 

Hübner, among others). North American Philotiella Mattoni, 

the monotypic Philotes Scudder, and the highly subspeciated 

Euphilotes Mattoni are all related to Asian genera. 

CA 

CA 

CA 

CA 

CA 

CA 

CA 

38 

Food Plant 

Eriogonum nudum 

Astragalus trichopodus var. lonchus 

Eriogonum parvifolium, E. cinereum, E. fasciculatum 

Eriogonum latifolium, E. parvifolium 

Lupinus albifrons, L. formosus, L. variicolor 

Lotus formosissimus 

Sedum spathulifolium 

Regional numbers indicate that lycaenids make up a higher 

proportion of butterfly species richness in the far western 

portions of the continent. For example, they comprise 29 of the 

162 (18%) butterfly and skipper species in Georgia (Harris 

1972) and 46 of the 240 (19%) species in Colorado (Brown 

1957, corrected for recent taxonomic changes). For southern 

California, Emmel and Emmel (1973) list 167 species of which 

54 (32%) are lycaenids. On a narrower geographic scale, 

lycaenids make up 37 of the 122 (30%) butterfly species from 

the San Francisco Bay area (Tilden 1965) and 30 of the 91 

(33%) species from Orange County in southern California 

(Orsak 1978). 

Perhaps not surprisingly, it is in western North America – 

with its diverse topography, elevation, and vegetation types – 

where differentiation at or below the species level is greatest 

(see Scott 1986). Additionally, it is in this same region, which 

harbours the most diverse temperate-zone lycaenid genera at 

the species level (i.e. Mitoura, Callophrys, Incisalia, Lycaena, 

Plebejus, and Euphilotes), that geographically restricted taxa 

are receiving the most attention from conservation biologists. 

Lycaenid characteristics and 

susceptibility to endangerment 

Subspecific differentiation 

As can be seen in Tables 1 and 2, 100% of the lycaenids 

currently protected under the U.S. Endangered Species Act 

(ESA), and 88% of the candidates for this status, are not full 

species. In a striking show of appreciation for the process of 

evolution, the U.S. Congress included both full species and 

subspecies as protectable taxonomic units (for vertebrates, 

even distinct populations at risk of endangerment or extinction 

are protected). Indeed, the most publicised extinction of a North

Table 2. Lycacnid taxa that are on the list of candidates for endangered status with the U.S. government. 

Data are current as of 21 November 1991 (Federal Register 1991b). Abbreviations of U.S. states are as follows: California (CA), Florida (FL), Illinois (IL), Indiana 

(IN), Maine (ME), Massachusetts (MA), Michigan (MI), New Hampshire (NH), Nevada (NV), New York (NY), Ohio (OH), Oregon (OR), Pennsylvania (PA), and 

Wisconsin (WI). Status categories are as follows: 1 = sufficient information to support a proposal for listing as endangered or threatened; 2 = current information 

suggests that taxon is 'possibly appropriate' for listing as endangered or threatened; 3A = best available information indicates that the taxon is extinct; 3C = current 

information indicates that the taxon is more abundant or widespread than previously thought; S = a taxon known to have stable numbers, U = additional information 

is required to determine the taxon's current abundance trend, D = taxon with declining numbers and/or which is being subjected to increasing threats. 

Species 

Eumaeus atala florida 

(Florida Atala) 

Euphiloles battoides spp. 

(Baking Powder Flat Blue) 

Euphilotes enoptes spp. 

(Dark Blue) 

Euphilotes rita spp. 

(Sand Mountain Blue) 

Euphilotes rita mattoni 

(Mattoni Blue) 

Hemiargus thomasi bethunebakeri 

(Miami Blue) 

Icaricia (=Plebejus) icarioides spp. 

(Point Reyes Blue) 


(White Mountains Icarioides Blue) 


(Spring Mountains Icarioides Blue) 

Icaricia (=Plebejus) icarioides fenderi 

(Fender's Blue) 

Icaricia (=Plebejus) icarioides moroensis 

(Morro Bay Blue) 

Icaricia (=Plebejus) icarioides pheres 

(Pheres Blue) 

Incisalia lanoraieensis 

(Spruce-Bog Elfin) 

Incisalia mossii ssp. ('hikupa') 

(San Gabriel Mountains Blue) 

Incisalia mossii ssp. 

(Marin Elfin) 

Lycaeides melissa samuelis 

(Karner Blue) 

Lycaeides dorcas claytoni 

(Clayton's Copper) 

Lycaena hermes 

(Hermes Copper) 

Lycaena rubidus spp. 

(White Mountains Copper) 

Mitoura gryneus sweadneri 

(Sweadner's Olive Hairstreak) 

Mitoura thornei 

(Thome's Hairstreak) 

Philotiella speciosa bohartorum 

(Bohart's Blue) 

Plebulina emigdionis 

(San Emigdio Blue) 

Plebejus saepiolus spp. 

(San Gabriel Mountains Blue) 

Status 

2,S 

2,U 

2,U 

2,U 

2,U 

3C,U 

2,U 

2,U 

NV 

2,U 

NV 

2,U 

2,U 

3A 

3C 

2,U 

2,U 

1,D 

2,S 

2,U 

2,U 

2,U 

2,U 

2,U 

2,U 

2,U 

39 

Range 

FL 

NV 

NV 

NV 

NV 

FL 

CA 

CA, 

CA, 

OR 

CA 

CA 

ME,NY,NH, 

Canada 

CA 

CA 

IN,MI,NH,NY,OH 

MA,IL,WI,PA. 

ME 

CA, 

Mexico 

CA.NV 

FL 

CA 

CA 

CA 

CA 

Food plant 

Zamia pumila 

Eriogonum sp. 

Eriogonum sp. 

Eriogonum sp. 

Eriogonum sp. 

Eriogonum sp. 

Lupinus sp. 

Lupinus sp. 

Lupinus sp. 

Lupinus sp. 

Lupinus chamissonis 

Lupinus sp. 

Picea mariana 

Sedum sp. 

Sedum sp. 

Lupinus perennis 

Potentilla sp. 

Rhamnus crocea 

Rumex sp. 

Juniperus silicicola 

Cupressus forbesii 

Chorizanthe membranacea 

Atriplex canescens 

Lupinus sp. 

Continued…

Table 2 (cont). Lycaenid taxa that are on the list of candidates for endangered status with the U.S. government. 

Species 

Plebejus saepiolus spp. 

(While Mountains Saepiolus Blue) 

Plebejus shasta charlestonensis 

(Spring Mountains Blue) 

Satyrium auretorum fumosum 

(Santa Monica Mountain Hairstreak) 

Slrymon acis bartrami 

(Bartram's Hairstreak) 

Status 

2,U 

NV 

2,D 

American butterfly was a putative full species, the Xerces Blue 

(Glaucopsychexerces (Boisduval)), a lycaenid that was probably 

a well-differentiated subspecies of the still widely distributed 

and highly variable Glaucopsyche lygdamus (Doubleday). 

Lycaenids appear to have a high degree of subspecific 

variation and these differentiated populations (many formally 

recognized as subspecies) are particularly susceptible to 

endangerment and extinction. Analysis of Scott's (1986) 

taxonomically conservative treatment provides empirical 

support for this view. According to his classifications, the ratio 

of subspecies to species in North America is 2.4 times greater 

for lycaenids than for non-lycaenid butterflies (187/142 and 

303/537 respectively). In addition, this ratio for those lycaenids 

largely restricted to western North America (i.e. Mitoura, 

Callophrys [including Incisalia], Lycaena, Plebejus, 

Glaucopsyche and Euphilotes) is 4.7 times higher than that for 

non-lycaenid butterflies (138/52). 

Dispersal abilities 

Dispersal is essential for the persistence of isolated populations. 

First, input of individuals from neighbouring areas can bolster 

populations whose numbers are dwindling, thereby preventing 

their extinction (the 'rescue effect' sensu Brown and Kodric- 

Brown 1977). Second, dispersal can provide an influx of 

genetically different individuals into a population, thereby 

increasing genetic diversity and presumably resulting in greater 

fitness and population viability (see review by Vrijenhoek 

1985). Thus, taxa with limited dispersal abilities should be far 

more susceptible to local extinction events than taxa with welldeveloped 

dispersal abilities. 

Although there have been numerous mark-recapture studies 

of North American butterflies (e.g. Ehrlich 1965, Arnold 1983, 

Murphy et al. 1986, Reid and Murphy 1986), few have focused 

on lycaenids. Despite the fact that many studies tend to 

underestimate mean dispersal distances, research to date 

indicates that lycaenids tend to move short distances between 

captures. For example, studies of the endangered Mission Blue 

(Plebejus icarioides missionensis Hovanitz) indicate that most 

adult movements are highly restricted: the majority of captures 

were in the immediate vicinity of larval hostplants and nectar 

resources (see Arnold 1983, Reid and Murphy 1986). However, 

studies have also shown a limited number of movements in the 

2,U 

2,U 

40 

Range 

CA, 

NV 

CA 

FL 

Food plant 

Lupinus sp. 

Trifolium, Astragalus, 

Lupinus 

Quercus sp. 

Croton linearis 

order of hundreds of metres, as well as a single dispersal event 

between habitat patches (and distinct demographic units) of 

nearly 2 kilometres. Nevertheless, the tendency for lycaenids to 

be comparatively sedentary should result in less frequent 

recolonisation and rescue events as well as reduced gene flow 

between populations, leading to greater interpopulation 

differentiation. 

Host specificity and successional stages 

All lycaenids currently protected in the U.S. are specific to one 

or just several related hostplant species (Tables 1 and 2). Many 

of these plant species, particularly the commonly used 

Eriogonum and Lupinus, are found largely in early successional 

communities that are temporary and unpredictable. Butterflies 

that specialise on such plants must track an ephemeral resource 

base that itself may be dependent on unpredictable and perhaps 

infrequent ecosystem disturbances. For such species, suitable 

habitat can be a limited, ever-shifting fraction of a greater 

landscape mosaic. As a result, local extinction events are both 

frequent and inevitable. 

The endangered Karner Blue (Lycaeides melissa samuelis 

Nabokov) is a prime example of the costs associated with such 

specialisation. Larvae of this subspecies feed exclusively on a 

lupine (Lupinus perennis) that is an early successional species 

restricted to pine-barren habitats (Zaremba 1991). The existence 

of these habitats is highly dependent on the occurrence of 

intermittent fires. However, in New York State, fire suppression 

and habitat loss have significantly reduced the size and number 

of this butterfly's patchily distributed populations. 

Even when appropriate larval hostplants and successional 

stages are available, conditions may still be insufficient to 

sustain lycaenid populations. For example, the federally listed 

San Bruno Elfin (Incisalia mossiibayensis (Brown)) is not only 

restricted to a few rocky outcrops that support its narrowly 

distributed hostplant (Sedum spathuliifolium), but the butterfly 

apparently can only complete its life cycle on individual plants 

growing under highly exacting and uncommon topoclimatic 

environments (Weiss and Murphy 1990). 

Sedentary behaviour combined with high levels of sitespecific 

hostplant adaptation are likely to place many genetically 

distinct lycaenids at great risk of local and regional extinction. 

While no studies document the existence of genetically based

host races in North American lycaenids, recent work details a 

genetic basis for larval hostplant preferences by adult butterflies 

in the rather sedentary nymphalid genus Euphydryas. Singer 

and colleagues have found significant differences in patterns of 

oviposition preference and tolerances for hostplants among 

phenotypically and geographically distinct populations, 

suggesting distinct adaptation at the population level (see 

Singer 1971; White and Singer 1974; Rausher 1982). More 

recently, Singer and colleagues have documented the existence 

of genetically based (i.e. heritable) differences between adjacent 

populations and within polyphagous populations (Singer 1983; 

Singer et al. 1988). 

Species in the lycaenid genus Euphilotes exhibit similar 

patterns in the use of larval hostplants, thereby suggesting the 

possibility of genetic differentiation among populations for 

hostplant tolerance (see Pratt and Ballmer 1986). In southern 

California, Euphilotes enoptes (Behr) has been found to feed on 

five species of Eriogonum; the butterfly is monophagous at 

some locations, while it is polyphagous in others, with clear 

host preferences. 

Association with ants 

Roughly half of the lycaenid species world-wide associate with 

ants, and their larvae possess numerous distinctive structures 

that facilitate these interactions (Downey 1962; Atsatt 1981; 

Pierce 1987). Although a few of these associations are 

antagonistic, with butterfly larvae preying on ant brood (Cottrell 

1984), the majority appear to be mutualistic (Pierce 1987). 

Lycaenids vary greatly in terms of their degree of dependence 

on ant associates and their degree of specificity for particular 

ant species. 

We propose that butterfly species which associate with ants, 

and particularly those species with strong dependence on them, 

are far more sensitive to environmental changes and thus more 

prone to endangerment and extinction, than species that are not 

tended by ants. While this hypothesis remains untested, it seems 

probable because of two factors. First, such species 

simultaneously require the right food plant and the presence of 

particular ant species – a combination that occurs infrequently. 

These dual requirements of tended species should result in 

spatial distributions that are patchier than those for untended 

species. The degree of patchiness should increase as dependence 

and/or the species specificity of lycaenids increase. Second, we 

suspect that selection will favour reduced dispersal by 

myrmecophilous lycaenids, because of the difficulty associated 

with locating patches that contain the appropriate combination 

of food plants and ants. Thus, in addition to occurring as 

isolated populations of variable sizes, ant-tended species may 

express genetic traits associated with reduced outcrossing. 

At this time, we cannot evaluate whether North American 

lycaenids that associate with ants are more vulnerable to 

endangerment or extinction than those without such 

dependencies. This is because there have been almost no 

studies of the ant associations of endangered lycaenid taxa. 

However, Downey (1962) observed that the Mission Blue was 

41 

tended by the ant Formica lasioides, and suggested (but did not 

demonstrate) that ants may protect caterpillars from natural 

enemies and even transport them to their food plants. D.A. 

Savignano has studied the ant associations of the Karner Blue, 

but this work has yet to be published. 

Numerous studies of non-endangered taxa in North America 

suggest that ants could be an important factor in the persistence 

of lycaenid populations. For example, parasitism levels of 

Glaucopsyche lygdamus oro (Scudder) (the Rocky Mountain 

subspecific relative of the federally listed Palos Verdes Blue, G. 

lygdamus palosverdesensis Perkins and Emmel) were 45–84% 

lower for ant-tended larvae than for untended larvae (Pierce and 

Mead 1981; Pierce and Easteal 1986). In Michigan, the work of 

Webster and Nielson (1984) also suggested that ant associates 

were beneficial for the Scrub-oak or Edward's Hairstreak, 

Satyrium edwardsii (Grote & Robinson). Clearly, we need to 

know much more about the ant associations of endangered 

lycaenids, as these interactions will be important considerations 

in management plans. 

Although not from North America, the Large Blue 

(Maculinea arion (L.)) provides an important, sobering, example 

of the often dire consequences associated with a dependence on 

ants (see Thomas 1980; Cottrell 1984; New 1991). Despite 

considerable efforts to prevent its loss, in 1979 the Large Blue 

became extinct in its native Britain. While many factors 

undoubtedly contributed to this demise, the most prominent 

appears to have been the species' extreme dependence on ants. 

During early instars, M. arion larvae fed on wild thyme (Thymus 

drucei praecox) and, at the fourth instar, were carried by 

Myrmica ants into their nests, where the lycaenids fed on ant 

brood. The level of grazing in the blue's grassland habitats was 

progressively reduced from around 1950, largely due to changing 

agricultural practices and attempts to protect habitat of this 

endangered species. However, due to unforeseen complexities 

of the system, these altered grazing regimes had drastic effects 

on the lycaenid populations. The primary ant-species host (M. 

sabuleti) could persist only in fields that were closely cropped 

by livestock. Thus, even slight reductions in grazing allowed M. 

scabrinodis, a low-quality host, to exclude M. sabuleti from the 

area, thereby leading to the butterfly's subsequent demise. 

Conservation planning in North America 

Lycaenids have played a central role in the development of 

environmental interests over land-use policy. Although much 

less publicised than the Large Blue another lycaenid provides 

a further example of the kind of conservation efforts that are 

required to protect endangered butterflies. 

The Mission Blue was conferred protection under the ESA 

in 1976, when the U.S. Fish and Wildlife Service formally 

recognized that encroaching urbanisation had virtually encircled 

the known distribution of this subspecies. More than half of the 

grassland habitat of the largest remaining known population on 

California's San Bruno Mountain had been lost during the 50 

years preceding the listing. Furthermore, half of this remaining 

habitat (a quarter of the total) had been overtaken by invasive

shrub and tree species. In 1978, developers and local government 

became aware that pending development would be prohibited 

by Section 9 of the ESA. 

Given that the Mission Blue occurred primarily on private 

land and the ESA only offered remedies for taxa on public 

lands, there was a pressing need for an innovative plan that 

would balance biological and economic concerns. Such an 

approach was engineered by a committee composed of 

developers, environmentalists, government officials, and 

biological consultants. Using size estimates of the butterfly 

populations, distributional records for three lupine (Lupinus) 

larval hostplants, and information on the butterfly's natural 

history, the committee designed the first 'Habitat Conservation 

Plan' (HCP). This plan protected 80% of remaining habitat on 

San Bruno Mountain, provided funds for the management and 

restoration of this habitat, and allowed for the development of 

the remaining land. In 1982, the U.S. Congress institutionalised 

habitat conservation planning with amendments to the ESA, 

pointing to the Mission Blue conservation program as the 

model for this new process. Several dozen HCPs (many of them 

controversial) have been initiated in the ensuing decade. 

In a second noteworthy case, a similar impasse between 

developers and environmentalists has focused on the Karner 

Blue in upstate New York. As mentioned earlier, this highly 

threatened subspecies, protected by the state but not the U.S. 

government, is restricted to fire-maintained gaps in early 

successional pitch-pine and scrub-oak barrens. The rather 

sedentary blue exists as a limited suite of metapopulations, 

consisting of collections of local populations that are dependent 

on a shifting mosaic of suitable habitat. 

There has been an extensive, broad-based effort to conserve 

the remaining Karner Blue population (see review by Zaremba 

1991). Primarily through the joint efforts of The Nature 

Conservancy and the New York State Department of 

Environmental Conservation, approximately 800ha of Karner 

Blue habitat have been protected as part of the Albany Pine 

Bush Preserve. In addition, the New York State Legislature 

established the Albany Bush Commission and charged the 

group with managing the remaining habitat. This and other 

ongoing programmes have involved prescribed burning, 

hostplant propagation, creation of effective dispersal corridors, 

and development of land-use practices to promote butterfly 

dispersal in areas adjacent to the preserve. Using the Karner 

Blue, Givnish et al. (1988) provided a model study for application 

of population viability analysis to conservation planning. 

Virtually identical in structure to an independently generated 

analysis of the well-studied nymphalid Euphydryas editha 

(Boisduval) (Murphy et al. 1990), this study broke from the 

traditional treatments of genetic threats and demographic 

stochasticity, and instead targeted environmental perturbations 

and metapopulation dynamics in an integrated scheme of reserve 

design and management. 

Conservation planning for lycaenids has focused primarily 

on habitat management. All of the listed taxa, and many of the 

candidates, survive as remnant populations in small 'garrison' 

reserves embedded within largely urban areas with long histories 

42 

of human settlement (Murphy 1987b). In such circumstances, 

opportunities to expand current distributions are few and 

conservation is less a matter of reserve design than reserve 

management. Heroic management efforts have brought Lange's 

Metalmark (Apodemia mormo langei Comstock) back from the 

brink of extinction. In 1976, the subspecies was listed as 

endangered by the U.S. government, as its remaining habitat 

was being threatened by sandmining and industrial development 

(see Opler 1991). The subspecies is restricted to the riparian 

sand dunes along the Sacramento and San Joaquin Rivers in 

central California. To prevent a further decline in numbers, in 

1980, the U.S. Fish and Wildlife Service acquired all of the 

lycaenid's remaining habitat. While seriously degraded by 

sandmining and invasive plants, this 24ha region was 

incorporated into the San Francisco National Wildlife Refuge, 

and there have been considerable efforts to restore the habitat. 

After a number of unsuccessful attempts to increase butterfly 

numbers, managers found that disking portions of the habitat 

resulted in dramatic growth of the larval and adult foodplant 

(Eriogonum nudutri). Since this action, the metalmark's 

abundance is estimated to have more than tripled, from fewer 

than 200 individuals in 1986 to more than 650 in 1989. 

Conclusions 

Extreme environmental events (such as drought, deluge, 

wildfire) can lead to dramatic fluctuations in the size of local 

butterfly populations, and in some well-documented cases, this 

has resulted in their extinction (e.g. Ehrlich et al. 1980; Murphy 

and Weiss 1988). For example, Ehrlich et al. (1972) reported 

that an early summer snowstorm caused the extinction of at 

least one subalpine population of Glaucopsyche lygdamus 

when it destroyed the entire standing crop of its larval food 

plant. Biotic factors, including the impact of natural enemies 

and ant associates, can also lead to significant variation in the 

size of lycaenid populations. However, as stressed throughout 

this volume, a long history of human-induced habitat alteration 

and destruction is responsible for the vast majority of declines 

and extinctions of lycaenids worldwide. 

This has certainly been the case for North American 

lycaenids, as already discussed for a number of taxa in this 

chapter. The El Segundo Blue (Euphilotes battoides allyni 

(Shields)) is yet another example. This small lycaenid is restricted 

to the El Segundo sand dunes along the coast of southern 

California. While most of its habitat, perhaps more than 10,000ha 

in extent, has been destroyed by housing and commercial 

developments, small portions of the sand-dune ecosystem (which 

support its preferred larval hostplant Eriogonum parvifolium) 

still remain near the Los Angeles International Airport and a 

Standard Oil refinery (Arnold 1983). While noteworthy efforts 

have been made by Standard Oil to establish populations of the 

El Segundo Blue on their lha property, comparatively little has 

been done by the City of Los Angeles on the inhabitable 

sections of their 80ha airport site. Since the mid 1970s, there

have been a number of unsuccessful attempts to develop much 

of the remaining site, several with revenue-generating activities 

linked to habitat management plans. 

Pressure to develop and disturb lycaenid habitat in North 

America is likely to intensify in coming decades, and more 

listings of endangered and threatened species should be expected. 

Only an increase in the already considerable efforts to conserve 

lycaenids (and insects in general) will suffice to keep more of 

them from suffering the fate of the recently extirpated Palo 

Verdes Blue in southern California (see Arnold 1987). 

Establishment of the Xerces Society in 1971 has been an 

important step, as it has greatly increased the attention paid to 

insect conservation. The ESA of 1973 and the use of HCPs have 

also been instrumental in facilitating the protection of threatened 

and endangered lycaenids on both public and private lands. 

Unfortunately, the U.S. government has added only six insects 

to its list since 1981 (Opler 1991) – despite the fact that habitat 

degradation and the list of candidates have increased 

considerably during the intervening years. This reluctance to 

list species must be reversed, so that the power of the ESA and 

HCPs can have their designed effect (Murphy 1991). 

Although the ESA and HCPs are powerful instruments of 

conservation planning and management, they are nevertheless 

'stop-gap' measures designed to take effect after taxa are in 

trouble. As attention continues to be focused on these essential 

management efforts, considerable effort must also be directed 

towards more long-term objectives associated with dramatically 

reducing the environmental degradation that is leading to the 

endangerment and extinction of additional taxa. As outlined by 

Ehrlich and Ehrlich (1990), Gore (1991) and others, these longterm 

objectives include stabilisation and then reduction of the 

growth of human populations, rapid development and 

deployment of environmentally appropriate technologies, 

comprehensive changes in the system of economic accounting 

so as to accurately reflect the effects of our actions on the 

environment, and the development of a detailed scheme for 

environmental education and research. 

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289–302.

Neotropical Lycaenidae: an overview 

Keith S. BROWN, JR. 

Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109 Campinas, 

Sao Paulo 13.081, Brazil 

Introduction 

The neotropical Lycaenidae are still only partly known and 

very little studied. They are here taken to include the Riodininae 

which are possibly closer to Nymphalidae than to other Lycaenids 

(Robbins 1988).The total of approximately 2300 species (Tables 

1–3; Robbins 1982,1992; Harvey 1987; Callaghan and Lamas 

in press) includes as many as 400 still to be described, and 

probably an equal number of well-known names which will be 

synonymised or joined with others as subspecies. With the 

exceptions of a single copper and about 60 blues, the species are 

divided almost equally between hairstreaks (all in the single 

tribe Eumaeini of the subfamily Theclinae) and metalmarks 

(Riodininae, with four neotropical tribes containing 1, 1, 137 

and about 1100 named species; the last tribe is separable into at 

least eight subtribes (Harvey 1987)). 

The neotropical fauna includes some of the most exquisite 

colours and bizarre patterns known in butterflies (Figure 1). It 

is perhaps fortunate that they attract little attention from collectors 

and dealers although as a result of this the body of biological 

and distributional information available is far less than for the 

swallowtails (Papilionidae, see Collins and Morris 1985) or 

some groups of Nymphalidae. A reasonable cross-section of 

phenotypes can be seen in colour in Hewitson's original 

descriptions and illustrations (1852–1872, 1863–1878), in 

Barcant's book on Trinidad butterflies (1970, Plates 5, 9, 11, 

12, 27, 28 plus 22 and 23 in black and white), in Lewis's 

'Butterflies of the World' (1973), and in Seitz' 

'Macrolepidoptera of the World' (1916–1920, Plates 121–159, 

110A, 113B, 193) but with many names outdated. Only minimal 

information can be unearthed in most general butterfly books 

(Smart, 1975, illustrates only 103 species) or local lists for the 

neotropics (de la Maza, 1988, shows only 118 species). The 

endemic Chilean lycaenid fauna is ignored in most publications, 

and many genera of South American hairstreaks have no name 

as yet. 

Less than 20% of the neotropical Theclinae (Robbins 1993) 

and 10% of the Riodininae (Harvey 1987) have been subject to 

any biological study (e.g. population censuses, juvenile biology 

and host plants, myrmecophily, behaviour, voltinism). While a 

45 

few species feed on plants of economic importance, especially 

Orchidaceae, Bromeliaceae, Leguminosae, Sapotaceae, 

Solanaceae, Myrtaceae, Anacardiaceae, Rubiaceae and 

Compositae, even these are rarely reared by entomologists. 

Highly diversified faunas in the high mountains (Theclinae) 

and lowland Amazonia (Riodininae) are so poorly sampled that 

even fundamental questions – on species relationships, generic 

assignments, distributions, resource partitioning, optional or 

obligatory relationships with ants, association of dimorphic 

females with their males, flight habits, seasonal variation in 

pattern and abundance, migration – remain to be answered. 

In such a frame, the picture is ill-defined, begging for much 

new work by biologists who are not averse to meeting the small, 

the little-known, the variable and complex; all too few have 

accepted this challenge. Although it may be valid that 'A true 

connoisseur of neotropical Lepidoptera can always be 

distinguished by his love for the Lycaenids' (Brown 1973), this 

is still an affair destined for frustration. 

Thus, the following attempts at generalisation and 

particularisation are very fragile, begging for more field work, 

laboratory study and experimentation. Very preliminary answers 

can be attempted for the following questions: 

(1) Are neotropical Lycaenidae 'typical' members of the 

family, or do they show their own biological styles and 

syndromes? 

(2) Do neotropical Lycaenidae show clear patterns of 

distribution, variation, behaviour, community structure, and 

ecological interaction? 

(3) Are neotropical Lycaenidae useful as indicators of other 

animal and plant species, historical and ecological factors, 

system characteristics including degree of disturbance, and 

general community structure and function? 

(4) Is it possible to identify threatened species or groups? Do 

they co-occur with endangered communities of other animals 

and plants? Can they be saved? 

Most of the answers must be sought by patient and diligent 

field work in the neotropics. In the past few years, a few 

scientists have come to terms with the systematic tangles of the 

group and have begun to do careful studies of the biology of 

some species. Hopefully, their spectacular and fascinating 

results (see De Vries 1990, 1991a) will attract more workers to

the family, to learn more about these small but disproportionately 

impressive, beautiful and varied insects (Figure 1). 

Systematics and ecology 

A summary of the recognized divisions of neotropical 

Lycaenidae down to the generic level (partial for the Theclinae) 

is presented in Tables 1–3, along with distributional, biological 

and bibliographic data. A number of outstanding and salient 

46 

facts characterise the neotropical Lycaenidae: 

• the small number of blues (Polyommatinae) in the region, 

with very widespread ocurrence of only three common 

species; 

• the predominance of forest groups, with very few species 

dependent on non-forest, open or successional habitats as is 

more common in the northern hemisphere; 

• the relatively low proportion of myrmecophilous species, 

and almost complete absence of lichen or fungus-eating 

species (although Calycopis larvae may often be detritivorous 

(Johnson 1985; Robbins 1992) and Sarota larvae feed on 

Figure 1. A pot-pourri of lycaenid diversity in the neotropics, all from 

Japi, São Paulo: (a) Theclinae. 

Key to names: 

1 'Thecla' phydela; 2 Atlides polybe; 3 Evenus regalis; 4 Arcas ducalis; 

5 Panthiadesphaleros; 6 Erora campa; 7 Chalybs Hassan; 8 Chalybs chloris; 

9 Arawacus tarania; 10 Cyanophrys acaste; 11 Cyanophrys bertha; 

12 Cyanophrys remits; 13 Brangas ca. didymon; 14 Chlorostrymon simaethis; 

15 Erora ca. opisena; 16 Ocaria cinerea; 17 'Thecla' deniva; 18 Ipidecla 

schausi; 19 Rekoa melon; 20 Ministrymon No.l.; 21 Rekoa meton Ventral; 

22 Brangas silumena; 23 Ocaria thales; 24 Magnastigma hirsuta; 

25 Chlorostrymon telea; 26 Parrhasius orgia; 27 'Thecla' elika; 28 

Ministrymon No. 2.; 29 Thereus cithonius; 30 Parrhasius selika. Ventral side 

shown, except 19, 25, 26, 27, 28 dorsal.

epiphylls – liverworts and blue-green algae (De Vries 

1988a)), and with only one case of carnivorous larvae 

known to date; 

• the rarified distribution of most species – widespread but 

very sporadic; 

• the small number of serious pest species considering the 

wide range of host plants; 

• the presence of two monotypic tribes (Stygini and 

Corrachiini) in the Riodininae; 

• the large number of monotypic genera and the existence of 

a few gargantuan genera (Strymon, Calycopis, Euselasia, 

47 

Mesosemia) (both perhaps due to insufficient study and the 

complexity of the family). 

Some of these patterns may be altered when the groups are 

better known, but at the moment the 'flavour' is strongly that of 

typical forest butterflies. This may be due to the predominance 

of forest biomes in the neotropics, but even open-vegetation 

genera (Rekoa, Strymon, Electrostrymon, Chlorostrymon, 

Calephelis, Ematurgina, Audre, Apodemia, Lemonias, Aricoris) 

show biological syndromes much like those of their forestinhabiting 

relatives and in contrast with other Lycaenidae 

which characterise successional habitats. 

Figure 1. A pot-pourri of lycaenid diversity in the neotropics, all from 

Japi, São 

Paulo: (b) Riodininae and Polyommatinae. 

Key to names: 

1 Lemonias glaphyra; 2 Calydna sp. n. nr. hemis; 3 Xenandra heliodes; 

4 Symmachia arion Female; 5 'Mesosemia' acuta; 6 Mesene pyrippe; 

7 Anteros lectabilis; 8 Napaea phryxe; 9 Emesis fastidiosa Male Ventral; 

10 Chorinea licursis; 11 Emesis fastiosa Female Ventral; 12 Barbicornis 

basilis; 13 Notheme erota; 14 Charts cadytis; 15 Lasaia agesilas; 16 Caria 

plutargus; 17 Synargis brennus; 18 Calephelis brasiliensis; 19 Pterographium 

sagaris satnius; 20 'Everes' cogina Female; 21 Zizula cyna tultiola; 

22 Baeotus johannae; 23 Adelolypa bolena; 24 Emesis fatimella; 25 Parcella 

amarynthina Female; 26 Theope thestias ca. discus; 27 Leucochimona matalha; 

28 Panara soana trabalis; 29 Lemonias zygia epona; 30 Eurybia pergaea var.; 

31 Riodina lycisca; 32 Mesosemia odice Female. Dorsal aspect shown, except 

1, 2, 7, 8, 9, 11, 14, 18, 24, 28, 32 ventral.

Table 1. Synopsis of neotropical Riodininae a ; systematics and biology b . 

Taxonomic groups: 

SUBFAMILY (TRIBE), 

Sublribe or Group, 

"Genus; 

RIODININAE g (STYGINI) 

S,yx 

(CORRACHIINI) 

Corrachia 

(EUSALASHNI) 

* † Euselasia 

Hades 

Methone 

(RIODININI) 

Mesosemiiti h 

Pcrophlhalma 

Mesophthalma 

*Leucochimona 

*Semomesia 

* † Mesosemia 

* † Eunogyra 

*Eurybia 

† 

Aleesa 

Mimocaslnia 

* Teralophthalma 

*Ithomiola 

Vollinia 

Hyphilaria 

Hermathena 

Cremna 

* † Napaea 

Eucorna 

Riodiniti i,p 

*Lyropleryx 

Necyria 

Cyrenia 

*Ancyluris 

Nirodia j 

Rhetus 

Chorinea 

'Nahida 

*lthomeis 

*Panara 

Isapis 

*Brachyglenis 

*Themone 

Plate 

numbers 

(Lewis 

1973) 

78:33 

71:42 

73:4–32 

74:1–2 

76:3 

77:22 

75:16 

74:24 

78:19 

72:4–5 

75:14–37 

72:29 

72:30–4 

73:1–3 

70:1 

76:4 

79:7–8 

74:12–3 

79:31 

74:5–7 

74:9 

71:43 

76:7–8 

:10–12 

– 

75:1–2 

76:5 

:13–4 

72:1 

70:4–12 

– 

78:8,12 

71:37 

76:6 

74:10–1 

77:18 

75:3 

71:2 

79:10 

Approx. 

no. of 

spp. 

1 

1 

134 

2 

1 

1 

1 

9 

8 

120 

1 

20 

6 

1 

6 

3 

2 

6 

2 

5 

13 

1 

4 

6 

1 

21 

1 

3 

7 

4 

9 

6 

1 

5 

4 

Typical or 

well-known 

species 

infernalis 

leucoplaga 

mys, geon 

noctula 

Cecilia 

tullius 

idotea 

mathata 

capanea 

cippus, 

telegone 

satyrus 

nicaea 

prema, amesis 

rothschildi 

phelina 

floralis 

radiata 

nicia 

candidata 

actoris, thasus 

eucharila 

sanarila 

apollonia 

bellona 

martia 

aulestes 

belphegor 

periander 

octauius 

coenoides 

astraea 

episatnius 

agyrtus 

esthema 

pais, poecila 

Distribution 

of 

nouns c 

Peru And 

CR 

NT, sp.AM 

TRAn 

AM–CR 

NT 

AM 

NT 

AM–BA 

NT 

AM-BA 

NT 

SAm 

AM 

AM–And 

AM 

TRAn 

SAm 

NT 

NT 

NT 

BR–SM 

NT 

And–CR 

AM 

NT 

BR–MG 

NT 

NT 

EcAnd 

AM–CR 

AM All 

NT 

NT 

AM 

48 

Mim d 

? 

– ? 

+– 

+ 

++ 

– 

– 

+ 

– 

_ 

– 

_ 

– 

– 

+–? 

++ 

– 

+– 

+? 

– 

_ 

– 

+ 

+? 

– 

+– 

– 

– 

+ 

++ 

++ 

+– 

– 

++ 

++ 

Myr e 

– 

– 

– 

– 

– 

– 

– 

– 

_ 

+–? 

+ 

+ 

+ 

– 

? 

– 

– 

_ 

? 

–? 

–? 

–? 

– 

–? 

– 

_ 

–? 

–? 

–? 

–? 

–? 

–? 

Time 


activity 

PM 

? 

AM.PM 

AM 

AM 

PM 

MD 

PM 

PM 

AM.PM 

AM 

LPM 

MD 

7 

PM 

MD 

LPM 

AM 

PM 

PM 

PM 

PM 

MD 

MD 

MD 

AM 

MD 

AM.MD 

MD 

? 

PM 

PM 

AM 

AM 

PM 

Habitat 

(usual) c 

CloudF 

CloudF 

HumidF 

RainF 

RainF 

HumidF 

HumidF 

HumidF 

RainF 

HumidF 

HumidF 

RiverF 

RainF 

RainF 

CloudF 

HumidF 

CloudF 

HumidF 

OpenF 

RainF 

HumidF 

CloudF 

RainF.Cd 

CloudF 

HumidF 

HumidF 

CmpRup 

HumidF 

HumidF 

CloudF 

RainF 

RainF 

RainF 

HumidF 

RainF 

Abn f 

1 

1 

2–5 

3 

2 

3 

2 

4 

3 

2–5 

3 

4 

2 

1 

3 

3 

2 

3 

2 

3–4 

3 

1 

2 

4 

2 

3 

2 

4 

3 

2 

3 

3 

3 

2 

2 

Larval 

host 

plants') 

Unknown 

Unknown 

Myrt, Clusi. 

Anacardi. 

Unknown 

Rubi. 

? 

Rubi. 

? 

Rubi. 

Marant. 

Solan. 

? 

? 

? 

? 

Orchid. 

? 

? 

Orchid., Bromeli. 

? 

? 

? 

? 

Melastomat. 

? 

? 

Flacourti., 

Celastr. 

? 

? 

? 

? 

? 

? 

Bibliography 

1,2,3,23 

1 

23 

23 

23 

4 

23 

23 

23 

23

Table 1 (cont). Synopsis of neotropical Riodininae a ; systematics and biology b . 



Subtribe or Group, 

* † Genus; 

Notheme 

Monethe 

Paraphthonia 

† 

Colaciticus 

Meiacharis 

† 

Cariomothus 

* † Lepricornis 

Pheles 

Barbicornis 

Syrmatia 

*Chamaelimnas 

Carlea 

Crocozona 

† *Baeotis 

Caria 

"Chalodeta 

Parcel/a 

† *Charts 

*Calephelis 

Amarynthis 

Amphiselenis 

* † Lasuia 

† 

Exoplisia 

Riodina 

* † Melanis 

:29–32 

* † Siseme 

† 

Comphotis 

Symmachiiti 

Luciliella 

*Mesene 

:10–4 

† 

Mesenopsis 

* † Xenandra 

79:35,38 

† 

Xynias 

* † Esthemopsis 

Chimaslrum 

† *Symmachia 

† 

*Pterographium 

† 

*Phaenochitonia 

Plate 

numbers 

(Lewis 

1973) 

76:19 

76:9 

– 

71:39 

76:1–2 

71:19–20 

74:23 

77:29 

70:38 

71:1 

79:9 

71:25–8 

71:21 

71:44 

70:23 

:35–7 

71:15–8 

71:22–3 

:34 

77:20 

72:23 

71:29–31 

:33 

(20:45–6) 

70:2 

70:3 

74:14–6 

76:15–7 

78:13–5 

74:26–7 

78:24–8 

71:41 

74:25 

75:5–6 

75:15 

74:28 

79:36 

72:25–8 

71:36 

78:10–1 

:22,31 

:34 

79:1–6 

78:21 

77:25–6 

Approx. 


spp. 

1 

3 

2 

2 

9 

4 

11 

1 

1 k 

4 

11 

1 

4 

14 

14 

9 

1 

15 

32 

1 

1 

11 

4 

3 

39 

10 

3 

3 

28 

3 

9 

4 

14 

1 

45 

9 

12 

Typical or 

well-known 

species 

erota 

alphonsus 

mitlone 

johnstoni 

ptolemaeus 

erythromelas 

atricolor 

heliconides 

basilis 

nyx, aethiops 

tircis, briola 

vilula 

caecius 

hisbon, zonata 

ino, trochilus 

jessa, theodora 

amarynthina 

auius, cleonus 

nilus 

meneria 

chama 

agesilas 

cadmeis 

lycisca 

xarife, pixe 

arisioteles 

irrorata 

camissa 

phareus 

bryaxis 

heliodes 

cynosema 

inaria 

argenteum 

probetor 

sagaris 

cingulus 

Distri- Mimd 

bution 


genus c 

NT – 

NT – 

Peru ? 

AM – 

NT – 

SAm – 

SAm +? 

SAm + 

All +– 

SAm +– 

NT + 

AM 

UpAM, – 

BR–SM 

++ 

NT + 

NT – 

NT – 

NT – 

NT – 

NT(+NA) – 

AM – 

Venez – 

NT – 

SAm – 

SAm – 

NT +– 

And – 

AM – 

TRAn – 

NT – 

NT + 

SAm – 

AM ++ 

NT ++ 

TRAn +–? 

NT +– 

SAm – 

NT – 

49 

Myr e 

–? 

– 

– 

– 

–? 

– 

–? 

– 

– 

– 

–? 

–? 

– 

– 

–? 

_ 

– 

–? 

–? 

–? 

–? 

–? 

– 

–? 

–? 

–? 

– 

– 

–? 

– 

– 

– 

– 

Time 


activity 

AM 

MD 

? 

PM 

AM,PM 

PM 

AM 

MD 

AM 

EAM 

AM 

PM 

AM 

AM 

MD 

MD 

AM 

MD 

MD 

MD 

MD 

MD 

MD 

MD 

MD 

MD 

PM 

PM 

PM 

PM 

MD 

PM 

PM 

PM 

PM 

PM 

PM 

Habitat 

(usual) c 

HumidF 

HumidF 

? 

RainF 

HumidF 

HumidF 

HumidF 

HumidF 

HumidF 

RainF 

HumidF 

RainF 

HumidF 

F, Cd 

HumidF 

HumidF 

HumidF 

HumidF 

Cmp 

HumidF 

CloudF 

RiverF 

RainF 

RiverF 

HumidF 

RiverF 

RainF 

CloudF 

HumidF 

RainF 

RainF 

RainF 

RainF 

RainF 

RainF 

HumidF 

HumidF 

Abn f 

3 

3 

? 

1 

4 

2 

2 

2 

4 

4 

2 

4 

3 

2 

4 

3 

3 

5 

5 

4 

? 

4 

2 

4 

4 

4 

2 

2 

3 

1 

1 

1 

1 

2 

1–2 

3 

3 

Larval Bibliohost 

graphy 

plants q 

? 

? 

? 

? 

Flacourti., 5 

Loranth. 

? 

Combret. 

? 

Sapor., Ulm. 6 

Zinziber 

? 

? 

? 

? 

Ulm. 2,3,7 

Sterculi., Aster. 

? 

? 

Aster. 2,3,8 

? 

? 

? 

? 

? 23 

Legumin., Aster. 3 

? 

? 

? 

Sapind. 

? 

? 

? 

? 

? 

? 

Melastom. 5,23 

?

Table 1 (cont). Synopsis of neotropical Riodininae a ; systematics and biology b . 




* † Genus; 

† *Stichelia 

Helicopiti 1 

† 

*Sarota 

† 

*Anteros 

Ourocnemis 

Helicopis m 

Emesiti 

* † Argyrogrammana 

Callistium 

† *Calydna 

* † Emesis 

Pixus 

† *Pachythone 

Pseudonymphidia 

Roeberella 

l.amphiotes 

Apodemia 

Zabuella 

Dinoplolis 

Echenais 

Imelda 

Astraeodes 

Dianesia 

Mycasior 

Petrocerus 

Lemoniiti n 

*L.emonias 

Thisbe 

Uraneis 

Catocyclolis 

* † Juditha 

* † Synargis 

Thyranota 

* † Ematurgina 

† *Audre 

† 

Aricoris 

Eiseleia 

Plate 

numbers 

(Lewis 

1973) 

77:24,28 

78:18 

71:32–3 

70:13–8 

77:14 

74:3–4 

70:21–2 

:24–5 

:27 

71:5 

71:7–14 

:38,40 

72:18–22 

:24 

– 

77:15–7 

– 

78:17 

– 

70:20? 

79:33 

72:2 

– 

74:8 

70:26 

70:19 

– 

74:19–20 

:22 

79:27–30 

79:32,37 

71:23 

74:17–8 

78:7 

72:17 

76:26–31 

77:1–4 

79:34 

72:14 

70:28–34 

77:13 

– 

Approx. 


spp. 

7 

11 

14 

2 

4 

18 

1 

25 

46 

1 

14 

1 

3 

1 

12 

1 

1 

1 

3 

1 

1 

3 

1 

10 

4 

3 

1 

6 

26 

1 

5 

22 

2 

1 

Typical or 

well-known 

species 

bocchoris 

gyas, chrysus 

formosus 

archytas 

cupido 

holosticta 

cleadas 

thersander 

cerea, lucinda 

corculum 

gigas 

clearista 

calvus 

velazquesi 

mormo 

tenella 

orphana 

thelephus 

mycea 

areuta 

carteri 

leucarpis 

catiena 

zygia 

irenea, molela 

hyalina 

aemulius 

azan, molpe 

tytia, abaris 

galena 

axenus 

epulus 

tutana 

terias 

Distribution 


genus c 

SAm 

NT 

NT 

AM 

AM–PB 

NT 

AM 

NT 

NT 

Colomb 

NT 

TRAn 

And 

Mexico 

Mexico 

Argent 

AM 

AM 

Andes 

AM–PE 

Bah,Cuba 

SAm 

BR–SM 

NT 

NT 

AM 

NT 

NT 

NT 

CBR 

SAm 

SAm 

SBR 

Argent 

50 

Mim d Myr e 

– –? 

– 

– 

– –? 

+ – 

– 

– –? 

– –? 

+– +– 

– 

–? 

– –? 

– –? 

– –? 

– 

– –? 

– –? 

– –? 

–? 

– –? 

– –? 

– 

+? 

– + 

– + 

+ + +? 

+? 

+– + 

+– + 

+ ? 

+? 

+– + 

+ ? 

+ ? 

Time 


activity 

PM 

N,AM 

AM 

? 

AM 

PM 

? 

MD 

MD 

PM 

PM 

PM 

? 

9 

MD 

9 

? 

PM 

PM 

? 

AM,PM 

PM 

PM 

PM 

PM 

PM 

PM 

PM 

PM 

MD 

MD 

MD 

MD 

Habitat 

(usual) c 

F.Cd.Cmp 

HumidF 

HumidF 

RainF 

SwampF 

HumidF 

? 

HumidF 

F,Cd 

RainF 

HumidF 

HumidF 

CloudF 

HumidF 

Cd,Cmp 

? 

? 

HumidF 

CloudF 

? 

Coast 

Scrub 

HumidF 

CloudF 

Cd,Cmp 

F,Cd 

HumidF 

HumidF 

HumidF 

HumidF 

Cd,Cmp 

Cd,Cmp 

Cd,Cmp 

Cd,Cmp 

Chaco 

Abn f 

4 

4 

2 

? 

4 

2 

? 

3 

4 

3 

2 

2 

2 

? 

3 

? 

? 

? 

3 

? 

3 

2 

2 

4 

3 

2 

2 

3 

3 

4 

3 

4 

2 

2 

Larval 

host 

plants q 

? 

Epiphylls 

Euphorbi. 

? 

Marant., Ar. 

Clusi. 

? 

Legumin., 

Myrt. 

? 

? 

? 

? 

? 

Ros., Legumin. 

? 

? 

? 

? 

? 

? 

? 

? 

Euphorbi. 

Legumin., 

Euphorbi. 

? 

? 

Legumin., 

Euphorbi. 

Simaroub., 

Legumin., 

Sterculi., 

Euphorbi. 

Legumin. 

? 

Legumin., Ros., 

Turner. 

? 

? 

Bibliography 

3 

2,3,23 

17 

17 

17 

2,3 

10 

21 

20 

11 

18 

12 

23 

14 

19

Table 1 (cont). Synopsis of neotropical Riodininae a ; systematics and biology 1 *. 




* † Genus; 

Nymphidiiti 

Parnes 

Periplacis 

† 

Menander 

*Zelolaea 

Pandemos 

† *Dysmalhia 

Joiceya 

Rodinia 

Calociasma 

† *Calospila 

† *Adeloiypa 

*† Selabis (=Orimba) 

† *Theope 

† *Nymphidium 

Stalachtiti 

Stalachtis 

Plate 

numbers 

(Lewis 

1973) 

77:21 

77:23 

75:4,7–9 

78:9 

79:39 

77:19 

72:6 

– 

78:16 

71:6 

71:3–4 

:21,74 

77:27 

:30–5 

78:1–6 

72:3,7–12 

:15–6 

77:5–12 

78:20,33 

79:11–26 

76:18 

:20–5 

78:29–30 

:32,35 

Approx. 


spp. 

2 

1 

8 

2 

2 

7 

1 

1 

4 

40 

31 

29 

45 

34 

8 

Typical or 

well-known 

species 

philotes 

glaucoma 

hebrus 

phasma 

pasiphae 

portia 

praeclara 

calpharnia 

lilina 

emylius, 

zeanger 

senta, bolena 

epitus, lagus, 

cruentata 

terambus 

lisimon 

phaedusa 

Distribution 


genus c 

AM–Atl 

SAm 

NT 

BR 

NT 

AM 

BR–MT 

AM 

NT 

NT 

NT 

NT 

NT 

NT 

SAm 

Notes: 

* Genus in which 'species' names will probably be united, reducing total 

number of species. 

† Genus in which new species will be described, increasing total number of 

species (If both symbols present, the first one should predominate - 

overall increase or reduce). 

a Does not include primarily Nearctic species extending into Mexico. 

b Information on Riodininae from Callaghan and Lamas (1994) and Harvey 

(1987) 

c AM = Amazon; And = Andes; Ant = Antilles; Argent = Argentina; Atl = 

Atlantic; BA = Bahia; Bah = Bahamas; BR = Brazil; CBR = Central 

Brazil; Cd = Cerrado; C\ = Cloud; Colomb = Colombia; Cmp = Campo; 

CR = Costa Rica; EcAnd = Ecuadorian Andes; F = Forest; Hu = Humid; 

MG = Minas Gerais; MT = Mato Grosso, Brazil; NA = North America; 

NT = Neotropics; PE = Pernambuco, NE Brazil; SAm = South America; 

SBR = Southern Brazil; Sec = Secondary forests; SM = Serra do Mar; 

sp.AM = especially the Amazon; TRAn = TransAndean; upAM = Upper 

Amazon basin; Venez = Venezuela. 

d Mim = mimicry of distasteful or venomous species; '++' = strongly 

present or characteristic,' +' = present,' +-' = weak, variable or sometimes 

present. 

e Myr = larva myrmecophilous. 

f Abn = usual abundance when found: 1 = very scarce, 2 • scarce, 3 = 

uncommon, 4 = common, 5 = abundant. 

g Maintained at family level by Harvey and others; subfamilies = Styg, 

Corrach, Hamear, Eus, Rio. 

h Divided by Harvey into Mesosemiini (first 5 genera), Eurybiini (nos 

7,8,9), 'incertae sedis' (Eunogyra and rest). 

51 

Mini d 

– 

– 

– 

+ 

– 

– 

– 

– 

– 

+– 

– 

++ 

+– 

+– 

++ 

Myr e 

+? 

+? 

+ 

+ 

+? 

+? 

+? 

+? 

+? 

+ 

+ 

+? 

+ 

+ 

+ 

Time 


activity 

PM 

PM 

PM 

PM 

PM 

PM 

? 

? 

PM 

PM 

PM 

PM 

PM 

PM 

MD 

Habitat 

(usual) c 

RainF 

HumidF 

HumidF 

RainF 

HumidF 

RainF 

Cd 

? 

OpenF 

HumidF 

HumidF 

RainF 

F,Cd 

HumidF 

F,Cd,Cmp 

Abn f 

2 

2 

2 

1 

1 

1 

1 

1 

2 

4 

4 

2 

2 

4 

4 

Larval 

host 

plants') 

? 

? 

Marcgravi. 

? 

? 

? 

? 

? 

? 

Malpighi. 

? 

Slerculi. o 

Legumin., 

Sterculi. 

Legumin., 

Sterculi. 

Simaroub., 

Legumin. 

Bibliography 

15 

16 

22 

5 

23,24 

i Divided by Harvey into 'Ancylurissection' (first 17 genera) and 'Riodina 

section' (remaining 23). 

j Probably part of Rhetus, isolated in high mesic rockfields (see Figure 3 

and species account). 

k United into a single species by Azzará (1973); probably 3 species at most. 

1 Divided by Harvey into Sarotini and Helicopini (this tribe monotypic). 

m Callaghan maintains 20 species, but 4 (Harvey) is more likely. 

n Divided by Harvey into a'Lemonias-secuon' (first 3 genera), a 'Synargissection' 

(next 4), and an'Audre-section' (last 5 genera). 

0 Larvae have been reported feeding on Membracidae (carnivorous), the 

only case of aphytophagy known in the neotropical Lycaenidae to date. 

(Harvey, pers. comm., indicates probable aphytophagy in Mimocastnia). 

p A single fossil genus (near Rhetus) and species have been described: 

Riodinella nympha Durden and Rose 1978, from the middle Eocene, 

about 48 million years ago. 

q Plant family names abbreviated by omission of standard ending. 

Bibliography cited in right-hand column: 

1 Harvey 1989; 2 Downey and Allyn 1980; 3 Kendall 1976; 4 Horvitz et al. 1987; 

5 Callaghan 1989; 6 Azzara 1978; 7 Clench 1967; 8 McAlpine 1971; 

9 Clench 1972; 10 Harvey and Clench 1980; 11 Ross 1964, 1966; 

12 Callaghan 1982b; 13 Callaghan 1986b; 14 Schremmer 1978; 15 Callaghan 

1977, 1978; 16 Harvey and Gilbert 1988; 17 Callaghan 1982b; 18 De Vries 

1988b, 1991b; 19 Miller and Miller 1972; 20 Callaghan 1979; 21 Callaghan 

1983b; 22 Talbot 1928; 23 Dias 1980; 24 Callaghan 1986a.

Table 2. Synopsis of systematics and biology of neotropical coppers and blues. 

SUBFAMILY (TRIBE) 

Section 

Genus 

LYCAENINAE 

Heliophorus section 

lophanus 

Plate 

nos 

(Lewis) 

62:34 e 

POLYOMMATINAE (POLYOMMATINI) b 

Leplotes section 

Leptotes 

Zizula section 

Zizula 

Brephidium section 

Brephidium 

Everes section 

Everes 

'Everes' c 

Lycaenopsis section 

Celastrina 

Polyommatus section 

Hemiargus 

(+2 subgenera) 

Pseudochrysops 

Madeleinea d (Itylos auctt.) 

Paralycaeides d 

Pseudolucicia d 

Nabokovia d 

Itylos d (Parachilades auctt.) 

Polytheclus d 

(19:34) 

69:51 

(19:17) 

(19:13) 

– 

(19:11) 

(19:23) 

67:18 

– 

– 

– 

67:25–6 

– 

67:23 

– 

No. 


spp. 

1 

8 

2 

1 

1 

2 

1 

9 

1 

>8 

4 

19 

2 

4 

2 

Typical 

species 

pyrrhias 

cassius 

cyna 

exilis 

comyntas 

cogina 

ladon 

hanno 

bornoi 

pelorias 

inconspicua 

chilensis 

faga 

titicaca 

sylphis 

Distribution* 

CAm 

NT 

NT 

CAm–Ven 

CAm 

BR–SM 

CAm 

NT 

Ant 

SAnd 

SAnd 

SAnd 

SAnd 

SAnd 

SAnd 

Habitat a 

CloudF 

Cmp, Sec 

Swamp 

Estuary 

Cmp 

Cmp 

Cmp, Forest 

Cmp, Scrub 

Scrub 

Scrub 

Scrub 

Puna 

Scrub 

Scrub, Puna 

Scrub 

Abundance* 

2 

5 

3 

4 

3 

3 

4 

4 

2 

2 

2 

3 

2 

2 

2 

Larval Hosts 

? 

Legumin. 

Legumin.? 

Various 

Legumin. 

Legumin.? 

Legumin. 

Legumin. 

Notes 

a The conventions and abbreviations in these categories follow those in Table 1. 

b Does not include North American (Nearctic) species invading northern Mexico. 

c Dr. Heinz Ebert gave a new name to this genus, still unpublished; may be true Everes. The species griqua Schaus was renamed as Pseudolucia parana by 

Hálint (1993), but Dr. Ebert regarded this taxon as belonging to the Everes section and congeneric with cogina. 

d Following Bálint 1993. 

e This figure number from de la Maza 1988. 

General references for neotropical Polyommatinae include Nabokov 1945; Clench 1964; and the very useful catalogue of Bridges 1988. 

Notes to Table 3 (opposite). 

a Table composed with the help of Dr. R.K. Robbins. A number of well-known genera of Eumaeini are either little-known biologically, or with infrageneric 

relationships still poorly defined, and thus are not included in this Table. Genera briefly described by Johnson (1991 a) are also omitted, for lack of complete 

information on their scope and position; the same applies to the 20 new neotropical genera and 88 new species proposed by Johnson (1992) 'based on adult 

wing pattern, tergal morphology and male and female genitalia' (all of these show great variation in some populations), and to the two new genera and 29 

new species added by Johnson and coauthors, in numbers 23 (1992, with five separate papers) and 24–29 (1993, including a review of Pseudolucia and 

two new genera of Polyommatinae, see Table 2. The well known genera not included here include: Theritas, Mithras, Allosmaitia, Thereus (=Noreena), 

Ocaria, Parrhasius, Michaelus, Oenomaus, Symbiopsis (Nicolay 1971b), Calycopis s.l. (=Calystryma, Femniterga, Tergissima and 15 of the other genera 

described in Johnson 1991a; the other seven new 'Outgroup' genera also include several segregates from well-known genera mentioned here), 

Electrostrymon, Lamprospilus, Theclopsis, Siderus, Contrafacia (=Orcya), Ipidecla, Hypostrymon, Nesiostrymon (see Johnson 1991b, including its 

citations of most previous papers of that author), Paiwarria and Theorema. 

b The conventions and abbreviations in these categories follow those in Table 1. 

c References: 1 Robbins 1987; 2 Nicolay 1971a; 3 Robbins 1980,1985; 4 Robbins 1991; 5 Nicolay 1980 and Johnson 1989; 6 Nicolay 1977; 7 Clench 1944, 

1946; 8 Nicolay 1980 and Callaghan 1982; 9 Nicolay 1982; 10 Miller 1980; 11 Robbins and Venables 1991. 

52

Table 3. Synopsis of the systentatics and biology of selected neotropical hairstrcaks (better defined genera, including about 30% of known species) a . 


Genus 

Atlides 

Areas 

Pseudolycaena 

Arawacus (=Polyniphes, 

Dolymorpha) 

Rekoa (=Heterosmailia) 

Chlorostrymon 

Magnastigma 

Cyanophrys 

Panthiades (=Cycnus) 

Olynthus 

Strymon 

Tmolus 

Ministrymon 

Brangas 

Chalybs 

Erora 

Trichonis 

laspis 

Janlhecla 

Plate 

numbers 

(Lewis) 

THECLINAE (EUMAEINI) 

Eumaeus 

67:17 

Theslius 

69:7 

Micandra 

67:47,51 

69:8 

Evenus 

69:13,19 

:25,45 

66:12 

:14–6 

68:8 

69:44,46 

68:39 

66:9–11 

67:24 

68:26 

69:17,21 

67:45 

68:12,44,48 

69:9 

67:30–1 

67:29,36 

68:15 

:23–4 

:34 

67:22,44 

68:5 

– 

66:18 

67:1,3 

:28 

68:19,53 

69:40 

:47–8 

69:49,52 

67:2,15 

:27 

68:41 

69:38 

66:13 

68:27 

68:20 

– 

69:18,27 

:41 

68:38 

No. 


spp. 

6–7 

1–3 

8–10 

13 

13 

7 

4–6 

16–20 

7 

5 

6 

19–22 

8 

10 

40–70 

10–11 

18–22 

18–20 

3 

>40 

2 

11–12 

10 

Typical 

species 

minijas 

pholeus 

platyptera 

regalis 

polybe 

imperialis 

marsyas 

aetolus 

dumenilii 

melon 

marius 

palegon 

simaethis 

julia, hirsuta 

herodotus 

bertha 

acaste 

bitias, phaleros 

punctum, fancia 

ziba 

mulucha 

echion 

azia 

una 

getus, silumena 

janias 

aura, phrosine 

hyacinthinus 

temesa, talayra 

janthina, rocena 

Distribution 

b 

Ant–AM 

AM 

CAm–And 

NT 

NT 

NT 

NT 

NT 

NT 

NT 

NT 

NT 

NT 

CR–SBR 

NT 

NT 

NT 

NT 

NT 

NT 

AM 

NT 

NT 

53 

Habitat b 

F, Scrub 

HumidF 

CloudF 

Cl, HumidF 

Cl, Humid 

RainF 

Cl, Humid 

RainF 

F, Cd, Scrub 

F, Cd, Scrub 

F, Cd, Cmp 

F, Cd, Cmp 

Hu, RainF, Cd 

HumidF 

Cl, Hu, RainF 

Cl, RainF 

F, Cd, Cmp 

Hu, RainF, Cd 

HumidF, Scrub 

Cl, Hu, RainF 

HumidF 

Cl, Hu, RainF 

RainF 

Forest 

Cl, Hu, RainF 

Abundance 

3 

2 

2 

2 

2 

3 

4 

4 

4 

2 

2 

3 

3 

2 

4 

3 

2 

2 

2 

2 

1 

2 

3 

Larval 

hosts 

Cycad. 

Legumin. 

Sapot. 

Loranth. 

Laur., 

Anacardi. 

Polyphagous 

Solan. 

Polyphagous 

Sapind. 

Polyphagous 

Polyphagous 

Lecythid. 

Polyphagous 

Polyphagous 

Legumin. 

Myr. b 

? 

– 

– 

– 

– 

– 

– 

+ 

+ 

+ 

+? 

+ 

+ 

+ 

+ 

Loranth., Sapind. 

Legumin. 

Polyphagous 

? 

Sterculi. 

? 

+ 

Refs c 

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

1 

11

Figure 2. Biogeographical patterns of neotropical Lycaenidae and other butterflies. 

54

On the basis of four foreleg characters, Robbins (1988) has 

advocated separation of the Riodininae from the Lycaenidae, 

suggesting affinity with the Nymphalidae. The considerable 

variation of some leg characters leads to difficulty and 

disagreement about their interpretation in many groups and in 

this case further integration with other sets of characters would 

be helpful before a final decision is reached. 

Distribution and variation 

As noted above, three species of neotropical blues (Hemiargus 

hanno, Leptotes cassias and Zizula cyna) are very widespread 

indeed despite their minute size and weak flight, especially 

Zizula cyna which struggles to rise above the grasses in the sites 

it inhabits throughout tropical America. Leptotes and Hemiargus 

are also known from distant oceanic islands (Galapagos and 

Fernando de Noronha), as well as all islands in the Caribbean. 

Robbins and Small (1981) have shown that most Theclinae are 

also widespread and undifferentiated, and suggest that this is 

due to mass, long-range, community dispersal during the dry 

season, like Pieridae and Hesperiidae, ranging over the entire 

region. Riodininae may be at the opposite extreme of the 

spectrum, most being extremely local (even in more widespread 

species) and often well-differentiated geographically (especially 

in Eurybia, Mesosemia, Nymphidium and the mimetic groups). 

Largely endemic thecline faunas are found in high mountains 

in southeastern Brazil, the Andean cloud forests and the Central 

American highlands, while local riodinine assemblages inhabit 

wet lowland forests (Callaghan 1983a, 1985). A largely endemic 

fauna (Table 2) inhabits the extremely long, narrow 'island' 

forming the Republic of Chile, isolated by desert (N), highaltitude 

rocks (E), ice (S) and ocean (W). There are contiguous 

areas in southwestern Bolivia and Peru extending up the semiarid 

'puna' as far as southern Ecuador in some cases. In the 

Brazilian Atlantic region (Figure 2) much endemism is seen in 

cerrado species (central plateau) in addition to montane and 

coastal forms; in the overall region, 49% of the Theclinae, 44% 

of the Riodininae, and two congeneric blues are endemic. These 

levels are slightly higher than for skippers, nymphalids or 

pierids (all 39%) or papilionids (42%), but they may drop when 

proper links can be discovered between Atlantic and Amazonian 

or Andean sister-groups (species or subspecies). 

Robbins (pers. comm.) has suggested a list of 25 thecline 

species or superspecies which are 'widespread, common in 

many regions and habitats, and likely to be recorded on a quick 

trip to the neotropics': Theritas mavors/triquetra, Pseudolycaena 

supersp. marsyas, Arawacus supersp. aetolus, Rekoa meton, R. 

marius, R. palegon, Tmolus echion, Oenomaus ortygnus, 

Panthiades bitiaslhebraeus, Parrhasiuspolibetes, Cyanophrys 

herodotus, Electrostrymon supersp. endymion, Calycopis 

isobeon, C. cerata, Chlorostrymon simaethis, Ministrymon 

una, M. azia, Strymon mulucha, S. bazochii, S. ziba, and 

'Thecla' (catch-all genus for the majority of species still not 

assigned to existing generic names) hemon, hesperitis, syllis, 

55 

celmus and tephraeus. In South America, Arcas imperialis, 

Panthiades phaleros and some other forest species could be 

added to this list. 

At the other extreme, one or two specimens are known from 

a single locality (such as two new Atlides species from a single 

2200m hilltop west of Cali, Colombia), and perhaps over a 

hundred (about 10%) are known from fewer than ten specimens 

in museum collections. In April 1991, Robbins took the fourth 

known specimen of a striking thecline, without generic or 

specific identification, on Mikania flowers invadingthis author's 

backyard; the first three were in a single Brazilian collection, 

awaiting description. So for these species, and perhaps for over 

50% of Riodininae, much information is still waiting in the 

wings before any idea of distribution, biology, affinities or 

importance can be assigned. The general patterns of 

biogeography of neotropical Lycaenidae are very similar to 

those known in other insects, as shown in Figure 2. 

Mimetic species that join 'rings' of similar and distasteful 

heliconians, ithomiines, arctiids, dioptids and other Lepidoptera 

vary in parallel with these (mostly in 16 genera of Riodininae, 

Table 1) and are just as useful as their models for identifying 

centres of endemism at the infraspecific level in the lowland 

neotropical forests (Figure 2). Many other Lycaenidae show 

strong geographic variation, often regarded today as at the 

species rather than the subspecies level. Probably about half of 

any local lycaenid fauna will contain some information about 

region and habitats. 

Behaviour and juvenile host plant 

relations 

Most neotropical blues and hairstreaks have the usual rapid, 

darting flight of these groups. Especially impressive are the 

larger hilltopping species of Arcas, Evenus, Atlides and Brangas, 

but they are surely no faster than some of the small species 

which are completely lost by the eye on each flight and are only 

seen again when they land back on the same perch. 

Riodinine flight varies from the same rapid, darting pattern 

(Theope, many Euselasia, Charis, Mesene, Symmachia, 

Stichelia, Calydna, Calospila) through a more usual and slower, 

erratic flight (always returning to a chosen perch) to the very 

casual, dipping or fluttering flight of mimetic species imitating 

their distasteful models (Methone, Ithomiola, Ithomeis, 

Brachyglenis, Themone, Chamaelimnas, Cartea, Uraneis, 

Stalachtis, and females of Esthemopsis and Setabis). A few 

species may imitate wasps with a 'buzzy' flight (Chorinea, 

Syrmatia, some Rhetus), while ground-perching species may 

rise in spirals when disturbed. Styx has a very weak, almost 

falling flight (fide G. Lamas). 

Adults usually visit small flowers, and many riodinines 

(principally males) are also found on damp sand or mud 

(Lyropteryx, Riodina, Rhetus, Necyria, Barbicornis, Parcella, 

Notheme, Monethe, Lasaia, Chamaelimnas, Caria, Siseme). 

All vertical levels of the habitat, and also adjacent non-forest

areas, are included in usual foraging surveys. 

Adults of the genus Eumaeus, locally common from southern 

Florida to central-west Brazil, incorporate toxic cycasins from 

their larval foodplants (mostly Zamia but often other 

Cycadaceae) (Rothschild et al. 1986; Bowers and Larin 1989). 

They are brightly marked models for mimicry rings, flying very 

slowly and liberating cyanide gas when crushed; their larvae 

are gregarious and also highly aposematic, having a delicate 

relationship with the phenology of their hosts (Clark and Clark 

1991). No other adult neotropical Lycaenidae have yet been 

shown to be similarly protected against predators. 

Mass movements of Riodininae communities have not been 

reported, though populations often appear suddenly in a new 

habitat as a few worn individuals and are later replaced by a 

large endogenous generation if foodplants are found. Theclinae 

seem to move over large regions in loose, multispecific groups 

(hundreds of individuals of dozens of species), or sometimes 

migrate unidirectionally or erratically as dense populations of 

a single species (Pseudolycaena and Calycopis), principally 

during the dry season when flowers are scarce. Sunny clearings 

can attract such concentrations, which can at times contain 

nearly a hundred species. Especially interesting areas to find 

these assemblages are sheltered sites on low or high ridges, on 

dry, sunny or windy days (Robbins and Small 1981). 

Territorial perching and area defence by males are widespread 

and easily demonstrated by marking neotropical Lycaenidae 

(see also Bates 1859). In open fields, males perch on the tips of 

the highest grass blades, driving off with their rapid forays other 

males, many other butterflies and even insect predators, bird 

predators and collectors. In forests, perching may be in the 

canopy (only in the morning since canopy species descend to 

the cooler understorey at noontime), on forest edges, in small 

clearings, on tree trunks in the sun or shade, on top of or under 

leaves at all levels (with distinct partitioning, Callaghan 1983a), 

along rivercourses or trails, or on hilltops and rocks (especially 

in mountains), almost always in an exposed, often sunlit spot. 

Territories defended vary from less than one to many dozens of 

cubic metres. As might be expected, hilltopping (or ridgetopping 

or edge-seeking) is very common in male Lycaenidae 

in wait for females of their sparse populations. More than one 

hundred species may accumulate at a favoured hilltop in late 

morning to mid-afternoon, colouring the sky in their territorial 

battles. Early morning 'leks' of many males have been seen in 

Euselasia, Barbicornis, Sarota and Syrmatia. 

When not exhibiting territorial behaviour, theclines perch 

on the tops of leaves or on twigs in the forest undergrowth, in 

shade or sun-flecks, 'rubbing' their hindwings in the usual 

manner to call attention to their 'false head' (Figure 3 and 

Robbins 1980,1985) (this also occurs in the nodminzs Anteros 

and Sarota). Whilst doing this, they fly upon the slightest 

provocation, usually landing quite far away. 

Riodinines usually perch under leaves with wings flat open 

(most genera) or closed (Euselasiini, Sarota, Anteros, Themone, 

Helicopis, Theope, most Setabis); some perch on tops of leaves 

with wings half-open (Mesosemia, Semomesia and relatives, 

also Eurybia at dusk). Particular underleaf perches are used 

56 

repeatedly and insistently by the same individual, and by 

members of the same population. If the resident individual is 

removed from a leaftop territorial perch or an underleaf resting 

station, a series of conspecific males will then occupy the same 

perch; depending upon the genus, these can be progressively 

younger and more splendid, or older and more worn than the 

first resident. 

Courtship can have both aerial and perched components, 

either of which can be very complex. They are presumably 

accompanied by diverse pheromonal signals and responses 

which are produced by scent-spots or pads, androconial brushes 

or patches. In Helicopis cupido (observed in Guyana) the male 

alternates rapid wing flutters (to 40° open) with short 220° wing 

opened phases (one-second cycles), whilst perched right behind 

the female (who maintains her wings closed) on top of a sloping 

large leaf of the larval foodplant (Montrichardia, Araceae); he 

then moves directly in to initiate mating. 

Oviposition is not often observed, but as with most butterflies, 

occurs in short bouts which involve appreciable searching, 

inspection, and tactile evaluation (foreleg 'drumming') of the 

exact site. Off-hostplant egg-laying is not common but has been 

observed, as in other butterflies. Eggs are often placed in axils 

and other tight places, and are very rarely found; they are often 

pincushion-shaped and highly sculptured (Downey and Allyn 

1979,1980,1981,1984), but in four small tribes of Riodininae, 

they are smooth and barrel-shaped (Harvey 1987). 

Myrmecophilous species may lay their eggs along ant foraging 

routes (Stalachtis, Lemonias, Mycastor, Nymphidium) or even 

visually encourage ants to directly remove eggs as they are 

expelled (Adelotypa senta;Audre domina, Robbins and Aiello 

1982). 

Larvae may be smooth (especially if myrmecophilous), 

tubercled, spiny or hairy, often in relation to their habitat. They 

are usually highly cryptic, even when on flowers (a habit 

common in Theclinae, rare in Riodininae), and may be 

polymorphic, probably incorporating pigments from the flowers 

on which they feed (Monteiro 1991). Some larvae and many 

pupae are found in 'nests' inside rolled leaves. Pupae are quite 

variable in shape, with systematic significance in the Riodininae 

(Harvey 1987). These biological characters are quite similar to 

those of other Lycaenidae, though some riodinine larvae are 

unique. 

Myrmecophily is quite frequent in Theclinae, perhaps in 

50% of species (De Vries 1990; Malicky 1970), but in Riodininae 

has been reported in only three subtribes with 283 species: 

Eurybiiti; Lemoniiti; and Nymphidiiti (Harvey 1987; De Vries 

1991a). Stalachtis, in a fourth monogeneric subtribe, is optionally 

myrmecophilous, even in a single population (susanna, phlegia, 

lineosa, on Simaroubaceae; Callaghan 1986; Benson, Francini 

and Brown, unpublished). 

Ant-associated larvae have easily recognisable specialised 

glands (Cottrell 1984). In Lycaenidae these include a dorsal 

nectary organ on the seventh abdominal segment and a pair of 

tentacle organs on the eighth segment. All known ant-associated 

riodinine larvae have a pair of glands (tentacle nectary organs) 

on the eighth segment. These secrete a solution which may be

57 

Figure 3. Pictures of Riodinines and Theclines in life, to show behaviour 

and habitat. 

(a) Paiwarria telemus (Manaus) [top left]; (b)Arawacus meliboeus (Japi, Sao 

Paulo) [top right]; (c) Strymon oreala (Santa Teresa, Espírito Santo) [middle 

left]; (d) Nirodia belphegor (Serra do Cip6, Minas Gerais) [middle right]; 

(e) Helicopis acis (Belém, Para) [bottom]. (Photos: K.S. Brown Jr., except 

(d) Ivan Sazima.)

very rich in amino acids (De Vries 1988b; De Vries and Baker 

1989). Larvae of Lemoniiti and Nymphidiiti have two additional 

myrmecophilous organs (Harvey 1987): a pair of anterior 

tentacle organs on the metathorax which produce chemicals 

affecting ant behaviour (De Vries 1988b); and a pair of vibratory 

papillae on the anterior margin of the prothorax, which produce 

sound to call ants (De Vries 1988b, 1990), also seen in other 

lycaenids (De Vries 1991a, 1991b). 

Food resources of larvae are exceedingly varied, with at 

least 50 plant families recorded more than once; no pattern can 

be discerned except for certain tendencies for given species, 

genera or tribes to use the same resource over a wide geographical 

range. In contrast, some species are extremely polyphagous, 

especially those that feed on flowers (mostly Theclinae: Robbins 

and Aiello 1982; Monteiro 1991). 

The normally Myrtaceae-feeding and rare Euselasia eucerus 

in Brazil has taken to imported Eucalyptus leaves and becomes 

extremely abundant and destructive in commercial plantations 

of these Australian trees. The riodinine Audre campestris, the 

thecline Michaelus jebus, and some blues may be pests of 

cultivated beans and other legumes, and Strymon ziba can 

become a serious pest on pineapples, but in general few 

Lycaenidae have achieved such notoriety. 

Lycaenids as indicator species 

The extremely sporadic distribution of most neotropical 

Lycaenidae, not only in collections but also in the field in both 

time and space, was noted by early naturalists (Bates 1859) and 

amply confirmed by all later observers. Riodinines are especially 

local, confined to a very narrow microhabitat, and active only 

at a particular time of day and level of forest; they are often 

greatly reduced in diversity and numbers in highly variable or 

unpredictable climates. Some are even nocturnal: Euselasia 

clesa was seen flying in typical territorial behaviour under a 

black light at 4 a.m., one hour before dawn and Sarota chrysus 

flies during the night and often comes to light (see also Miller 

1970). Most species, however, pick a time slot in the early 

morning or mid-afternoon to be active, with the latest species 

often being those of Nymphidium and Eurybia, the earliest 

Euselasia, Syrmatia and Sarota. While many species are present 

year round, great seasonal variation in both presence and 

abundance is the rule (as for Theclinae), with species often 

peaking either in mid-summer, late fall or late spring in higher 

to lower elevations. At least one species in perhumid tropical 

forest (Euselasia zara in southeastern Brazil) has been seen 

only in November (in lowlands) to February (in uplands) and 

seems to be univoltine, as has also been suggested for Audre 

domina in Panama (Robbins and Aiello 1982). 

All this natural fluctuation leads to serious problems in 

establishing baseline data for lycaenids, or recognising any 

significant tendencies to change or vary coherently in different 

habitats. When added to the great difficulty in collecting and 

identifying most species, the tendency to move about over the 

58 

landscape, and the chaotic state of the systematic and biological 

data, these characters greatly diminish the utility of neotropical 

Lycaenidae as ecological indicators at the present time. 

This does not mean that they are not potentially very useful 

in surveys and monitoring of natural and altered systems. 

Riodinines compose nearly half the daily list of butterflies in 

most parts of the Amazon Basin, and each species seems to be 

delicately tuned into a large number of physical and biological 

factors in its environment, which are therefore faithfully 

indicated by the species' presence (though not excluded by its 

absence). Some species may feed on a single plant genus or 

family as larvae or may be associated with certain ant species, 

thus necessitating the presence of these resources which may be 

harder to find and recognize than flying adult butterflies. The 

overall richness and diversity of the local lycaenid fauna offers 

excellent opportunities for correlation with different systems in 

varying stages of naturalness. Even a partial list can lead to 

hypotheses about ecological factors, history of the region, 

resources present, soils and climate, primary productivity, 

importance of other communities, and stability of the whole 

system. The routine and very propitious use of Lycaenidae as 

indicators in the neotropics only awaits the resolution of the 

systematic picture in order to produce useable manuals for 

identification, and wider biological studies in order to expand 

the knowledge of indicator parameters possible with diverse 

species and groups. 

Threatened neotropical lycaenids 

Insects that are highly stenoecious – with many narrow 

environmental requirements or specialised interactions 

necessary for their survival – may be easily reduced or eliminated 

locally by minor habitat alteration through natural or human 

disturbance. This applies to many neotropical Lycaenidae 

populations. With this local extinction, local adaptive genes 

will disappear, reducing the biodiversity and biosynthetic 

capacities of life. 

On the other hand, winged insects are often migratory and 

accustomed to seeking out the narrow environmental conditions 

needed for growth and reproduction, and are already adapted to 

normal levels of natural disturbance; they are likely to be 

widespread even though local, and very persistent in the regional 

species pool. Thus, a definition of 'threat' to a neotropical 

lycaenid should consider its ecological characteristics, 

geographical distribution, aptitude for mobility and colonisation, 

density of colonies, usual population size, voltinism (are adults 

always around to be able to flee destruction and colonise 

elsewhere?), and the kind, degree and extent of local or regional 

disturbance patterns – both natural (on long and short timescales) 

and artificial. 

In principle, essentially all lycaenid species should survive 

transformation of primitive mosaic habitats (in areas of complex 

topography) into anthropic mosaics with 10–30% of the original 

vegetation maintained in large patches. Such is the case in the

Brazilian Atlantic forests, where reduction of the original forest 

to 12% of its former area has not caused detectable extinction 

of lycaenids or indeed of any other insects or vertebrates 

monitored (Brown 1991; Brown and Brown 1991). In 

ecologically more homogeneous areas, reduction of natural 

habitat to 10% or less of its original area may still not lead to 

overall species extinction, though much genetic variation will 

be lost with local populations (and less vagile species in some 

groups can be eliminated). Most lycaenids will survive even in 

smallish habitats (0.1–100ha) as long as the edge effects (Lovejoy 

et al. 1986; Janzen 1984,1986) do not overwhelm the essential 

habitat characteristics: isolated areas of 10ha may be dramatically 

transformed within a year, and appreciable effects can be seen 

up to 250m from an edge in larger patches of experimental 

fragments in the central Amazon (Brown 1991). However, 

local disturbance can have a positive effect: it often brings in 

many new species of sun-loving lycaenids to the more diverse 

successional community (Hutchings 1991). 

In regions subjected to large-scale commercial conversion 

(to pasture, monoculture, silviculture, agrosilviculture or timber 

production), many lycaenids may not find new habitat and more 

large-scale regional extinction is possible. If a species is confined 

to the region, it might become extinct, especially if the habitat 

is very specialised and different from neighbouring areas (a 

basin, mountain top, headwater system, lakeshore or marsh, 

seasonally flooded region, or other natural 'island'). Thus, 

land-use patterns could make a large difference in the genetic 

erosion and threat to lycaenid species in tropical forests (Brown 

and Brown 1991). 

Brazil includes half the forest area, two of the four speciesendemic 

regions and 19 of the 45 subspecies-endemic centres 

in the neotropics; it has about 425 species of Theclinae and 700 

species of Riodininae (Brown 1982,1991). A new official list 

of fauna threatened with extinction in Brazil was prepared by 

the Zoology Society in 1989 (Bernardes et al. 1990). It includes 

23 butterflies, only one of which is a lycaenid: Joiceya 

praeclarus, an inconspicuous monotypic genus known only 

from a small area in central Mato Grosso, and not seen there 

since its original discovery in the 1920s. This would represent 

the syndrome of 'restricted to a limited area, isolated, under 

intense large-scale human occupation', though it is probably 

preserved in the four conservation units established recently in 

the area. However, at least one population of the species must 

be found before its survival can be assured or even dealt with. 

A second list, prepared by the Zoology Society, 

recommending 'further study', includes another five species of 

Lycaenidae and also six genera of little-known Riodininae 

which include mostly rare, extremely local and sometimes 

geographically limited species – the syndrome of 'widespread 

but extremely sporadically recorded' (Alesa, Colaciticus, 

Esthemopsis, Mesenopsis, Symmachia and Xenandra). To the 

generic list could be added Petrocerus, Erora, and other groups 

of riodinines and theclines confined to high-elevation habitat 

islands in southeastern Brazil (same syndrome as Joiceya, but 

somewhat more widespread). 

The five species of Lycaenidae on the second list represent 

59 

further syndromes of rarity and threat, which need some more 

study before inclusion on the official Brazilian list of threatened 

fauna. They include the theclines Arcas ducalis (widespread 

but local in hills and mountains, mostly on hilltops) and Arawacus 

aethesa (highly restricted geographically to a lowland area 

under intensive deforestation); and the riodinines Eucorna 

sanarita (few localities in high mountains), Nirodia belphegor 

(a monotypic genus possibly part of Rhetus from natural 

rockfields in mountains of central Minas Gerais), and Helocopis 

cupido nr. lindeni (a few coastal swamps in the northeast). 

Other rare or restricted species may be added in the coming 

years to this list. 

Outside Brazil, the best candidates for threatened status may 

be island species or high-altitude groups, with most members 

still very little known. The primitive, taxonomically isolated 

Styx infernalis is but rarely seen at medium high elevations in 

Peruvian cloud forests. These and some further Brazilian cases 

are treated in the species accounts in this book. 


Dr. Robert K. Robbins of the U.S. National Museum 

(Washington) kindly criticised the text of this chapter and 

produced the final version of Table 3, as well as providing 

extensive unpublished information on Theclinae. Drs Donald J. 

Harvey of the same institution, and Curtis John Callaghan of 

Búzios, Rio de Janeiro and the Museu Nacional, furnished 

unpublished catalogues and extensive information on 

Riodininae. Reprints of papers and further notes on Lycaenidae 

were provided by Kurt Johnson of the American Museum of 

Natural History (New York) and Stan S. Nicolay of Virginia 

Beach. Dr. Olaf H.H. Mielke of the Departamento de Zoologia, 

Universidade Federal do Parana, Curitiba has helped greatly in 

knowledge of the Brazilian butterfly fauna over the past 30 

years, and curated a very rich and complete collection which 

has greatly aided in understanding systematics and 

biogeography. 

References 

AZZARÁ, M.L. 1978. Revisão do g6nero Barbicornis Godart, 1824 

(Lepidoptera, Lycaenidae, Riodininae). Acta Biol. Paran., Curitiba 7: 

23–69. 

BÁLINT, Z. 1993. A catalogue of the polyommatine Lycaenidae (Lepidoptera) 

of the xeromontane oreal biome in the Neotropics as represented in 

European collections. Repts Mus. Nat. Hist. Univ. Wisc. (Stevens Point) 

29: 1–36. 

BARCANT, M. 1970. Butterflies of Trinidad and Tobago. Collins, London, 

314pp. 

BATES, H.W. 1859. Notes on South American butterflies. Trans. ent. Soc. 

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Threatened Lycaenidae of South Africa 

Michael J. SAMWAYS 

Department of Zoology and Entomology, University of Natal, Pietermaritzburg 3200, South Africa 

Southern African geology and geography 

To appreciate the conservation biology of lycaenids in South 

Africa, it is necessary to introduce the characteristic geology 

and geography of the area. About 180 million years B.P., the 

great supercontinent of Pangaea began to split. By 135 million 

years B.P., the southern tip of Africa looked very much in 

outline as it does today. Since that time, uplifting and erosion 

has led to the appearance of 22 physiographic regions, each 

characterised by altitude and surface form (Figure 1). These 

regions fall naturally into two groups. The first group, of 12 

regions, is the Interior Plateau. The second group, of 10 regions, 

is the Marginal Zone, which is divided from the Interior Plateau 

by the great divide known as the Great Escarpment. 

Figure 1. The 22 physiographic regions of South Africa. 

1. Upper Karoo; 2. Highveld; 3. Kaap Plateau; 4. Southern Kalahari; 

S. Bushmanland; 6. Namaqualand Highlands; 7. Bushveld Basin; 8. Bankveld; 

9. Pietersburg Plateau; 10. Waterberg Plateau; 11. Soutpansberg; 12. Lesotho 

Tableland; 13. Lebombo Hills; 14. Lowveld; 15. Middelveld; 16. Eastern 

Midlands; 17. Winterberg Mountains; 18. Great Karoo; 19. Doring Karoo; 

20. Cape Folded Mountains; 21. Little Karoo; 22. Coastal Belt. 

Sub regions: NN = Northern Natal; SN = Southern Natal; TR = Transkei; 

EC = Eastern Cape; SC = Southern Cape; WC = Western Cape. 

62 

The Interior Plateau is the southern tip of the great African 

Plateau, and its altitude varies from a minimum of slightly 

under 900m in the Kalahari Desert, to almost 3500m in the 

Lesotho Highlands. Here, the thick Karoo sediments of the 

Carboniferous to Triassic ages were covered in the Jurassic by 

thick lava flows, which, on cooling to a hard layer of basalt, 

protected the underlying softer rocks from weathering, giving 

rise to the high mountains of southern Africa. 

The Marginal Zone between the Great Escarpment and the 

coast, varies in width from 60km in the west to 240km in the east. 

Its elevation varies from sea-level to a maximum of 2300m in the 

Swartberg Range south of the Great Escarpment. In turn, the 

Great Escarpment is composed of several distinct mountain 

ranges, giving southern Africa a distinctive and varied topography. 

Off the coast, the northward flowing cold Benguela Current 

moves up the west coast, and the southward flowing warm 

Agulhas Current flows down the east coast. These flow patterns, 

along with topography and global wind patterns, influence the 

area's rainfall regimes. There are three distinct rainfall regions 

in southern Africa: summer, winter and all-season rainfall 

areas. The varied topography and climate has resulted in nine 

climatic zones (Figure 2). 

Species richness and threatened species 

The long period of geographical isolation in southern Africa, 

lack of glaciation, varied topography and climate has produced 

a wide range of biotypes and high levels of endemism and 

species richness in both plants and animals (Huntley 1989). 

Vári and Kroon (1986) list 8300 species of Lepidoptera in 

southern Africa (i.e. south of the Zambezi River), while Pinhey 

(1975) estimates that the final total will exceed 10,000. 

In South Africa (i.e. south of the Limpopo River), 632 

species of butterfly (Papilionoidea and Hesperioidea) so far 

have been described. Of these, 102 (16%) are under some level 

of threat (Henning and Henning 1989). If subspecies are included, 

the total number of threatened taxa is 14%. 

Vari and Kroon (1986) list 389 lycaenid species in southern 

Africa. Of these, 310 occur in South Africa, and 105 species and

Fig. 2. The nine climatic zones in South Africa. 

1. Subtropical Lowveld; 2. Subtropical Coast; 3. Temperate Coast; 

4. Mediterranean; 5. Plateau Slopes; 6. Temperate Eastern Plateau; 

7. Subtropical Plateau; 8. Semi-arid Plateau; 9. Desert. 

subspecies are in some way threatened (Henning and Henning 

1989) (Table 1 and Figure 3). This is 75% of all threatened 

butterflies in the country. Two species of lycaenid are now 

Extinct, and another one species and one subspecies are 

Endangered (Table 1 and Figure 3). 

Particularly significant is that 96% (101) of the threatened 

species (71% species and 25% subspecies) are endemic to 

South Africa (Clark and Dickson 1971; Henning and Henning 

1989; Murray 1935). These figures are high for a portion of a 

continental land mass, but not unusual for other biotic groups in 

the subcontinent. Of the remaining four species, only one is 

widespread in Africa, while the other three have limited 

distributions or are on the edge of a range just extending into 

South Africa. 

Geographical distribution of threatened 

species 

Figure 4 illustrates that by far the richest area for rare endemics 

is the Cape Fold Mountains (physiographic areas 19,20 and 21 

in Figure 1). Most occur on mountain slopes, but some inhabit 

ridges (e.g. Poecilmitis wykehami, Thestor dicksoni dicksoni), 

peaks (e.g. Lepidochrysops outeniqua, P. endymion, P. balli) 

Figure 3. Status of South Africa's 105 threatened lycaenid species and subspecies using the official IUCN categories (Data from Henning and Henning, 1989). 

63

Table 1. Status of threatened lycaenid species and subspecies in South Africa (from the South African Red Data Book, Henning and Hcnning 1989). 

Lipteninae 

Alaena margaritacea Eltringham 

Deloneura immaculata Trimen 

Durbania amakosa albescens Quickelberge 

Durbania amakosa flavida Quickelberge 

Ornipholidotos peucetia penningtoni (Riley) 

Liphyrinae 

Aslauga australis Cottrell 

Miletinae 

Thestor brachycerus (Trimen) 

Theslor compassbergae Quickelberge & McMaster 

Theslor dicksoni calviniae Riley 

Thestor dicksoni dicksoni Riley 

Thestor dryburghi Van Son 

Thestor kaplani Dickson & Stephen 

Thestor montanus pictus Van Son 

Thestor pringlei Dickson 

Thestor rossouwi Dickson 

Thestor stepheni Swanepoel 

Thestor strutti Van Son 

Thestor swanepoeli Pennington 

Thestor tempe Pennington 

Thestor yildizae Koçak 

Theclinae 

Aloeides caledoni Tite & Dickson 

Aloeides carolynnae Dickson 

Aloeides clarki Tite & Dickson 

Aloeides dentatis dentatis (Swierstra) 

Aloeides dentatis maseruna (Riley) 

Aloeides egerides (Riley) 

Aloeides kaplani Tite & Dickson 

Aloeides lutescens Tite & Dickson 

Aloeides merces Henning & Henning 

Aloeides nollothi Tite & Dickson 

Aloeides nubilus Henning & Henning 

Aloeides pringlei Tite & Dickson 

Aloeides rossouwi Henning & Henning 

Aloeides trimeni southeyae Tite & Dickson 

Argyrocupha malagrida cedrusmontana 

Dickson & Stephen 

Argyrocupha malagrida malagrida (Wallengren) 

Argyrocupha malagrida maryae Dickson & Henning 

Argyrocupha malagrida paarlensis (Dickson) 

Bowkeria phosphor borealis Quickelberge 

Bowkeria phosphor phosphor (Trimen) 

Capys penningtoni Riley 

Chrysoritis oreas (Trimen) 

Chrysoritis cottrelli Dickson 

Erikssonia acraeina Trimen 

Hypolycaena lochmophila Tite 

Iolaus (Epamera) aphnaeoides Trimen 

Iolaus (Epamera) diametra natalica Vári 

Iolaus (Pseudiolaus) lulua Riley 

Oxychaeta dicksoni (Gabriel) 

Phasis pringlei Dickson 

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64 

Phasis thero cedarbergae Dickson & Wykeham 

Poecilmitis adonis Pennington 

Poecilmitis aureus Van Son 

Poecilmitis azurius Swanepoel 

Poecilmitis balli Dickson & Henning 

Poecilmitis brooksi learei Dickson 

Poecilmitis daphne Dickson 

Poecilmitis endymion Pennington 

Poecilmitis henningi Bampton 

Poecilmitis hyperion Dickson 

Poecilmitis irene Pennington 

Poecilmitis kaplani Henning 

Poecilmitis lyncurium (Trimen) 

Poecilmitis lyndseyae Henning 

Poecilmitis nigricans nigricans (Aurivillius) 

Poecilmitis nigricans zwartbergae Dickson 

Poecilmitis orientalis Swanepoel 

Poecilmitis pan Pennington 

Poecilmitis penningtoni Riley 

Poecilmitis pyramus Pennington 

Poecilmitis pyroeis hersaleki Dickson 

Poecilmitis rileyi Dickson 

Poecilmitis stepheni Dickson 

Poecilmitis swanepoeli Dickson 

Poecilmitis trimeni Riley 

Poecilmitis wykehami Dickson 

Trimenia wallengrenii (Trimen) 

Polyommatinae 

Anthene minima (Trimen) 

Cyclyrius babaulti (Stempffer) 

Lepidochrysops bacchus Riley 

Lepidochrysops badhami Van Son 

Lepidochrysops balli Dickson 

Lepidochrysops hypopolia (Trimen) 

Lepidochrysops jamesi classensi Dickson 

Lepidochrysops jamesi jamesi Swanepoel 

Lepidochrysops jefferyi (Swierstra) 

Lepidochrysops littoralis Swanepoel & Vari 

Lepidochrysops loewensteini (Swanepoel) 

Lepidochrysops lotana Swanepoel 

Lepidochrysops methymna dicksoni Tite 

Lepidochrysops oosthuizeni Swanepoel & Vari 

Lepidochrysops oreas oreas Tite 

Lepidochrysops outeniqua Swanepoel & Vari 

Lepidochrysops penningtoni Dickson 

Lepidochrysops pephredo (Trimen) 

Lepidochrysops poseidon Pringle 

Lepidochrysops pringlei Dickson 

Lepidochrysops quickelbergei Swanepoel 

Lepidochrysops swanepoeli Pennington 

Lepidochrysops titei Dickson 

Lepidochrysops victori Pringle 

Lepidochrysops wykehami Tite 

Orachrysops ariadne (Butler) 

Orachrysops niobe (Trimen) 

Tuxentius melaena griqua (Trimen) 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

R 

I 

R 

R 

R 

I 

R 

R 

R 

Ex 

R 

R 

R 

R 

R 

V 

E 

R 

R 

R 

R 

R 

I 

R 

R 

R 

I 

R 

I 

R 

V 

R

Figure 4. Distribution of South Africa's threatened lycaenid taxa across the 22 physiographic regions. (N.B. A few taxa occur in more than one region). 

and gullies (e.g. P. azurius). Some species, despite the harsh 

winter conditions of the Cape, can be found at high elevations 

(e.g. Argyrocupha malagrida cedrusmontana at 1900m and 

Aloeides pringlei at 2072m ). The lower elevations of the Cape 

(physiographic subregions 22SC and 22WC) are also rich in 

localised endemics (Figure 4) illustrating the wide range of 

biotopes occupied by this group of butterflies in the 

topographically varied and geologically stable area on the 

southern tip of Africa. 

The next richest area in rare, threatened lycaenids is the east 

coast and hinterland. Most of the blue butterflies in these areas 

are fairly abundant, and the mountain peaks do not have the 

same richness of localised endemics as in the Cape. One species 

(Lepidochrysops lowensteini) occurs at about 2800m in the 

Lesotho Tableland, while the middle altitudes support 14 

threatened taxa. The lower altitudes of the east coast are also 

relatively rich in threatened endemics. When the four east coast 

subregions of physiographic region 22 are added together 

(Figure 4), the coastal plain from Mozambique in the north to 

Port Elizabeth in the south hosts 13 taxa. 

The Central Plateau (regions 1, 2, 3, 4, and 5) and the 

Northern Transvaal savannah (regions 7, 8, 9, 10, 11, 13, and 

14) are poor in localised, threatened endemics, with regions 4, 

5, and 13 supporting none. The reason for this is uncertain, but 

with the monotonous landscape and little opportunity for 

topographic isolation, it is possible that competition from more 

vigorous species has taken place. This may even have happened 

in recent times with Lepidochrysops hypopolia. One of the two 

original localities for this species is Potchefstroom on the 

Highveld, where it was found in 1879: today the habitat is 

occupied by the closely-related L. praeterita, with L. hypopolia 

not having been recorded for over 110 years. 

65 

Threats leading to taxon extinction 

Overcollecting 

There appear to be no verified cases of butterfly species going 

extinct through overcollecting (New 1984, Pyle et al. 1981). 

Many of the lycaenids in South Africa are in remote rugged 

localities which are not readily accessible. Some of the lowland 

species are more easily collected, but there is no verified case 

of overcollecting affecting a population level permanently. 

Invasive species 

Many plant species of the Cape fynbos heaths and shrublands 

have their seeds dispersed and buried by ants. At the turn of the 

century, the exotic Argentine ant (Iridomyrmex humilis (Mayr)) 

appeared in South Africa and recently has invaded the fynbos 

and has begun to supplant indigenous ants (Bond and Slingsby 

1984). For blue butterflies of the Cape this is a serious 

development for two reasons. Firstly, I. humilis does not 

disperse and bury the seeds as do the native ants: this will 

eventually lead to the alteration of the habitat through loss of 

Cape Proteaceae species by gradual attrition of seed reserves. 

Secondly, because many lycaenids depend directly on certain 

indigenous ant species as hosts, the loss of native ants must 

inevitably have serious long-term repercussions for the survival 

of many of the lycaenids. 

By late 1978, I. humilis had reached the Highveld, but to 

date has not been recorded from Natal. Should further increase 

in the range of I. humilis occur, which is probable, many 

lycaenids throughout the country could be affected. This may

e inevitable as there are no known methods of arresting the 

expansion orcontrolling the population levels of this aggressive, 

invasive ant. 

Apart from the very serious threat from the Argentine ant, 

none of the other ant species or other hosts associated with 

lycaenids are under threat. In fact, most species are widespread 

and abundant (e.g. Anoplolepis custodiens (Smith), 

Crematogaster spp., Camponotus spp.). 

The invasive spread of alien trees and shrubs has also been 

significant for most South African biomes (Macdonald et al. 

1986). For blue butterflies, genera such as Pinus, Acacia, 

Eucalyptus, Solarium and Rubus pose the greatest threat by 

partial or total alteration of the habitat. Without containment, 

these weeds are likely to be increasingly threatening to lycaenid 

populations (Figure 5). The habitat of Argyrocupha malagrida 

malagrida is already being heavily invaded by alien vegetation, 

as is the habitat of one of the populations of Poecilmitis pan. 

There is no evidence that classical biological control 

programmes have had any adverse effect upon lycaenid 

conservation, or even upon any other threatened insect species 

in South Africa (Samways 1988). 

Change in landscape use 

Loss of habitat is a complex issue and involves more than 

simply loss of natural areas to agriculture and urbanisation. For 

example, the increased subdivision of the landscape by vehicle 

tracks and roads prevents the spread of natural fires which are 

an important natural influence preventing the succession from 

grassland to scrub (Tainton and Mentis 1984). The savannah is 

burnt to simulate the effects of lightning strikes and maintain 

habitats in a relatively pristine state. Conversely, accidental 

fires that are more intense or regular than would normally be the 

case in nature can be a threat in their own right. This is 

particularly the case adjacent to urban or commercially forested 

areas. Such a fire threat, as well as direct habitat loss, faces, for 

example, Thestor yildizae on Table Mountain in Cape Town. 

The majority of taxa are included in the butterfly Red Data 

Book (Henning and Henning 1989) because of their extremely 

limited natural distribution (Figure 5) particularly in 

mountainous areas. However, agricultural and urban 

developments are major threats for many lycaenids, particularly 

those at low elevations. In the case of Orachrysops ariadne, 

Figure 5. Major threats, in decreasing magnitude, to South Africa's 105 rare lycaenid taxa. (N.B. Some lycaenids are threatened by more than one 

category.) 

Many are not threatened but simply have strictly limited distributions. All are threatened by global warming and increased radiation from the southern hole in 

the ozone layer. 'Extinct' is the IUCN categorisation; other categories are the actual threats. It is not known what the threats were to the Extinct species. (Data 

calculated from identified and listed threats for all red-listed species in Henning and Henning, 1989). 

66

although its site is privately protected, absence of large herbivores 

and lax management of vegetation is posing a threat to the longterm 

security of the species. 

Many of the threat categories are interrelated, e.g. urban 

expansion and fire hazards or dam construction. The overall 

root cause is the same: increased human population pressure 

which is particularly threatening to the lowland species but not 

so much to those of the rugged, relatively inaccessible, Cape 

mountains. 

Global warming and thinning of the ozone layer 

In the long term, these threats may outweigh all others. The 

earth's temperature may rise by 3°C by the year 2050, and 

models predict that although there will be little change at the 

equator, the poles may be TC warmer (Pearman 1988). Besides 

a rise in sea level, there may also be a shift in seasons. For South 

Africa, projections are for a generally warmer, drier situation, 

where soil moisture conditions in the Highveld might be 11–18% 

drier than at present (Huntley et al. 1989). The exceptionally 

species-rich Cape fynbos would be particularly affected, with 

the possible risk of a collapsing domino effect with many 

localised invertebrates disappearing along with their plant 

hosts. Additive upon these general effects will be an increase in 

inclement conditions such as droughts, hailstorms and 

hurricanes. Adding further stress will be the growing hole in the 

ozone layer of the stratosphere over the South Pole, with 

depletion in parts of up to 50% (Brunke 1988). South Africa, 

with its southerly geographical position, is likely to be 

particularly affected by the increased levels of ultraviolet 

radiation. Already, in 1989, there was a 45% depletion 

(Scourfield et al. 1990). 

All these adverse environmental pressures bode ill for many 

lycaenids with their localised distributions and specialised lifecycles 

(many of which are still to be determined). Temperature 

falls by 0.6° C for every 100m rise in altitude, which suggests 

that to maintain the same local thermal environment, species 

must move up the mountainside by at least 500m by the middle 

of next century. Such movement would be difficult for species 

whose habitat has been partitioned by roads, buildings, etc. 

(Siegfried 1989). Further, as many of the South African lycaenids 

occur on isolated ridges and peaks, they cannot go higher and 

must adapt or die. In the short term, genetic adaptation is 

unlikely, and these ecological pressures may be too great for 

survival. 

Conservation measures 

The production of the Red Data Book on butterflies (Henning 

and Henning 1989) has been an important step forward 

(Samways 1989a). Although not all the species and subspecies 

listed may be truly threatened (apart from the effects of significant 

global events) with many further populations awaiting discovery, 

the book has focused on the rarity of many butterflies, particularly 

67 

lycaenids, and is a major stepping stone for further research. 

Few insects are protected by law in South Africa, and each 

province has its own ordinances. To date no specific species or 

subspecies is protected by law in Natal or the Orange Free State. 

In the Cape Province, according to Ordinance 19 of 1974, 

Schedule 2, Protected Wild Animals may not be hunted, killed, 

captured or kept in captivity without a permit from the 

Department of Nature and Environmental Conservation. 

Amendment of Schedule 2 (13 February 1976) gives full 

protection to: Aloeides egerides, A. lutescens, Argyrocupha 

malagrida malagrida, Trimenia wallengrenii, Oxychaeta 

dicksoni, Lepidochrysops bacchus, Poecilmitis endymion, P. 

lyncurium, P. nigricans nigricans, P. rileyi, Thestor dicksoni 

dicksoni and T. kaplani. In the Transvaal, by Ordinance 12 of 

1983, Section 45 (Schedule 7) (Protected Wild Animals), 

Poecilmitis aureus is protected. 

The 582 nature reserves in southern Africa total 7 x 10 6 ha, 

5.8% of the land surface (Huntley 1989). Siegfried (1989) 

estimated that 74% of vascular plant species and over 90% of 

each of the vertebrate groups are represented in nature reserves. 

Such comparative data is not available for insects, but of the 

threatened lycaenids listed by Henning and Henning (1989), 

two are extinct, and only 24 (23%) occur in nature reserves or 

wilderness areas, all but one state owned. Apart from the 

protection ordinances, this means that 75% are not 

geographically protected in nature reserves. For many of these, 

apart from global atmospheric threats, they are relatively safe as 

their localities are in remote terrain. Given the extent of remote 

terrain there is also the possibility of the occasional new species 

being found from time to time. 

In recent years there has been an increase in invertebrate 

conservation awareness in South Africa. Aloeides dentatis 

dentatis in particular has become quite a celebrity as the 

Roodepoort City Council has established the 12ha Ruimsig 

Entomological Reserve specifically for the butterfly (Henning 

and Henning 1985). 

Although such authorities as the National Parks Board, the 

Natal Parks Board, the Defence Force, the Universities of 

Natal, Pretoria and Stellenbosch have a strong interest in 

conservation and support invertebrate conservation projects, 

only the Transvaal Provincial Administration has a full-time 

Invertebrate Conservation Officer. This is encouraging, but 

still inadequate bearing in mind that there are about 80,000 

described species of insect in South Africa (Prinsloo 1989) and 

that this may only be a quarter of the total percentage, including 

a large proportion of endemics. The emphasis to date has been 

on the conservation of large vertebrates, while the Wildlife 

Society principally supports the conservation of habitats and 

whole natural areas. It is well known that vertebrates are not 

good indicator or umbrella species for invertebrate conservation. 

Other invertebrates with specific habitat requirements and 

appropriately sized home ranges are better as flag species 

(Samways 1989b). 

Although the setting aside of game and nature reserves has 

given refuge to the great majority of vertebrates, it applies to 

less than one-quarter of the blue butterflies. Nature reserves

nevertheless are playing a role. Acquisition of much more land 

for state-owned nature reserves is unlikely, as most land is 

firmly accounted for. This means that invertebrate zoologists, 

entomologists and naturalists must monitor areas where blue 

butterfly populations are still maintaining a foothold, and lobby 

for conservation of that patch of land. As these habitat patches 

are indeed small (tennis court size in the case of Argyrocupha 

malagrida malagrida) it is a feasible proposition, as shown by 

the Henning brothers and the Roodepoort City Council in the 

setting aside of the Ruimsig Entomological Reserve. 

Summary 

Southern Africa, with its long stable geological history and 

highly varied topography and climate, is rich in insect species. 

In South Africa, 632 species of butterfly have been described, 

of which 102 (16%) are under some sort of threat. There are 389 

lycaenid species in southern Africa, with 310 species occurring 

in South Africa. A total of 105 species and subspecies of 

lycaenid are threatened, 75% of all threatened butterfly taxa in 

the country. Of the 105 threatened species two lycaenid species 

are Extinct; one species and one subspecies are Endangered; 

seven taxa are Vulnerable; 71 are Rare; and 23 are Indeterminate. 

A high proportion of the threatened lycaenid taxa (71% species 

+ 25% subspecies = 96% total) are endemic. Most of the 

threatened taxa occur in the Cape fold mountain area, usually on 

mountain slopes, and sometimes at high elevations (up to 

1,800m). Many of the rare lycaenids also occur along the east 

coast and hinterland, including one at 2800m. The Central 

Plateau and Northern Transvaal savannah are poor in localised 

endemics. 

Overcollecting is not a threat. Of great seriousness is the 

invasive Argentine ant (Iridomyrmex humilis) which is 

supplanting native ant hosts for lycaenids, the key species in 

maintaining the character of the habitats through seed burial. 

Invasive exotic plant species are also a threat for several 

lycaenids, as is fire, mining, dam building, forestry, and, for one 

species, neglectful management. For the lowland species, 

agricultural and urban expansion are the greatest threats, but for 

many of the species, which generally inhabit mountainous 

areas which are rugged or remote, it is simply that they have an 

extremely limited distribution. 

Of great concern in southern Africa is the overall effect of 

global warming and the hole in the ozone layer. Projections 

indicate that most species will need to shift in elevation by 

500m by the year 2050 if they are to remain at the same 

temperature. For most species, especially those that inhabit 

hilltops, such a shift would be ecologically and genetically 

impossible. 

Twelve lycaenid taxa are protected by law in the Cape 

Province, and one species in the Transvaal. None are protected 

in the Orange Free State or Natal. Only 23% of threatened 

lycaenids occur within nature reserves or wilderness areas, 

compared with over 90% for each of the vertebrate groups and 

68 

74% for vascular plants. One species, Aloeides dentatis, has 

been allocated a 12ha reserve of its own. Further acquisition of 

patches of land for specific lycaenid populations is a feasible 

short-term approach to further the cause of invertebrate 

conservation. 


The Foundation for Research Development and the University 

of Natal Research Fund provided financial assistance. Messrs. 

Stephen and Graham Henning kindly criticised the text and Mrs 

Ann Best and Mrs Myriam Preston kindly processed the 

manuscript. 

References 

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Argentine ant (Iridomyrmex humilis) and myrmecochorous Proteaceae. 

Ecology 65: 1031–1037. 

BRUNKE, G.C. 1988. Tropospheric background measurements of CFCI3(F- 

II) conducted at Cape Point, South Africa, since 1979. In: Macdonald, 

I.A.W. and Crawford, R.J.M. (Eds), Long-term Data Series relating to 

southern Africa's Renewable Natural Resources. South African National 

Scientific Programmes Report No. 157. Foundation for Research 

Development, Council for Scientific and Industrial Research, Pretoria. pp. 

434–435. 

CLARK, G.W. and DICKSON, C.G.C. 1971. Life Histories of South African 

Lycaenid Butterflies. Purnell, Cape Town. 272pp. 

HENNING, S.F. and HENNING, G.A. 1985. South Africa's endangered 

butterflies. Quagga 10: 16–17. 

HENNING, S.F. and HENNING, G.A. 1989. South African Red Data Book- 

Butterflies. South African National Scientific Programmes Report No. 

158. Foundation for Research Development, Council for Scientific and 

Industrial Research, Pretoria. 175pp. 

HUNTLEY, B.J. (Ed.) 1989. Biotic Diversity in Southern Africa: Concepts 

and Conservation. Oxford University Press, Cape Town. 380pp. 

HUNTLEY, B.J., SIEGFRIED, R. and SUNTER, C. 1989. South African 

Environments into the 21st Century. Human and Rousseau Tafelberg, 

Cape Town. 127pp. 

MACDONALD, I.A.W., KRUGER, F.J. and FERRAR, A.A. (Eds) 1986. The 

Ecology and Management of Biological Invasions in southern Africa. 

Oxford University Press, Oxford. 324pp. 

MURRAY, D.P. 1985. South African Butterflies. A Monograph of the Family 

Lycaenidae. Staples Press, London. 195pp. 

NEW, T.R. 1984. Insect Conservation – An Australian Perspective. Junk, 

Dordrecht. 184pp. 

PEARMAN, G.I. (Ed.) 1988. Greenhouse: Planning for Climate Change. 

CSIRO, Melbourne. 

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273pp. 

PRINSLOO, G.L. 1989. Insect identification services in South Africa. 

Proceedings of the Seventh Entomological Congress organized by the 

Entomological Society of southern Africa, Pietermaritzburg 10–13 July 

1989. p.107. 

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Ann. Rev. Entomol. 26: 233–258. 

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Are they compatible? Env. Conserv. 15: 349–354. 

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of bush crickets (Tettigoniidae) in southern France. Env. Cons. 16: 

217–226. 

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SALTER, L.F. 1990. Ozone: the South African connection. S. Afr. J. Sci. 

86: 279–281. 

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reserves. In: Huntley, B.J. (Ed.) Biotic Diversity in Southern Africa: 

69 

Concepts and Conservation. 

TAINTON, N.M. and MENTIS, M.T. 1984. Fire in grassland. In: Booysen, P. 

de V. and Tainton, N.M. (Eds). Ecological Effects of Fire in South African 

Ecosystems. Springer-Verlag, Berlin. pp. 115–147. 

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the Transvaal Museum, Pretoria. 198pp.

Introduction 

Australian Lycaenidae: conservation concerns 

T.R. NEW 


The Lycaenidae of Australia comprise about 140 described 

species (Table 1, Figure 1). The family is most diverse in the 

northern tropical parts of the country, where the fauna of 

northern Queensland has strong relationships with that of New 

Guinea and parts of the Oriental Region. Indeed, Eliot (1973) 

chose to follow Gressitt (1956) in considering the northern tip 

of Queensland, Cape York Peninsula, as part of the Oriental 

Region in delimiting areas for considering lycaenid distribution 

patterns. 

Taxonomic appraisal of the fauna, at least of the adult 

stages, is relatively complete (Common and Waterhouse 1981) 

and it is possible to provide a reasonably sound appraisal of the 

status and broad scale distribution of all species. Dunn and 

Dunn (1991) give maps of the documented distribution of 

Australian butterflies. A few further species undoubtedly remain 

to be discovered, but most future changes in taxonomy are 

likely to be made at the species/subspecies interface, with some 

subspecies being elevated in status as more information becomes 

available. Not surprisingly for a large continent extending from 

the humid tropics to cold temperate regions and with large arid 

and semiarid zones, the distribution of many lycaenid species is 

circumscribed, and there is room for considerably more research 

to clarify the precise ranges and status of many of the more 

elusive resident taxa. 

Only one riodinine (Praetaxila segecia punctaria 

(Fruhstorfer)) extends into Australia, where it is confined to the 

Table 1. Taxonomic summary of the Australian Lycaenidae (data from 

Dunn and Dunn 1991). 

Subfamily 

Liphyrinae 

Riodininae 

Theclinae 

Polyommatinae 

Total 

No. genera 

1 

1 

15 

22 

39 

No. species 

1 

1 

75 

65 

142 

70 

extreme north of Queensland; its close relatives are found in 

New Guinea. Intriguingly, four species of Lycaeninae occur in 

New Zealand, but none in Australia. In general, the Australian 

Lycaenidae show some attenuation in diversity from those of 

New Guinea and Malaysia, and the number of subfamilies is 

smaller. This is particularly obvious at the tribal level: Australia 

has 32 species of Luciini compared with about a hundred in 

New Guinea, and only four Arhopalini compared with 38 in 

New Guinea and slightly more than a hundred in Malaysia. Of 

the generally widespread subfamilies four are absent: Miletinae; 

Curetinae; Lycaeninae; and Poritiinae. The two predominant 

subfamilies in Australia are Polyommatinae andTheclinae, and 

endemism is high in both (Polyommatinae: 4/22 genera, 22/65 

species; Theclinae: 6/15 genera, 25/75 species). Theclini, in 

particular, are a major radiation in the Australian butterfly 

fauna (Kitching 1981). These figures for endemism do not 

include numerous putative subspecies which are clearly restricted 

to Australia and often have very limited distributions. The 

biological status of most of these is by no means clear and some, 

at least, may constitute sibling species complexes. 

Many endemic taxa are very rare. Several, indeed, are 

known only from one or two localities. For example, 

Hypochrysopspiceatus Kerr, McQueen & Sands has only been 

found in two small colonies in southern Queensland and only 

one of these now remains. Jalmenus aridus Graham & Moulds, 

one of very few butterflies believed to be endemic to Australia's 

interior arid zone, is known from one colony in Western 

Australia. A number of subspecies of Ogyris Westwood and 

Candalides Hiibner, inter al., also have very limited distributions. 

Some of these, such as the 'O. idmo group' in Western Australia, 

are taxonomically more complex than currently documented 

(Field 1992), and perhaps of greater conservation concern than 

suspected at present. 

It is notable that ant-dependence to differing extents occurs 

in virtually all the genera which are diverse and widespread. 

This habit may have been instrumental in leading to 

diversification of Lycaenidae in some semiarid regions where 

plant growth is very irregular and the spectrum of food plants 

limited. Larval feeding habits differ considerably between 

various genera. Additional foodplant records continue to be 

accumulated (e.g. Valentine and Johnson 1988). In Ogyris

Figure 1. Australia: main political regions and lycaenid fauna. 

Regions denoted by inilial letters: ACT, Australian Capital Territory; NSW, New South Wales; NT, Northern Territory; SA, South Australia; T, Tasmania; 

V, Victoria; WA, Western Australia. Lycaenid subfamilies: L, Liphyrinae; P, Polyommatinae; R, Riodininae; T, Theclinae. Figures are no. of included genera/ 

species. 

Figure 2. Australia: main specific places mentioned in the text. 

71

caterpillars of some species may live in ant nests throughout 

their lives. 

By contrast many other Ogyris larvae feed on mistletoes 

(Loranthaceae, Viscaceae), and this unusual host plant group 

may have been a major basis for speciation in this genus. In 

contrast, Hypochrysops has exploited a taxonomically diverse 

range of flora, so that diversification has occurred in these 

genera by using contrasting ecological strategies. However, 

very few lycaenids feed directly on one dominant tree genus, 

Eucalyptus. 

The Australian fauna 

Some highlights of the Australian fauna are described below 

with much of the information gleaned from the extensive data 

summarised by Common and Waterhouse (1981). 

• The widespread oriental species Liphyra brassolis major 

Rothschild is rather rare in northern tropical Australia, 

where it occurs in association with Oecophylla ants. 

• In the Luciini, many taxa are very local. For example, Lucia 

limbaria Swainson occurs in isolated localities in the south 

and east of mainland Australia. 

• Several species of Acrodipsas Sands (formerly referred to 

Pseudodipsas C. & R. Felder) are also rare and localised: A. 

arcana (Miller & Edwards) is known from only one hilltop 

in New South Wales, and A. hirtipes Sands from a hilltop 

near Coen (northern Queensland) (Figure 2) where it occurs 

with A. melania Sands. The latter is known also from a 

single specimen captured at the very tip of Cape York 

Peninsula. A. brisbanensis cyrilus (Anderson & Spry) is a 

local subspecies in Victoria and A. b. brisbanensis (Miskin), 

although more widely distributed along the east coast, is 

also regarded as rare. Indeed, none of the seven species of 

Acrodipsas is common. 

• Paralucia Waterhouse & Turner contains three species of 

which two are rare: P. spinifera Edwards & Common (see 

Dexter and Kitching, this volume) is one of our rarest 

lycaenids, and is known only from one restricted area of 

New South Wales; and P. pyrodiscus lucida Crosby has 

recently aroused considerable conservation interest in 

Victoria (see New 1992, and New, this volume). 

• Hypochrysops C. & R. Felder is the most speciose lycaenid 

genus in Australia, and many of the 18 species are rare and 

local (Sands 1986 and Sands, this volume). It is absent from 

Tasmania, and most of the species are northern or east 

central in distribution. Several New Guinea (or closely 

related) species are known only from northern Cape York, 

and the only species in Western Australia is H. halyaetus 

Hewitson. As with Philiris Rober and Arhopala Boisduval, 

Australian subspecies of some New Guinea species have 

developed. 

• Ogyris, with 12 endemic Australian species, also occurs in 

New Guinea but is of considerable biological interest in 

Australia. Several species include a number of named 

72 

infraspecific taxa and many of these are local and rare forms 

attractive to collectors. Both subspecies of O. idmo Hewitson 

are extremely rare, as are O. otanes C. & R. Felder, O. 

ianthus Waterhouse, and O. iphis Waterhouse & Lyell. All 

merit strenuous conservation measures, in common with 

several more widespread forms. 

• Jalmenus Hubner contains 10 species. J. pseudictinus is 

regarded as very local in parts of eastern Queensland, as is 

J. lithochroa Waterhouse in southern South Australia. J. 

dementi Druce is known from a very small region of 

northwestern Australia, and J. aridus from a colony near 

Kalgoorlie (Figure 2). 

• The single species of Pseudalmenus Druce, P. chlorinda 

(Blanchard) is confined to southeastern Australia, and seven 

subspecies have been described. Four of these are from 

Tasmania (see Prince, this volume), where populations 

separated by only a few kilometres have very different wing 

markings. P. c. fisheri Tindale, from Western Victoria, is 

also very local. 

• Some other lycaenids are not regarded as rare but are very 

clearly restricted to particular geographical regions or 

ecological communities: Neolucia hobartensis (Miskin), 

with two subspecies, is an alpine/subalpine species of 

southeastern Australia. 

Distributional and diversity patterns 

The distribution patterns (Dunn and Dunn 1991) reveal several 

trends relevant to the consideration of lycaenid conservation in 

Australia. The northern part of Queensland, as for many other 

biota, supports a large number of taxa on the southernmost 

fringes of their Oriental/New Guinea distributions. Remnant 

rainforests in this region are particularly important butterfly 

habitats (Monteith and Hancock 1977) and the individual 

patches may be viewed as an 'archipelago' of these in northern 

Queensland. 

Lycaenid diversity is greatest around the eastern and 

southeastern fringe of the Australian mainland, with a smaller 

number of species in the west or southwest. Very few species 

occur in the climatically inhospitable inland. 

A very high proportion of Australian Lycaenidae are forest 

and open woodland-frequenting taxa. A few herb/shrub 

community taxa, such as Lampides boeticus (L.), Theclinesthes 

serpentata (Herrich-Schaffer) and Zizina labradus (Godart), 

are amongst the most widely distributed lycaenids in Australia. 

Habitat relationships in the southeast are exemplified by an 

analysis for the Australian Capital Territory (Table 2: Kitching 

et al. 1978). 

Most of the non-endemic species belong to oriental genera, 

and essentially constitute the 'younger northern element' of the 

fauna. Endemism at the generic level is distinctively more 

southern (and, especially, southeastern), with the implication 

that some, at least, may represent speciation from earlier 

colonisers than those taxa which occur solely in the north; a

Table 2. Habitat relationships of 25 species of I.ycaenidac recorded from 

the Australian Capital Territory. Figures given are numbers of species. 

Data from Kitching et al. 1978. 

No. Habitat 

1 

2 

3 

4 

5 

Lowland savannah 

Savannah woodland 

Dry sclerophyll forest 

Wet sclerophyll forest 

Alpine zone 

Total 

5 

16 

15 

11 

4 

No. species 

shared with habitat 

2 3 4 5 

5 5 

12 

2 

5 

6 

1 

1 

2 

4 

Not 

shared 

similar situation occurs in Australian Satyrinae (New in press). 

Couchman and Couchman (1977) considered Tasmanian forms 

of P. chlorinda to be 'evidently of very ancient origin'. Centres 

of endemism, local 'critical faunas', can thus be delimited with 

some degree of reliability. 

Conservation 

Lycaenid diversity is highest in those parts of Australia which 

are subject to substantial, rapid and largely irreversible changes 

by European people. Thus diversity is highest in the eastern and 

southeastern fringe of the Australian mainland, with smaller 

numbers of species in the west or southwest and very few in the 

climatically inhospitable inland. Environmental changes through 

human activities have accelerated in recent decades and show 

little sign of abating or slowing in the near future: assessing 

threats to taxa at both local and national levels becomes essential 

if the loss of species and notable subspecies is to be avoided. 

For some parts of the country it is not possible to determine 

if some taxa were formerly more widespread than they are at 

present. Inferences on past endangering processes, including 

habitat change and loss, thus contain a large element of historical 

supposition. 

As stated earlier, local 'critical faunas' can be delimited 

with some degree of reliability. In addition, there are a number 

of 'critical habitats' for Lycaenidae which can be identified. 

These occur on a macroscale (isolated pockets of rainforest – as 

in northern Queensland, alpine or mallee vegetation) and also 

as specific sites which support one or more narrow endemics 

and which may be especially vulnerable–several of the hilltops 

referred to in the taxonomic summary are good examples of 

this. Because of their restricted or sporadic distributions over 

this vast island continent, a substantial number of endemic 

lycaenids on the Australian mainland must be considered 

vulnerable to continuing habitat change, even though they are 

not directly or imminently threatened. 

– 

4 

2 

3 

– 

73 

Threatened taxa 

In a preliminary survey of threatened insects in Australia, Hill 

and Michaelis(1988) included nine species of Lycaenidae, five 

of which were known only from one or two sites. The survey 

was based on a questionnaire circulated widely to entomologists 

in Australia, seeking information on priorities for insect 

conservation. In all, 24 taxa of Lycaenidae (Table 3) were noted 

by respondents and these included three subspecies of P. 

chlorinda, two of which occurred in Tasmania. Tasmanian taxa 

were noted as of concern by other respondents, so that 

conservation concern for native Lycaenidae has a broad 

geographic base within Australia. 

It is notable that the Hill and Michaelis list includes threatened 

Lycaenidae from all mainland states except the Northern 

Territory, where documentation is relatively incomplete. 

Nadolny (1987) noted that the most endangered butterfly in 

New South Wales is likely to be Paralucia spinifera, which is 

Table 3. Taxa of Lycaenidae in Australia which may be threatened. 

Data from Hill and Michaelis (1988), digested from respondents to survey 

conducted by Australian National Parks and Wildlife Service, closing February 

1985. States abbreviated as in Figure 1. 

Taxon 

Acrodipsas arcana (Miller & Edwards) 

A. brisbanensis brisbanensis (Miskin) 

A. illidgei (Waterhouse & Lyell) 

A. n.sp. 

A. myrmecophila (Waterhouse & Lyell) 

Hypochrysops apollo Miskin 

H. clean Grose-Smith 

H. epicurus Miskin 

H. hippuris Hewitson 

H. ignitus ignitus (Leach) 

H. piceatus Kerr, Macqueen & Sands 

Jalmenus lithochroa Waterhouse 

Jamides cytus claudia (Waterhouse & Lyell) 

Ogyris amaryllis meridionalis Bethune-Baker 

O. idmo halmaturia Tepper 

O. idmo idmo Hewitson 

O. otanes C. & R. Felder 

Paralucia spinifera Edwards & Common 

Philiris azula Wind & Clench 

P. ziska titeus D'Abrera 

Pseudalmenus chlorinda chlorinda (Blanchard) 

P. c. barringlonensis Waterhouse 

P. c. conara Couchman 

Theclinesthes albocincta (Waterhouse) 

State(s) 

Q,NSW 

Q.NSW 

Q 

WA 

Q,NSW,V 

Q 

Q 

Q,NSW 

Q 

SA,NSW,V 

Q 

SA 

0 

SA 

SA,V 

WA 

SA,NSW,WA 

NSW 

0 

Q 

T 

NSW 

T 

SA

of considerable phylogenetic importance in possibly linking 

Paralucia with related genera. P. spinifera, the Bathurst Copper, 

has recently been the subject of more detailed study in the State 

(Dexter and Kitching, this volume). 

However, in general there are few detailed accounts of 

range contractions of taxa other than on a very local scale or by 

inferences from older collectors who claim that some species 

are now scarcer or more restricted than in the past. More survey 

work, particularly in poorly documented areas, is needed to 

identify all threatened taxa of the Australian Lycaenidae. 

Major threats to Australian lycaenids 

The survey by Hill and Michaelis (1988) identified some of the 

major threats to Australian lycaenids and these are listed below, 

together with other threats specified in the literature: 

• land clearing, sometimes by fire, for agriculture and urban 

development; 

• urbanisation and tourist resort development; 

• pest fly control; 

• roadworks; 

• mining; 

• collecting (individual and commercial); 

• agricultural practices, e.g. pasture improvement. 

Examples of species threatened by the ecological processes 

follow; others can be found in the literature. Any of these 

species might also be susceptible to overcollecting. 

(i) The best known and most accessible site for H. piceatus 

is a small patch of mistletoe-bearing Casuarina along some 

200m of road verge, which could be eliminated easily and 

inadvertently by road widening. 

(ii) O. otanes, recently (July 1989) the first butterfly to be 

nominated for listing under the Victorian Flora and Fauna 

Guarantee Act, is known in Victoria from a single hilltopping 

site in the arid northwest of the state which has been the subject 

of physical disturbance during the establishment of a 

trigonometric survey point, by vehicles on sand-dunes. Other 

populations of O. otanes were known in the 1970s but were 

destroyed by collecting. 

(iii) One of the very few sites at which Acrodipsas 

myrmecophila occurs, and the only one known in Victoria at 

present, is currently the target of mining exploration. 

Additionally, it is not clear whether the two species of Acrodipsas 

described from a hill near Coen breed on the hill or in nearby 

lowland vegetation and then hilltop. In both cases, much nearby 

land is currently subject to mining exploration and this could 

pose a threat. Several such hilltops in Australia are among the 

classic collecting localities favoured by butterfly collectors, 

and such rare species could be rendered additionally vulnerable 

by uncontrolled collecting. 

(iv) The several species associated with coastal mangrove 

vegetation are vulnerable as drainage occurs for urban or tourist 

resort development. 

(v) The detailed appraisal of P. chlorinda in Tasmania 

(Couchman and Couchman 1977), which has recently been 

updated (Prince 1988), claims that it has been eliminated over 

74 

whole areas where one or other of its required major resources 

(a eucalypt, an Acacia – normally A. dealbata – and an 

Iridomyrmex ant) have been destroyed. Couchman and 

Couchman located the species in more than 50 localities in the 

years following 1945 but at the time of writing their account 

found it difficult to think of more than 10 localities where P. 

chlorinda might then survive. 'Pasture improvement' with 

removal of all mature eucalypts and acacias in paddocks badly 

affected P. c. conara, and most of the habitat of the eastern P. 

c. chlorinda has been clearfelled for woodchips. Two other 

forms are extinct, one because of local tree clearing and one 

because of housing development and clearing/burning, and 

some existing forms are highly localised (see Prince, this 

volume). 

(vi) Of the three species of Acrodipsas listed as threatened 

by Hill and Michaelis (1988), A. arcana is threatened by fires 

and clearing, A. illidgei (a mangrove-frequenting species, see 

Samson this volume) by urbanisation and use of insecticides 

against mosquitoes, and a third (undescribed) species is subject 

to habitat destruction by clearing for agriculture. 

Geographical areas of importance for lycaenids 

While it is essential to identify threatened species as foci for 

conservation attention, some larger geographical areas also 

merit particular attention by harbouring diverse or unusual 

faunal groups. Worthy of note are: 

i) Cape York Peninsula. This is a major region of faunal 

interchange between Australia and New Guinea, with many 

northern Lycaenidae not extending further into Australia. This 

northern element includes some 75 species (more than half the 

Australian Lycaenidae) and, whereas a number of these extend 

further down the east coast or elsewhere, many species are both 

rare and restricted to the 'far north', often to forested areas. The 

number of rainforest Lycaenidae decreases latitudinally from 

about 32 species at Iron Range to none in the cool temperate 

forests of Victoria and Tasmania. Tropical rainforests are a key 

habitat for maintaining the integrity of tropical fauna in 

Australia. 

ii) Southern Queensland. Kitching (1981) highlights the 

very high diversity of Lycaenidae in southern Queensland, and 

suggests that this reflects the confluence of northern and southern 

faunas. Thus, many of the northern taxa reach their southern 

limits here and the putatively older southern forms, their most 

northern range extensions. There are also a number of rare 

species found only in this region, which supports around half 

the Australian lycaenid species. 

iii) Western Victoria, particularly the Grampians 

('Gariwerd') mountains and the 'Mallee region'. A number of 

rare inland forms occur in these areas, and some taxa from 

semiarid regions are scarce or non-existent elsewhere. 

These regions include substantial areas of National Park or 

other reserves: Iron Range and the Grampians ('Gariwerd') 

National Parks are two examples. In situations where the prime 

conservation need is security of habitat and our state of 

knowledge of particular species does not permit intervention to

markedly influence management, protection of these reserves 

and their enhancement is the most important step which can be 

taken. Presence of rare Lycaenidae, such as P. chlorinda fisheri 

in the Grampians ('Gariwerd') can here augment pressures for 

habitat protection and maintenance of the integrity of reserves. 

The role of insects in such planning in Australia is in its infancy, 

and the limited work on lycaenid conservation to date has been 

almost entirely species-targeted. 

In the current Australian political climate, demands for 

multiple land use, sometimes involving substantial intrusion 

into National Parks, are not uncommon. 

Legislation 

None of the threatened lycaenids identified by Hill and Michaelis 

(1988) is formally listed as endangered in any state other than 

Victoria and Queensland, so there is no legal mechanism for 

their protection other than their fortuitous and largely 

undocumented/unmonitored incidence in nominally safe 

'Reserves'. 

The issue of 'species listing' is a controversial one in 

Australia. In the past a number of insect species have been 

gazetted as 'protected' in various States, with little apparent 

reason. This practice has in some instances alienated concerned 

collectors and others who are in a position to help very positively 

with the documentation needed to clarify the status and well 

being of thetaxa involved. In no case until 1989 had 'listing' of 

an insect species been accompanied by any form of formal 

undertaking or provision for study of the biology of the species. 

The single species listed in Queensland legislation, 

Acrodipsas illidgei (see Samson, this volume) was designated 

in 1990 as 'permanently protected fauna', an extraordinarily 

high level of nominal protection placing it on a par with some 

charismatic vertebrates. 

The recent legislation enacted in Victoria in 1988 and 

known as the 'Flora and Fauna Guarantee' is pioneering in 

scope. Taxa can be nominated for listing, and made subject to 

an 'interim conservation order' – a legal hiatus which provides 

the opportunity for a more formal appraisal of the status of the 

species during a period when it is nominally protected from 

further intensification of the threatening processes such as 

habitat destruction. Essentially, the onus then falls on the State 

Department for Conservation and Natural Resources to 

investigate the species, and to clarify its need for conservation. 

If identified as threatened, a management plan must be produced 

to clarify the major steps needed to protect the species. Clearly, 

this is limited to within Victorian State boundaries, but many 

conservationists are hoping that despite its Utopian and possibly 

impracticable (because of restricted logistic capability) ideals, 

the Guarantee will tangibly safeguard rare Victorian endemics 

and isolated remnant or outlying populations of taxa more 

common in other parts of the country. 

As noted earlier, O. otanes was the first species to be listed 

in Victoria and this was followed by nominations for a number 

of other lycaenids (Acrodipsas myrmecophila, A. brisbanensis, 

Paraluciapyrodiscus lucida, as further candidates. The isolated 

75 

hill on which A. myrmecophila occurs in Victoria, Mt Piper, 

also supports A. brisbanensis and has been listed (as Butterfly 

Community No. 1) for investigation as a 'threatened community', 

again a pioneering step for butterfly conservation in Australia, 

on the basis of this unique co-occurrence. Very few lycaenid 

species have thus been accorded consideration for legal 

protection in Australia. 

Both O. otanes and O. idmo have been placed, for some 10 

years, on a 'voluntary restricted collecting code' list of the 

Entomological Society of Victoria. This 'code', heeded by the 

great majority of responsible collectors, restricts the numbers 

of adults which can be taken by any person to two each year, and 

deters the collection of early stages. Because Ogyris larvae 

pupate in groups under the loose bark of eucalypts, this stage is 

undoubtedly vulnerable to overcollecting: it is the easiest one to 

obtain in order to procure first class cabinet specimens, and 

collectors tend to take a surfeit of pupae to counter losses due 

to parasitoids. 

Although particular lycaenids are highly sought after by 

collectors in Australia, the extent of commercial dealing in 

butterflies is rather low. A few local forms have appeared in 

dealers' lists in Australia in recent years, including various 

subspecies of Pseudalmenus and Ogyris. Prices have been in 

the order of $5–10/specimen, rarely more, and the market does 

not appear to be large. No information is to hand on numbers of 

specimens sold to overseas collectors as opposed to those in 

Australia, but a number of the rarer taxa have been listed as 'expupa', 

implying their wild origin. 

The future for the Australian Lycaenidae 

Habitat alteration in many parts of Australia has continued at an 

accelerating pace in recent years. In some documented cases 

habitat degradation has resulted in serious range contractions 

for lycaenids (Hill and Michaelis 1988; Couchman and 

Couchman 1977, updated by Prince 1988). There are 

undoubtedly other undocumented cases amongst the Australian 

lycaenids. There is clearly still a very long way to go to improve 

knowledge of biology and local distributions. Even for Victoria, 

perhaps the best documented mainland state, distribution maps 

are incomplete on a fine scale and progress towards improving 

this condition is slow. 

However, there are some encouraging signs. There is, for 

example, the improved legislation pioneered by Victoria, which 

has produced the legal mechanisms necessary for protection of 

invertebrate species in this State and has taken this beyond mere 

prohibition of collecting to provision for sound scientific 

management. Another encouraging sign in recent years is that 

an increasing number of biologists and others are becoming 

aware of the effects of habitat alteration on insects, are seeking 

to counter them at all levels, and to document more fully the 

distribution of butterflies in Australia. Finally, there is evidence 

of an increasing awareness of the importance of invertebrates in 

ecosystems and natural community dynamics. Butterflies are

playing their part as 'invertebrate ambassadors' in promoting 

this awareness: the Eltham Copper (see New, this volume) has, 

more than any other single invertebrate species, helped to bring 

the plight of many insects to wide public and government 

attention in Victoria. A. illidgei in Queensland (see Samson, 

this volume) has also been a key factor in saving some important 

coastal areas from poorly planned development. 

The topic of insect conservation is now widely discussed in 

Australia, and many of the themes of concern are addressed by 

New (1984, 1992) and Greenslade and New (1991). A more 

general appraisal of the Australian environment and the 

widespread changes that have occurred during only 200 years 

of European settlement are included in Jeans (1986). The 

factors and processes exemplified above for lycaenids are of 

much wider concern in affecting much of the endemic Australian 

biota. 

References 

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. 

2nd ed. Angus and Robertson, Sydney. 

COUCHMAN, L.E. and COUCHMAN, R. 1977. The Butterflies of Tasmania. 

Tasmanian Year Book 1977: 66–96. 

DUNN, K.L. and DUNN, L.E. 1991. Review of Australian Butterflies: 

Distribution, Life History and Taxonomy. Part 3. Lycaenidae. Power 

Press, Bayswater, Victoria. 

ELIOT, J.N. 1973. The higher classification of the Lycaenidae: a tentative 

arrangement. Bull. Brit. Mus. nat. Hist. (Ent.) 28: 373–506. 

FIELD, R. 1992. [Research Grant Report on 'Life history studies and species 

determination of the Ogyris idmo Hewitson (Lepidoptera: Lycaenidae) 

complex in Western Australia']. Myrmecia 28 (4): 12–17. 

GREENSLADE, P. and NEW, T.R. 1991. Australia: conservation of a 

76 

continental insect fauna. In: Collins, N.M. and Thomas, J.A. (Eds) 

Conservation of Insects and their Habitats, pp. 33–70. Academic Press, 

London. 

GRESSITT, J.L. 1956. Some distribution patterns of Pacific Island Fauna. 

Syst. Zool. 5: 11–32. 

JEANS, D.N. (ed.) 1986. Australia – a Geography. Vol. 1. The Natural 

Environment. Sydney University Press, Sydney. 

HILL, L. and MICHAELIS, F.B. 1988. Conservation of insects and related 

wildlife. Occasional Paper No. 13. Australian National Parks and Wildlife 

Service, Canberra. 

KITCHING, R.L. 1981. The geography of the Australian Papilionoidea. In: 

Keast, A. (Ed.) Ecological Biogeography of Australia. W. Junk, The 

Hague, pp. 979–1105. 

KITCHING, R.L., EDWARDS, E.D., FERGUSON, D., FLETCHER, M.B. 

and WALKER, J.M. 1978. The butterflies of the Australian Capital 

Territory. J. Aust. ent. Soc. 17: 125–133. 

MONTEITH, G.B. and HANCOCK, D.L. 1977. Range extensions and notable 

records of butterflies of Cape York Peninsula, Australia. Aust. ent. Mag. 

4: 21–38. 

NADOLNY, C. 1987. Rainforest Butterflies in New South Wales: their 

Ecology, Distribution and Conservation. NSW National Parks and Wildlife 

Service, Sydney. 

NEW, T.R. 1984. Insect Conservation: an Australian Perspective. W. Junk, 

Dordrecht. 

NEW, T.R. 1992. Conservation of butterflies in Australia. J. Res. Lepid. 29: 

237–253. 

NEW, T.R. (in press). The evolution and characteristics of the Australian 

butterfly fauna. In: Jones, R.E., Kitching, R.L. and Pierce, N.E. (Eds) 

Biology of the Australian Butterflies. CSIRO, Melbourne. 

PRINCE, G.B. 1988. The conservation status of the Hairstreak Butterfly 

Pseudalmenus chlorinda Blanchard in Tasmania. (Report to Department 

of Lands, Parks and Wildlife. Hobart, Tasmania). 


(Lepidoptera: Lycaenidae). Entomonograph No. 7. E.J. Brill, Leiden. 

VALENTINE, P.S. and JOHNSON, S.J. 1988. Some new larval food plants 

for north Queensland Lycaenidae (Lepidoptera). Aust. ent. Mag. 14: 

89–91.

PART 3. ACCOUNTS OF PARTICULAR TAXA 

OR COMMUNITIES 

The following examples complement and extend points raised 

in the previous section. The series of case-histories presented 

includes some which are classics in insect conservation and 

which have necessitated considerable research over many years, 

and some which are regarded as priorities for future study and 

appraisal. They range from the well-known to the speculative. 

Most are taxon-based, but several neotropical assemblages 

regarded as 'threatened communities' are included also. 

For two of the taxon studies which have been of critical 

importance in advancing the knowledge and practice of lycaenid 

conservation, authors were not found. Rather than ignore these 

and impoverish the perspective which I hope this book will 

provide, I have included accounts of these abstracted from 

published papers and reports. The 'species accounts' are, 

therefore at two levels: those attributed to particular authors 

who have usually played leading roles in the study of the 

particular species and non-attributed accounts which give a 

more historical perspective from an 'outsider'. These latter 

accounts may be open to update and revision. 

Several species are discussed in substantial detail in the 

previous section, and accounts of the Mission Blue and Karner 

Blue (see Cushman and Murphy, this volume) are of the utmost 

importance in insect conservation in North America. Likewise, 

Bálint (this volume) provides short accounts of 17 further 

eastern European taxa, mostly little known but which can act as 

foci for future attention in that region. Despite lack of current 

detailed information on such taxa, appraisals such as those 

presented for these taxa are invaluable in demonstrating the 

scope of regional needs. Some taxa have been especially 

significant in raising public awareness of butterfly conservation 

(New 1991). The Large Blue (Maculinea arion) in Britain and 

Introductory comment 

77 

the Mission Blue (Plebejus icarioides missionensis) in the 

United States are, perhaps, the most widely known. The Xerces 

Blue (Glaucopsychexerces) became extinct in California shortly 

before the Second World War, and is commemorated in the 

name of the Xerces Society, a leading body for the promotion 

of invertebrate conservation in North America and elsewhere. 

Foundation of the Society in December 1971 was, indeed, 

stimulated by the plight of the Large Blue in Britain (Pyle 

1976). 

Many of the most informative and influential cases of 

lycaenid conservation in the northern hemisphere have involved 

conservation of subspecies, sometimes of remnant populations 

or those close to the edge of a species' range. 

As far as possible, a standard sequence of subheadings is 

adopted in this section, to facilitate comparison between taxa. 

Comments on' Status', unless otherwise made clear, refer to the 

geographical range or country indicated, and not necessarily 

the entire species range. ('Red List') denotes that the species is 

listed by IUCN (1990). The abbreviation 'UTM', used in 

several European species accounts, refers to 'Universal 

Transverse Mercator' projection. 

References 

IUCN 1990. IUCN Red List of Threatened Animals. IUCN, Cambridge. 

NEW, T. R. 1991. Butterfly Conservation. Oxford University Press, Melbourne. 

PYLE, R.M. 1976. Conservation of Lepidoptera in the United States. Biol. 

Conserv. 9: 55–75.

The mariposa del Puerto del Lobo 

Agriades zullichi Hemming (= nevadensis Zullich) 

M.L. MUNGUIRA and J. MARTIN 

Departamento de Biologia (Zoologia), Facultad des Ciencìas, Universidad Autonoma de Madrid, Madrid, Spain 

Country: Spain (southeast). 

Status and Conservation Interest: Status – vulnerable. 

This species is probably one of Europe's rarest butterflies. 

Restricted to the high altitude schist screes of the Sierra Nevada 

(Granada Province), it is threatened in part of its range by a 

planned redevelopment of the ski station already built within its 

range. Its collection has only been reported five times although 

the area in which it lives is a classical collecting site. Nevertheless, 

a recent survey (Munguira 1989) found it was still abundant in 

one locality. 

Taxonomy and Description: The species was regarded as a 

subspecies of Agriades glandon (de Prunner) for almost 50 

years after it was described as a new species. This fact has 

prevented it from being listed separately by Heath (1981) and 

Viedma and Gomez (1976, 1985). 

Recent studies consider it to be a true species (Kudrna 1986; 

Munguira 1989) based on features of larval and adult 

78 

morphology, geographical isolation and its distinct ecology. 

There are two other species of this genus in Europe including A. 

glandon which is common in the Pyrenees, Alps and 

Scandinavian tundra and A. pyrenaicus Boisduval which lives 

in several high altitude ranges in southern Europe. These two 

species have been listed as 'endemic' (Viedma and Gomez 

1976, 1985) and 'vulnerable' (Heath 1981) respectively. 

Distribution: A. zullichi has only been found in three localities 

in the Sierra Nevada (southern Spain), each one in a different 10 

x 10km UTM square (Munguira 1989). One suitable area for the 

species is a 40km long and 5–10km wide area on the higher 

altitudes of the Sierra, but the butterfly is only present in small 

scattered patches due to the distribution of its foodplant. A 

thorough study of the whole area has not been made, and all the 

available information comes from two classic sites (Puerto del 

Lobo and Veleta). The altitudinal range of the species is 2600 

to 2900m. 

Population Size: Population numbers seem to be high in the 

type locality (Puerto del Lobo) where at least several hundred 

adults were collected in 1968 (Fernandez-Rubio 1970). In a 

larval survey from this locality we recorded 56 larvae in 0.5ha 

giving a rough estimate of 3000 butterflies for the total population 

in the area. On the other hand, the most endangered colony in 

the Veleta probably only supports c. 100 adults, and in a thorough 

larval survey we only found 12 larvae. 

Habitat and Ecology: The species is restricted to schist screes 

on wind-exposed hill ridges where the vegetation cover is poor. 

The climatic vegetation is a grassland of the Erigeronto frigidi 

– Festuceto clementei series. The foodplant (Vitaliana 

primuliflora) grows in tight cushions 10 to 40cm across and 

5cm high, in three patches in the Veleta (of 25, 900 and 1500 

square meters respectively) and is abundant over an area of 8ha 

in the Puerto del Lobo. 

The female lays eggs singly inside the leaf rosettes of the 

foodplant. The first instars feed on the parenchyma of the 

needle-like leaves and are of a purple colour resembling the 

dead leaves of the plant, among which they usually rest. 

Overwintering takes place inside the plant cushions at the third

Hahitat of A. zullichi: S. Juan, Sra Nevada, 2760 m, May 1987 (photo by M.L. Munguira). 

instar. The last two instars feed on flowers, especially on the 

corolla and developing fruits. Their colour is green with yellow, 

black and white markings, making them difficult to see in their 

colourful environment. They reach their full grown condition 

(fifth instar) at the end of May after 10 months at the larval 

stage. The species is never associated with ants. This fact was 

reported by Chapman (1911) for A. glandon that, like A. 

zullichi, lacks the dorsal nectary organ found in other lycaenids. 

Pupation takes place under stones and adults emerge a month 

later (mid-July). 

Threats: Development of tourist resorts clearly threatens the 

survival of the Veleta colony: redevelopment of the existing ski 

station could easily cause the extinction of this population. 

While the other populations are not threatened by this particular 

development they are unprotected at the moment, making 

future developments in their areas possible. Collecting could 

probably damage the Veleta colony, whose very low population 

numbers make it sensitive to any aggression. Large scale 

collecting should be banned, because the low total population 

numbers of the species make any reduction in numbers dangerous 

for its future. 

Conservation: The declaration of Sierra Nevada as a Man and 

Biosphere Reserve and Natural Park has proved to be ineffective 

in protecting the Veleta area from tourist developments. We 

79 

suggest that the area be declared a National Park. This would 

conserve not only its five endemic butterflies (Munguira and 

Martin 1989), but also the unusual richness of exclusive insects 

and plants of the area. 

The distribution of the food plant should be carefully 

mapped, in order to identify other possible areas where the 

butterfly might be found or introduced if necessary. 

Access to the species' habitat should be limited. Particularly, 

the construction of new roads in the proximity of the main 

population should be controlled if at all permitted. 

Other management practices for habitat improvement are 

not needed because of the climatic character of the plant 

communities in which it lives. Therefore the correct policy for 

the species conservation is to reduce impacts to a minimum, and 

leave the habitat as undisturbed as possible. 

References 

CHAPMAN, T. A. 1911. On the early stages of Latiorina (Lycaena) orbitulus, 

an amyrmecophilous Plebeiid Blue butterfly. Trans, ent. Soc. London, 

1911: 148–159. 

FERNANDEZ-RUBIO, F. 1970. Redescubrimiento de una rara mariposa en 

Sierra Nevada. Nota sobre la captura del Lycaenido: Plebejus glandon 

zullichi Hemming, 1933 (=nevadensis Zullich y Reisser, 1928). Arch. 

Inst. Aclimatacion Almeria 15, 161–167. 

HEATH, J. 1981. Threatened Rhopalocera (butterflies) in Europe. Council of 

Europe, Strasbourg.

KUDRNA, O. 1986. Butterflies of Europe. 8. Aspects of the Conservation of 

Butterflies in Europe. Aula Verlag, Wiesbaden. 

MUNGUIRA, M.L. 1989. Biologia y biogeografia de los licenidos ibericos en 

peligro de extincion (Lepidoptera, Lycaenidae). Ediciones Univ. 

Autonoma de Madrid, Madrid. 

MUNGUIRA, M.L. and MARTIN, J. 1989. Biology and conservation of the 

80 

endangered lycaenid species of Sierra Nevada, Spain. Nota lepid., 12 

(suppl. 1): 16–18. 

VIEDMA, M.G. and GOMEZ, M.R. 1976. Libro Rojo de los lepidopteros 

ibericos. ICONA, Madrid. 

VIEDMA, M.G. and GOMEZ, M.R. 1985. Revision del Libro Rojo de los 

lepidopteros ibericos, ICONA, Monografias no. 42, Madrid.

The Large Copper (Dutch – Grote Vuurvlinder), Lycaena dispar 

Haworth 

Area: Europe to eastern Asia. 

E. DUFFEY 

Cergne House, Church Street, Wadenhoe, Peterborough PE8 5ST, U.K. 

Status and Conservation Interest: Status – L. d. dispar: 

extinct; L. d. batava: rare; L. d. rutila: not threatened at present, 

(Red List). 

A widely distributed wetland species with several described 

subspecies. It was first recorded in 1795 in the Huntingdonshire 

fens, England. This distinctive and very local subspecies, L. d. 

dispar, became extinct in about 1851, at least partly due to 

excessive collecting of the very locally distributed larvae (Duffey 

1968). The Dutch race, L. d. batava, which is very similar to L. 

d. dispar, was introduced to Woodwalton Fen Nature Reserve, 

England, in 1927 and still survives there. 

Taxonomy and Description: L. dispar was first described by 

Haworth (1803) and the much more widely distributed L. d. 

rutila by Werneburg in 1864. The Dutch race, L. d. batava, was 

discovered in 1915 and described by Oberthür in 1923. However, 

Higgins and Hargreaves (1983) combine the extinct British 

race with the Dutch race under the name L. d. dispar, presumably 

because these two races are very difficult to distinguish unless 

a series of each is available. L. d. dispar and L. d. batava are 

generally larger and have more brilliant colours than L. d. 

rutila. (See Bink 1970 for further discussion about the 

subspecies.) 

In the following account the generally used name for the 

Dutch population, L. d. batava, is retained. 

Distribution: The extinct British population appeared to be 

confined to a few fen areas in Huntingdonshire, Cambridgeshire 

and Norfolk, although specimens were occasionally taken 

elsewhere. The Dutch population of L. d. batava is confined to 

a few localities in the provinces of Friesland and Overijssel 

(Bink 1972 and pers. comm. 1991) and has declined in recent 

years so that very few colonies survived in 1991. L. d. rutila is 

widely but locally distributed in Europe (eastern Germany, 

Poland, Baltic countries, Hungary and Russia) but is said to be 

declining in the west of its range due to drainage and reclamation 

of wetlands (Higgins and Hargreaves 1983). 

81 

Population Size: The Dutch population of L. dispar has declined 

markedly in recent years and only one strong colony is known, 

although small numbers occur elsewhere (F.A. Bink, pers. 

comm.). The decline is said to be due to lack of management of 

the habitat which becomes unfavourable as succession proceeds. 

If this situation does not improve, the Dutch population may 

become seriously endangered. 

The introduced L. d. batava in Britain is very insecure as it 

is confined to only about 30ha of about 214ha at Woodwalton 

Fen. It has been shown that it is vulnerable to excessive summer 

flooding (Duffey 1977) and that without protection for the 

larvae the population gradually declines to extinction over a 

number of years. This population has survived since 1927 either 

by protecting a large proportion of the larvae from predators by 

rearing in muslin cages, or else by maintaining a captive 

population so that reintroductions can be made when numbers 

become dangerously low. 

Although the L. d. rutila population has declined in recent 

years, it is still widely distributed and not thought to be in 

danger. 

Habitat and Ecology: Bink (1970,1972,1986) has shown that 

L. d. batava in the Netherlands occurs in an area of overgrown 

peat cuttings where the female lays her eggs on the great water 

dock Rumex hydrolapathum in open reed or sedge beds. At 

Woodwalton Fen in England it has been shown (Duffey 1977) 

that when the foodplants become hidden in taller vegetation the 

ovipositing females often fail to find them. The preferred type 

of Rumex hydrolapathum is often of moderate size, and large 

plants are avoided, especially if growing by open water. 

Nevertheless, in years when the butterflies are numerous, eggs 

may be laid on all size groups. L. d. batava is single-brooded but 

may produce a second brood in warm summers. L. d. rutila is 

normally double-brooded. A large female L. d. batava is capable 

of producing up to 700 eggs but under natural conditions in the 

field, production averages 60 per female (Bink 1986) and, at 

Woodwalton Fen, 114 per female (Duffey 1968). Bink (1986) 

has shown that L. d. batava reaches its highest pupal weight on 

host plants growing at pH 5.5–6.5. Below pH 5.5 the foodplants 

suffer from acidity stress, resulting in a reduction in the protein 

content of the leaves. The insects reared on such plants are

smaller and lay fewer eggs. Nevertheless, there is no convincing 

evidence that ovipositing females select plants of the best 

quality. 

Eggs are laid singly or in lines, usually along a leaf midrib, 

on the underside. However, scattered single eggs are not 

infrequent on the upper side of the leaves of the foodplant. The 

larvae hatch after 7–10 days, and graze the leaf surface, forming 

'windows'. They hibernate as third-instar larvae in dead leaves 

around the base of the water dock plants. During hibernation the 

larvae are unaffected by winter flooding. In the spring, depending 

on weather and the regrowth of the docks, the larvae emerge in 

late April or early May to feed until they pupate about mid-June. 

In July the adults are on the wing, with males usually being the 

first to emerge. When a second brood is recorded in L. d. batava 

the adults are smaller than the first brood and fewer eggs are 

produced. The double broods of L. d. rutila have flight periods 

of May-June and August-September. 

Threats: The decline of L. dispar, especially in central and 

western Europe, is due to the loss of wetland habitats where the 

preferred foodplant grows. In the Netherlands Bink (1986) has 

shown that succession in the fens leads to oligotrophic conditions 

which reduces growth and nutrient quality of the foodplant. 

Fenland reserves in the Netherlands require effective 

management to preserve the best conditions for L. dispar. In 

England the species is at risk because it occurs at only one 

locality and a captive population has to be maintained as an 

insurance against extinction. 

Conservation: Vigorous efforts to protect wetlands are being 

made throughout Europe by the International Union for the 

Conservation of Nature and the International Council for the 

Protection of Birds, but not specifically for invertebrates. The 

British population of L. d. batava is the responsibility of the 

Nature Conservancy Council for England (English Nature) and 

although a permanent wild population cannot survive in the 

82 

small area available without larval protection or reintroductions, 

no serious attempt has yet been made to assess whether other 

suitable areas can be found in the East Anglian wetlands. The 

most secure population in the Netherlands is in the Weerribben 

fen area. This region could provide a nucleus for re-establishment 

in neighbouring fens providing they are maintained in a 

favourable condition. There is no doubt that the best prospects 

for L. d. batava are in the Netherlands, if the appropriate 

authorities could be persuaded to manage suitable fenland areas 

more effectively. 

Throughout most of continental Europe where conservation 

work is effective, the main emphasis is on habitat protection for 

plants and for vertebrates. More attention should be given to 

invertebrates, particularly declining populations such as L. d. 

batava and L. d. rutila. The former may soon be endangered and 

although the latter is not threatened at present, it may become 

so in the future. 

References 

BINK, F.A. 1970. A review of the introductions of Thersamonia dispar Haw. 

(Lep., Lycaenidae) and the speciation problem. Ent. her. Amst. 30: 

179–83. 

BINK, F.A. 1972. Het onderzoek naar de grote vuurvlinder (Lycaena dispar 

batava (Oberthur)) in Nederland (Lep., Lycaenidae). Ent. Ber. Amst. 32: 

225–39. 

BINK, F.A. 1986. Acid stress in Rumex hydrolapathum (Polygonaceae) and 

its influence on the phytophage Lycaena dispar (Lepidoptera: Lycaenidae). 

Oecologia, Berl. 70: 447–451. 

DUFFEY, E. 1968. Ecological studies on the large copper butterfly Lycaena 

dispar Haw. batavus Obth. at Woodwalton Fen National Nature Reserve, 

Huntingdonshire. J. appl. Ecol. 5: 69–96. 

DUFFEY, E. 1977. The re-establishment of the large copper butterfly Lycaena 

dispar batava Obth. on Woodwalton Fen National Nature Reserve, 

Cambridgeshire, England, 1969–73. Biol. Conserv. 12: 143–258. 

HIGGINS, L. and HARGREAVES, B. 1983. The Butterflies of Britain and 

Europe. Collins, London.

Country: England. 

The Adonis Blue, Lysandra bellargus Rottemburg 

Status and Conservation Interest: Status – locally extinct in 

Britain, some rapid decline of other colonies: threatened. 

Intensive surveys of Lysandra (or Polyommatus) bellargus 

in the early 1970s revealed that this local species had declined 

substantially, and had become rare over much of southern 

England, the northern fringe of its European range. Rapid 

losses occurred in the 1950s and the late 1970s and the butterfly 

had become extinct on many sites. These included some which 

continued to support the Chalkhill Blue, Lysandra coridon 

(Poda). In some cases the sites had been destroyed or the 

foodplant eliminated by agricultural practices, but disappearance 

also from areas of unimproved farmland implied that other 

effects – perhaps related to grazing regimes – might be involved. 

Cool weather was also suggested to be a factor inducing 

decline. A study of the ecology of the species (Thomas 1983) 

revealed some unexpected subtleties relevant to the conservation 

and management of open grassland species, and of taxa in 

marginally suitable climatic regimes. 

Distribution: L. bellargus occurs over much of Europe, where 

populations have generally not declined as conspicuously as in 

Britain. In England, it is confined to calcareous grassland in the 

south. 

Population Size: This species forms discrete colonies which 

commonly contain from about 150–850 individuals. Most 

adults do not stray far from the colonies and the populations are 

effectively closed; many are isolated from their nearest neighbour 

colony by tens of kilometres, and no interchange is likely to 

occur between colonies even a kilometre or so apart. 

Population size within a colony can vary greatly. Heath et 

al. (1984) note one Dorset population increasing from fewer 

than 50 adults to more than 60,000 between 1977 and 1982. 

Such variations provide evidence of resilience of populations to 

extinction (Morris and Thomas 1989) but can also reveal 

likelihood of extinction: one colony declined from 3400 adults 

to extinction in only three years. In the past, L. bellargus has 

been recorded from all calcareous formations in southern England. 

Extinctions include several colonies in nature reserves. 

83 

Habitat and Ecology: L. bellargus has two generations each 

year. Eggs laid in late August or September hatch into 

overwintering caterpillars which mature to adults by around 

late May to early June. Offspring of this spring generation 

develop more rapidly to reach the adult stage in only around 2–3 

months. Eggs are laid singly on the foodplant foliage. Larvae 

are day-feeders, and all stages from the second instar onward 

are tended by ants, mainly Myrmica sabuleti and Lasius alienus. 

Larvae and pupae are often buried by ants, which continue to 

tend them. 

Larvae feed only on one foodplant species, the horseshoe 

vetch Hippocrepis comosa, which occurs much further north in 

England than the butterfly does, and is still present in many 

southern areas from where L. bellargus has disappeared. 

Nearly all populations occur on steep south-facing slopes, 

mainly on closely cropped, unimproved pasture. Females prefer 

to oviposit in short turf and in sheltered sun-spots. In sites 

where there is sward of varying heights, oviposition is restricted 

almost entirely to short (1–4 cm) vetch, areas which (because of 

high insolation) are both warm and support numerous ants. 

Threats: Colony extinction has been due mainly to habitat 

change. About one-third of colonies were lost because of loss 

of Hippocrepis due to ploughing or 'agricultural improvement'. 

However, Hippocrepis persists abundantly on some other sites, 

and grazing incidence and intensity are important factors in the 

butterfly's well being. Closely cropped sites were very suitable, 

although very heavy grazing is harmful. Some major extinctions 

coincided with the onset of myxomatosis in the 1950s, which 

resulted in massive loss of rabbits and a resultant decline in 

grazing intensity on much chalk grassland. Reduction in grazing 

intensity appeared to be a major factor leading to extinction, 

and many surviving colonies are on ground grazed by cattle. 

'Improved' or lightly grazed sites have only small populations. 

However, it is not profitable to graze unimproved pasture 

closely, and some hillsides have been abandoned or are grazed 

very irregularly – factors likely to lead to a further decline of L. 

bellargus on such sites. Cessation of grazing can lead to 

development of coarse grasses and 'choking out' of Hippocrepis. 

The butterfly's close association with short swards is evident

from its oviposition behaviour, but the reasons behind this are 

not clear. L. bellargus is common in tall pasture in parts of 

Europe, for example, and dependence on hotter areas on the 

fringe of its range might also be a factor influencing site 

suitability. Warmth might be important both for the butterfly 

itself and for its influence on ants, so that details of their 

association with L. bellargus' early stages might be very subtle. 

Pasture improvement by drilling, herbicides and fertilisers 

remains a threat. Until recently, steeper hillsides have not been 

ploughed, but some are now cultivated, despite their very thin 

soil. 

Conservation: The above ecological observations (Thomas 

1983) emphasise that there is little alternative for practical 

conservation of the Adonis Blue but to manage sites actively for 

its specialised ecological requirements – either on nature reserves 

or on commercial farmland, in which case subsidy agreements 

may be necessary to ensure site security. Merely reserving 

habitat of L. bellargus is not sufficient, although reserves are 

clearly recommended as a basis for management regimes. For 

new reserves, preference may be accorded to ones which 

support more than one colony. Management, probably involving 

rotational grazing or mowing, must seek to ensure the availability 

of short sward on south-facing slopes, and that habitats are not 

overly fragmented or have barriers (such as valleys or areas of 

tall scrub) imposed between them. Much Hippocrepis has been 

converted into a form suitable for L. bellargus during the last 

decade by increased rabbit and stock grazing and, if accessible 

to a founder population, some such sites have been colonised 

successfully. 

84 

Thomas (1983) also suggested that the carrying capacity of 

many sites for L. bellargus could be increased by creating more 

south-facing 'sun-spots', perhaps by using explosives or earthmoving 

equipment, and that such methods could be used to 

construct sites in previously unsuitable areas. 

Another consideration is to introduce L. bellargus to new 

areas, or to sites from which it had earlier disappeared, once 

these have been rendered suitable again by management. Thus, 

L. bellargus was re-introduced in 1981 to Old Winchester Hill 

National Nature Reserve, where it had become extinct in the 

1950s (Thomas 1989). It increased rapidly in numbers and was 

still present after 16 generations (Thomas 1991). Only the 

shortest Hippocrepis were utilised, and the breeding sites 

circulated in pattern with successive paddocks being grazed 

heavily in rotation. 

References 

HEATH, J., POLLARD, E. and THOMAS, J.A. 1984. Alias of Butterflies in 

Britain and Ireland. Viking Books, Harmondsworth. 

MORRIS, M.G. and THOMAS, J.A. 1989. Reestablishment of insect 

populations, with special reference to butterflies, pp. 22–36 In Emmet, 

A.M. and Heath, J. (Eds) The Moths and Butterflies of Great Britain and 

Ireland Vol. 7, Part 1. Harley Books, Great Horkesley. 

THOMAS, J.A. 1983. The ecology and conservation of Lysandra bellargus 

(Lepidoptera: Lycaenidae) in Britain. J. appl. Ecol. 20: 59–83. 

THOMAS, J.A. 1989. Ecological lessons from the re-introduction of 

Lepidoptera. Entomologist 108: 56–68. 

THOMAS, J.A. 1991. Rare species conservation: case studies of European 

butterflies. pp. 149–197 In: Spellerberg, I.F., Goldsmith, F.B. and Morris, 

M.G. (Eds) The Scientific Management of Temperate communities for 

Conservation. Blackwell, Oxford.

Area: England, France, elsewhere in Europe. 

Status and Conservation Interest: Status – M. arion, M. 

alcon, M. teleius: vulnerable (Wells et al. 1983); M. nausithous: 

endangered (Wells et al. 1983); M. teleius: endangered (Red 

List). 

Large Blues were noted by Wells et al. (1983) as 'some of 

the most rapidly declining butterflies in Europe, and probably 

in Asia too'. All are threatened with extinction in Europe, 

because of land use changes (Elmes and Thomas 1992). The 

Large Blue, M. arion (L.), became extinct in Britain in 1979 

despite valiant long-term efforts to save it, and is extinct also in 

the Netherlands, Belgium and parts of northern France. The 

Alcon Large Blue, M. alcon Denis & Schiffermueller, is also 

extinct in many former European localities. The Scarce Large 

Blue, M. teleius Bergstrasser, is apparently declining throughout 

its European and east Palaearctic range, and is extinct in 

Belgium and the Netherlands. The Dusky Large Blue, M. 

nausithous Bergstrasser, is also undergoing local extinctions. 

Elmes and Thomas (1992) assessed the five European species 

as 'Endangered'. Thomas (1984) referred to M. teleius and M. 

nausithous as 'among the world's rarest butterflies' and it has 

been recommended that research into Maculinea biology be 

given top priority in butterfly conservation programmes (Heath 

1981). See also Bálint (this volume), for notes on Carpathian 

taxa. M. arion in Britain is the best documented conservation 

case, and a European subspecies is currently the subject of 

translocations into Britain. This is one of very few such 

international translocation programmes (see also Duffey, this 

volume, on translocation of L. dispar). M. arion has been the 

target of conservation efforts in Britain since the 1920s, latterly 

coordinated by a Large Blue Committee, and full-time ecological 

work on the British M. arion has been pursued since 1972. 

Substantial biological information relevant to conservation of 

some other species has also accumulated (Thomas 1984; Elmes 

and Thomas 1992). 

Maculinea species are protected legally in Britain, Belgium 

and France. 

Taxonomy and Description: The British form of M. arion was 

subspecies eutyphron (Fruhstorfer), and recent successful 

Large Blues, Maculinea spp. 

85 

translocation attempts (Thomas 1989) involve the Swedish M. 

a. arion. Rebel's Large Blue,M. rebeli Hirschke, has sometimes 

been treated as a subspecies of M. alcon, but is distinct 

biologically (Elmes and Thomas 1987). 

Distribution: Maculinea is Palaearctic. M. arion occurs from 

western Europe to southern Siberia, Armenia, Mongolia and 

China; M. teleius occurs from Spain to China and parts of Japan; 

M. nausithous is confined to Europe. The Greater Large Blue, 

M. arionides Staudinger, occurs only in China and Japan and its 

status is unclear. Wells et al. (1983) list it as 'Vulnerable' but 

surveys have not been undertaken in the alpine forest regions it 

frequents. 

Because several species have been studied in detail they are 

treated separately below. Much recent work and synthesis on 

the conservation needs of Maculinea in western Europe is 

included in Elmes and Thomas (1992). 

(i) M. arion 

Population Size: About 90 sites for M. arion were known in the 

southern half of England, mainly concentrated in six areas. The 

colonies were all circumscribed and most comprised a few tens 

to a few hundred adults: the largest probably contained up to 

2000–5000 adults in their 'best' years (Thomas and Emmet 

1989). In general, colonies were isolated from each other and 

closed, as adult dispersal ability is poor. 

Extinction of the colonies in the six main English areas 

occurred as follows: colonies in Northamptonshire died out 

around 1860; the last definite record from south Devon was in 

1906; colonies in Somerset survived until the late 1950s; 

periodic declines in the Cotswolds culminated in the last known 

colony disappearing in 1960–1964; the last colony from the 

Atlantic coast of Devon and Cornwall died out in 1973; and 

those in Dartmoor disappeared in the 1970s. Only two sites 

remained by 1972, and the butterfly finally became extinct in 

Britain in 1979 when the reared female offspring of the last 

remaining female died before any males emerged which could 

mate with them (Thomas 1980).

Habitat and Ecology: M. arion occurs on unimproved grassland 

and is ecologically specialised. It is univoltine and adults are 

relatively short-lived. Females lay eggs on the flower buds of 

wild thyme, Thymus praecox. The first three caterpillar instars 

feed on the thyme flowers and the last (fourth) drops to the 

ground and thereafter depends on the attentions of Myrmica 

ants. As with other Maculinea species, caterpillars are carried 

into Myrmica nests, where they feed on ant eggs, larvae and 

prepupae; this is its major growth stage. Caterpillars hibernate 

and pupate inside ant nests, and the adult M. arion emerges in 

late June to mid July. 

Any species of Myrmica ant will tend the larvae when they 

leave the foodplant, but M. arion is essentially specific to 

Myrmica sabuleti for successful rearing. Two species of 

Myrmica, sabuleti and scabrinodis, are common where Thymus 

grows, but a high density of sabuleti with thyme within 2m of 

their nest entrances is needed for the well-being of M. arion, 

and the size of a Large Blue colony was correlated with the 

number of sabuleti nests present. 

The largest colonies in England covered 10–20ha with a few 

thousand thyme plants and sabuleti nest densities of one every 

l–2m 2 . Small colonies occurred on areas of less than lha if 60% 

of the ground was occupied by sabuleti. Thomas (1991) believed 

that a 'safe' population of 400–1000 M. arion adults could be 

supported on lha under ideal conditions: populations with less 

than 400 adults (reflecting around 2500 usable sabuleti nests) 

might undergo periodic extinctions. Suitable colony sites were 

south-facing areas with short-grazed (to about 2cm) turf so the 

ground could be sun-baked (Thomas 1980,1991). Sward height 

was important: if grazing was removed so that the height 

exceeded about 4cm, sabuleti declined substantially, and 

scabrinodis became relatively more abundant. 

The precise biotope of M. arion varies slightly in different 

parts of Europe, but is always narrow (Thomas 1991). 

Threats: Site alienation through improvement for agriculture 

(by treatment with herbicides or fertilisers, or more direct 

conversion by ploughing or drilling) destroyed about half the 

sites and exterminated the colonies on them. The other sites 

were mainly abandoned for agriculture, with the resultant 

cessation of domestic stock grazing, aided in the 1950s by the 

spread of myxomatosis and removal of rabbit grazing. Although 

M. sabuleti disappeared rapidly from high sward the thyme 

could persist in sward up to 10cm, but it declined in abundance 

and few seedlings became established. Several populations of 

M. arion on nature reserves disappeared because of lack of 

appreciation of the need for grazing management. In hindsight, 

with the knowledge now available, it is very likely that extinction 

in Britain could have been prevented. 

Collecting could have played a role in extinction of some 

small colonies over the years. 

Conservation: With knowledge gained in recent years, through 

the studies of Thomas (1991 and references therein), sites in 

Britain suitable for M. arion have been prepared by prescription 

grazing and M. sabuleti populations have thereby been increased 

86 

substantially on several sites as a basis for attempts to reintroduce 

M. arion. This commenced in the early 1980s, using Swedish 

stock. This was chosen because of its phenological suitability: 

female butterfly emergence had to coincide with the development 

of Thyntus flower buds in Britain for oviposition. It was in fact 

the only suitable stock available since the Thymus-feeding 

races of M. arion are all rare in northern Europe. A trial 

introduction in 1983 was followed by a major release on one site 

in 1986. Seven butterflies emerged in 1984 from the 1983 

release and small numbers were present also in 1985 and 1986. 

About 200 additional larvae were imported in 1986: some 75 

adults emerged in 1987 and 150–200 in 1988. By 1991, it was 

estimated (Thomas 1991) that the main site could support 

around 600–750 adult butterflies. Introductions have now been 

made to other sites and populations will be monitored for 

several years. Changes in farming practices (EEC farming 

subsidies in the 1980s) and return of rabbit grazing have 

rendered some of the former sites again suitable for M. arion. 

The reintroduction has been adjudged successful (Morris and 

Thomas 1989; Elmes and Thomas 1992). 

Before its extinction in 1979, it had already been anticipated 

that reintroduction might be needed, and the work was 

coordinated through the 'Joint Committee for the Conservation 

of the Large Blue', with several commercial firms providing 

sponsorship. The butterfly had already became a familiar emblem 

for much other conservation work on British butterflies. It 

appeared on stamps and received wide media coverage. One 

possible role for a reintroduced colony in due course may be to 

serve as a tourist attraction, with controlled access, thus serving 

as an important avenue for education on butterfly conservation. 

(ii) M. nausithous and M. teleius 

Population Size: The small amount of published information 

suggests that colonies are discrete, closed and generally small 

with no more than a few hundred individuals. 

Habitat and Ecology. These two species coexist on some sites. 

They both breed in marshland, and females oviposit on 

flowerbuds of the same foodplant, great burnet, Sanguisorba 

officinalis. Because of the observed coexistence, it had been 

assumed generally that the ecology of the two species was very 

similar (Thomas 1984) but each has specialised individual 

requirements (Elmes and Thomas 1987). As in other species of 

Maculinea, larvae feed on the plant for the first three instars and 

then feed on Myrmica brood in ant nests. Larval sizes of the two 

species differ markedly at the time of leaving the plant: an 

average larva of M. nausithous weighs 1.15mg, and that of M. 

teleius, 4.32mg. The principle host ant species differ (Thomas 

et al. 1989): for M. nausithous it is Myrmica rubra and for M. 

teleius, M. scabrinodis, and this segregation helps to explain 

why populations of Maculinea species have always been 

localised in areas where Myrmica ants and foodplants are 

abundant. In areas where the butterflies coexist (mainly wetlands 

around bogs or in swampy fields) high densities of both host 

Myrmica species occur.

Threats: Habitat alienation has been attributed to development 

and drainage of wetlands (Wells et al. 1983); even if the main 

reedy areas survive, the drier fringes (where breeding occurs) 

may be lost. All known sites for both species in the Rhone 

Valley were damaged by reservoir construction in 1981. More 

subtle site changes, affecting the abundance and well-being of 

the ants, are also likely to occur: extinction occurs if the density 

of the particular ant becomes too low and this is occurring 

because traditional methods of hay and reed cutting are being 

abandoned (Thomas 1991). Myrmica scabrinodis is abundant 

only in short vegetation, and M. teleius is relatively common 

there. M. nausithous gradually replaces it as succession proceeds 

and the vegetation becomes taller (4–7 years since 

establishment). It may later disappear. 

Conservation: Reserve establishment to safeguard habitat is a 

priority for both species, with management to conserve 

plagioclimax conditions. Thomas (1991) notes that high densities 

of both M. scabrinodis and Sanguisorba can be maintained in 

moist hay meadows cut once a year in parts of France and 

Poland, and there is little doubt that habitats suitable for both 

species can be created easily. Translocation may well be a 

practical conservation option in the absence of natural 

colonisation of new habitats. Large populations of the butterflies 

can be supported on small land areas, and vegetation cutting on 

a 3-year rotation may be sufficient to maintain site suitability. 

(iii) M. alcon and M. rebeli 

Population Size: Confusion in the past over the relative status 

of these two taxa means that much of the historical and detailed 

distribution of each is not wholly clear. Both species occur in 

small colonies: many colonies of M. alcon contain fewer than 

100 individuals. 

Habitat and Ecology. The two species can coexist, but M. 

rebeli can occur at much higher altitudes than M. alcon and 

extends to 1000m in the Swiss alps (Elmes and Thomas 1987). 

Eggs of both species are laid on Gentiana – those of M. alcon 

on G. pneumonanthe and G. asclepiadea, and of M. rebeli on G. 

cruciata and G. germanica. Only large, vigorous plants are 

suitable, and the general life history is similar to that of other 

Maculinea. The main host ant of M. alcon is Myrmica ruginodis, 

and for M. rebeli, M. schencki. 

Threats: Major threats are various forms of habitat destruction, 

predominantly through agricultural changes, but M. alcon is 

87 

also threatened to some extent from urbanisation. Elmes and 

Thomas (1987) cite specific threats for Swiss populations. 

Conservation: As for other Maculinea, habitats can be created 

by particular mowing or grazing regimes, and reservation with 

correct management is needed. A survey of the species' range 

to detect the best habitats is needed, especially to detail the 

distribution of M. rebeli in the alps. Recent work by Elmes et al. 

(1991a, b) has led to greater understanding of the interaction 

between the M. rebeli caterpillar and its ant hosts. 

References 

ELMES, G.W. and THOMAS, J.A. 1987. Le genre Maculinea. pp. 354–368 

In: Les Papillons de Jour et leurs Biotopes. Ligue Suisse pour la Protection 

de la Nature, Basle. 

ELMES, G.W. and THOMAS, J.A. 1992. Complexity of species conservation 

in managed habitats: interaction between Maculinea butterflies and their 

ant hosts. Biodiv. and Conserv. 1: 155–169. 

ELMES, G.W., THOMAS, J.A. and WARDLAW, J.C. 1991(a). Larvae of 

Maculinea rebeli, a large-blue butterfly, and their Myrmica host ants: 

wild adoption and behaviour in ant-nests. J. Zool. 223: 447–460. 

ELMES, G.W., WARDLAW, J.C. and THOMAS, J.A. 1991(b). Larvae of 

Maculinea rebeli, a large-blue butterfly and their Myrmica host ants: 

patterns of caterpillar growth and survival. J. Zool. 224: 79–92. 

HEATH, J. 1981. Threatened Rhopalocera in Europe. Council of Europe, 

Strasbourg. 

MORRIS, M.G. and THOMAS, J.A. 1989. Re-establishment of insect 

populations, with special reference to butterflies. In: Emmet, A.M. and 

Heath, J. (Eds) The Moths and Butterflies of Great Britain and Ireland. 

Vol. 7, Part 1. Harley Books, Great Horkesley, pp. 22–36. 

THOMAS, J.A. 1980. Why did the large blue become extinct in Britain? Oryx 

15: 243–247. 

THOMAS, J.A. 1984. The behaviour and habitat requirements of Maculinea 

nausithous (the dusky large blue butterfly) and M. teleius (the scarce large 

blue) in France. Biol. Conserv. 28: 325–347. 

THOMAS, J.A. 1989. The return of the Large Blue butterfly. Brit. Wildlife 1: 

2–13. 


butterflies. In: Spellerberg, I.F., Goldsmith, F.B. and Morris, M.G. (Eds) 

The Scientific Management of Temperate Communities for Conservation, 

Blackwell, Oxford, pp. 149–197. 

THOMAS, J. A., ELMES, G. W., WARDLAW, J.C. and WOYCIECHOWSKI, 

M. 1989. Host specificity among Maculinea butterflies in Myrmica ant 

nests. Oecologia 79: 452–457. 

THOMAS, J.A. and EMMET, A.M. 1989. Maculinea. In: Emmet, A.M. and 

Heath, J. (Eds) The Moths and Butterflies of Great Britain and Ireland. 

Vol. 7, Part 1. Harley Books, Great Horkesley, pp 171–175. 


Red Data Book. IUCN, Gland.

Polyommatus humedasae (Toso & Balletto) 

E. BALLETTO 

Dipàrtimento di Biologia Animale, Università di Torino, V. Accademia Albertina 17, Torino, Italy –10123 

Country: Italy (northwest). 


This is one of the most well known, endemic species living 

in a very rare and endangered type of habitat, known to be 

occupied by a number of other restricted insects (Magistretti 

and Ruffo 1959, 1960) and plants (Peyronnel 1964). Even 

though one or two additional biotopes will probably be 

discovered in the future it is doubtful whether those populations 

will be as well represented as they are in the species' type 

locality. 

Taxonomy and Description: Polyommatus humedasae is a 

species of the subgtnus Agrodiaetus that has been described in 

comparatively recent times (Toso and Balletto 1976). Even 

though doubts on its species-level identity were initially 

expressed by some authors (Higgins and Riley 1983), these 

were finally dispelled (Higgins and Hargreaves 1983) by the 

study of its haploid chromosome complement (n = 38: Troiano 

et al. 1979). 

The most closely related species, judging by external 

morphology, is the Greek Polyommatus aroaniensis (Brown 

1976), an endemic of the mountains of Peloponnesos (Aroania 

Ore, i.e. Mt. Chelmos) and characterised by a lower number in 

its haploid chromosome complement (n = 15–16). 

Distribution: Polyommatus humedasae lives in a particularly 

small area of a dozen hectares by Pondel (= Pont d'Ael) in the 

Val d'Aosta region (northwest Italy). The average altitude of 

the biotope is 1100m and it lies in the montane vegetational 

zone. 

Population Size: The population structure is probably closed. 

Even though the biotope currently known may not be the only 

one where this species lives, there is but scanty evidence that 

this is the case, mostly based on a single and old museum 

specimen and 'entomological gossip'. 

Surveys of adult specimens demonstrated a reasonably high 

population vigour with an average density of 11 specimens/ 

hectare and at least 110 animals instantaneously present at the 

site over a period of one month. Although no studies focused on 

88 

the estimation of the total population number are available, it 

may be represented by a few thousand adults. 

Habitat and Ecology: Polyommatus humedasae lives in xerothermophilous 

environments of the Festucetalia vallesiacae 

(Stipo-Poioncarniolocae) vegetational formations; the detailed 

association was never investigated to a sufficient detail at the 

phyto-sociological level. The geological substrate is represented 

by ophiolitiferous calco-schists of a Jurassic age (Balletto et al. 

1982). As these schists are very fissured and fragmented, they 

play a fundamental role in providing a xeromorphic character 

to this biotope, irrespective of the orientation, which is to the 

northeast. 

Adults are often concentrated on what appear to be the 

remnants of some now abandoned alfalfa fields, whose flowers 

represent a good nectar source for all 'blues' of the genus 

Polyommatus. No male territorial behaviour has been described 

or observed. 

Eggs are laid singly on the lower surface of the leaves of 

Onobrychys montana Lam. & DC. In a laboratory study in 

1982, oviposition started in mid-August and peaked between 

August 15–21. Hatching starts in the last half of September 

(Manino et al. 1987). Newly-hatched larvae, lmm long, feed 

for a few weeks and then shelter for hibernation. Hibernation 

takes place at the first instar, in the litter at the plant base. 

Feeding starts again in mid-April, and the second moult takes 

place in the first half of May. Moults are carried out within the 

litter at the base of the foodplant. Each moult takes a few days 

to perform. Each of the successive instars lasts about 15 days. 

Full-grown larvae (mid-June) are green and 15mm long. 

Pupation also takes place in the litter and lasts about 20–25 

days. 

No relationship with ants has, so far, been described. 

Adults fly from mid-July to mid-August. The butterfly 

community of Pondel is particularly rich and includes about 30 

other species flying synchronously in the same biotope (Balletto 

etal. 1982). 

Apart from Medicago sativa, adult nectar sources include 

Sedum ochroleucum spp. montanum and Onobrychys montana, 

the larval food plant. Actual and potential nectar sources are 

very abundant throughout the period when imagines are flying

and nectar does not seem to represent a limiting factor of 

population size. 

Threats: Potential threats are of a rather varied and contrasting 

nature. Since the only biotope known for this species is situated 

in the montane ecological zone its climax is represented by 

woodland. No adult specimens of Polyommatus humedasae, 

however, have been observed in the woods on any of five 

surveys carried out in different years. A few small alfalfa fields 

that used to be cultivated in this biotope until about ten years 

ago have now been abandoned. It seems most likely, therefore, 

that if natural succession was to continue towards the climax 

vegetation the biotope would disappear, probably together with 

the animal. As with many other butterflies, in fact, this appears 

to be an ecotonal species, taking advantage of the intermediate 

stages of the recolonisation of sun-exposed landslides. It seems 

unlikely, however, that in the present situation this 'blue' will 

colonise new biotopes by a natural process. 

As with many other endemics, another threat is overcollecting. 

Even though the exact location of the site was not 

divulged in the original description in order to prevent such a 

threat, collectors soon discovered the place and many of them 

can often be met in a single day on the biotope. 

Conservation: The biotope lies a few hundred metres from the 

outer edge of the Parco Nazionale del Gran Paradiso, Italy's 

largest protected area and one of the most strictly regulated. It 

seems therefore obvious that a first measure for the conservation 

of Polyommatus humedasae would be to extend the northern 

boundary of the Gran Paradiso National Park to include this 

biotope (Balletto and Kudrna 1985). The reason why this step 

has not yet been taken is apparently that the general area is 

heavily under pressure by the conflicting interests of the 

inhabitants of the valleys of Cogne and Aosta, who do not want 

to renounce rights for agricultural and chamois-stalking 

89 

practices. Site management, even though not yet needed, may 

become necessary in the future, to interrupt the natural trend of 

the vegetation towards a closed woodland. 

References 

BALLETTO, E., BARBERIS, G. and TOSO, G.G. 1982. Aspetti dell'ecologia 

dei Lepidotteri ropaloceri nei consorzi erbacei delle Alpi italiane. Collana 

'Promozione della qualita dell'ambiente': Quaderni sulla 'struttura 

delle Zoocenosi terrestri' II.2: 11–96. 

BALLETTO, E. and KUDRNA, 0.1985. Some aspects of the conservation of 

butterflies in Italy, with recommendations for a future strategy. Boll. Soc. 

ent. ital. 117 (1–3): 39–59. 

BROWN, J. 1976. Notes regarding previously undescribed European taxa of 

the genera Agrodiaetus Hubner, 1822 and Polyommatus Kluk, 1801. 

Entomologist's Gaz. 27: 27–83. 

HIGGINS, L.G. and HARGREAVES, B. 1983. The butterflies of Britain and 

Europe. Collins, London. 

HIGGINS, L.G. and RILEY, N.D. 1983. A field guide to the butterflies of 

Britain and Europe, 5th ed., 384pp. Collins, London. 

MAGISTRETTI, M. and RUFFO, S. 1959. Primo contributo alla conoscenza 

della fauna delle oasi xerotermiche prealpine. Mem. Mus. civico St. nat. 

Verona 7: 99–125. 

MAGISTRETTI, M. and RUFFO, S. 1960. Secondo contributo alla conoscenza 

della fauna delle oasi xerotermiche prealpine. Mem. Mus. civico St. nat. 

Verona 8: 223–240. 

MANINO, Z., LEIGHEB, G., CAMERON-CURRY, P. and CAMERON- 

CURRY, V. 1987. Descrizione degli stadi preimmaginali di Agrodiaetus 

humedasaeToso & Balletto, 1976 (Lepidoptera, Lycaenidae). Boll. Mus. 

regionale Sci. nat. Torino 5(1): 97–101. 

PEYRONNEL, B. 1964. Escursione della Societa Botanica Italiana in Val 

d'Aosta. Giorn. bot. ital. 71: 183–196. 

TOSO, G.G. and BALLETTO, E. 1976. Una nuova specie del genere 

Agrodiaetus Hubn. (Lepidoptera: Lycaenidae). Annali Mus. civico Sty. 

nat. G. Doria, Genova 81: 124–130. 

TROIANO, G., BALLETTO, E. and TOSO, G.G. 1979. The karyotype of 

Agrodiaetus humedasae Toso & Balletto (Lepidoptera: Lycaenidae). 

Boll. Soc. ent. ital. 111(7–8): 141–143.

Polyommatus galloi (Balletto & Toso) 

E. BALLETTO 

Dipàrtimento di Biologia Animale, Università di Torino, V. Accademia Albertina 17, Torino, Italy — 10123 

Country: Italy (south) 

Status and Conservation Interest: Status – rare. 

This is one of the few relatively well known species living 

in a very rare and endangered type of habitat, known to be 

inhabited by a number of exclusive insects and plants (Gavioli 

1936; Avena and Bruno 1975). 

Taxonomy and Description: Polyommatus galloi is another 

species of the subgenus Agrodiaetus Hübner described in 

comparatively recent times (Balletto and Toso 1979). Its specieslevel 

distinction from Polyommatus ripartii (Freyer) (south 

France, northwest Italy) was confirmed by the study of the 

haploid chromosome complement (n = 66: Troiano 1979, 

instead of n = 90 as in P. ripartii: Lesse 1960). 

The most closely related species, from both external 

morphology and haploid chromosome complement, is 

apparently Polyommatus demavendi (Pfeiffer), distributed from 

Turkey to north Iran and characterised by a slightly larger 

number of chromosomes (n = 70–71: Lesse 1960). 

Distribution: Polyommatus galloi lives over an area of several 

square kilometres shared between the southern Italian regions 

of Calabria and Lucania, on Mt Pollino and on the Orsomarso 

mountain range. The altitude of biotopes inhabited by this 

species ranges from 1800 to 2200m. 

Population Size: The population structure is relatively closed 

in that specimens at high altitudes on Mt Pollino are unlikely to 

be able to reach the Orsomarso chain, and vice versa. Each of 

the two main sets of biotopes contain an apparently small 

number of metapopulations. Even though the biotopes currently 

known may not be the only ones where this species lives, there 

is currently no evidence that this is the case. 

Surveys of adult specimens demonstrated a reasonably high 

population density with an average 6–7 specimens/hectare 

simultaneously present at the same site over a period of about 

one month. On the whole this species may be represented by 

several thousand adults/year. 

90 

Habitat and Ecology: Polyommatus galloi lives in xeromorphic 

environments, at the highest elevations reached in the southern 

Italian Apennines by the vegetational formations of the Alliance 

Bromion erecti. The particular Sub-Alliance represented here 

(Seslerio-Xerobromenion apennium) is considered somewhat 

transitional to the Seslerietalia apenninae present at the high 

elevations of the central Apennines in otherwise similar 

ecological conditions (Avena and Bruno 1975). The geological 

substrate is represented by Lower Cretaceous grey limestones, 

or by Jurassic calcitutites and limestones (Balletto et al. 1982). 

Adults are generally concentrated on the flowerheads of the 

nectar sources. No male territorial behaviour has been described 

or observed. No particular study has been devoted to the 

reproductive biology of this species but females have been 

observed to lay eggs on the lower surface of the leaves of 

Onobrychis caputgalli. (L.) Lam. Adults fly from mid-July to 

mid-August. 

No relationship with ants has, so far, been described or 

observed. 

Apart from Lavandula angustifolia ssp. angustifolia, a good 

nectar source for most xerophilous species of Polyommatus, 

adults feed on the flowerheads of Sedum album, Picris 

hieracioides, Echinops ritro, Cirsium afrum and Onobrychis 

caputgalli. The latter also represents the larval foodplant. 

Actual and potential nectar sources are very abundant throughout 

the period when imagines are flying and nectar is not likely to 

represent a limiting factor for population size. 

Threats: A study conducted in 1977 and again in 1980–81 

(Balletto et al. 1982) has shown that this species is particularly 

sensitive to the adverse influence of grazing. Even though the 

effect of sheep overgrazing is particularly severe, even slight 

grazing by domestic stock can result in considerably diminished 

population densities, which soon fall well below 50% of normal. 

A potential threat is over-collecting although for the time being 

this is not an issue. 

Conservation: The most important thing would be to reverse 

the negative effects of sheep grazing.

Mt. Pollino is included in a Natural Park, one of the largest 

in southern Italy, but unfortunately not one having particularly 

strict regulations with regard to stock rearing practices (Balletto 

and Kudrna 1985). The main reason why, up until now, steps in 

this direction have not yet been taken, is that the general area is 

under pressure from the conflicting interests of the local 

shepherds, who do not want to renounce, or even reduce their 

grazing rights. 

In the Mediterranean area, where summer rainfall is episodic, 

overgrazing causes the soil moisture to evaporate very quickly 

and in the long run can easily elicit severe transformations of the 

habitat of this species. 

Site management, even though not yet needed, may become 

necessary on a local basis, to interrupt the natural trend of the 

vegetation towards a closed woodland. 

91 

References 

AVENA.G. and BRUNO, F. 1975. Lineamenti della vegetazione del Masiccio 

del Pollino (Appennino calabro-lucano). Not. Fitosoc. 10: 131–153, 

Roma. 

BALLETTO, E. and KUDRNA, O.1985. Some aspects of the conservation of 

butterflies in Italy, with recommendations for a future strategy. Boll. Soc. 

ent. ital. 117(1–3): 39–59. 

BALLETTO, E. and TOSO, G.G. 1979. On a new species of Agrodiaetus 

(Lycaenidae) from Southern Italy. Nota lepid. 2(1/2): 13–25. 

BALLETTO, E., TOSO, G.G. and BARBERIS, G. 1982. Le comunita di 

Lepidotteri ropaloceri nei consorzi erbacei dell'Appennino. Collan 

'Promozione della qualita dell'ambiente': Quaderni sulla 'struttura 

delle Zoocenosi terrestri' II.l: 77–143. 

GAVIOLI, O.1936. Ricerche sulla distribuzione altimetrica della vegetazione 

in Italia: 3°. Limiti altimetrici delle formazioni vegetali nel gruppo del 

Pollino (Appennino calabro-lucano). N. Giorn. bot. ital., n.s. 43. 

LESSE, H. (de) 1960. Les nombres de chromosomes dans le groupe 

d'Agrodiaetus ripartii Freyer (Lepidoptera Lycaenidae). Revue fr. Ent. 

27(3): 240–264. 

TROIANO, G. 1979. Karyotype. In: Balletto E., & Toso G.G., On a new 

species of Agrodiaetus (Lycaenidae) from Southern Italy. Nota lepid. 2(1/ 

2): 13–25.

The Sierra Nevada Blue, Polyommatus golgus (Hübner) 


Deparlamento de Biologia (Zoologia), Facultad des Ciencìas, Universidad Autonoma de Madrid, Madrid, Spain 

Country: Spain (southeast). 

Status and Conservation Interest: Status – vulnerable (listed 

as endangered (Viedma and Gomez 1985) or vulnerable (Heath 

1981)). 

This species, also known as the Nina de Sierra Nevada, and 

Agriades zullichi are flagship representatives of a very peculiar 

and endangered habitat. They live in an area with 53 endemic 

plant species and at least 50 exclusive insects (including three 

endangered Orthoptera). Several authors have supported the 

idea of protecting the area as a National Park (Gómez-Campo 

1987; Munguira and Martin 1989a), but part has been developed 

as a ski station, creating a conflict between conservation and 

development in an area where job creation and investment are 

badly needed. 

P. golgus is listed in Appendix II of the Berne Convention 

of which Spain is a Contracting Partner. 

Taxonomy and Description: Hübner described golgus as a 

distinct species, but it was considered by many authors to be a 

subspecies of P. dorylas (Denis & Schiffermuller) (Agenjo 

1947). Lesse (1960) studied the chromosome numbers of the 

dorylas group and stated that golgus had a lower number (n = 

c. 131–134) than the other species. This has been used by other 

authors (Gomez and Arroyo 1981) as evidence in favour of it 

being a true species. 

Another closely related species, P. nivescens (Keferstein), 

with n = c. 190–191 (one of the highest chromosome numbers 

in the animal kingdom) is endemic to Spain and also lives in the 

Sierra Nevada at lower altitudes; and an ecologically similar 

species (P. atlantica (Elwes)) is endemic to the Atlas Mountains 

in Morocco. 

Distribution: Restricted to four 10x10km UTM squares in the 

Sierra Nevada (Granada Province, southeast Spain). It lives at 

heights ranging from 2500 to 2900m in the oromediterranean 

and crioromediterranean zones (the Mediterranean equivalents 

of subalpine and alpine zones). 

Population Size: Population structure is not closed and small 

numbers of adults can be seen all around the suitable area. 

92 

Larval surveys revealed a very low population density, but the 

populations occupied a very extensive area. Adult males are 

concentrated in wet places where they defend perching sites 

against other conspecific males in a behaviour similar to lek 

behaviour (Munguira and Martin 1989b). The population studied 

probably had several thousand adults, but no accurate estimate 

has been made. Records of the species from 1838 to 1986 show 

it as abundant in the habitat to which it is restricted. 

Habitat and Ecology: Present in grassland communities 

growing among dwarf junipers (Genisto baeticae-Junipereto 

nanae) and at higher altitudes on climax grasslands (Erigeronto 

frigidi-Festuceto clementei) growing among schist screes 

(Munguira 1989; Munguira and Martin 1989b). Due to the very 

harsh weather conditions (with snow cover for nine months) the 

plants growing in the area are strong rooted perennials with 

aerial parts growing close to the ground. One of these plants is 

Anthyllis vulneraria arundana, the butterfly's foodplant, which 

is also endemic to the area. 

P. golgus, in cop., Sierra Nevada, July 1985 (photo by M. Munguira).

Figure 2. Habitat of P. golgus, Sierra Nevada, Veleta, September 1986 (M. Munguira). 

Eggs are laid singly on the upperside of curled leaves of the 

plant, whose parenchyma is used as food by the caterpillars. 

The species overwinters in its third larval instar. Larvae are 

regularly tended by Tapinoma nigerrimum ants, that often have 

their nests close to the foodplants. Pupation takes place in June, 

after five larval instars, in the ground near the foodplant. Adults 

fly in July in a single generation. 

Nectar sources include Arenaria tetraguetra, Silene 

rupestris, Jasione amethystina and Hieracium pilosella. The 

flowers of these plants and many others are abundant in the area 

during the flight period, and therefore nectar does not seem to 

be a limiting factor for the species. 

Threats: Tourism-related development is the major threat for 

the butterfly. A metalled road crosses one of the areas and a ski 

station actually exists in part of the butterfly's habitat. 

Redevelopment of the ski station poses the greatest threat, 

involving construction of new roads and buildings and reshaping 

of slopes for ski courses. These would have great impact on the 

butterfly's habitat and make other further impacts (pollution, 

refuse accumulation) more likely. 

Conservation: The climax character of the plant communities 

in which the butterfly lives is a great advantage for conservation 

93 

practice: the only necessary action is to protect the area and try 

to reduce impacts to their minimum. The declaration of a 

National Park in the area seems necessary, at least to protect one 

of the richest mountain ranges in Europe as far as endemic 

plants is concerned (Blanco 1989), as well as the five endemic 

butterflies (Parnassius apollo nevadensis Oberthur, Erebia 

hispaniaBut\cT,Polyommatusgolgus,Agriadeszullichi,Aricia 

morronensis ramburi) and other insects. New developments in 

the Monachil and Dilar valleys should be stopped and any jobcreating 

alternatives studied so that conservation does not 

necessarily involve refusal to develop a depressed area. Sierra 

Nevada was declared in 1986 a reserve of the Man and Biosphere 

(MAB) project, and in 1989 it became a Natural Park, but this 

conservation status has not prevented the area from being 

damaged. 

The inclusion of P. golgus in the appendices of the Berne 

Convention probably assures its protection in the long term but 

no short-term actions have been undertaken, and the 

developments currently taking place have not been stopped 

despite legislation against them. 

This is one of the most obvious cases in which the importance 

of education should be stressed. Unlike other National Parks, in 

which the conservation interests are geological formations, 

remarkable forests or vertebrate faunas, the principal values of

Sierra Nevada are its small endemic invertebrates and plants. Its 

high altitude and geographical location makes it the southernmost 

limit for many northern and alpine plants and animals. Its 

populations are therefore of key importance in conserving the 

genetic diversity of these species. It is therefore necessary to 

enhance awareness among the general public of the great 

scientific and conservation interest of this apparently unattractive 

area. 

References 

AGENJO, R. 1947. Catálogo ordenador de los lepidópteros de España. 

Sexagesima novena familia. Graellsia 5. 

BLANCO, E. 1989. Areas y enclaves de interés botánico en España (Flora 

silvestre y vegetación). Ecologia 3: 7–21. 

G6MEZ, M.R. and ARROYO, M. 1981. Catálogo sistemático de los 

94 

lepidópteros ibéricos. Ministerio de Agricultura y Pesca, Madrid. 

G6MEZ-CAMP0, C. (Ed.) 1987. Libra rojo de especies vegetates amenazadas 

de España peninsular e Islas Baleares. ICONA, Madrid. 



LESSE, H. De. 1960. Spéciation et variation chromosomique chez les 

lépidoptères rhopaloceres. Ann. Sci. Nat. Zool. Biol. Anim. 2: 1–223. 

MUNGUIRA, M.L. 1989. Biologiaybiogeografla de los licénidos ibéiricos en 

peligro de extincién (Lepidoptera, Lycaenidae). Ediciones Universidad 

Autónoma de Madrid, Madrid. 

MUNGUIRA, M.L. and MARTIN, J.1989a. Biology and conservation of the 

endangered lycaenid species of Sierra Nevada, Spain. Nota lepid. 12 

(suppl. 1): 16–18. 

MUNGUIRA, M.L. and MARTIN, J. 1989b. Paralelismo en la biologia de tres 

especies taxonómicamente próximas y ecológicamente diferenciadas del 

género Lysandra: L. dorylas, L. nivescens y L. golgus (Lepidoptera, 

Lycaenidae). Ecologia 3: 331–352. 

VIEDMA, M.G. and GOMEZ, M.R. 1985. Revisión del Libro Rojo de los 

lepidópteros ibéricos. ICONA, Madrid.

Le Faux-Cuivre smaragdin*, Tomares ballus F. 

Henri A. DESCIMON 

Laboratoire de Systématique evolutive, Université de Provence, 3 place Victor Hugo, 13331 Marseille Cedex 3, France 

Country: Western Mediterranean (France). 

Status and Conservation Interest: Status – not thought to be 

threatened at the present time. 

This species may be a test one, facing both the impetus of 

economic development in the Mediterranean region, especially 

in the coastal belt where its habitats occur, and the abandonment 

of traditional land occupation, which provides its chief haunts. 

Taxonomy and Description: The Genus Tomares Rambur is 

limited to the western Palaearctic region. Three of the six 

species recognized are distributed around the Mediterranean 

basin. T. ballus occurs as two subspecies: the nominal one, from 

north Africa and southern Spain, and catalonica Sagarra, from 

Spanish Catalonia and southeast France. The latter is 

characterised by a yellowish-green hue of the hindwing underside 

while the nominal subspecies is bluish-green. 

Distribution: T. ballus is widely distributed in Maghreb and 

the southern half of Spain (Gomez-Bustillo and Fernandez- 

Rubio 1974). In France, it is limited to a nucleus disjunct from 

the main area in the littoral region of the Var and Alpes- 

Maritimes, with some recently discovered colonies in Bouchesdu-Rhone. 

Although still widespread in the Var department and 

able to colonise available habitats rapidly, the species has 

undergone some restriction of its geographical range in France. 

It no longer exists in the Alpes Maritimes, where it was present 

until the 1970s at Cannes and Vallauris. It has also been 

eliminated in its classical localities close to Hyeres. 

Population Size: In southern Spain, the species is often very 

abundant (Jordano et al. 1990a) and widespread. In France, the 

colonies are patchy and unstable, although they appear to be 

dense (several hundreds/ha, according to the author's visual 

estimations). However, the species displays obvious colonising 

abilities, and starts thriving in new available habitats within a 

very few years. Episodical colonisation of localities outside its 

normal breeding range is occasionally reported (de Laever 

1954). 

* A name contrived recently by G.C. Luquet. 

95 

Habitat and Ecology. Throughout its area of distribution, the 

species is confined to open landscapes, either steppe vegetation 

formation, or forest clearings of a sufficient size, where the 

vegetation covering is sparse. In France, it is confined to 

calcareous substrates. In this country, there are two main kinds 

of habitat: forest clearings of the live oak-Aleppo pine forest; 

and semi-neglected orchards of olive trees and vines, where by 

far the more abundant colonies are to be found. 

Foodplants differ according to the geographical region but 

always belong to Fabaceae: in northern Africa, mainly Erophaca 

baetica (Powell in Oberthur 1910); in southern SpainAstragalus 

lusitanicus (Sierra Morena) and Medicago polymorpha 

(Guadalquivir Valley) (Jordano et al. 1990a, 1990b); in France, 

either Bonjeana hirsuta (Chapman 1904) or predominantly 

Anthyllis tetraphylla, which is also used in northern Morocco 

(Descimon and Nel 1986). B. hirsuta is chiefly confined to the 

woodland clearings habitat, while A. tetraphylla used to thrive 

in cultivated areas where weeding was regular enough to 

prevent scrub invasion but sparse enough to allow annuals, 

therophytes and hemicryptophytes to grow. 

Feeding behaviour appears to be highly opportunistic and 

depends upon the compatibility between the vegetative cycle of 

the plant and the developmental stage of the butterfly. Fine 

adjustment of the latter to the constraints imposed by the 

various host-plant cycles is evident. 

In France, eggs are laid singly or in very small groups, while 

in Sierra Morena, they are laid in clumps of several tens. The 

caterpillars prefer to consume flowers and seedpods, but can 

also attack leaves. In France, the duration of the caterpillar stage 

is 50–60 days, depending on yearly climate. Pupation occurs 

under stones present in the habitat. Pupal duration is about 10 

months. 

Threats: In France, near the seashore, the only cause of 

disappearance of T. ballus is the rampant urbanisation of the 

strip of land bordering the crowded beaches of the Mediterranean 

sea. On the mainland, improved cultivation techniques quickly 

cause the disappearance of'weeds' such as Anthyllis tetraphylla. 

In woodland areas, the dramatically increased frequency of

fires (of course due to urbanisation) is also a cause of the 

extinction of some colonies, in particular those close to Hyeres. 

However, in this case, the return of the butterfly following 

resprouting of its foodplants can be rapid if nearby colonies are 

preserved. In all zones, nibbling ('moth-eating' in French!) of 

the countryside by private houses and their gardens specifically 

attacks the chief haunt of T. ballus: sunny, terraced, sparsely 

cultivated landscapes. In the zones not yet struck by building 

speculation (and they become scarce, since, under the southern 

sun, every piece of land is under threat), land abandonment 

causes the open therophytic landscapes to be invaded by scrub 

and then continuous pine forest with no more suitable habitats 

available to T. ballus. 

The low yield of traditional Mediterranean cultivation 

practices does not now allow for continuing a landscape 

maintenance regime which, in the past, provided a high floristic 

and faunistic diversity. 

Collecting does not play any role in the decline of the 

species: even in the limited areas where overcollecting was 

exercised by collectors when the habitats remained ecologically 

preserved, a decrease in the species' abundance was scarcely 

observed. 

Conservation: At present, no legal measures for protection are 

taken in France. Should such measures be taken, they would 

probably be limited to prohibiting collection. Such provisions 

prove extremely inefficient – especially in a country of Latin 

and Mediterranean tradition – and can even be 

counterproductive: professional entomologists waste much time 

in satisfying administrative formalities; honest amateur 

entomologists are discouraged and give up butterfly watching; 

dishonest collectors ignore the laws completely; and 

unscrupulous dealers continue to enjoy the increased prices of 

'black market' specimens. 

The problem of conservation of a large, semi-continuous 

and widespread population of Tomares ballus in southern 

France is closely linked to the general problem of overall 

96 

conservation of biological diversity and even of human living 

quality in this region. Up to the present time, in spite of some 

pungent attacks through urbanisation of the seashore region, 

southern France has been relatively safe from the 'economic 

development syndrome' which rages in northern Europe and 

involves intensified exploitation of profit-earning zones, 

abandonment of other zones, overurbanisation and general 

pollution (Descimon 1990). 

At the present time, perhaps the most effective strategy to 

conserve T. ballus habitat would be an educational effort 

directed towards private landowners in 'moth-eaten' 

countryside: such efforts could maintain parts of the traditional 

Meditteranean landscape, such as olive orchards, with a low 

level of 'cleaning'. 

In Spain, where the species is more abundant, and Maghreb 

(Thomas and Mallorie 1985), the problems are less serious but 

probably basically the same. 

References 

DE LAEVER, E. 1954. Tomares ballus Fabr. dans les Basses Alpes. R. fr. 

Lépidoptérologie 14: 165. 

DESCIMON, H. and NEL, J. 1986. Tomares ballus F. est-il une espéce 

vulnérable en France? Alexanor 14: 219–231. 

DESCIMON, H. 1990. Pourquoi y a-t-il moins de papillons aujourd'hui? 

Insectes 77: 6–10. 

GOMEZ-BUSTILLO, M.R. and FERNANDEZ-RUBIO, F. 1974. Mariposas 

de la Peninsula iberica. Ropaloceros (II). Ministerio de Agriculture, 

Madrid. 258pp. 

JORDANO, D., HAEGER, J.F. and RODRIGUEZ, J. 1990a. The effect of 

seed production by Tomares ballus (Lepidoptera: Lycaenidae) on 

Astragalus lusitanicus (Fabaceae): determinants of differences among 

patches. Oikos 57: 250–256. 

JORDANO, D., HAEGER, J.F. and RODRIGUEZ, J. 1990b. The life-history 

of Tomares ballus (Fabricius, 1787) (Lepidoptera: Lycaenidae): phenology 

and host plant use in southern Spain. J. Res. Lepid. 28: 112–122. 

THOMAS, CD. and MALLORIE, H.C. 1985. Rarity, species richness and 

conservation: butterflies of the Atlas mountains in Morocco. Biol. Conserv. 

33: 95–117.

Country: U.K. 

The Silver-studded Blue, Plebejus argus L. 

CD. THOMAS 

Centre for Population Biology, Imperial College at Silwood Park, Ascot, Berks SL5 7PY, U.K. 

Status and Conservation Interest: Status – P. a. masseyi: 

extinct; P. a. cretaceus: not threatened but few colonies survive; 

P. a. caernensis: not threatened; P. a. argus: not threatened but 

few colonies survive. 

P. argus has virtually disappeared from four-fifths of its 

former British range (Heath et al. 1984; Thomas and Lewington 

1991). Studies have been undertaken in north Wales (CD. 

Thomas 1985a,b, 1991; Thomas and Harrison 1992), Devon 

(Read 1985), Suffolk (Ravenscroft 1986,1987,1990), Norfolk 

(N.Armour-Chelu, pers. comm.), and Dorset (J.A. Thomas 

1991). P. argus is regarded as an important indicator of vigorous 

heathland habitats, and has been severely affected by habitat 

fragmentation, and by the cessation of traditional management 

which maintains the heathland successions required by this 

insect. Many of the British populations of P. argus are already 

on reserves, and English Nature, the Countryside Council for 

Wales, The Royal Society for the Protection of Birds (RSPB), 

the National Trust, the County Naturalist Trusts, and Local 

Councils actively foster this species. 

Taxonomy and Description: P. argus has formed several local 

races in Britain, and although these probably do not warrant the 

formal status of subspecies (CD. Thomas 1985a), they do 

provide extra conservation interest. Race masseyi was found on 

the mosslands at the southern margin of the Lake District, and 

had blue females (de Worms 1949). Scottish populations were 

similar. Race caernensis is restricted to limestone grasslands in 

north Wales. They are very small, and have blue females. 

Heathland populations in north Wales look similar to caernensis, 

but are slightly larger, and a mossland population in north 

Wales is similar but distinctly larger (CD. Thomas 1985a). 

Race cretaceus occupies calcareous grasslands in southern 

England; it is large with relatively pale males. Race argus is 

found elsewhere. 

Distribution: Europe and temperate Asia. In Britain, P. argus 

survives in Wales and southern England but is extinct in 

Scotland. 

97 

Population Size: Race cretaceus has suffered a reduction in 

range and is only recorded now at Portland Bill (Dorset), where 

the remaining colonies are in good health (Warren 1986; 

Thomas and Lewington 1991). Race caernensis is thriving, 

despite its restricted range in north Wales. In 1983, the peak 

emergence was about 250,000 in 10 colonies on the Great 

Orme, and about 30,000 in 16 colonies in the Dulas valley: the 

total emergence was perhaps three times greater (CD. Thomas 

1985b). Numbers were similar at both localities in 1990. An 

introduced population was established near Prestatyn in 1983, 

and was vigorous by 1990 (Thomas and Harrison 1992). 

Heathland populations in north Wales are mostly smaller than 

those on the limestone, but one large population (about 40,000 

at peak in 1983) occurs on the RSPB Reserve of South Stack 

Range. P. argus did not decline on heathlands in north Wales 

between 1983 and 1990 (Thomas and Harrison 1992). The 

mossland population in north Wales contained about 5000 

adults at peak in 1983, and was similar in 1990. 

Race argus populations occur predominantly on heathland, 

where it is mostly a story of continuing attrition. Only one 

population is left in the Midlands, and just a handful survive in 

East Anglia and south Wales. On the Sandlings of Suffolk, six 

of nine colonies contained more than 500 individuals at peak in 

1986, yet only one or possibly two still contained this number 

by 1990, and two colonies were on the verge of extinction: the 

total 1990 Sandlings population was estimated to be just 23% 

of that in 1986, flying over 53% of the 1986 area (Ravenscroft 

1986,1987,1990, pers. comm.). Similarly, in Devon P. argus 

is practically confined to one area of heathland (Read 1985). It 

is only in the Poole Basin, New Forest, and on the west Surrey 

heaths that race argus remains numerous (Thomas and 

Lewington 1991). A few populations survive on sand dunes in 

Cornwall. 

Habitat and Ecology: P. argus occurs on heathlands, calcareous 

grassland, sand dunes and mossland, and the larvae feed on a 

wide variety of hosts in the Leguminosae, Ericaceae and 

Cistaceae (CD. Thomas 1985a,b; Jordano et al. in press). 

Despite the apparent breadth of biotopes and host plants, P. 

argus is local because it has other specialised requirements. In 

the north, particularly, P. argus is restricted to warm

microclimates: it occurs on sites that contain plenty of hot, bare 

ground, mostly on south-facing slopes. The eggs are laid at the 

margins of bare ground and vegetation, and the larvae feed on 

the tender growth of their host plants. Further south, vegetation 

edges seem to be less important, but relatively short successional 

vegetation is still favoured (Ravenscroft 1990; Thomas and 

Lewington 1991; Jordano et al. in press, N. Armour-Chelu, 

pers. cotnm.). 

Eggs are laid in midsummer, in response to ants (N. Armour- 

Chelu, pers. comm.), and these overwinter and hatch in spring. 

The hatchling larvae are attractive to, and are usually picked-up 

by, the workers of Lasius niger or Lasius alienus, and they are 

taken back to the nest (Ravenscroft 1990; Jordano and CD. 

Thomas in press). Quite what happens to the first instar larvae 

in the nest is unknown, but the later instars feed on foliage above 

ground, constantly tended by Lasius (CD. Thomas 1985a; 

Ravenscroft 1990; Jordano and CD. Thomas in press). The ants 

are pugnacious in defence of larvae and, if provoked, will pick 

up and retreat with any that are small enough to carry. Pupae can 

be found under stones, where they are always tended by Lasius, 

and as often as not they are inside Lasius nests. The emerging 

adults are tended by Lasius. 

In different populations, P. argus eggs are significantly 

associated with different species of plants or combinations of 

plants and microhabitats (CD. Thomas 1985a; Read 1985; 

Ravenscroft 1990; J. A. Thomas 1991; Jordano et al. in press; N. 

Armour-Chelu, pers. comm.): indeed, there are almost as many 

different plant associations as populations studied. In contrast, 

four studies all show an association of P. argus with Lasius ants 

(e.g. Ravenscroft 1990; N. Armour-Chelu, pers. comm.). 

Suitable conditions occur in Britain where there is a coincidence 

of young host plant foliage, warm conditions, and adequate 

densities of Lasius. 

Threats: P. argus is threatened by biotope loss and 

fragmentation, caused by modern agriculture, afforestation, 

urbanisation, etc. For example, in one of the best remaining 

regions for heathland, the Poole Basin in Dorset, only 14.6% of 

the original heathland survived to 1978: 62% of the remaining 

fragments are of

nearly 50 years in a series of limestone habitat patches in the 

Dulas valley in north Wales. 


I thank N. Armour-Chelu, D. Jordano and N. Ravenscroft for 

access to unpublished material. 

References 

DE WORMS, C.M.G. 1949. An account of some forms of Plebejus argus L. 

Rpt. Raven Ent. Nat. Hist. Soc. 1949: 28–30. 

HEATH, J., POLLARD, E. and THOMAS, J.A. 1984. Atlas of Butterflies in 

Britain and Ireland. Viking, Harmondsworth. 

JORDANO, D., RODRIGUEZ, J., THOMAS, CD. and FERNANDEZ 

HAEGER, J. (in press). The distribution and density of a lycaenid 

butterfly in relation to Lasius ants. Oecologia. 

JORDANO, D. and THOMAS, CD. (in press). Specificity of an ant-lycaenid 

interaction. Oecologia. 

RAVENSCROFT, N.O.M. 1986. An investigation into the distribution and 

ecology of the silver-studded blue butterfly (Plebejus argus L.) in Suffolk: 

an interim report. Suffolk Trust for Nature Conservation, Ipswich. 

RAVENSCROFT, N.O.M. 1987. Plebejus argus colony population estimates 

and changes in status in Suffolk 1985–1986. Suffolk Trust for Nature 

99 

Conservation, Ipswich. 

RAVENSCROFT, N.O.M. 1990. The ecology and conservation of the silverstudded 

blue butterfly Plebejus argus L. on the Sandlings of East Anglia, 

England. Biol. Conserv. 53: 21–36. 

READ, M. 1985. The silver-studded blue conservation report. Msc thesis, 

Imperial College. 

THOMAS, CD. 1985a. Specialisations and polyphagy of Plebejus argus 

(Lepidoptera: Lycaenidae) in North Wales. Ecol. Ent. 10: 325–340. 

THOMAS, CD. 1985b. The status and conservation of the butterfly Plebejus 

argus L. (Lepidoptera: Lycaenidae) in North West Britain. Biol. Conserv. 

33: 29–51. 

THOMAS, CD. 1991. Spatial and temporal variability in abutterfly population. 

Oecologia 87: 577–580. 

THOMAS, CD. and HARRISON, S. 1992. Spatial dynamics of a patchily 

distributed butterfly species. J. Anim. Ecol. 61: 


butterflies. In: I.F. Spellerberg, F.B. Goldsmith and M.G. Morris (Eds) 

The scientific management of temperate communities for conservation. 

BESSymp. 31: 149–197. 

THOMAS, J. and LEWINGTON, R. 1991. The Butterflies of Britain and 

Ireland. Dorling Kindersley, London. 

WARREN, M.S. 1986. The status of the cretaceus race of the silver-studded 

blue butterfly, Plebejus argus L., on the Isle of Portland. Proc. Dorset 

Nat. Hist. Arch. Soc. 108: 153–155. 

WEBB, N.R. and HASKINS, L.E. 1980. A ecological survey of heathlands in 

the Poole Basin, Dorset, England, in 1978. Biol. Conserv. 17: 281–296.

The Zephyr Blue, Plebejus pylaon (Fischer-Waldheim) 


Departamento de Biologia (Zoologia), Facultad de Ciencias, Universidad Autonoma de Madrid, Madrid, Spain 

Area: Spain, Switzerland, Italy, Hungary, Bulgaria, Romania, 

Yugoslavia, Albania, Greece, Turkey, Russia and Asia Minor. 

Status and Conservation Interest: Status – Hungary: 

endangered; Spain, Switzerland, Italy: vulnerable; other 

countries: indeterminate, (see Heath 1981; Munguira et al. 

1991). 

The species, known also as the 'nina del astragalo', is 

present over a wide area, but always local and isolated in several 

populations some of which form different subspecies (species 

for some authors, Kudrna 1986, Bálint and Kertész 1990). 

Nevertheless, young stages of all these different forms are 

dependent on Astragalus plants and live on dry steppe-like 

habitats. This makes the butterfly rare throughout its range. The 

species' biotopes are at present being altered as a result of the 

change from traditional land uses to more aggressive agricultural 

practices, or are being abandoned. The conservation of the 

species cannot be achieved by simply protecting the sites, due 

to the seral character of P. pylaon habitats, and would need 

certain management practices in order to maintain desirable 

ecological features. 

Taxonomy and Description: The taxonomic level of the forms 

under consideration is not clear at the moment and needs further 

study. Plebejus vogelii Oberthür and particularly P. martini 

Allard, are true species for most of the authors due to their 

morphological and ecological features (Higgins and Hargreaves 

1983, Bálint and Kertész 1990). Nevertheless, the taxa included 

under the name pylaon are treated by some authors as distinct 

species. Kudrna (1986) for example splits the group into four 

different species in Europe: hesperica Rambur,pylaon, sephirus 

and trappi Verity. Bálint and Kertész (op. cit.) group the 

European forms in these same four taxa, but they split each 

group into several biogeographical sub-groups without naming 

them as separate species. Within these so-called species a large 

number of subspecies have been described, but at least with the 

Spanish ones (group hesperica above) we consider all the 

subspecies as synonyms (Munguira 1989), and this certainly 

may be the case in other groups. 

100 

Distribution: The species is present from central Spain to the 

surroundings of the Baikal Lake. In Spain it lives in the central 

Plains, Iberian Mountains and near Sierra Nevada in Granada 

Province in a total of 37 UTM squares of 10 x 10km (Munguira 

1989, Munguira and Martin 1989). In Italy and Switzerland 

subspecies trappi is also very local. In the last country it is only 

found in 16 UTM squares (Gonseth 1987) in the Valais. It is also 

very local in Hungary (Bálint and Kertész 1990) and only 

present in 22 UTM squares in Yugoslavia (Jaksic 1988). 

Population Size: In Spain the populations are local but the 

number of adults is high. Many plants can have up to 20 eggs, 

giving adult yields in each population of several thousand 

individuals. After overwintering, larval numbers may reach the 

level of 2–3 per plant. Due to the local character of the 

populations, some localities are sensitive to extinction events. 

In Sierra de Alfcar, the type locality of Spanish hesperica, the 

butterfly has never been collected since its description in 1839. 

In central Spain a colony was partially destroyed by a limestone 

Plebejus pylaon, male, Camporeal (photo by M. Munguira).

pit (Gomez-Bustillo 1981) but at present the species is still on 

the site although in reduced numbers. Other localities still 

support important numbers of the species but are vulnerable to 

human impacts. 

Habitat and Ecology: The species always occurs on disturbed 

Quercus rotundifolia forests (encinares) in the Iberian Peninsula. 

Serai communities are maintained on dirt road verges, quarries, 

or on land disturbed by overgrazing. The geological substrate 

is clay or limestone. The soil is exposed in approximately 75% 

of the surface of the places where the foodplant (Astragalus 

alopecuroides) grows. The vegetation is formed by strong 

rooted perennials and shrubs specialised to live in habitats with 

very poor soil conditions. Altitude ranges from 400 to 1400m, 

and rainfall is always scarce (between 400 and 500mm per year) 

in every locality studied. 

Eggs are laid on the leaflets of the foodplant in May or early 

June. After a week the first instar larvae begin to feed on the 

parenchyma of the leaves, leaving characteristic eye-like damage 

on the plant. At the beginning of July the larvae undergo the 

second moult and build a silken refuge on the base of the 

foodplant in which they spend all the rest of the summer and the 

winter (aestivating and overwintering). In the following March 

the larvae begin to feed on the young shoots of the plant and 

develop quickly until reaching the full-grown condition (fifth 

Habitat of P. pylaon, Camporeal, May 1991 (photo by M. Munguira). 

101 

instar) in March–April. They pupate in the ground and emerge 

as adults 20 to 30 days later. 

The last two larval instars are invariably tended by ants 

belonging to several species at each locality. We have found 

attending ants of eight different species of the genera Formica, 

Camponotus, Crematogaster and Plagiolepis. Larvae have 

dorsal nectary organs (Newcomer's gland) and tentacle organs. 

Tentacles are displayed when the larva is in danger, and the 

attending ants become excited and attack any potential enemy 

when this happens. This behaviour probably defends larvae 

against parasitoids and in fact, despite having reared almost 40 

larvae to adults, we never obtained parasitoids from the species. 

Relationships with ants are facultative, and the whole life cycle 

can be completed without the ants' presence. 

The butterfly is present in all the localities where we have 

seen its foodplant. Nevertheless, the plant also exists in southern 

France where pylaon has never been recorded. In the high 

altitude localities of the Iberian Mountains the foodplant is the 

Iberian-African A. turolensis (Sheldon 1913). The biology and 

ecology of subspecies trappi and sephirus are similar (SBN 

1987; Balint and Kertész 1990). The foodplant of trappi is 

Astragalus exscapus, and the butterfly appears a month later 

than in Spain, making the whole life cycle a month late. The 

habitat in the Valais consists of dry grasslands on rocky limestone 

slopes.

Threats: Most lowland populations are endangered by 

urbanisation, because they live on flat areas suitable as building 

sites. In the Tagus Valley the species lives in surrounding small 

olive groves, and modern agriculture practices can also destroy 

this habitat, reducing headlands and hedges to a minimum as a 

consequence of management intensification. While small fields 

maximise the border area, increases in the grove area are 

reducing the headlands and hedges. 

Some of the highland populations are threatened by road 

developments (SBN 1987), or because they are potential areas 

for pine plantations in the Spanish Iberian Mountains (Munguira 

1989). 

Most of the areas in which the butterfly lives have seral 

vegetation communities. Rabbit or sheep grazing is probably 

necessary to maintain the proper habitat in Spain (Munguira 

1989). On the other hand in Switzerland, sheep overgrazing 

probably endangers the foodplant suitability (SBN 1987). 

Conservation: The species is present at the moment in several 

Natural and National Parks: Tshatkal Reserve (Uzbekistan), 

Galicica National Park (southern Yugoslavia), Gran Paradiso 

National Park (Val d'Aosta, Italy) and El Regajal Reserve 

(Madrid, Spain). This by no means assures proper protection 

for the butterflies, because National Parks and Reserves are 

normally created to protect the larger mammals and birds. At 

least with these reserves, the four European subspecies are 

protected in theory. The real conservation of the species can 

only be achieved when proper management is maintained on at 

least several suitable hectares on each reserve. This management 

involves continuing with traditional land uses such as extensive 

grazing. Nevertheless, the exact grazing regime and the required 

sheep and goat stocking rates are not known properly, and 

require further study. 

The Regajal Reserve, one of the Spanish localities where 

pylaon is present, illustrates the conflicts between insect 

conservation and development. The area was suggested as a 

Lepidoptera Reserve in the mid-seventies (Viedma and Gomez- 

Bustillo 1976). Later on a project to build a roundabout crossing 

the area was conceived, receiving the criticism of amateur and 

professional entomologists alike. The site is not only valuable 

102 

for its rarities, but also because it has always been a collecting 

site for most of Spain's leading entomologists. The motorway 

was finally built in 1989, crossing the reserve through its very 

centre. The impact of the motorway is certainly serious: for 

example the rivers have changed their regime, causing several 

serious floods in 1990. The damage to insect populations will 

never be known because a suitable impact assessment was not 

done before building the motorway. After all these events, the 

area is receiving a protected status by regional authorities, and 

will probably be the first Spanish Reserve created mainly to 

protect Lepidoptera. 

References 

BÁLINT, Z. and KERTÉSZ, A. 1990. A survey of the subgenus Plebejides 

(Sauter, 1968) – preliminary revision. Linn. Belg. 12: 190–224. 

GOMEZ-BUSTILLO, M.R. 1981. Protection of Lepidoptera in Spain. Beih. 

Veroff. Natursch. Landschaft. Bad.-Wurt. 21: 67–72. 

GONSETH, Y. 1987. Atlas de distribution des papillons diurnes de Suisse 

(Lepidoptera Rhopalocera). CSCF, Neuchatel. 



HIGGINS, L.G. and HARGREAVES, B. 1983. The Butterflies of Britain and 

Europe. Collins, London. 

JAKSIC, P. 1988. Provisional distribution maps of the butterflies of Yugoslavia 

(Lepidoptera, Rhopalocera). Societas Entomologica Jugoslavia, Zagreb. 

KUDRNA, 0.1986. Butterflies of Europe. Vol. 8. Aspects of the Conservation 

of Butterflies in Europe. Aula-Verlag, Wiesbaden. 

MUNGUIRA, M.L. 1989. Biologia y biogeografia de los licenidos ibericos en 

peligro de extincion (Lepidoptea, Lycaenidae). Ediciones Universidad 

Autonoma de Madrid, Madrid, Spain. 

MUNGUIRA, M.L. and MARTIN, J. 1989. Biology and conservation of the 

endangered lycaenid species of Sierra Nevada, Spain. Nota lepid. 12 

(Suppl. 1): 16–18. 

MUNGUIRA, M.L., MARTIN, J. and REY, J.M. 1991. Use of UTM maps to 

detect endangered lycaenid species in the Iberian Peninsula. Nota lepid. 

Suppl. 2: 45–55. 

SBN. 1987. Tagfalter und ihre Lebensraume. Schweizerischer Bund fur 

Naturschutz, Basel. 

SHELDON.W.G. 1913. Lepidoptera at Albarracin in May and June, 1913. 

Entomologist 46: 283–9, 309–13, 328–32. 

VIEDMA, M.G. and GOMEZ-BUSTILLO, M.R. 1976. Libro Rojo de los 

lepidopteros ibericos. ICONA, Madrid.

The Pannonian Zephyr Blue, Plebejus sephirus kovacsi Szabó 

Z. BÁLINT 

Zoological Department, Hungarian Natural History Museum, Baross utca 13, Budapest, H–1008, Hungary 

Country: Hungary. 

Status and Conservation Interest: Status – rare. 

The butterfly was found in 1949, in the close vicinity of 

Budapest and it was thought that this population was the only 

Hungarian one. Several new colonies were discovered in the 

1980s, but all were within a 30km radius of the first one. These 

colonies represent the most western as well as the most northern 

occurrences of the species. Very recently a new population has 

been found in northeast Hungary showing continuity towards 

the Transylvanian (Romania) colonies. 

Taxonomy and Description: P. sephirus is a member of a 

western palaearctic group of lycaenids of central Asian 

xeromontane origin (Báiint and Kertész 1990a; Bálint 1991a). 

Several described subspecies of sephirus exist in the Carpathian 

Basin (central Hungary: ssp. kovacsi Szabó, 1954 (= foticus 

Szabó, 1956); Banat (Voivodina): ssp. uhryki Rebel, 1911; 

Transylvania: ssp. proximus Szabó, 1954), some which are 

most probably synonyms of the nominate subspecies sephirus 

Frivaldszky, 1835 occurring in the Balkans. 

Distribution: Very restricted in two isolated parts of the 

country. Eight small colonies can be found north of Budapest, 

one in Tokaj, northeast Hungary, found very recently (1991) by 

Professor Varga. 

Population Size: All colonies are strongly isolated, and the 

structure of the populations is closed. There is no possibility of 

interchange of adults even between the closest populations, 

whilst they are separated by heavily cultivated agricultural 

regions or human settlements. One small population which is 

divided by a double rail track was studied by the capturerecapture 

method. The results show that the adults are strongly 

stenochorous, with the life history of the species closely 

associated with the larval foodplant. 

Two of the larger populations have been monitored during 

the last two years. All known colonies in the vicinity of 

Budapest have been estimated by counts of adults during the 

last three years. The major two colonies contained about 700–900 

103 

butterflies. Three smaller colonies had fewer than 100 

individuals. 

Habitat and Ecology: Colonies are strongly confined to the 

association Astragalo-Festucetum rupicolae, a typical habitat 

of forest steppe on loose calciferous soil. There is one generation 

each year with a very short flight period of adults from the 

middle of May to the beginning of June. Adults are active only 

when the air temperature is above 25°C, and the wind is not 

strong. Eggs are laid singly on the larval foodplant, Astragalus 

exscapus L. Caterpillars hatch after about ten days. After a short 

feeding period they retreat to ant chambers at the base of the 

plants. The diapause lasts from about the end of June (because 

of the long hot summer) to the middle of the following spring, 

about the middle of April. All older instars are tended by ants, 

so Plebejus sephirus is a steadily myrmecophilous species. The 

following ant species are involved: Tetramorium (caespitum 

gr.); Formica pratensis; Camponotus aethiops; and Lasius 

(alienus gr.) (Fiedler 1991). Caterpillars pupate in the upper 

end of the ant chambers. The pupal stage lasts about ten days 

and the pupae are also tended by ants. Adults of P. sephirus take 

nectar mainly from Dianthus giganteiformis Borb. ssp. 

pontederae Kern., endemic to the Pannonian region. 

Threats: The present scattered Hungarian distribution of the 

species shows that current distribution is mainly the result of 

human activity: the loose soil of the central Carpathian Basin 

has been cultivated from early historical times. The major 

recent threats have been: (1) the establishment of new housing 

estates; (2) illegal rubbish heaps; (3) motor-cross activities; (4) 

afforestation; (5) natural succession; (6) overcollecting. 

Conservation: The first discovered habitat of P. sephirus, 

namely Somlyóhegy near Fót, has been a nature reserve since 

1953. Two larger areas, where the strongest colonies exist, will 

be protected in the very near future. The butterfly and its larval 

hostplant are protected by Hungarian law (Bálint and Kertész 

1990b). A paper on the conservation and management of P. 

sephirus was ordered by the Hungarian authorities and was

drawn up (Bálint 1991b), but there is no financial support to 

undertake the practical measures suggested to ensure good 

management. The recommendations for the P. sephirus 

populations in the Budapest area are as follows: 

1. Protection of all colonies during the flight period of the 

species; 

2. Monitoring of all populations; 

3. Two important habitats with strong P. sephirus colonies 

could be attached to the Somlyohegy Nature Reserve, which 

could be a special reserve for P. sephirus, where it is 

necessary to arrest natural succession and negative human 

influences; 

4. Stronger publicity for conservationist activities. 

104 

References 

BÁLINT, Zs. 1991a. (A xeromontane lycaenid butterfly: Plebejus pylaon 

(F.W., 1832) and its relatives), I. Jan. Pann. Muz. Évk. 35: 33–69 (in 

Hungarian). 

BÁUNT, ZS. 1991b. (The ecology and conservation of Plebejus sephirus 

(Frivaldszky, 1835) in Hungary). Budapest (in Hungarian). 

BALINT, Zs. and KERTÉSZ, A. 1990a. A survey of the subgenus Plebejides 

Sauter, 1968 – preliminary revision. Linneana Belgica 12: 190–224. 

BÁLINT, Zs. and KERTÉSZ, A. 1990b. The conservation of Plebejus sephirus 

(Frivaldszky, 1835) in Hungary. Linneana Belgica 12: 254–273. 

FIEDLER, K. 1991. Systematic, evolutionary, and ecological implications of 

myrmecophily within the Lycaenidae (Insecta: Lepidoptera: 

Papilionoidea). Bonn. Zool. Monog. 31, 210 pp.

The threatened lycaenids of the Carpathian Basin, east-central 

Europe 

Zsolt BÁLINT 

Hungarian Natural History Museum, Zoological Department, Budapest, Baross utca 13. H–1008 

The following species are notable Lycaenidae which occur in 

the region. Each is treated as a discrete account and these 17 

taxa collectively indicate the main lycaenid conservation needs 

in the Carpathian Basin, and the eastern adjacent region of 

Romania. The references are given for the whole account rather 

than each species separately. 

Aricia macedonica isskeutzi Balogh 



A subspecies confined to a very small region of northern 

Hungary. 

Taxonomy and Description: Close to Aricia allous (Geyer) 

(see Geiger 1988) but differing in some minor genitalic and 

superficial characters (Varga 1968). 

Distribution: Aricia macedonica Verity distributed in the 

Balkans is an allopatric group of taxa, distinct from that of A. 

artaxerxes (Fabricius) (Britain), A. allous (Alps and central 

Europe) and A. inhonora (Jachontov) (Scandinavia, Russia). 

The subspecies issekutzi is the most northern and western 

representative of the macedonica-complcx, and it can be found 

in the Karst of Torna (northern Hungary and southern Slovakia) 

and in the Bükk Mountains (northern Hungary). 

Population Size and Status: Not known. 

Habitat and Ecology: Xerothermophilous, univoltine (from 

late June to early August). Stenotopic, inhabiting clearings of 

forest steppes, rocky and dry grasslands. Caterpillar steadily 

my rmecophilous (Fiedler 1991). Larval foodplants Helianthemum 

ovatum (Viv.) Dunal and Geranium sanguineum L. 

Threats: The populations are strongly isolated and threatened 

by intensification of grassland management, afforestation, 

urbanisation and tourism. 

105 

Conservation: Most of the Hungarian habitats can be found in 

the territories of Aggtelek National Park and Bükk National 

Park. Thorough ecological studies are necessary to continue the 

work of Varga (1968). 

Aricia eumedon (Esper) 



A protected species, which is only known from northern 

Hungary. The example recorded from Transdanubia (W. 

Hungary), collected at the beginning of the century, has never 

been confirmed. 

Taxonomy and Description: All the central European 

populations are identical (see Geiger 1988). 

Distribution: Transpalearctic. The species is widely distributed 

in the brook or river valleys of the Carpathians (Slovakia), but 

it is absent from the central part of this range. 


Habitat and Ecology: Hygrophilous, univoltine (mainly July). 

Stenotopic, the colonies can be found in wetlands along water 

courses with luxuriant vegetation of the larval foodplant 

Geranium palustre (L.). Caterpillars are steadily 

myrmecophilous (Fiedler 1991). The main nectar source of the 

adults is also the above-mentioned purple flowered Geranium 

species. 

Threats: Natural succession, overcollecting. 

Conservation: All the Hungarian colonies, which constitute a 

single metapopulation, are in a small mountain stream valley 

situated in the Aggtelek Natural Park. Patrolling of the known 

sites is needed along with a study of the ecosystem of the stream 

valley.

Cupido osiris (Meigen) 


Status and Conservation Interest: Status – endangered. 

A protected species. Only three Hungarian sites are known, 

and these are the most northern permanent populations of this 

species. 

Taxonomy and Description: The Hungarian populations do 

not differ from the central European ones (see Geiger 1988). 

Distribution: Mediterranean. Very rare in Slovakia, several 

populations in Transylvanian Basin, Romania (Bálint 1985a). 


Habitat and Ecology: Xerothermophilous, stenotopic, probably 

bivoltine. The Hungarian habitats include abandoned orchards 

and vineyards. Caterpillars are steadily myrmecophilous (Fiedler 

1991). Larval foodplants are Colutea arborescens (L.) and 

Onobrychis viciaefolia Scop.. 

Threats: Only one of the three populations has been monitored 

in the last years, most probably the most endangered one. 

Factors giving concern include: earthworks as a result of openpit 

mining; air pollution from a lime factory; and the recultivation 

of abandoned orchards and vineyards. 

Conservation: The conservation of the species in Hungary has 

not yet been achieved. Patrolling and study of the known 

populations, together with the monitoring of suitable habitats 

for new sites are urgently required. 

Jolana iolas (Ochsenheimer) 



A protected species, J. iolas was discovered in Hungary. 

The habitat of the original specimens was recently destroyed by 

urbanisation. Many known colonies have disappeared in recent 

decades. 

Taxonomy and Description: The Hungarian populations do 

not differ from the Central European ones (Geiger 1988). 

Distribution: Mediterranean. The northern limit of the species 

range can be found in Hungary. The southern Slovakian records 

have not been confirmed recently (Kulfan and Kulfan 1991). 


106 

Habitat and Ecology: Xerothermophilous, univoltine 

(May-July), stenotopic. The known habitats are mainly 

abandoned vineyards or Pannonian karst oak-scrubs. Caterpillars 

are presumed to be moderately myrmecophilous (Fiedler 1991) 

and the larval foodplant is Colutea arborescens (L.) (Uhrik- 

Mészáros, 1948; Szabó, 1956). Adults fly around the bushes of 

the larval foodplant, which is also their main nectar source. 

Threats: Afforestation (planting of Pinus nigra Arn.), 

urbanisation (a lot of habitats have been built upon) and 

overcollecting. 

Conservation: Some colonies can be found in Landscape 

Protection Areas. Their habitats need full protection. The 

ecology of the species must be studied. 

Maculinea alcon (Denis & Schiffermuller) 



The Hungarian populations have almost totally disappeared 

and only a very few colonies remain in the western part of the 

country. 

Taxonomy and Description: The Hungarian populations are 

identical with the European ones (see Geiger 1988). 

Distribution: West Palaearctic. The most easterly populations 

can be found in Hungary and in Romania (Transylvania). 


Habitat and Ecology: Hygrophilous, univoltine (late 

July-beginning of August), stenotopic. Larval hostplant is 

Gentiana pneumonanthe (L.). Caterpillars are obligately 

myrmecophilous (Fiedler 1991). Other details concerning the 

Hungarian populations are not known. 

Threats: Wetland drainage. 

Conservation: All the remaining populations must be 

discovered and studied. Some already known colonies can be 

found in Landscape Protection Areas. The whole ecosystem 

(incorporating the effects of traditional agricultural practices) 

of the habitats must be protected.

Maculinea nausithous (Bergstrasser) 



A protected species. Serious loss of habitat through wetland 

drainage and intensification of grassland management has 

taken place. 

Taxonomy and Description: The Hungarian populations are 

identical with the European ones (see Geiger 1988). 

Distribution: West Palaearctic. The most easterly populations 

can be found in the western part of the country (Transdanubia). 


Habitat and Ecology: Hygrophilous, univoltine (late 

July-beginning of August), presumed stenotopic. Larval 

hostplant is Sanguisorba officinalis(L.). Caterpillars are 

obligately myrmecophilous (Fiedler 1991). Other details 

concerning the Hungarian populations are not known. 

Threats: Much of the suitable habitat for this species has 

become the victim of wetland drainage and intensification of 

grassland management. 

Conservation: All the remaining populations must be 

documented and measured and the ecology of the species in 

Hungary must be studied. Some populations are in the Landscape 

Protection Area and in the Ferto-tó National Park, where the 

whole ecosystem must be protected effectively. 

Maculinea sevastos limitanea Bálint 

Country: Romania. 


Endemic to the eastern Carpathians. The populations are 

very scattered. 

Taxonomy and Description: Resembles Maculinea alcon, but 

the wing shape is more expanded, and the underside darker 

brown somewhat similar to M. nausithous (Bálint 1986). 

Distribution: An east Mediterranean species, its most western 

and northern populations can be found in Transylvania 

(Romania) (see Kudrna 1986). 


Habitat and Ecology: Xerothermophilous, univoltine (July). 

Caterpillars are presumed to be obligately myrmecophilous. 

107 

The larval hostplant is Gentiana crutiata (L.). The species 

occurs in the same habitats as Parnassius apollo transylvanicus 

(Schweitzer, 1912), and Polyommatus dory las magna (Bálint 

1985). Ecologically very similar to the transpalaearctic M. 

xerophila Berger. 

Threats: Afforestation, intensification of grassland management 

and tourism. 

Conservation: The unique ecosystem of Békás-szoros (Cheile 

Bicazului, the type locality of the subspecies) with its 

surroundings in the eastern Carpathians could be a National 

Park in Romania, where the traditional grassland and forest 

management could be maintained. Tourists could be restricted 

to indicated paths rather than be allowed to ramble freely. 

Plebejus (Lycaeides) idas (L.) 



The species is very little known in Hungary, and only 

scattered faunistic records exist. Its typical habitat (heath covered 

with Calluna vulgaris (L.) Hull. and Vaccinium myrtillus L.) is 

very rare. 

Taxonomy and Description: The Hungarian populations 

belong to one of the central European subspecies described as 

stempfferschmidti Beuret, which is most probably identical 

with the German or the other central European populations 

(Geiger 1988). The species needs a comprehensive taxonomic 

revision. 

Distribution: Holarctic. Most probably the Mediterranean 

(Balkans, Serbia, Croatia)'idas' populations represent another 

species, so the Hungarian colonies seem to be boundary 

populations. 


Habitat and Ecology: Xerothermophilous, bi- or trivoltine, 

presumed stenotopic. Caterpillars are regularly or obligately 

myrmecophilous (Fiedler 1991). Its typical habitat is heath 

covered with Calluna vulgaris (L.) Hull. and Vaccinium myrtillus 

L.). Other details (such as larval foodplants in other localities, 

etc.) are unknown from Hungary. Sarothamnus scoparius (L.) 

Wimm. is known as the larval foodplant in northeastern Hungary, 

on a site with atlantic influences. 

Threats: Intensification of grassland and forestry management. 

Conservation: Only one stable and strong population is known. 

It is situated on a xerophytic, silicate grass steppe of an acid 

mountain slope, partly covered with scrub (Quercus cerris L.

and Q. petraea (Mattuschka) Lieblein) in northeast Hungary 

(Bálint 1991). This unique habitat was strongly disturbed by the 

opening of a new forestry service road, the changing of the 

water-course system and the partial destruction of the grassland. 

Field research on this species is urgently required in Hungary 

to determine if it exists at other sites and to gather basic 

information on the species. 

Polyommatus (Agrodiaetus) admetus 

(Esper) 



The species was described on the basis of Hungarian 

specimens. A typical forest steppe species, many of its habitats 

were destroyed in the last decade. Very local in the country and 

in the Carpathian Basin. 

Taxonomy and Description: The species is a member of the 

ripartii (Freyer)-group, which consists of many very closely 

related allopatric species (Geiger 1988). The group was analysed 

by De Lesse (1960). 

Distribution: An east Mediterranean species. The species has 

the western and northern European limits of its range in Hungary. 


Habitat and Ecology: Xerothermophilous, univoltine (late 

June–July), stenotopic. Caterpillar presumed steadily 

myrmecophilous (Fiedler 1991). Larval hostplant in Hungary: 

Onobrychis arenaria (Kit.) DC. (Szabó 1956). Other details are 

not known. 

Threats: Major threats are afforestation, recultivation, 

urbanisation and air pollution. 

Many habitats have been lost through the recultivation of 

abandoned orchards and vineyards or were destroyed when 

built over very recently (especially in the area surrounding 

Budapest where the species is now extinct). Some colonies 

have disappeared because of artificially planted Pinus nigra 

Arn. (Pest County: Esztergom). 

Conservation: Only two populations have been confirmed 

recently; one of them is situated in the territory of Aggtelek 

National Park. The second population is strongly threatened by 

a lime factory (Pest County: Vac). Since the ecology of the 

species is totally unknown, field studies are urgently required. 

Field surveys are also essential since it seems very likely that 

more populations exist. 

108 

Polyommatus (Agrodiaetus) damon (Denis 

& Schiffermüller) 



The species has been recorded only from two Hungarian 

localities (Bálint 1991), and one of these has not been confirmed 

in recent years. The single remaining population is the only 

known confirmed one in the whole Carpathian Basin. 

Taxonomy and Description: The Hungarian population is 

identical to those of central Europe (Geiger 1988, p.388). 

Distribution: Transpalearctic (from Iberian peninsula to 

Mongolia), but the range consists of very scattered populations. 


Habitat and Ecology: Xerothermophilous, univoltine (July, 

beginning of August), stenotopic. Caterpillar nocturnal feeder, 

steadily myrmecophilous (Fiedler 1991). Larval hostplant 

Onobrychis viciaefolia L. (pers. obs.). Other details are not 

known. 

Threats: The single remaining population is in the territory of 

Budapest, and this site is the most popular centre for picnics, 

weekend activities and winter sports. These recreational activities 

represent a serious threat to the species. 

Conservation: Fencing off the habitat to safeguard the species 

is not possible in this area and the only long-term solution for 

the species in Hungary seems to be translocation to another 

suitable habitat (possibly in a protected area). Suggested field 

studies in conjunction with a proposal for the conservation of 

P. damon are under preparation. 

Polyommatus (Vacciniina) optilete (L.) 

Country: Slovakia. 


The only recently known population of the species in the 

Carpathian Basin can be found in northwest Slovakia. 

Taxonomy and Description: The Slovakian populations do 

not differ from those of central Europe (Geiger 1988). 

Distribution: Holarctic. It has been recorded from several 

northern Carpathian localities, but only one site has been 

confirmed recently (Kulfan and Kulfan 1991). 

Population Size and Status: Not known.

Habitat and Ecology: Tyrpophil, univoltine (July), stenotopic. 

Caterpillar myrmecoxenous (Fiedler 1991). The larval hostplant 

in the Alps is mainly Vaccinium L. species (Geiger 1988). No 

details concerning the Slovakian populations are known. 

Threats: Natural succession and wetland drainage. 

Conservation: It would be important to protect the peat bogs 

and to study the ecology of the Slovakian populations. 

Polyommatus eroides (Frivaldszky) 

Country: Slovakia. 


In the Carpathians, records of the species exist only from 

Slovakia. The species has not been reported from Hungary, or 

even from Romania. 

Taxonomy and Description: Resembling Polyommatus eros 

(Ochsenheimer) (Geiger 1988), but larger with a deeper blue 

upperside ground colour and a somewhat wider black border. 

Distribution: An east Mediterranean species, perhaps endemic 

for the Balkans. The Slovakian populations were connected 

with the Moravian ones, but both of them have become strongly 

isolated in recent times and are very close to extinction, most 

probably by human influences (see Králicek and Povolny 

1957). 


Habitat and Ecology: Xerothermophilous, univoltine (July), 

stenotopic. Caterpillar presumed to be steadily my rmecophilous 

(Fiedler 1991). Other aspects of the ecology of the species are 

totally unknown. 

Threats: The records are very scattered, which suggests that 

the populations are on the way to disappearing. Kulfan and 

Kulfan (1991) grouped the species into a complex of 

xerothermophilous butterflies which are threatened by several 

harmful factors, including recultivation of their habitats 

(abandoned orchards and vineyards), burning fields, 

overgrazing, building-over, illegal rubbish-dumping, 

earthworks, afforestation and natural succession. 

Conservation: Ecological and taxonomic investigations are 

urgently required. 

109 

Polyommatus dorylas magna (Bálint) 



Endemic to the eastern Carpathians. 

Taxonomy and Description: Much larger than the nominate 

race, with a very wide and conspicuous antemarginal white 

border (Bálint 1985b). 

Distribution: P. dorylas is distributed in the western Palaearctic 

region. This subspecies is known only in the mountain system 

of the eastern Carpathians (Romania and Carpat-Ukraine, 

?Slovakia). 


Habitat and Ecology: Xerothermophilous, univoltine (July 

and beginning of August). Caterpillar steadily myrmecophilous 

(Fiedler 1991). Larval hostplant and main nectar source of the 

imagines: Anthyllis vulneraria L. (pers. obs.). The species lives 

in the same habitats as Parnassius apollo transylvanicus 

(Schweitzer, 1912) and Maculinea sevastos limitanea (Bálint 

1985). 

Threats: Afforestation, intensification of grassland 

management, tourism. 

Conservation: The same suggestions as those advanced for M. 

sevastos limitanea apply equally to this taxon. 

Lycaena helle (Denis & Schiffermuller) 



The species is endangered in the western part of its range. 

Taxonomy and Description: The Romanian populations are 

identical to those of central Europe (Bálint and Szabó 1981). 

Distribution: Transpalearctic. The European populations are 

very scattered (Meyer 1981–1982). Only three sites are known 

in Transylvania (western part of Romania). One of them was 

reported at the beginning of this century and has never been 

confirmed. The second record originates from the 1970s and 

this, also, has not been confirmed recently. Only the third, the 

most recently discovered site (in 1979, Szabo 1982), seems to 

be strong enough to survive. 

Population Size and Status: Not known.

Habitat and Ecology: Hygrophilous, bivoltine (April, early 

May and July), stenotopic. Caterpillars myrmecoxenous (Fiedler 

1991). The ecology of the Transylvanian populations is not 

known. 

Threats: Wetland drainage, overcollecting. 

Conservation: The habitat of the strongest population should 

urgently be declared a nature reserve and the whole ecosystem 

also needs protection. A part of the swamp has been destroyed 

already, and Maculinea alcon L. has disappeared. The butterfly 

must be protected by law, because it is the subject of intensive 

collecting. 

Lycaena tityrus argentifex Bálint 



Endemic to the alpine and subalpine zones of the eastern and 

southern Carpathians. 

Taxonomy and Description: Somewhat similar to the taxon 

subalpinus Speyer, but the female often with submarginal 

orange lunules, and the underside ground colour deep silver 

grey with large spots. 

Distribution: West Palaearctic species. The subspecies 

argentifex is confined to the eastern and southern Carpathians, 

occurring at the same elevations as subalpinus in the Alps 

(800–2000m). 


Habitat and Ecology: Alpicol, univoltine (July–August), 

presumed stenotopic. Caterpillar myrmecoxenous (Fiedler 

1991). The habitats are mesophilous meadows, but other details 

concerning the ecology of the subspecies are not known. 

Threats: Intensification of grassland management and wetland 

drainage. The known habitats are traditionally used by the 

native inhabitants as grassland. Any modification of the system 

by changes in the regime or the use of the meadows could be 

dangerous for the species. 

Conservation: Establishment of natural reserves and the study 

of the ecology of the species are required. 

110 

Pseudophilotes bavius hungaricus 

(Diószeghy) 



An endemic subspecies of the Transylvanian Basin (West 

Romania). 

Taxonomy and Description: Close to P. bavius Eversmann, 

but differing in some minor characters (König 1988). 

Distribution: An east Mediterranean species. The western 

limit of its range is represented by the populations of hungaricus. 

Only very few populations are known and they are very scattered 

(Szabó 1982). Very recently the species (most probably an 

undescribed subspecies) was found in Dobrogea, south Romania 

(Székely, pers. comm.). 


Habitat and Ecology: Xerothermophilous, univoltine (April), 

stenotopic. Caterpillar presumed weakly myrmecophilous 

(Fiedler 1991). Larval hostplant Salvia nutans L. The early 

stages were described and the ecology was partly studied by 

König (1988). 

Threats: Intensification of grassland management, afforestation, 

earthworks, air pollution and overcollecting. Some habitats 

were overgrazed (Cluj Napoca-Kolozsvárl: Szénafiivek), or 

planted over by Pinus nigra Arn. or Robinia pseudacacia L. 

(Teuis-Tövis). Others are endangered by a chemical factory 

(close to Sibiu-Nagyszeben). A single colony in a nature 

reserve (Suat-Magyarszovát: Csigla domb) was most probably 

exterminated by the activities of the native collectors. 

Conservation: Some habitats are protected by law, but the 

protection remains only on paper and has not been enforced. 

Very strict dispositions would be needed. Thorough ecological 

studies are necessary to continue the initial work of König 

(1988). 

Tomares nogelii dobrogensis (Caradja) 



Only a single population is known from Romania. 

Taxonomy and Description: see Higgins and Riley (1983) 

and Hesselbarth and Schurian (1984).

Distribution: An east Mediterranean species. The Romanian 

locality in Dobrogea represents the most westerly one known. 


Habitat and Ecology: Xerothermophilous, univoltine (May). 

Caterpillar steadily myrmecophilous (Fiedler 1991). Other 

details concerning the Romanian populations are not known. 

Threats: Overgrazing, overcollecting. 

Conservation: The known site is very small and strongly 

disturbed by human activity. The habitat is well known amongst 

the Romanian lepidopterists, so overcollecting of the species is 

a real danger in spite of the fact that the habitat is a nature 

reserve. The Romanian population of nogelii has never been 

studied from the ecological point of view, and field studies of 

nogelii dobrogensis are urgently required. 

References 

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(Lepidoptera: Lycaenidae). Folia ent. hung, 46: 256–257 (in Hungarian). 

BÁLINT, Zs. 1985b. Plebicula dorylas magna nov. ssp. (Lep.: Lycaenidae) 

from the eastern Carpathians, Romania. Neue Ent. Nachr. 14: 14–20. 

BÁLINT, Zs. 1986. Further studies on Maculinea alcon (Den. & Schiff., 

1775) (Lepidoptera: Lycaenidae). Galathea, Nürnberg 2: 92–108. 

BÁLINT, Zs. 1991. Conservation of butterflies in Hungary. Oedippus 3: 

5–36. 

BÁLINT, Zs. and SZABÓ, Gy. 1981. The occurrence of Lycaena helle (Den. 

et Schiff., 1775) in the Szatmár lowland. Folia ent. hung. 42: 235–236 (in 

Hungarian). 

111 

DE LESSE, H. 1960. Les nombres de chromosomes dans la classification du 

groupe d'Agrodiaetus ripartii Freyer (Lepidoplera, Lycaenidae). Rev. 

franc. Ent. 27: 240–264. 

FIEDLER, K. 1991. European and North West African Lycaenidae 

(Lepidoptera) and their associations with ants. J. Lepid. Soc. 28:239–257. 

GEIGER, W. (Ed.) 1988. Tagfalter und ihre Lebensräume. Arten. Gefährdung. 

Schutz. Schweizerische Bund für Naturschutz, Basel, 2nd edn., xi + 516 

pp. 

HESSELBARTH, G. and SCHURIAN, K.G. 1984. Beitrag zur Taxonomie, 

Verbreitung und Biologie von Tomares nogelii (Herrich-Schäffer, 1851) 

in der Türkei (Lepidoptera, Lycaenidae). Entomofauna 5: 243–250. 

HIGGINS, L.G. and RILEY, N.D. 1983. A Field Guide to the Butterflies of 

Britain and Europe. Collins, London, 5th edn., 384 pp. 

KÖNIG, F. 1988. Morphological, biological and ecological data on Philotes 

bavius hungarica Diószegtiy, 1913 (Lepidoptera, Lycaenidae). 4thNation. 

Conf. Entomol. Cluj-Napoca, pp. 175–182. (in Romanian). 

KRÁLICEK, M. and POVOLNŸ, D. 1957. Polyommatus eros eroides 

(Frivaldsky, 1837) (sic) in Czechoslovakia. Acta Soc. ent. Czech. 53: 

193–201 (in Czech.). 

KUDRNA, 0. 1986. Aspects of the Conservation of Butterflies in Europe. 

Butterflies of Europe. Volume 8. Aula-Verlag Wiesbaden, 323 pp. 

KULFAN, J. and KULFAN, M. 1991. Die Tagfalterfauna der Slowakei und 

ihr Schutz unter besonderer Berücksichtigung der Gebirgökosysteme. 

Oedippus 3: 75–102, 4 figs. 

MEYER, M. 1981–1982. Revision systematique, chorologique et ecologique 

des populations europeennes de Lycaena (Helleia) helle Denis & 

Schiffermüller, 1775. 

SZABÓ, A. 1982. Data to the Romanian distribution of the species Lycaena 

helle Schiff. and Philotes bavius Ev. (Lepidoptera, Lycaenidae). Studii si 

Comunicari, Reghin 2: 299–306 (in Romanian). 

SZABÓ, R. 1956. The Lycaenids of Hungary. Folia ent. hung. 9 (SN): 

235–262. 

UHRIK-MÉSZÂROS, 1948. Data on the knowledge of the life history of 

Lycaena iolas O. Folia ent. hung. 3 (SN): 5–8 (in Hungarian). 

VARGA, Z. 1968. Bemerkungen und Ergänzungen zur taxonomishen 

Beurteilung und Ökologie der im Karpatenbecken vorkommenden 

Populationen von Ariciaartaxerxes Fabr.Acta biol. Debrecina 6:171–185.

Country: Japan. 

'Chosen-aka-shijimP, Coreana raphaelis Oberthur 

T. HIROWATARI 

Entomological Laboratory, College of Agriculture, University of Osaka Prefecture, Sakai, Osaka 591, Japan 


In Japan, this species occurs only in very local areas of 

northern Honshu, and all the populations seem to be threatened 

with extinction. In most districts, positive actions by the residents 

in support of conservation are evident. This is one of the model 

cases of conservation in Japan (Sibatani 1989), and activities 

have included: (a) monitoring population size; (b) volunteer 

patrolling; (c) breeding of butterflies; (d) planting of foodplants; 

and (e) campaigns for public education in conservation 

awareness. 

Taxonomy and Description: The genus Coreana Tutt is 

monotypic and one of the most primitive members of the tribe 

Theclini (sensu Eliot 1973). Representatives from Iwate 

Prefecture are treated as ssp. yamamotoi Okano (broader orange 

marking on upperside), and from Yamagata Prefecture as ssp. 

ohruii Shirozu (broader black border on hindwing upperside). 

Distribution: Confined to northern Honshu Iwate Pref.: Kuji 

(40°ll'N, 141°46'E) to Miyako (39°38'N, 141°59'E), around 

Sizukuishi (39°41'N, 140°59'E); Yamagata Pref.: Shinjo 

(38°45'N, 140°18'E) to Kawanishi (37°59'N, 140°02'E); 

Niigata Pref.:Asahi(38°15'N,139°35'E),Sekikawa(36°06'N, 

139°36'E). 

Outside Japan, this species is distributed in some disjunct 

areas of Amur and southern Ussuri (southeastern Russia), 

northern China and the Korean Peninsula. 

Habitat and Ecology: In northern Honshu the species occurs 

on the borders of woods, along creeks, in marshes near low hills 

where Fraxinus spp. (Oleaceae) grows, the foodplant of this 

species. There are many rural houses in this area which are 

surrounded by hedges of this plant, and such hedges are now the 

main habitat of this species. 

The larvae mainly feed on Fraxinus japonica, and 

occasionally on F. lanuginosa and F. mandshurica. Eggs are 

laid in batches of several to 20 on the trunks of foodplants. 

Larvae hatch in April. Second instar larvae make simple shelters 

by spinning leaves. Third and 4th instar larvae nibble the stems 

112 

of the leaves and sit on drooping leaves. Fourth (last) instar 

larvae are sometimes tended by ants (Lasius spp.). Pupae are 

found under fallen leaves, pieces of decayed wood or under 

stones (Fukuda et al. 1984). 

There is one generation each year. Adults emerge in mid- 

June, and are seen until early August. The species overwinters 

as eggs. 

Threats: Habitats of this species are being progressively 

destroyed by the development of housing sites, etc. Felling 

foodplants in the area along creeks or marshes near low hills is 

one of the main factors which reduces the population size. One 

further threat is the decrease in the number of hedges made of 

Fraxinus spp. which now seem to be essential for the species. 

Conservation: This species has been strongly associated with 

rural human life in its range in Japan and its survival seems to 

depend on the future activities of the local communities. 

In Kawanishi (Yamagata Pref.), this species was designated 

as a protected species of the Prefecture in 1977. However, in the 

absence of effective conservation measures, its habitats have 

been progressively destroyed. Nevertheless, local bodies began 

to promote conservation and urged the local government to take 

conservation measures. For example, in 1989, at the time of 

creek improvement in Kawanishi, the foodplants of C. raphaelis 

were transplanted to unoccupied ground (c. 600m) that had 

been taken over by the local government, and some of the plants 

were also transplanted to school grounds (Nagaoka amd Kusakari 

1989). 

In Iwate Pref., the butterfly is now designated a protected 

species in all the cities, towns and villages (Ogata et al. 1989). 

As in the case of Yamagata Pref., local governments as well as 

local bodies do recognize that prohibition of collecting without 

adequate management measures are not effective for real 

protection of butterflies, and they are now devising the following 

measures: (1) continuous monitoring of population size by 

local bodies and lepidopterists; (2) environmental protection by 

patrolling and by checking the urbanisation plans of the local 

governments; (3) breeding butterflies and planting of their 

foodplants in public places, such as school grounds or private 

housing sites; and (4) education for conservation awareness by

holding lecture meetings and by producing public relations 

leaflets or videotapes. 

References 

FUKUDA, H., HAMA, E., KUZUYA, T., TAKAHASHI, A., TAKAHASHI, 

M., TANAKA, B., TANAKA, H., WAKABAYASHI, M. and 

WATANABE, Y. 1984. The Life Histories of Butterflies in Japan Vol. 3. 

Hoikusha, Osaka (in Japanese). 

NAGAOKA, H. and K. KUSAKARI. 1989. Habitat status of Coreana raphaelis 

113 

and conservation practice in Kawanisi-shi, Yamagata Prefecture. 

In: Hama, E., Ishii, M. and Sibatani, A. (Eds) Decline and 

Conservation of Butterflies in Japan, I. pp. 65–73. 

Lepidopterological Society of Japan, Osaka (in Japanese). 

OGATA, Y., MIURA, H. and Y. MUROYA. 1989. Population status 

of Coreana raphaelis and the conservation practices in Iwate 

Prefecture. In: Hama, E., Ishii, M. and Sibatani, A. (Eds) Decline 

and Conservation of Butterflies in Japan, I. pp. 74–82 (in Japanese). 

SIBATANI, A. 1989. Decline and conservation of butterflies in Japan. 

In: Hama, E., Ishii, M. and Sibatani, A. (Eds) Decline and 

Conservation of Butterflies in Japan 1, pp. 16–22. 

Lepidopterological Society of Japan, Osaka.

Country: Japan. 

'Oruri-shijimi', Shijimiaeoides divinus Fixsen 

T. HIROWATARI 

Entomological Laboratory, College of Agriculture, University of Osaka Prefecture, Sakai, Osaka 591, Japan 


In Japan, this species occurs in very local and disjunct areas 

in Honshu and Kyushu. The populations of Honshu are feared 

extinct, especially in the northern district. In fact, the ones in 

Aomori Prefecture and Iwate Prefecture are thought to have 

been extinct since the late 1970s. No effective conservation 

measures have been taken by local governments in Kyushu 

except for the prohibition of collecting. 

Taxonomy and Description: The genus Shijimiaeoides Beuret, 

related to Maculinea van Eecke and Glaucopsyche Scudder, 

contains two species, lanty Oberthur and divinus Fixsen, both 

occurring in eastern Asia. Representatives from Honshu and 

Kyushu are treated as ssp. barine Leech and asonis Matsumura, 

respectively of S. divinus. The nominotypical subspecies occurs 

in the Korean Peninsula, and also seems to be endangered 

(Fukuda et al. 1984). 

Distribution: Very local, in disjunct areas of Honshu and 

Kyushu. The localities of known habitats consist of three 

groups: (1) northern Honshu: Aomori Pref., Iwate Pref. Morioka 

(39°41 'N, 141 °08'E); (2) central Honshu: Niigata, Nagano and 

Gunma Prefs.; and (3) Kyushu: Aso, Kuju. Outside Japan, this 

species is distributed in local areas of the Korean Peninsula. 

Population Size: Population size of this species has never been 

estimated systematically. However, the decline of populations 

in northern Honshu seems to be obvious. In Aomori Pref. they 

began to decrease in the 1960s and became extinct by the 

second half of the 1970s (Muroya 1989). In Azumino, Nagano 

Pref., this species was widespread in the 1970s, but it is now on 

the brink of extinction (Kobayashi 1989). Exceptionally in 

Kyushu, populations of this species and their habitats seem to 

have been conserved during these decades. 

Habitat and Ecology: Usually occurs on sunny grassland, 

river banks, degraded areas along railroads and, especially, on 

volcanic slopes. 

114 

Larvae obligately feed on the flowers and flowerbuds of 

Sophora flavescens (Leguminosae). In Kyushu (Aso, Kuju), 

volcanic slopes are used for pasture, and S. flavescens grows 

well in such grasslands, because cattle avoid grazing it. 

There is one generation each year, with adults present in 

May in Kyushu, and in June in Honshu. Eggs are laid on a 

flowerbud. The larva is yellowish and milky white, attended by 

ants (not identified). The pupa is blackish and found on the 

ground near the foodplant. 

Threats: Extinction of the northern Honshu population (Aomori, 

Iwate) is attributable to degradation of habitat caused by 

cultivation, alteration in land management practices, and 

urbanisation such as construction of golf courses, airport, and 

so on. The habitats of S. divinus are mostly grasslands, which 

may be easily cultivated or used in urban development. For 

example, at the foot of Mt. Iwaki (Aomori Pref.), exploitation 

by the national and local governments was carried out for seven 

years (1962–1968) during which time a total of 2155ha of 

grassland were cultivated, covering most of the habitats of the 

species. After that small populations that had survived in 

grasslands along creeks also became extinct in the late 1970s. 

Alterations in land management practices, another of the 

main factors, has involved a decrease in the number of cattle 

and a resultant cessation of grass-cutting: this has led to 

successional changes which are not suitable for the growth of 

the foodplant. 

Conservation: In spite of the decline in the 1960s in northern 

Honshu, no campaign was undertaken to save the species. 

Consequently, the populations of Aomori were eradicated in 

the late 1970s. The populations of central Honshu seem to be at 

a crucial stage at the present time but no conservation project 

has been undertaken, either. In Kyushu, the local governments 

prohibit collecting of the species, and local bodies are cooperating 

on a volunteer patrolling system. An essential measure would 

be to conserve the grassland and prevent any further alterations 

in land management practices. However, at this moment nothing 

has been done as a result of inadequate financial support.

References 

FUKUDA, H., HAMA, E., KUZUYA, T., TAKAHASHI, A., TAKAHASHI, 

M., TANAKA, B., WAKABAYASHI, M. and WATANABE, Y. 1984. 

The Life Histories of Butterflies in Japan, Vol. 3. Hoikusha, Osaka (in 

Japanese). 

115 

KOBAYASHI, Y. 1989. Decline of Shijimiaeoides divinus populations in 

Azumino, Nagano Prefecture, pp. 97–98, in Hama, E., Ishii, M. and 

Sibatani, A. (Eds) Decline and Conservation of Butterflies in Japan I. 

Lepidopterological Society of Japan, Osaka (in Japanese). 

MUROYA, Y. 1989. Decline ofShijimiaeoides divinus populations in Aomori 

Prefecture. Ibid.: 90–97 (in Japanese).

Lange's Metalmark, Apodemia mormo langei Comstock 

Jerry A. POWELL 1 and Michael W. PARKER 2 

1 Department of Entomological Sciences, 201 Wellman Hall, University of California, Berkeley, 

California 94720-0001, U.S.A. 

2 U.S. Fish and Wildlife Service, San Francisco Bay National Wildlife Refuge Complex, Newark, CA 94560, U.S.A. 

Country: U.S.A. 

Status and Conservation Interest: Status – possibly out of 

danger; endangered listing (USFWS). 

This local subspecies was one of the first eight insects to be 

listed as an endangered species in 1976 under the Federal 

Endangered Species Act. Its remnant habitat was purchased by 

the U.S. Fish and Wildlife Service (USFWS) in 1980 and 

designated as the Antioch Dunes National Wildlife Refuge, the 

first such refuge in the country established to protect insects and 

plants. Population numbers of Apodemia mormo langei declined 

over a period of 50 years to a few hundred individuals in the 

early 1980s. During the past decade they have increased tenfold 

or more in response to recovery actions, which include exclusion 

of vehicular and most foot traffic, removal of exotic vegetation, 

and extensive outplanting of the larval host plant. 

General accounts of the unique insect and plant communities 

and destruction of the Antioch dunes by sand mining and 

Female Lange's Metalmark on flowers of Senecio douglasii. 

116 

industrial development have been recorded (Howard 1983; 

Howard and Arnold 1980; Farb 1964; Powell 1981, 1983). 

Taxonomy and Description: Considerable polytypy is 

expressed, often discordantly, in hostplant species, seasonal 

phenology and voltinism, and in size and colour pattern of the 

adults (Opler and Powell 1962; Powell 1975). 

Distribution: Apodemia mormo (Felder & Felder) (Riodininae) 

is widely distributed in the western Nearctic and occurs in 

scattered, often isolated colonies. Some forms of the butterfly 

are quite limited in geographic distribution and may be separated 

by distances of only 15–20km from other populations that are 

easily distinguished phenotypically. Apodemia mormo langei 

was discovered in 1933, on the riverine sand dune system just 

east of Antioch, Contra Costa Co., California (Comstock 1938). 

This subspecies is represented by just one population (Opler 

and Powell 1962). The nearest known colonies of Apodemia 

mormo are located on the northwest slope of Mt. Diablo and on 

the hills northwest of Vallejo, Solano Co., which are respectively 

about 15km SW and 39km WNW of the langei site. These 

populations have appreciable differences in colour pattern from 

langei. 

Population Size: U.S. Geological Survey topographical maps 

and aerial photographs that span 1905 to 1969, document the 

destruction of the Antioch sand dunes. At the turn of the 

century, after agricultural development of the region, they 

extended for 3–4km in a narrow band along the San Joaquin 

River and reached heights of 20–35m. Sand mining operations 

began prior to and during the 1920s (Howard and Arnold 1980). 

Although the installation of two powerline towers, in 1909 and 

1927, resulted in habitat disturbance including building 

construction and introduction of exotic plants, today these 

Pacific Gas and Electric (P.G. & E.) towers stand on the two 

remaining remnants of sand hills. 

By 1931, five huge sand pits, each with a railroad spur, were 

in operation, for the developing San Francisco Bay area. During 

post World War II years, massive industrial developments 

replaced the eastern half of the dunes, and in the 1950s sand 

mining moved to the western sector. A Kaiser Gypsum plant

was completed in 1956, and this isolated the two remaining 

remnants of the habitat, the Stamm Property (SP) to the west 

and the 'Little Corral' (LC) flanked by the P.G. & E. properties 

to the east (Figure 1). 

Although its habitat was gradually restricted during the 40 

years following discovery, the local abundance of Apodemia 

mormo langei remained high. Lepidopterists observed the 

butterflies by the hundreds and collected specimens at rates of 

25–30 per hour in various seasons during 1947–1972. Even as 

late as 1972, three observers took 70 specimens one day and 50 

more seven days later and estimated sighting 150–200 on each 

date. 

Soon thereafter, however, the colonies were severely affected 

by increased sand-mining at the western parcel (SP), by 

rototilling on the P.G. & E. properties in compliance with a 

county ordinance for fire prevention, and by overgrazing by 

horses. Thus by 1976, when Powell began population census by 

transect counts, numbers of langei had dropped alarmingly. 

During mark-recapture monitoring by Arnold and Powell from 

1977–1982, the maximum number of langei two persons could 

observe in 3.5–4.5 hour periods at the two sites was 45–55 

(Arnold and Powell 1983; Powell unpubl. data). Total population 

numbers, calculated from daily Jolly-Seber and Manly-Parr 

estimates and the survival rates of individual butterflies, declined 

from more than 2000 in 1977 to fewer than 600in 1982 (Arnold 

1985 and unpubl. reports to USFWS). 

The butterfly numbers stabilised and increased slightly in 

1983–84 and then rose significantly as langei began to occupy 

Eriogonum in peripheral areas that had been both planted and 

colonised naturally. The estimated total population exceeded 

1200 in 1985, the final year of Arnold's mark-recapture studies 

(Arnold, unpublished report to USFWS). Subsequently, single 

day sighting counts, made weekly by USFWS personnel, indicate 

that the numbers of langei have continued to climb dramatically. 

Seasonal peak numbers have risen from 168 in 1986 to more 

than 1900 in 1991. It is likely that the population has 10 to 20 

times the number of butterflies that it was estimated to have 

contained 10 years ago (e.g. 6000 to 12,000). 

Habitat and Ecology: Populations occur in close association 

with the larval foodplants, species of Eriogonum (Polygonaceae), 

typically in well drained semiarid sites, such as rocky desert 

slopes, sand dunes, or chaparral-covered hills, ranging from sea 

coast to timberline at 2750m. Colonies of A. m. langei are 

limited to dense or moderately dense patches of the larval 

foodplant, Eriogonum nudum auriculatum (Arnold and Powell 

1983). Arnold and Powell believed that isolated or spindly, 

scattered plants fail to support colonies of the butterfly because 

early instar larvae derive insufficient protection for 

overwintering. 

A. m. langei is univoltine, with adults flying for about 30 

days beginning in early August. Males precede females by a 

few days, and peak numbers occur about two weeks after 

emergence begins. The eggs are deposited in clusters of 2–4 on 

withering foliage on the lower half of the plant. Eclosion of 

larvae takes place during winter, after rainfall and foliation 

begin. Feeding occurs by skeletonizing the leaf surfaces and the 

inflorescence stalks by later instars in June and July. Larvae 

feed in early morning and presumably evening and retreat to the 

base of the plants during the day. Pupation occurs in the litter 

Figure 1. Map of the Antioch Dunes National Wildlife Refuge, just east of Antioch, Contra Costa Co., California. 

Redrawn from Antioch North Quadrangle, U.S. Geological Survey topographic map, 1953. Shaded areas delineated by dotted lines indicate construction sites, 

1953–1968, compiled from aerial photographs. Sparsely doited areas depict sand-mining excavations during 1953–1967. The two corridors traversed by P.G. 

& E. powerlines are the only unexcavated hills that remain. Bold lines define the two parcels of the Refuge, "Stamm Property" (SP) and "Little Corral" (LC), 

the latter flanked by P.G. & E. properties. 

117

at the base of the plant, in late July and early August 

(Arnold and Powell 1983). 

Males tend to occupy restricted areas day after day, 

while females move greater distances. Mark-recapture 

data suggested that males live an average of about 12 

days. Adults of both sexes forage for nectar; Eriogonum 

serves as the primary nectar source, at least in recent 

years, while occasional visits are made to Gutierrezia 

and Senecio (Asteracecae), both of which were formerly 

more abundant, and other plants (Arnold and Powell 

1983). 

Under natural circumstances, this variety of 

Eriogonum nudum probably lived as an edge species, 

occupying slip slopes of active sand in the hills that were 

stabilised by scattered oaks and a rich flora of desert 

affinities (Howard 1983). There is no doubt that the 

active sand habitat greatly increased during the 1920–1940 

era of sand-mining. Thus, it is likely that, along with its 

larval host, A. m. langei had increased in population 

numbers by the time of its discovery in the early 1930s. 

Conservation: Rehabilitation of habitat suitable for 

colonisation by A. m. langei begore before acquisition of 

the Refuge lands by USFWS. Propagation and outplanting 

on the P.G. & E. properties began in 1979, following 

inadvertent rototilling of one of the primary stands of 

Eriogonum, despite efforts by the company to prevent 

such an accident. P.G. & E. contracted with Biosystems 

Analysis Inc. to develop a restoration plan, and about 450 

seedlings were planted in March 1980 (Howard and 

Arnold 1980). Collective efforts during 1980–1984, which 

were orchestrated by Howard and Arnold, were financed 

by P.G. & E., grants from the California Native Plant 

Society, a USFWS contract, and assistance by the 

University of California, Berkeley undergraduate botany 

association and other volunteers (Arnold 1985). During 

the same years, the USFWS began planting Eriogonum at 

the western parcel (SP) by scarifying and seeding the 

excavated surface left bare by a final surge of sandmining 

in 1978–79. In 1985 the USFWS entered a 

Cooperative Agreement with P.G. & E. that allows the 

Service to manage the additional 5ha owned by P.G. & 

E., which are contiguous with LC (Figure 1). In 1987, 

fencing of the lands was completed, and this virtually 

eliminated further human degradation of the habitat. 

Also in 1987 a vineyard to the south of the Apodemia 

colony at SP was removed and subsequently planted with 

7000 Eriogonum seedlings. Small numbers of langei 

have begun to occupy that area (58 were observed in one 

day in 1991). 

In 1991 a more ambitious cooperative restoration 

project was initiated to create new sand dunes in previously 

mined areas occupied by weedy vegetation. Low hills, 

consisting of sand mined from another P.G. & E. property 

up river, have been deposited and contoured on the LC 

and P.G. & E. western parcels. These were subsequently 

118 

seeded and planted and now bear mantles of Eriogonum 

and Senecio douglasii seedlings, the latter a major nectar 

source for the butterflies, as well as two endangered 

plants, Oenothera deltoides var. howelli (Onagraceae) 

and Erysimum capitatum var. angustatum (Brassicaceae). 

Altogether, we estimate that outplanted Eriogonum 

colonies that have reached densities believed to be 

sufficient to support Apodemia occupy areas of 9600m 2 

at SP and 3200m 2 on the P.G. & E. properties, a total of 

nearly 1.3ha. 

In 1979 the maximum distribution of the foodplant 

range (range width x range length, right angle) was 

estimated to be 2.3ha at LC and 1.5ha at SP, not more than 

one-third of which was suitable for Apodemia as judged 

by occurrence of adults (Arnold and Powell 1983). Hence, 

there was only about 1.3ha of viable Eriogonum habitat 

at its lowest ebb, and subsequent efforts have at least 

doubled that area. Several additional patches of the host 

plant, planted in recent years, are expected to develop 

into viable habitat within a few seasons. 

Threats: Effects of weediness and possibility of fire are 

the principal threats to continued existence of A. m. 

langei. A standing crop of annual weeds poses a fire 

danger each year during the long dry season 

(May-October). The refuge is bordered inland by 

industrial development, and there are several small 

beaches that are accessible to recreational boat visits, so 

possible sources of human-initiated fires cannot be 

controlled. 

Conservation and recovery efforts have been a 

dramatic success, increasing the population numbers 

from a perilously low level a decade ago. However, after 

10–15 years Eriogonum host plants senesce, and they fail 

to reproduce in the absence of open, active sand. We have 

witnessed the growth and decline of Eriogonum and 

associated langei colonies in several areas during the past 

15 years. For example, a robust colony developed in 

association with the foundations of a building that was 

removed after 1972 on P.G. & E. west; it was the home 

of a strong colony of langei during 1978–1982, but these 

plants were senescent by 1988 and have died out by 1992. 

The foundations protected the plants from rototilling 

during the 1970s, but competition with weeds prevented 

seedling growth. A similar fate can be predicted for most 

of the existing and recently planted patches of Eriogonum. 

Once a colony of the food plant is established, it is 

impractical to prevent a ground cover of weeds, especially 

annual exotic grasses, yellow star thistle, Russian thistle, 

and vetch, which does not kill mature Eriogonum but 

prevents development of its seedlings. Thus, it is easier 

to clear a site and plant new patches of the host plant than 

to maintain existing ones. 

To be successful on a long term basis, the management 

plan needs to prescribe replacing senescent patches of 

Eriogonum on a continuing basis.


We thank Richard Arnold, Alice Howard, John Steiner, and 

Jean Takekawa for discussions and information regarding the 

recent and past history of the Antioch sand dunes and the flora; 

Arnold and Steiner reviewed a draft of the manuscript and 

offered useful criticisms. 

References 

ARNOLD, R.A. 1985. Private and government-funded conservation programs 

for endangered insects in California. Nat. Areas J. 5: 28–39. 

ARNOLD, R.A. and POWELL, J.A. 1983. Apodemia mormo langei. pp. 

99–128. In: Arnold, R. A., Ecological studies of six endangered butterflies 

(Lepidoptera: Lycaenidae): Island biogeography, patch dynamics, and 

the design of habitats preserves. Univ. Calif. Publns Entomol. 99. 

COMSTOCK, J.A. 1938. A new Apodemia from California. Bull. South Calif. 

119 

Acad. Sci. 37: 129–132. 

FARB, P. 1964. Insect city in the dunes, pp. 44–49. In: The Land and Wildlife 

of North America. Life Nature Library, Time-Life Books, New York. 

HOWARD, A.Q. (Ed.) 1983. The Antioch dunes. A report to the U.S. Fish and 

Wildlife Service. Prepared under P.O. No. 11640-0333-1; 115 pp. + 

appendices. 

HOWARD, A.Q. and ARNOLD, R.A. 1980. The Antioch dunes – safe at last? 

Fremontia 8(3): 3–12. 

OPLER, P.A. and POWELL, J.A. 1962. Taxonomic and distributional studies 

on the western components of the Apodemia mormo complex (Riodinidae). 

J. Lepid. Soc. 15: 145–171. 

POWELL, J.A. 1975. Riodinidae. The Metalmarks. pp. 259–272. In: Howe, 

W.H. (Ed.). The Butterflies of North America. Doubleday; Garden City, 

New York. 

POWELL, J.A. 1981. Endangered habitats for insects: California coastal sand 

dunes. Atala 6: 41–55 (1978). 

POWELL, J.A. 1983. Changes in the insect fauna of a deteriorating riverine 

sand dune community during 50 years of human exploitation. Report to 

U.S. Fish and Wildlife Service. Prepared under P.O. no. 11640-0333-1; 

78 pp.

The Hermes Copper, Lycaena hermes (Edwards) 

D.K. FAULKNER and J.W. BROWN 

Entomology Department, San Diego Natural History Museum, P.O. Box 1390, San Diego, California 92112, U.S.A. 


Status and Conservation Interest: Status–rare; indeterminate 

(Red List). 

The Hermes Copper is a remarkably distinct species, differing 

considerably from its congeners in both morphology and 

ecology. It has a highly restricted geographical distribution in 

the southwestern United States and adjacent northwestern Baja 

California, Mexico. Because of the widespread loss and 

fragmentation of its habitats, associated with urbanisation and 

other development, this species has lost a significant portion of 

its former range. Owing to its vulnerable nature, the Hermes 

Copper is recognized by the United States Fish and Wildlife 

Service as a 'category 2 candidate' species for listing as 

threatened or endangered. 

Taxonomy and Description: The Hermes Copper was described 

as Chrysophanus hermes by W.H. Edwards (1870) from 

' California'. Wright (1906) later described Chrysophanus delsud 

from San Diego County, California; the latter is a subjective 

synonym of hermes. The species has been included in the 

genera Tharsalea Scudder (e.g. Comstock 1927; Wright 1930) 

and Lycaena Fabricius (dos Passos 1964; Howe 1975). Miller 

and Brown (1979) placed hermes in the monotypic genus 

Hermelycaena Miller and Brown on the basis of its unique 

morphological and ecological characteristics. Currently, most 

taxonomists consider hermes to be a member of the Holarctic 

genus Lycaena. 

Distribution: The Hermes Copper is a narrowly endemic 

species, restricted to western San Diego County, California, 

and a small portion of adjacent Baja California, Mexico (Brown 

1980; Ehrlich and Ehrlich 1961; Emmel and Emmel 1973; 

Garth and Tilden 1986; Orsak 1977; Rindge 1948; Scott 1988). 

Its total range is approximately 250km from north to south 

(from about Fallbrook in San Diego County, California, to 

slightly south of Santa Tomas in Baja California, Mexico), and 

50km from east to west (from near the coast, inland to about 

Pine Valley, California). Within this range it occurs in small, 

disjunct colonies. 

120 

In San Diego County, the Hermes Copper has been recorded 

from El Cajon, Santee, Flynn Springs, Blossom Valley, Tecate, 

Pine Valley, Guatay, and numerous other localities, many of 

which no longer support native vegetation. 

Population Size: In the absence of focused studies on population 

size and vagility, information on these parameters is mostly 

anecdotal. The Hermes Copper has been collected at about 35 

localities in the United States and four localities in Mexico. 

Most colonies are isolated from each other. Hence, gene flow 

between populations probably is rather low. In addition, adults 

exhibit limited vagility: they do not hilltop and they seldom are 

observed beyond the immediate vicinity of the larval host. 

Thorne (1963) indicated that colonies vary little in size from 

year to year, but there are few quantitative data to support this 

observation. It is likely that few colonies exceed 50 individuals. 

Habitat and Ecology: This species occurs in coastal sage scrub 

and open southern mixed chaparral communities in which the 

larval host plant, redberry (Rhamnus crocea Nutt. in T. & G., 

Rhamnaceae) is a common component. In San Diego County, 

these habitat types range from near sea level along the coast to 

about 1250m at the western edge of the Laguna Mountains. The 

foodplant is a fairly common species, extending well beyond 

the range of the butterfly. Hence, the restricted distribution of 

the Hermes Copper is difficult to explain. 

The Hermes Copper is univoltine, with adults present from 

mid-May through early July, depending upon elevation. The 

primary adult nectar source at most localities is flat-top 

buckwheat (Eriogonumfasciculatum Bentham; Polygonaceae), 

but L. hermes also has been observed to nectar on slender 

sunflower (Helianthus gracilentus) and a few other composites 

(Asteraceae). Males perch on vegetation along trails and 

openings, and confront other butterflies that pass by. 

Eggs are laid singly on stems of the foodplant and overwinter 

until the following spring. The egg is white, echinoid, and 

covered with deep pits between high, irregular walls. The fullygrown 

larva is apple green, with a mid-dorsal band of darker 

green bordered with yellowish-green. Pupation occurs on the 

foodplant, and the pupa is attached by a cremaster and a silken

girdle. Full details of the life history are presented by Comstock 

and Dammers (1935). Ballmer and Pratt (1989) indicate that the 

larvae of Lycaena hermes differ greatly from those of other 

Lycaena species in host preference and morphology. No 

parasitoids or predators are recorded. 

Threats: Although declines have not been documented 

quantitatively, the Hermes Copper undoubtedly has suffered 

from loss and fragmentation of habitat as a result of urbanisation. 

As long ago as 1930, Wright (1930) reported 'Its trysting places 

are being rapidly taken over by realtors and the species may 

soon become extinct...' Indeed, much of the former range of L. 

hermes is presently occupied by urbanised portions of San 

Diego. As development in San Diego County extends eastward 

from the coast, the Hermes Copper is further threatened by 

habitat loss. 

Conservation: Currently, the Hermes Copper receives minimal 

protection under the California Environmental Quality Act. 

Under this legislation, impacts to sensitive plants and animals 

and sensitive habitat types must be assessed to determine 

whether the adverse affects of habitat loss and fragmentation 

resulting from development are 'significant'. If they are 

determined to be significant, mitigation measures are required 

to reduce impacts below a level of significance. Unfortunately, 

because invertebrates typically receive little attention in the 

environmental review process, impacts to these species usually 

are undocumented. 

Management recommendations for the Hermes Copper 

include increased awareness of this species, particularly for 

those individuals and agencies involved in the environmental 

review process, and protection of existing colonies from habitat 

loss and fragmentation associated with urbanisation. The Hermes 

Copper is presently recognized by the United States Fish and 

Wildlife Service as a category 2 candidate species for listing as 

endangered or threatened; the Service recently received a 

121 

petition to list the species as threatened. Such a listing would 

increase significantly the protection afforded Hermes Copper. 

References 

BALMER, G. and PRATT, G. 1989 (1988). A survey of last instar larvae of 

the Lycaenidae of California. J. Res. Lepid. 27: 1–70. 

BROWN, J.W. 1980. Hermes copper. Environment Southwest 491: 23. San 

Diego Natural History Museum. 

COMSTOCK, J.A. 1927. The Butterflies of California. Published by the 

author. 227 pp. 

COMSTOCK, J.A. and DAMMERS, CM. 1935. Notes on the early stages of 

three butterflies and six moths from California. Bull. South. Calif. Acad. 

Sci. 34: 120–141. 

DOS PASSOS, C.F. 1964. A synonymic list of the Nearctic Rhopalocera. 

Mem. Lepid. Soc. 1: 1–145. 

EDWARDS, W.H. 1870. Description of a new species of Lepidoptera found 

within the United States. Trans. Amer. Entomol. Soc. 3: 21. 

EHRLICH, P.R. and EHRLICH, A.H. 1961. How to Know the Butterflies. 

Wm. C. Brown Co. Publ., Dubuque, Iowa. 262 pp. 

EMMEL, T.C. and EMMEL, J.F. 1973. Butterflies of Southern California. 

Nat. Hist. Mus. Los Angeles County, Sci. ser. 26: 1–148. 

GARTH, J. and TILDEN, J.W. 1986. California Butterflies. University of 

California Press, Berkeley. 246 pp. + plates. 

HOWE, W.H. 1975. Butterflies of North America. Doubleday and Co., Inc., 

Garden City, New York. 633pp. 

MILLER, L.D. and BROWN, F.M. 1979. A revision of the American coppers 

(Lepidoptera: Lycaenidae). Bull. Allyn Mus. 51: 22–23. 

ORSAK, L.J. 1977. Butterflies of Orange County. Univ. Calif. Irvine, Mus. 

Syst. Biol. Res. ser. 4. 349 pp. 

RINDGE, F.H. 1948. Contributions toward a knowledge of the insect fauna of 

Lower California. No. 8. Lepidoptera: Rhopalocera. Proc. Calif. Acad. 

Sci. 24: 303. 

SCOTT, J.A. 1986. The butterflies of North America. Stanford University 

Press, Stanford, Calif. 583 pp. 

THORNE, F.T. 1963. The distribution of an endemic butterfly, Lycaena 

hermes. J. Res. Lepid. 2: 143–150. 

WRIGHT, W.G. 1906. Butterflies of the West Coast of the United States. 

Whitaker and Ray Co., San Francisco, Calif. 257 pp. 

WRIGHT, W.G. 1930. An annotated list of the butterflies of San Diego 

County, California. Trans. San Diego Soc. Nat. Hist. 6: 1–40.

Thome's Hairstreak, Mitoura thornei Brown 

John W. BROWN 

Entomology Department, San Diego Natural History Museum, P.O. Box 1390, San Diego, California 92112, U.S.A. 


Status and Conservation Interest: Status – possibly threatened. 

Thome's Hairstreak is a geographically isolated and 

ecologically distinct taxon that is restricted to a single mountain 

in southwestern San Diego County, California. Owing to its 

highly restricted distribution and the potential threats of habitat 

loss and degradation, Thorne's Hairstreak is recognized as a 

'category 2 candidate' for listing as endangered or threatened 

by the United States Fish and Wildlife Service. 

Taxonomy and Description: Thorne's Hairstreak was 

described by Brown (1983) as Mitoura thornei. Although most 

authors have treated it as a distinct species (e.g. Brown 1983; 

Garth and Tilden 1986; Ferris 1989; Ballmer and Pratt 1989), 

Shields (1984) suggests that it is a subspecies of Mitoura loki 

(Skinner), and Scott (1986) suggests that it is a subspecies of 

Mitoura grynea Huebner. Most likely, M. thornei is part of a 

'superspecies' complex in which the degree of morphological 

divergence and genetic isolation among taxa do not conform 

well with our fixed system of binomial (or trinomial) 

nomenclature. Regardless of taxonomic opinion, Thome's 

Hairstreak is ecologically distinct and geographically isolated 

from its nearest congeners. 

The nearctic genus Mitoura Scudder frequently is considered 

a subgenus of Callophrys Billberg. Hence, Thome's Hairstreak 

occasionally is referred to as Callophrys(Mitoura) thornei. 

Distribution: Thome's Hairstreak is restricted to Otay Mountain 

(= San Ysidro Mountains) in the southwestern portion of San 

Diego County, California. On Otay Mountain it is confined to 

places where the larval foodplant, Tecate cypress (Cupressus 

forbesi; Cupressaceae), grows. Although a significant stand of 

Tecate cypress occurs to the north in Coal Canyon, Orange 

County, California, and small populations are found to the 

south in northwestern Baja California, Mexico, Thorne's 

Hairstreak has not been documented from any of these localities 

(e.g. Orsak 1977; Brown 1983). 

Population Size: Otay mountain undoubtedly supports an 

extensive, nearly contiguous population (or set of populations) 

122 

of Thome's Hairstreak, although no quantitative data are 

available. Most of the Tecate cypress on the mountain has not 

been subject to encroachment by development or other human 

activities that result in loss of habitat, although chaparral fires 

frequently reduce or eliminate stands of the trees. 

Habitat and Ecology: Thorne's Hairstreak occurs only in 

southern interior cypress forest (Holland 1986) or where this 

habitat blends into other habitat types. The larval foodplant is 

Tecate cypress (Cupressus forbesi), a closed-cone conifer that 

occurs on mesic slopes and drainages in chaparral, and with 

which the adults are intimately associated. Thorne's Hairstreak 

is at least double brooded, with adults flying in late February 

through March, and again in June. Capture records indicate that 

the second brood may be only a partial one, as is the case with 

the closely related Mitoura loki. The emergence of laboratory 

reared individuals in August suggests the presence of a third 

brood in the fall, but this is yet to be documented in the field. 

The early stages of M. thornei closely resemble those 

described for M. siva (Edwards) (Coolidge 1924), M. loki 

(Comstock and Dammers 1932a), and M. nelsoni (Boisduval) 

(Comstock and Dammers 1932b), (see Ballmer and Pratt 1989). 

The eggs are echinoid and light green, and are laid singly on the 

new growth of the host plant. The egg stage lasts 7 to 14 days. 

Newly hatched larvae initially bore into the young stems of the 

host but later become external feeders. The larvae closely 

resemble the terminal twigs upon which they feed. Complete 

larval development, from hatching to pupation, requires 26–35 

days under laboratory conditions (Brown 1983). Pupation 

generally occurs in the duff or debris at the base of the host trees. 

No parasitoids or predators are recorded. 

Threats: Fire is an integral element in the natural history of 

Tecate cypress as it is the major factor that initiates cone 

opening and subsequent seed dispersal (Zedler 1977). Chaparral 

fires undoubtedly have caused fluctuations in the populations 

of both the cypress and the associated butterfly. Zedler (1977) 

indicates that Tecate cypress requires approximately 25 years 

to reach reproductive maturity. Hence, an increase in the 

incidence of fire (i.e. a frequency of less than every 25 years) 

could severely affect the host trees. In recent years, chaparral

fires have been common in the Otay Mountain area, usually as 

a result of carelessness by people. Chaparral fires probably 

represent the greatest threat to the cypress and its associated 

insect fauna, including Thorne's Hairstreak. 

Conservation: Currently, Thome's Hairstreak receives minimal 

protection under the California Environmental Quality Act. 

Under this legislation, impacts to sensitive plants and animals 

and sensitive habitat types that result from development 

activities, must be assessed to determine whether the adverse 

affects of habitat loss and fragmentation are 'significant'. If 

they are determined to be significant, mitigation measures are 

required to reduce impacts below a level of significance. 

Unfortunately, because invertebrates typically receive little 

attention in the environmental review process, these impacts 

typically are undocumented. 

Management recommendations for Thorne's Hairstreak 

include increased awareness of this species, particularly to 

those individuals and agencies involved in the environmental 

review process, and increased fire control and fire management 

on Otay Mountain. Much of Otay Mountain is under the 

ownership of the United States Bureau of Land Management. 

Consequently, much of this land is likely to remain an open 

space. Thome's Hairstreak presently is recognized by the 

United States Fish and Wildlife Service as a category 2 candidate 

species for listing as endangered or threatened; the Service 

recently received a petition to list the species as threatened. 

Such a listing would increase significantly the protection 

afforded this species. 

123 

References 

BALLMER, G. and PRATT, G. 1989 (1988). A survey of last instar larvae of 

the Lycaenidae of California. J. Res. Lepid. 27: 1–70. 

BROWN, J.W. 1983. A new species of Mitoura Scudder from southern 

California.J. Res. Lepid. 21: 245–254. 

COMSTOCK, J.A. and DAMMERS, CM. 1932a. Metamorphosis of five 

California diurnals (Lepidoptera). Bull. South Calif. Acad. Sci. 13:33–45. 

COMSTOCK, J.A. and DAMMERS, CM. 1932b. Metamorphosis of six 

California Lepidoptera. Bull. South Calif. Acad. Sci. 13: 88–100. 

COOLIDGE, K.R. 1924. The life history of Mitoura loki Skinner (Lepid.: 

Lycaenidae). Entomol. News 35: 199–204. 

FERRIS, C. 1989. Supplement to the catalogue/checklist of the butterflies of 

America north of Mexico. Mem. Lepid. Soc. 3: 1–103. 

GARTH, J. and TILDEN, J.W. 1986. California Butterflies. University of 

California Press, Berkeley. 246 pp. 

HOLLAND, R. 1986. Preliminary descriptions of the terrestrial natural 

communities of California. State of California, Department of Fish and 

Game. 156 pp. 

ORSAK, L.J. 1977. Butterflies of Orange County. Univ. Calif. Irvine, Mus. 

Syst. Biol. Res. ser. 4. 349 pp. 

SCOTT, J.A. 1986. The Butterflies of North America. Stanford University 

Press, Stanford, Calif. 583 pp. 

SHIELDS, O. 1984. Comments on recent papers regarding western 

Cupressaceae-feeding Callophrys (Mitoura). Utahensis 4: 51–56. 

ZEDLER, P.H. 1977. Life history attributes of plants and fire cycles; a case 

study in chaparral dominated by Cupressus forbesi. pp. 451–458. In: 

Mooney, H.A. and L.E. Conrad (tech. coords.), Proceedings of the 

Symposium on the Environmental Consequences of Fire and Fuel 

Management on Mediterranean Ecosystems. Palo Alto, Calif.

Sweadner's Hairstreak, Mitoura gryneus sweadneri (Chermock) 


Thomas C. EMMEL 

Department of Zoology, University of Florida, Gainesville, Florida 32611, U.S.A. 

Status and Conservation Interest: Status – rare, threatened. 

This local butterfly was described from the city of St. 

Augustine on the northeastern coast of Florida, a site which 

remains the centre of this rarity's distribution in the state. 

Virtually everywhere, it is threatened by reduced available 

habitat due to land clearing for housing and other development. 

In late 1986, after learning of the trend towards irreversible 

habitat loss for the Sweadner's Hairstreak, the mayor of St. 

Augustine led a drive to enact a city ordinance to protect the 

butterfly and its only known native foodplant. St. Augustine 

thus became the second city in the United States to protect a 

threatened butterfly site by law. (Pacific Grove on the central 

California coast protects the overwintering colonies of the 

common Monarch butterfly, Danaus plexippus.) 

Taxonomy and Description: The butterfly was described in 

1944 by noted lepidopterist Frank H. Chermock. Chermock 

named it as a distinct species, Mitoura sweadneri. A.B. Klots 

(1951) listed it as a subspecies of M. gryneus in his popular field 

guide, and it has been referred to as a subspecies ever since. In 

1993, Emmel, Baggett and Fee will publish a paper elevating 

the taxon to full specific status again, based on life history 

characteristics, hybrid crossing studies, and genitalic differences. 

Distribution: Sweadner's Hairstreak is considered very local 

and usually very rare. It has been found in several sites in the 

present city limits of Jacksonville, south through St. Augustine 

to New Smyrna Beach on the east coast of Florida, and several 

colonies are now known on the Gulf coast of Florida around 

Crystal River north to Cedar Key. Each colony is quite separated 

geographically from the others. 

Population Size and Status: All colonies are very isolated and 

generally quite small in number (from 12 to 50 adults normally 

being present). The adults of both sexes appear to occupy a 

small home range, generally on a single large cedar tree or on 

several closely adjacent cedars. However, the males are strongly 

territorial and chase other adults that enter their defended areas. 

124 

Thus, a maximum of three or four adults may be found spaced 

around a single tree. 

Habitat and Ecology: The butterfly lives in sandy coastal 

habitat occupied by its only known native foodplant, the Southern 

Red Cedar (Juniperus silicicola). These coniferous trees support 

both the immature stages and the adults, although the adults 

depend on locally growing wildflowers near the base of the 

trees for nectar. 

The egg of Sweadner's Hairstreak is pale green, and the 

young larva is superbly coloured to match the plant and scalelike 

leaves of the cedar twig. The mature larva is flat and sluglike 

in shape, and has a deep green ground colour with pale 

green diagonal stripes high on the sides. The pupa is dark brown 

and serves as the hibernating stage for the species during the 

short north Florida winter. The species has three annual broods 

(spring, summer, fall). 

For unknown reasons, the butterfly seems to be rather 

tightly adapted to coastal climates and is rarely found very far 

inland, despite the much wider inland distribution of Southern 

Red Cedars. 

Threats: The greatest threat to the species is continued 

development of the coastal habitat for housing resulting from 

urban expansion and the desire for recreational beach homes. 

Other threats include road construction, dump clearing, and 

related land disturbances which have destroyed many of the 

formerly available habitats for the species. Finally, even if the 

red cedar trees are left in the area by the government ordinance 

requirements, the multiple-brooded adults may not survive if 

the proper wildflowers are not left in close association with the 

surviving trees. Without nectar, the adults die within a day or 

two, prior to reproducing. 

Conservation: The conservation needs of Sweadner's 

Hairstreak lie not only in legal protection of its native foodplant 

but also in protecting strips of native vegetation associated with 

those Southern Red Cedar trees. The butterfly can survive in a 

remarkably small area (even 20m 2 ), as long as nectar sources 

are left in association with the red cedar trees. Popular support

in the cities of Jacksonville, St. Augustine, and St. Augustine 

Beach has led to the passing of several civic ordinances since 

1986 by those urban areas to preserve the butterfly and its 

foodplant. Additionally, the cities are reducing pesticide spraying 

for mosquito control in butterfly colony areas. There is 

considerable hope now that the loss of developmentally attractive 

coastal areas having Southern Red Cedar and Sweadner's 

Hairstreak colonies may be arrested (Emmel 1987). 

125 

References 

EMMEL, T.C. 1987. Delicate balance: Sweadner's Hairstreak butterfly 

(Mitoura gryneus sweadneri). Florida Wildlife 41 (2): 39. 

KLOTS, A.B. 1951. A Field Guide to the Butterflies. Houghton Mifflin Co., 

Boston, Massachusetts. 349 pp.

Bartram's Hairstreak, Strymon acis bartrami (Corns tock & 

Huntington) 


Thomas C. EMMEL and Marc C. Minno 


Status and Conservation Interest: Status–rare; indeterminate 

(Red List). 

Bartram's hairstreak is classified as a threatened species 

because of its restricted distribution, low abundance, and recent 

loss of habitat. 

Taxonomy and Description: This subspecies is restricted to 

Florida and was described in 1943 by Comstock & Huntington. 

It is characterised by the heaviest white markings of all the 

subspecies of Strymon acis. The species ranges from Florida 

throughout the West Indies to Dominica (Riley 1975). Typical 

Strymon acis from Antigua and Dominica form the largest race. 

5. a. petioni Comstock & Huntington occurs on Hispaniola and 

has gray undersides. 5. a. mars Fabricius from the Virgin 

Islands, St. Kitts, and Puerto Rico has brown on the underside 

and a large orange area. S. a. gossei Comstock & Huntington 

from Jamaica and the Cayman Island resembles the last 

subspecies in the narrowness of the white bands, but the 

submarginal zone is broader. The Bahamas subspecies armouri 

Clench has narrow white bands and a narrow submarginal 

marking area. The Cuban subspecies casasi Comstock & 

Huntington is very similar to bartrami in Florida, with heavy 

white markings. The scientific and common name of Bartram's 

Hairstreak for the Florida population honours the memory of 

William Bartram, an early Florida naturalist whose journeys 

through the state first brought wide recognition of its unique 

natural history. 

Distribution: This attractive little hairstreak is found only in 

southern Dade County, including the Everglades National 

Park, and on Big Pine Key in the Florida Keys. Hurricane 

Andrew may have severely impacted the mainland population 

of this butterfly on 24 August 1992, as it moved directly through 

the known colonies of this butterfly when crossing southern 

Dade County. 

Population Size: Thirty field surveys, conducted by Hennessey 

and Habeck (1991) between 23 May and 16 December 1988, 

found an average of 0.5 adults per hectare in pineland habitats 

126 

of the Everglades National Park and 0.3,1.0 and 2.7 individuals 

per hectare at three locations on Big Pine Key. The status of 

mainland populations following Hurricane Andrew is not yet 

known at the time of writing (December 1992). 

Habitat and Ecology: Bartram's Hairstreak is a small grayish 

butterfly with two pairs of delicate tails on the hindwings. The 

undersides of the hindwings have a highly distinctive pattern of 

white spots and lines, plus a red eyespot at the base of the tail. 

It occurs in open tropical pinelands that have an abundance of 

the larval hostplant. Females lay their eggs singly on the 

flowers of Woolly Croton, Croton Hnearis. The larvae feed on 

the flowers and leaves of the host. Adults frequently perch on 

Woolly Crotons and visit nearby flowers for nectar. Several 

generations are produced each year. 

Threats: The primary threats to this species are housing 

developments and agricultural clearing in the heavily crowded 

southern Dade County area. However, the natural disaster 

presented by the tremendously destructive Hurricane Andrew 

winds, on 24 August 1992, may have destroyed or severely 

affected all the remaining habitat areas on the mainland. On 

both Big Pine Key in the Florida Keys and in southern Dade 

County, open tropical pinelands are also threatened periodically 

by fire. 

Conservation: Additional surveys are badly needed now to 

monitor the abundance and distribution of Bartram's Hairstreak 

in both southern Dade County and the Florida Keys. Ecological 

studies should be conducted to identify the species' habitat 

requirements. Prescribed burning of pinelands may be necessary 

to maintain habitat for this species (Hennessey and Habeck 

1991), but land managers should take care not to burn large 

tracts entirely, lest populations of Bartram's Hairstreak and 

other rare butterflies be destroyed by the fire. Careful 

management planning is needed for the remaining habitat in the 

Keys and in the Everglades National Park. The species is an 

attractive butterfly and could be used in publicity campaigns 

advocating preservation of these increasingly rare tropical 

pineland habitats in Florida (Minno and Emmel 1993).

References 

HENNESSEY, M.K. and HABECK, D.H. 1991. Effects of mosquito adulticides 

on populations of non-target terrestrial arthropods in the Florida Keys. 

Unpublished final report. U.S. Fish and Wildlife Service and University 

127 

of Florida Cooperative Wildlife Research Unit, Gainesville, Florida. 

MINNO, M.C. and EMMEL, T.C. 1993. Butterflies of the Florida Keys. 

Scientific Publishers, Gainesville, Florida. 168 pp. 

RILEY, N.D. 1975. A Field Guide to the Butterflies of the West Indies. 

Demeter Press, Inc., Boston, Massachusetts. 224 pp.

The Avalon Hairstreak, Strymon avalona (W.G. Wright) 

Thomas C. EMMEL and John F. EMMEL 

Department of Zoology, University of Florida, Gainesville, Florida 32611, U.S.A., and 26500 Rim Road, Hemet, California 

92544, U.S.A. 


Status and Conservation Interest: Status – insufficiently 

known (Wells et al. 1983, Red List). 

This hairstreak is one of the world's most restricted species, 

being found only on Santa Catalina Island just slightly more 

than 20 miles off the coast from Los Angeles, southern California. 

The limited land available for development in southern 

California, including Santa Catalina, means that ultimately this 

species is threatened by development. Proximate dangers include 

overgrazing and other destruction by cattle and feral livestock 

such as goats. 

Possible hybridisation with the related S. melinus (which 

was recorded from Santa Catalina for the first time in 1978) has 

been suggested, but such claims are probably inaccurate (Gall 

1985; Gorelick 1987). 

Taxonomy and Description: This species was described by 

W.G. Wright in 1905 from the vicinity of Avalon on Catalina 

Island. It does not occur on any of the other Channel Islands, 

where a related species, the Gray Hairstreak (Strymon melinus 

Hü+bner) may be found. No geographic variation has been noted 

in Strymon avalona on Santa Catalina Island. 

Distribution: This species occurs only on Santa Catalina 

Island, off the coast of southern California. 

Population Size: The Avalon Hairstreak is common in select 

localities such as the hills around Avalon. The butterfly is 

generally distributed in various localities from an elevational 

range of sea level to 65m. A number of colonies have been 

reported along the road to Renton Mine, Pebbly Beach, Jewfish 

Point, the Isthmus, and hills to the west of Avalon. 

Habitat and Ecology: The species prefers chaparral and grasscovered 

slopes at relatively low elevations. Captures have been 

reported in every month of the year, with the first brood primarily 

occurring from mid-February through April, a second brood in 

July and August, and a third brood flying from September to 

November. The adults frequently perch on bushes and grassy 

areas in chaparral, and visit the flowers of common sumac (Rhus 

alurind) and the giant buckwheat (Eriogonum giganteum Wats.). 

128 

Eggs are laid singly, usually in the terminal buds or on 

immature flowers of the foodplant, Lotus argophyllus (Gray) 

Greene var. ornithopus (Greene) Ottley, and Lotus scoparius 

(Nutt.) Ottley, in the Leguminosae. Mature larvae show 

considerable variation in ground colour, ranging from a pale 

apple green to pale pink. The entire body of the larva, except the 

cervical shield, is covered with short white fine hairs. The pupa 

is pale pinkish-brown or wood brown with markings of various 

shades of olive-green. Pupation occurs at the base of the 

foodplant, with the usual support of a delicate silk girdle 

(Emmel, Emmel and Mattoon in press). 

Threats: The species is not known to be endangered at the 

present time, although increased urbanisation or changes in 

vegetation cover due to overgrazing by domestic and feral 

livestock constitute potential threats every year. No special 

conservation measures have been taken on Santa Catalina 

Island for this butterfly. 

Conservation: The widespread continental species Strymon 

melinus has not yet become successfully established on Santa 

Catalina Island. Likewise, this rare endemic Strymon avalona 

has not established itself on any of the other Channel Islands or 

on the mainland of North America. The situation should be 

monitored yearly to better understand why Strymon melinus in 

particular, has not managed to establish itself yet on Santa 

Catalina Island where many of its potential foodplants are 

growing. 

References 

EMMEL, T.C., EMMEL, J.F. and MATTOON, S.O. In press. The Butterflies 

of California. Stanford University Press, Stanford, California. 

GALL, L.F. 1985. Santa Catalina's endemic Lepidoptera. II. The Avalon 

Hairstreak, Strymon avalona, and its interaction with the recently 

introduced Gray Hairstreak, Strymon melinus (Lycaenidae). In: Menke, 

A.S. and Miller, D.R. (Eds). Proc. 1st Symposium on Entomology of the 

California Islands. Santa Barbara Museum of Natural History, pp. 95–104. 

GORELICK, G.A. 1987. Santa Catalina's endemic Lepidoptera. II. The 

Avalon Hairstreak, Strymon avalona (Lycaenidae): an ecological study. 

Atala 14 (1986): 1–12. 


Red Data Book. IUCN, Gland, Switzerland.


The Atala Butterfly, Eumaeus atala florida (Röber) 

Thomas C. EMMEL and Marc C. MINNO 


Status and Conservation Interest: Status – out of danger; 

vulnerable (Red List). 

The Atala butterfly is one of the most strikingly coloured 

butterflies in Florida and it is desirable to collectors. Its 

conservation interest arises from the fact that the species was 

feared extirpated in the early 1970s after the few known 

colonies died out, but then made a spectacular recovery starting 

in 1979 from a single small colony rediscovered in the Miami 

area. The species has recovered much of its former range and is 

now even considered a pest of ornamental cycad plantings! It is 

assessed as relatively common in parts of southern Florida 

(Bowers and Larin 1989; Minno and Emmel 1993). 

This species was the subject of one of three full lycaenid 

entries in the IUCN Invertebrate Red Data Book (Wells et al. 

1983), and one of six butterflies discussed in the Invertebrate 

volume of Franz (1982). It gave its name 'Atala' to a journal 

published by the Xerces Society. 

Taxonomy and Description: The subspecies was described by 

Röber in 1926, on the basis of having more extensive bluegreen 

on the upper side and larger spots of that colour on the 

underside of the hindwings. The typical subspecies, Eumaeus 

atala atala Poey (1832), occurs in Cuba, on Andros Island, and 

on Great Abaco Island in the Bahamas (Riley 1975). Some 

lepidopterists have questioned if the Florida population is 

sufficiently distinct to be called a separate race (Clench 1977). 

Distribution: The Atala Butterfly is found in southern Florida 

from Broward County in the vicinity of Fort Lauderdale south 

to southern Dade County, and it is recorded historically from 

Elliott Key and Key Largo (the most recent record for the latter 

Key being 5 June 1960). Most of the populations in southern 

Dade County were probably severely impacted by Hurricane 

Andrew on 24 August 1992. 

Population Size: The Atala Butterfly was once abundant in the 

rimrock areas of the southern mainland of Florida, but largescale 

harvesting of the host plant, coontie (Zamia pumila, 

Cycadaceae) for starch in the late 1800s greatly reduced the 

129 

number of coontie plants. Urbanisation and development of the 

coastal habitat favoured by the Atala also had a large impact. By 

1965, the Atala had been reduced to a single known population 

in Hugh Taylor Birch State Park. After this colony died out, the 

Atala was feared extirpated from Florida. However, in the late 

1970s, another colony was found on Virginia Key in the Miami 

area. Conservationists such as Roger Hamler of Dade County 

Parks set out potted coontie plants on which females laid eggs. 

Plants with eggs were then moved to other locations and new 

colonies were started. Rawson (1961) demonstrated the 

possibility of translocating E. atala by liberating adults in a new 

site. 

The Atala has made a spectacular recovery and is now found 

in urban and natural areas around Fort Lauderdale and Miami, 

and has been successfully introduced into the Everglades 

National Park. Some plant nurseries and botanical gardens 

currently consider the Atala a pest species, as the larvae are 

capable of defoliating lamia species used in landscaping. It is 

not known whether our current population is of original Florida 

stock or the result of a new introduction from the Bahamas or 

Cuba. 

The very few records of Atala from the Florida Keys include 

only Elliot Key and Key Largo (Schwartz 1888). Small (1913) 

listed coontie among the plants found in the Keys, but we have 

not encountered it on any of the islands. The early pioneers 

probably extirpated coontie (and thereby the butterfly) from the 

Keys, as it was a readily available source of starch. The only 

modern record of the Atala from the Keys is the single capture 

of a male on 5 June 1960 in the City of Key Largo. The status 

of the populations on the mainland was probably severely 

impacted by Hurricane Andrew on 24 August 1992, and 

subsequent surveys have not yet been done to ascertain the 

species' status on the mainland. 

Habitat and Ecology: The Atala is found in tropical pinelands 

and hardwood hammocks in close association with the larval 

foodplant, coontie (Zamia pumila, Cycadaceae). The Atala 

adults are the largest lycaenids in Florida and occur all year 

round.It is one of the most strikingly coloured butterflies on its 

ventral surfaces, with jet black wings, iridescent blue spots and 

a red patch on the underside of the hindwings. The upper wings

are black with iridescent green in males and iridescent blue in 

females, while the abdomen is bright red. The adults have a 

slow fluttering flight pattern. Males perch on the leaves of 

shrubs and make circular flights around the perch site like other 

hairstreaks. Both sexes often visit flowers. 

The white eggs are laid in clusters on the young growth of 

coontie. Larvae are bright red with a yellow spot, and are 

probably distasteful to birds or other predators. Pupae are 

brown with small dark spots and hang from the substrate by a 

silk girdle. Droplets of bitter tasting liquid are exuded over the 

cuticle of the pupae. 

Threats: The Atala is listed as a species of special concern in 

Florida because of its restricted distribution and cyclic 

fluctuations in abundance. This butterfly is currently known 

from more colonies on the mainland than were recorded before 

1965, but the Atala has lost habitat areas formerly occupied in 

the Florida Keys. Hurricane Andrew probably severely impacted 

populations of the Atala in southern Dade County. 

Continued threats occur in both wild and urban areas. The Atala 

occurs in tropical hardwood hammocks and pinelands in close 

association with the host plant. It also now uses some urban 

areas such as gardens and nurseries where the foodplant is 

grown as an ornamental. Thus this species is exposed to both 

habitat clearing and burning in the wild (although prescribed 

burning of pinelands may be necessary to maintain habitat). 

Spraying or other pest control measures in urban habitats pose 

another threat. For example, before Hurricane Andrew, Fairchild 

Botanical Garden in southern Dade County regularly used Bti 

sprays to control the infestation of the Atala Butterfly on its 

valuable cycad collection (from the 1980s through summer 

1992). 

130 

Conservation: The unexpected occurrence of wide habitat 

destruction by Hurricane Andrew in the summer of 1992 

necessitates new surveys to monitor the distribution and 

abundance of the Atala in Florida. Outside of the affected urban 

areas, prescribed burning of pinelands may be necessary to 

maintain habitat for Atalas in natural settings. The species is 

probably permanently lost from the Florida Keys, unless 

restoration plantings of the coontie host plants are done on Big 

Pine Key where some substantial tropical pinelands remain 

(there are no pinelands left on Elliot Key or Key Largo, its 

originally recorded habitat in the Florida Keys). Taxonomic 

studies should be conducted to determine if the current taxon 

present in south Florida is the same as that present before 1965, 

and to define the relationship of E. atala florida to E. atala atala 

in the Bahamas. 

References 

BOWERS, M.D. and LARIN, Z. 1989. Acquired chemical defense in the 


CLENCH, H.K. 1977. A list of the butterflies of Andros, Bahamas. Ann. 

Carnegie Mus. 46: 173–194. 

FRANZ, R. (Ed.) 1982. Rare and Endangered Biota of Florida. 6. Invertebrates. 

University Presses of Florida. 

MINNO, M.C. and EMMEL, T.C. 1993. Butterflies of the Florida Keys. 

Scientific Publishers, Gainesville, Florida. 168 pp. 

RAWSON, G.W. 1961. The recent rediscovery of Eumaeus atala (Lycaenidae) 

in southern Florida. J. Lepid. Soc. 15: 237–244. 

RILEY, N.D. 1975. A Field Guide to the Butterflies of the West Indies. 

Demeter Press, Inc., Boston, Massachusetts. 224 pp. 

SCHWARTZ, E.A. 1888. Notes on Eumaeus atala. Insect Life 1: 37–40. 

SMALL, J.K. 1913. Flora of the Florida Keys. Published by the author, New 

York. 162 pp. 

WELLS, S.M., PYLE, R.M. and COLLINS, N.M. 1983. TheIUCNInvertebrate 

Red Data Book. IUCN, Gland.

Smith's Blue, Euphilotes enoptes smithi (Mattoni) 


Department of Zoology, University of Florida, Gainesville, Florida 32611, U.S.A., and 26500 Rim Road, Hemet, 

California 92544, U.S.A. 


Status and Conservation Interest: Status – endangered (Red 

List). 

This butterfly is known only from the coastal fog belt of 

Monterey County in California, where it inhabits the immediate 

coast of the Big Sur country. A member of a widely distributed 

western U.S. species, Euphilotes enoptes smithi is an endemic 

California subspecies whose coastal habitat has suffered a 

number of disturbances, including beach recreation and offroad 

vehicles. Montane habitats, in general, have suffered less 

(Arnold 1983a,b). 

Taxonomy and Description: The subspecies was described 

(Mattoni 1954) as different from the numerous other subspecies 

of Euphilotes enoptes in part because of the broad black 

marginal borders on the lustrous blue upper wings of the males. 

Females are brown above, with a band of red-orange marks 

across the hindwings. The overall distinguishing features from 

other subspecies are the light undersurface ground colour and 

prominent black markings with a faint black terminal line 

(Emmel, Emmel and Mattoon in press). 

The male is distinguished by the broad marginal border of 

the hindwings. The ventral surface of both males and females 

has a faint terminal line and a light ground colour with large 

prominent spots. 

Distribution: This species is confined to coastal Monterey 

County from Big Sur and the mouth of the Salinas River 

southward to Del Rey Creek and an area several miles north of 

the San Luis Obispo County line. It ranges from near sea level 

to approximately 65m elevation. The type locality is at Burns 

Creek, State Highway 1, in Monterey County. 

Population Size: Populations are small in number and probably 

have never been particularly large because of the relatively 

restricted habitat. 

Habitat and Ecology: The Smith's Blue butterfly occurs on 

cliffs, steep slopes, and road cuts along the immediate coast, 

131 

within the northern coastal scrub plant community. It also is 

found extensively on coastal and inland sand dunes, and 

occasionally on serpentine grassland. 

The adults emerge between mid-June and early September, 

corresponding with the blooming of the buckwheat plants on 

which they feed, rest, sun, and mate. While each adult lives for 

only about one week, individual emergences are scattered over 

the extended flight period. Females deposit eggs singly in 

buckwheat flowers. Larvae hatch 4–8 days later and go through 

five instars before pupating in flowerheads or in the litter and 

sand at the base of the plant. Pupae hang in place from September 

until adults emerge the following year. The verified host plant 

for the larvae is Eriogonum parvifolium Sm. in Rees. (Pratt and 

Ballmer 1989). Larvae also feed on E. latifolium, and differences 

in plant phenology at different sites may represent a potential 

isolating mechanism for populations. 

Threats: The primary threat to the conservation and recovery 

of the Smith's Blue butterfly is the wide variety of man-made 

disturbances of the coastal habitat. Dunes are widely threatened 

by beach recreation, off-road vehicles, housing developments, 

and road construction. Additionally, non-native plants such as 

iceplant and Holland dunegrass invade the dunes and displace 

native buckwheat. At Fort Ord, the sand dunes have been 

damaged by military vehicles and infantry exercises. At the 

Seaside-Marina dune system and the Del Monte Forest, their 

habitat has been destroyed by sand mining. More than half the 

dune habitat present at the turn of the century had been destroyed 

by about 1980 (Powell 1981). 

Conservation: A conservation plan for Smith's Blue designed 

by Arnold (1983b) was one of the first detailed prescriptions for 

a North American lycaenid. Prevention of further habitat loss or 

change was the prime need, and Arnold' s plan aimed to maintain 

known populations of E. e. smithi by coordinating habitat 

preservation, rehabilitation and management. 

Five categories of action were proposed: (1) preservation 

and protection of existing habitats; (2) implementation of shortterm 

and long-term management; (3) development of monitoring 

programmes to census selected populations annually to assess

the effects of management efforts; (4) promotion of public 

awareness of the butterfly and its habitat, and (5) enforcement 

of laws and regulations to protect the butterfly. 

This species was listed as Endangered on June 1, 1976. In 

1977, the U.S. Army established a butterfly preserve at Fort 

Ord, and some of the non-native plants have been removed 

there as well as native plants re-established. For remnant sand 

dune habitats at Sand City in Monterey County, the city agreed 

to complete a conservation plan before proceeding with any 

work in the dune areas that have been zoned for housing 

development. 

132 

References 

ARNOLD, R.A. 1983a. Ecological studies on six endangered butterflies 

(Lepidoptera: Lycaenidae): Island biogeography, patch dynamics, and 

the design of habitat preserves. Univ. Calif. Publns Entomol. 99: 1–161. 

ARNOLD, R.A. 1983b. Conservation and management of the endangered 

Smith's Blue Butterfly, Euphilotes enoptes smithi (Lepidoptera: 

Lycaenidae).J. Res. tepid. 22: 135–153. 

EMMEL, T.C., EMMEL, J.F. and MATTOON, S.O. In press. The Butterflies 

of California. Stanford University Press, Stanford, California. 

MATTONI, R.H.T. 1954. Notes on the genus Philotes. 1. Description of three 

new subspecies and a synoptic list. Bull. South. Calif. Acad. Sci. 53: 

157–165. 

POWELL, J.A. 1981. Endangered habitats for insects: California coastal sand 

dunes. Atala 6: 41–55. 

PRATT, G.P. and BALLMER, G.R. 1989. A survey of the last instar larvae of 

the Lycaenidae (Lepidoptera) of California. J. Res. Lepid. 27: 1–81.

The El Segundo Blue, Euphilotes bernardino allyni (Shields) 

Country: U.S.A.(California). 


List). 

This subspecies is a sand obligate ecotype found only on the 

El Segundo sand dune ecosystem of the coastal plain of western 

Los Angeles. Its remaining three discrete colonies are surrounded 

by a dense urban area that remains both a fast growing and fast 

denaturing area. As a federally listed endangered species, the El 

Segundo Blue (ESB) is significant in conferring an umbrella of 

protection on at least ten other species of plant and animals that 

are restricted to this sand dune system and to more than 20 

species restricted to the coastal sand dunes of southern California 

and northern Baja California. 


Shields (1975) from the El Segundo sand dunes of Los Angeles 

county just prior to its listing among the first group of butterflies 

to be legally recognised as endangered by the Endangered 

Species Act of 1973. The taxon was originally classified as a 

subspecies of E. battoides, but both Shields (Shields 1975, 

Reveal and Shields 1988) and Mattoni (1989) independently 

diagnosed allyni as more logically belonging to the bernardino 

group of four related but clearly separated subspecies which 

they recognised as a distinct species. 

Distribution: The ESB was historically restricted to the El 

Segundo sand dunes that covered about 1200ha. Today three 

discrete colonies exist on fragments that still maintain some 

characteristics of the natural community. 

Population Size: The largest population is on property of the 

Los Angeles International Airport (LAX), where 80ha were 

recently set aside by the city of Los Angeles as a biological 

preserve, in part to satisfy biotic requirements of the butterfly. 

When serious study started in 1984, total population size was 

about 500. Following restoration efforts the standing population 

in 1990 was about 4000 (Mattoni 1988, 1990a, b, 1992). The 

second largest colony is on a 0.6ha lot on the Chevron Oil 

refinery property. The population prior to study was about 2000 

and has since decreased to under 500 (Arnold 1983,1986). A 

R.H.T. MATTONI 

9620 Heather Road, Beverly Hills, California 90210, U.S.A. 

133 

small colony of a few hundred butterflies persists on a 0.2ha 

isolated dune fragment at Redondo Beach. The latter was 

discovered in 1984. 

Habitat and Ecology: The species is sand obligate and adapted 

to a single foodplant, the coastal buckwheat, Eriogonum 

parvifolium, (Polygonaceae). It has one generation per year, 

adults appearing from June through mid-August. Excepting the 

fossorial diapausing pupal stage, the entire life cycle of the 

butterfly is associated with flowerheads of the buckwheat 

including egg deposition, larval growth, nectaring, mating and 

dying (Mattoni 1991). The larvae are ant associated, but the 

relationship is facultative and the only ant now noted in 

association is the exotic Argentine ant, Iridomyrmex humilis. 

Adult butterflies were highly sedentary at the tiny Chevron site 

(Arnold 1983), but nothing is known of movements at LAX 

beyond observed concentrations near dense patches of foodplant. 

Threats: Pratt (1987) first recognised that the major threat to 

the ESB at LAX was the presence of a dense common buckwheat 

stand which had been planted during a poorly conceived 

restoration effort 20 years earlier. This earlier flowering species 

provided sustenance to a guild of non-diapausing Lepidoptera 

that later migrated to the flowering coastal buckwheat. 

Overwhelming numbers, cannibalism, and shared parasites 

devastated the ESB. Removal of part of the exotic plant biomass 

was believed a major cause of the recent resurgence of ESB 

populations (Mattoni 1991). 

The ESB at the Chevron site, while serving as a linchpin in 

the oil company's advertising campaign to show its concern for 

nature, precipitously declined (Arnold 1986). The cause was 

probably an intensive mark-release programme to assess 

population size, coupled with extreme trampling across a small 

area. The Redondo Beach site is completely untended, but is not 

suitable for building development and is thus not threatened by 

developers. 

Conservation: Listing the ESB under the federal Endangered 

Species Act provides one of the greatest success stories of that 

legislation. For the principal colony at LAX, the 1976 listing 

happened just in time to halt a plan to develop almost the entire

120ha of dunes as a golf course. Following a protracted planning 

and political process, the preserve area increased in size. Under 

leadership of the local Councilwoman, the Mayor and Airport 

Commission agreed to set aside 80ha of the highest quality land 

as a permanent preserve with the proviso that the golf course on 

the remaining 40ha be designed with all rough areas re-planted 

habitat. 

In the meantime, the Airport Commission provided $180,000 

to begin restoration of the dunes ecosystem. Work on the dunes 

is now in its third year. 

By contrast, the Chevron company has emphasised creating 

conditions to maximise survival of the ESB, at the expense of 

the ecosystem. The main effort has been directed towards 

creating a butterfly garden; because the site is so small, this 

approach has some validity. On the other hand long-term 

stability would be better insured with a community mimicking 

that of the historic system, and not a near monoculture of the 

foodplant with more glamorous advertising potential. Oppewall 

(1975) documents efforts by local collectors who successfully 

convinced Chevron to set aside the area as a preserve. For this, 

the company is to be commended. 

134 

References 


(Lepidoptera, Lycaenidae): Island biogeography, patch dynamics, and 

the design of habitat preserves. Univ. Calif. Publns Entomol. 99. 161pp. 

ARNOLD, R.A. 1986. Private and government funded conservation programs 

for endangered insects in California. Natural Areas Journal 5: 28–39. 

MATTONI, R.H.T. 1988. Captive propagation of California endangered 

butterflies. Report to Calif. Dept. Fish and Game. Contract C-1456. 

MATTONI, R.H.T. 1989. The Euphilotes battoides complex: Recognition of 

a species and description of a new subspecies. J. Res. Lepid. 27:173–185. 

MATTONI, R.H.T. 1990a. Habitat evaluation and species diversity on the 

LAX El Segundo sand dunes. Rept. to the LAX board of airport 

commissioners. 

MATTONI, R.H.T. 1990b. Unnatural acts: succession on the El Segundo 

sand dunes in California. In: Hughes, H.G. and T.M. Bonnickson (Eds.) 

Proc. first SER Conference, Berkeley, CA 1989. Soc. Ecol. Restoration, 

Madison WI 53711. 

MATTONI, R.H.T. 1992. The endangered El Segundo blue butterfly. J. Res. 

Lepid. 29: 277–304 (1990). 

OPPEWALL, J.C. 1975. The saving of the El Segundo Blue. Atala 3: 25–8. 

PRATT, G. 1987. Competition as a controlling factor of Euphilotes battoides 

allyni larval abundance. Atala 15: 1–9. 

SHIELDS, O. 1975. Studies on North American Philotes IV. Taxonomic and 

biological notes and new subspecies. Bull. Allyn Museum No. 28. 36pp. 

SHIELDS, O. and REVEAL, J. 1988. Sequential evolution of Euphilotes 

(Lycaenidae, Scolitantidini) on their plant host Eriogonum (Polygonaceae, 

Eriogonoideae). J. Linn. Soc. 33: 51–91.

The Palos Verdes Blue, Glaucopsyche lygdamus palosverdesensis 

Perkins and Emmel 

Country: U.S.A. (California) 

Status and Conservation Interest: Status – probably extinct 

(Red List). 

The subspecies, now certainly extinct, was a coastal bluff 

ecotype found only on the southern half of the Palos Verdes 

peninsula in southern Los Angeles county. The species has high 

conservation value, nonetheless, as it has not been officially 

delisted under the theory that extinction is not certain. Hence all 

building projects in the species distribution area – and there are 

many – must recognise habitat value. At least one major 

development is currently stalled waiting approval of a plan to 

protect foodplants. The great benefit of the situation is provision 

of protection to other endangered species, now unlisted, 

occurring in the habitat. 


Perkins and Emmel (1977) from Los Angeles county just prior 

to its listing among the second group of butterflies to be legally 

recognised as endangered by the Endangered Species Act. The 

taxon was diagnosed as a subspecies distinct from, Glaucopsyche 

lygdamus australis, the southern blue, by exclusive use of the 

milk vetch Astragalus leucopsis, very fast flight, and several 

wing characteristics. 

Distribution. By the time of its discovery by Perkins in the 

early 1970s, the Palos Verdes Blue (PVB) was already restricted 

to a few fragments retaining some natural characteristics. In 

1977 at least nine discrete colonies existed. The last known 

occurrence was in 1983. 

Population Size: The largest populations known during the 

brief time span the PVB was extant were at Atala Vista Terrace 

(type locality) and in the scrub extending from Palos Verdes 

Drive east to Friendship Park. The former locality was built 

over in 1978. Population sizes were never estimated and by the 

early 1980s numbers were extremely low, probably less than 

100 adults among all the remaining fragments at that time 

(Arnold 1985). In spring of 1982 at Hesse Park, I counted six 

adults on the best day, with some 20 plants. Each plant had at 



135 

least 100 eggs, and one plant over 500. Foodplant availability 

was limiting, probably due to spring disking for fire control. 

Habitat and Ecology: The PVB was a coastal, sage-associated 

ecotype, restricted in foodplant use to the milk vetch. The vetch 

is restricted to the fog belt across southern exposures at elevations 

between 100 and 300 metres. The historical area probably 

occupied by the PVB was no more than 4000ha. The flora of the 

northwest slopes of the peninsula included the low shrub 

legume, Lotus scoparious, foodplant of the sister subspecies, 

G. lygdamus australis. Whether australis was parapatric to 

palosverdesensis is unknown. 

The butterfly was single brooded with adult flight in February 

and March. Eggs were usually laid on flowerheads of the 

foodplant, but when foodplant numbers were diminished just 

prior to extinction, eggs were laid over the entire plant. Larvae 

usually fed on seed within developing seedpods of the vetch and 

were ant tended in the last two instars. The final known 

generation, observed at Hesse Park, had larvae feeding on 

leaves as well since the flowerheads and seeds were exhausted. 

Three other Lycaenid butterflies were associated with the 

flowerhead/seedpod guild: Strymon melinus; Leptotes marina; 

and Everes amyntula. The first two are polyphagous, have 

many alternative foodplants, and are widespread species. The 

latter, with the PVB is a monophage restricted to the vetch. It 

has been extirpated from the Palos Verdes peninsula, although 

the last specimens were sighted in 1986 (Jess Morton, Tony 

Leigh, pers. comm.). 

Threats Leading to Extinction. Arnold (1986) reported the 

decline of the species and speculated that it had become extinct. 

Intensive search has been conducted by several local 

lepidopterists every year since 1983 without success. The 

proximate cause of extinction was denaturing of the land by 

development and fire suppression tactics. The historic population 

must have been continuous over the 4000ha coastal scrub 

habitat. With intensive development from 1950 the habitat was 

fragmented, although one 500ha section remains. Clearing 

practice so degraded this that the construction at Hesse Park in 

1982, performed by the city of Rancho Palos Verdes in violation

of the federal Endangered Species Act, destroyed the last 

remaining colony. The city was subsequently sued by the 

federal government, but this legal action was dismissed under 

the theory that the city could not be held liable. 

Conservation. Ironically, a simple captive breeding method 

was developed in 1983 using the southern blue as a surrogate 

(Mattoni 1988). Had the technique been available the previous 

year, it would have been possible to have saved the species by 

captive breeding for later re-introduction into the habitat. 

Unfortunately the foodplant stands were continually being 

cleared for fire prevention and few plants now remain. It has 

been suggested (Mattoni unpublished) that an effort be 

implemented to release numbers of the southern blue, preadapted 

to feeding on vetch by captive mass rearing, into sites to be 

heavily restocked with vetch. This action would again grace the 

136 

area with a butterfly that may evolve characteristics of its 

extinct relative. In addition to restoring natural biodiversity the 

plan would provide a demonstration of adaptive processes in a 

restored environment. 

References 

ARNOLD, R.A. 1985. Palos Verdes blue butterfly management plan. U.S.F. 

and W.S. informal report. 37pp. 

ARNOLD, R.A. 1986. Decline of the endangered Palos Verdes blue butterfly 

in California. Biol. Conserv. 40: 203–217. 

MATTONI, R.H.T. 1988.1988 report to California Department of Fish and 

Game, Contract C-1456. 

PERKINS, E.M. and EMMEL, J.F. 1977. A new subspecies of Glaucopsyche 

lygdamus from California (Lepidoptera: Lycaenidae). Proc. Ent. Soc. 

Wash. 79: 468–471.

The Xerces Blue, Glaucopsyche xerces (Boisduval) 


Department of Zoology, University of Florida, Gainesville, Florida 32611, U.S.A., and 26500 Rim Road, Hemet, California 

92544, U.S.A. 


Status and Conservation Interest: Status – extinct (Red List). 

The Xerces Blue, a former resident of the sand dunes in San 

Francisco, was the first butterfly species in North America to 

become extinct through human interference. (Two satyrines in 

the genus Cercyonis were the first subspecies of wider ranging 

U.S. species to disappear in historic times.) Today, an 

international organization devoted to the conservation of 

invertebrates, The Xerces Society, is named for this diminutive 

creature, a butterfly which has achieved fame far beyond its size 

and former restricted geographic distribution. From a scientific 

viewpoint, it was one of the most variable butterflies known and 

it would have made a unique tool for research on the effects of 

population size and geographic distribution on infra-specific 

variation. But for its untimely extinction, G. xerces would have 

contributed greatly to human knowledge in ecology and 

evolutionary biology. 

Taxonomy and Description: The species was first described 

by Boisduval (1852) from one male and two females taken 

within the area now occupied by the city of San Francisco, in 

San Francisco County, California. These specimens were 

collected by Pierre Joseph Michel Lorquin, a French gold 

seeker and naturalist who arrived in San Francisco in late 1849 

or early 1850 and apparently made his first shipment of specimens 

to Boisduval in the fall of 1851. Thus he had only two seasons 

at the most to collect the first set of specimens that he sent to 

Boisduval. While the species was extraordinarily variable, and 

several forms were named (originally thought to be separate 

species), all are presently referred to the nominotypical 

subspecies (Emmel, Emmel and Mattoon in press) and no 

geographic subspecies are recognized. 

The closest living relative of the Xerces may well be a 

newly recognized subspecies of Glaucopsyche lygdamus 

residing on Santa Rosa Island, off the coast of southern California 

(Emmel and Emmel in press). The extraordinary phenotypic 

resemblance of the wing maculation of this new subspecies to 

that of G. xerces suggests a close phylogenetic relationship. 

The genitalia of G. xerces are very similar to those of G. 

lygdamus, and on the basis of male genitalia alone, the Xerces 

137 

Blue would have been assigned subspecific status under 

lygdamus. However, Downey and Lange (1956) pointed out 

considerable differences between these species in larval stages, 

adult wing maculation, and ecology. Additionally, the two 

species were once sympatric in San Francisco and hybrids were 

never detected. 

Distribution: All recorded specimens are from San Francisco, 

and include the area presently covered by the city. The 

distribution on the San Francisco Peninsula ranged from near 

Twin Peaks to North Beach, and from the Presidio on the Bay 

southward to Lake Merced. Lone Mountain, formerly an isolated, 

sandy hill, was the classical locality for Xerces in the early 

1900s. 

Population Size: The only known colonies were in the San 

Francisco Bay area. No estimates of population size have come 

down to us in the early literature, but by comparison with the 

closely related G. lygdamus colonies in the area, we can guess 

that the typical size was several hundred individuals in a 

colony. By the year 1919, the only known population remaining 

was flying in a limited area west of the Marine Hospital, at the 

Presidio, San Francisco. The same area was still inhabited on 

March 23, 1941. The butterflies were then limited to a small 

area 21m wide by 46m long, in which Lotus scoparius was 

found. The last known specimens of Xerces were collected at 

the Presidio during May 1941 (Downey and Lange 1956), and 

many later visits to the area in search of the butterfly were 

fruitless. 

Habitat and Ecology: The habitat of the PVB was sandy areas 

where a prostrate dune ecotype of the perennial legume, Lotus 

scoparius (Nutt.) Ottley, occurred. The butterfly typically 

occurred around patches of Lotus that grew in the partial shade 

of the Monterey Cypress (Cupressus macrocarpa Hartw.) in 

well-drained, sandy soil. Lupinus arboreus Sims, which also 

served the Xerces females as a hostplant for oviposition and 

larval growth, was widely distributed among the Lotus, and had 

a wider range than that of the Lotus and the butterflies. In 1956, 

the same association of plants at the last colony site remained 

essentially unchanged from 1941, and was also observed by the

authors in 1963–67 in essentially unchanged condition. To the 

north, similar sandy areas with these Lotus and Lupinus species 

occur in Marin County, but the Monterey Cypress does not 

occur there. It appears that the expansion of the city of San 

Francisco into the natural sand dune habitat formerly available 

to Glaucopsyche xerces (and the other insects of the region) had 

destroyed too much of the habitat for the butterfly to continue 

to support a sustainable population beyond 1941. 

The adults of Xerces flew in a single brood from about 

March 10 to April 15. However, specimens were taken from as 

early as late February to as late as early June. The flight, mating 

behaviour, and oviposition behaviour of Xerces were apparently 

similar to that of the other Glaucopsyche species of California. 

The female laid eggs singly in the small depression at the base 

of the leaflet on Lupinus arboreus, or laid eggs on new growth 

near the tips of the leaflets on Lotus scoparius. In 1939, it was 

observed that as many as nine eggs were laid per plant, and that 

the Lotus was slightly favoured by the females, with an average 

of 3–4 eggs per plant compared to 2 eggs per plant for the 

Lupinus. 

The butterflies were associated with ants in their natural 

habitat although the ant associates were never identified. In the 

laboratory, larvae were fed substitute foodplants (Lupinus 

micranthus Dougl. and Astragalus menziessii Gray), and the 

captive larvae were raised without any ants so the species was 

not completely dependent on ants for successful maturation and 

pupation. The pale green larvae had long hairs on each side of 

the dorsal surface, and the whole body was covered with a 

whitish pile. The general colouration and pattern were variable, 

a trait found in other nearctic blue species today. The pupal 

colouration was highly variable. Diapause was in the pupal 

stage, with the length of that stage averaging from 10 to 11 

months. 

138 

Threats: The decline of this fascinating butterfly and its final 

extinction in 1941 appears to be attributable solely to the 

expansion of the City of San Francisco in the preceding 60–80 

years. This resulted in the removal of the native vegetation in 

the dunes and reduction of the remaining habitat areas to an 

unsustainable size. In such small populations, a minor change 

in climate or other environmental factor could prove devastating, 

especially to a species that flew in a single annual brood and had 

no options to react, little time or space, and a small genetic pool 

of variability. Collecting might have been detrimental also as 

the populations declined. 

While a live Xerces Blue will never be seen again on Earth, 

the species has served as a remarkably effective symbol for the 

fragility of nature among American citizens and conservationists 

around the world. Scientific analysis of 344 historical specimens 

(Downey and Lange 1956) offer a hint as to the remarkable 

variation in this species and the annual changes of the frequency 

of pattern forms in the population. 

References 

BOISDUVAL, J.B.A.D. de. 1852. Lepidoptères de la Californie. Ann. Soc. 

Entomol. France (Series 2) 10: 275–324. 

DOWNEY, J.C. and LANGE, JR., W.H. 1956. Analysis of variation in a 

recently extinct polymorphic Lycaenid butterfly, Glaucopsyche xerces 

(Bdv.), with notes on its biology and taxonomy. Bull. South. Calif. Acad. 

Sci. 55 (3): 153–207. 

EMMEL, T.C. and EMMEL, J.F. In press. A new xerces-like subspecies of 

Glaucopsyche (Lycaenidae) from Santa Rosa Island, California. 

Systematics of Western North American Butterflies. Scientific Publishers, 

Gainesville, Florida. 

EMMEL, T.C, EMMEL, J.F. and MATTOON, S.O. In press. The Butterflies 

of California. Stanford University Press, Stanford, California.

The Mission Blue, Plebejus icarioides missionensis Hovanitz 

J. HALL CUSHMAN 

Center for Conservation Biology, Department of Biological Sciences, Stanford University, Stanford, CA 94305, U.S.A. 



List). 

In 1976, the Mission Blue was officially listed as an 

endangered species by the U.S. Fish and Wildlife Service. The 

subspecies is also listed as endangered with the California 

Department of Fish and Game and almost all of its habitat 

protected as park land. 

Taxonomy and Description: The Mission Blue is one of 12 

recognized subspecies of the highly variable Plebejus icarioides 

and was first described by Hovanitz (1937). 

Distribution: While Plebejus icarioides as a whole is patchily 

distributed throughout western North America, the present-day 

distribution of the Mission Blue subspecies is restricted to four 

known areas on the northern tip of San Francisco Peninsula. 

Although it is impossible to document the historical distribution 

of this subspecies, the Mission Blue almost certainly occurred 

throughout much of the coastal scrub habitat of the northern 

peninsula (Reid and Murphy 1986). 

Population Size: The largest known Mission Blue population 

is on San Bruno Mountain (San Mateo Co.) and is estimated to 

consist of 18,000 adults. A substantial population is also found 

at Fort Baker (Marin Co.), although a precise estimate of its size 

is not available. Significantly smaller populations are found on 

Twin Peaks (San Francisco Co.) and the Skyline Ridges (San 

Mateo Co.), with estimates of 500 and 2000 adults, respectively 

(Reid and Murphy 1986). 

Habitat and ecology: The Mission Blue has one generation per 

year, and adults fly from mid-April to mid-June. Adults live for 

up to one week, remain close to the larval host plant, and feed 

on nectar from a wide variety of plants, including Eriogonum 

(Polygonaceae) and numerous composites. Females oviposit 

on three lupine species, Lupinus albifrons, L. formosus, and L. 

varicolor, which are found in areas of recent disturbance and 

attain highest densities in grasslands on thin, rocky soils (Arnold 

1983; Reid and Murphy 1986). 

139 

Females lay eggs on the leaves, stems, flowers, and seed 

pods of Lupinus hosts. The eggs are usually deposited singly 

and hatch in 4–10 days. After about three weeks, the secondinstar 

larvae enter an obligatory diapause from which they 

emerge the following spring. These post-diapause larvae 

subsequently complete their development to adult in four to 

five weeks (Arnold 1983; Reid and Murphy 1986; Newcomer 

1911). 

The cryptically coloured larvae may exhibit significant agespecific 

differences in feeding behaviour. Downey (1962) 

noted that older larvae are most commonly found at the base of 

food plants while the younger larvae are substantially higher up 

on the host. Because of this, he postulated that the majority of 

mature larvae in this species are nocturnal feeders. 

Juvenile stages of the Mission Blue are attacked by a variety 

of natural enemies. Arnold (1983) reported that 'Approximately 

35% of field collected eggs were parasitised by an unidentified 

encyrtid wasp.' In addition, Newcomer (1911) and Downey 

(1962) both reported that the larval stages of P. icarioides were 

parasitized by various braconid and tachinid species. 

As with many other lycaenid species, Mission Blue larvae 

are tended by ants. These associations have not received much 

attention (although see Downey 1962), and are thus poorly 

understood. However, the larvae are known to possess abdominal 

nectary glands which become active in the third or fourth instar 

and attract ants. Downey (1962) found that the butterfly could 

be reared successfully in the laboratory without ants, but makes 

no statements about how often older larvae are ant-tended in the 

field. Clearly, the importance of ants to the development and 

survival of Mission Blue larvae is very much an open question 

and requires further study. 

Threats: Agricultural and urban expansion have resulted in the 

progressive loss of native grassland habitat and reduced exchange 

among existing Mission Blue populations. These major threats 

have been significantly reduced due to the habitat conservation 

plan mentioned in the following section. However, a remaining 

threat is the invasion of grassland habitats by non-native plant 

species. While lupine, the Mission Blue host plant, is relatively 

resistant to invasions of non-native grasses, it is quite susceptible 

to the invasion of woody species which create too much shade.

The major culprits in this regard are gorse (Ulex europeaus), 

blue gum (Eucalyptus globulus) and broom (Cytisus spp.), all 

of which are on the increase in grassland areas such as those on 

San Bruno Mountain. 

Conservation: Concern about this subspecies, as well as the 

San Bruno Elfin (Callophrys mossi bayensis) and Callippe 

Silverspot (Speyeria callippe), was instrumental in leading to 

the formation of the U.S.A.'s first habitat conservation plan in 

1983. The plan created the San Bruno Mountain County Park, 

an extensive area encompassing the largest existing butterfly 

population. Considerable effort has been made to annually 

monitor the size of adult populations and initiate management 

activities such as the control of invasive species (see Reid and 

Murphy 1986, and Bean et al. 1991 for details). 

140 

References 


(Lepidoptera, Lycaenidae): island biogeography, patch dynamics, and 


BEAN, M., FITZGERALD, S. and O'CONNOR, M. 1991. The San Bruno 

habitat conservation plan. In: Reconciling conflicts under the endangered 

species act World Wildlife Fund, Washington, D.C. pp. 52–65. 

DOWNEY, J.C. 1962. Myrmecophily in Plebejus (Icaricia) icarioides 

(Lepidoptera: Lycaenidae). Entomol. News 73: 57–66. 

HOVANITZ, W. 1937. Concerning the Plebejus icarioides Rassenkreis 

(Lepidoptera: Lycaenidae). Pan-Pacific Entomol. 13: 184–189. 

NEWCOMER, E.J. 1911. The life histories of two lycaenid butterflies. Can. 

Ent. 43: 83–88. 

REID, T.S. and MURPHY, D.D. 1986. The endangered Mission Blue butterfly, 

Plebejus icarioides missionensis. In: B.A. Wilcox, P.F. Brussard and 

B.G. Marcot (Eds). The management of viable populations: theory, 

applications, and case studies. Center for Conservation Biology, Stanford, 

California, pp. 147–167.

The San Bruno Elfin, Incisalia mossii bayensis (Brown) 

Stuart B. WEISS 

Center for Conservation Biology, Department of Biological Sciences, Stanford University, Stanford, CA 94305, U.S.A. 



List). 

This subspecies is endemic to the northern San Francisco 

Peninsula, California. It was one of the first butterflies protected 

under the U.S. Endangered Species Act in 1976. As a local 

endemic in a highly urbanised region, threats to the San Bruno 

Elfin include land development and invasive introduced plant 

species. 

Taxonomy and Description: The subspecies was discovered 

rather late, in 1962 (MacNeill 1963), and described by Brown 

(1969a). It was originally described as Callophrysfotis bayensis, 

but was later recognized to be in the species C. mossii (Edwards) 

(now genus Incisalia, which occurs from Vancouver Island 

along the coast range to near Los Angeles). Populations to the 

north in Marin County are recognized as a different subspecies, 

those to the south in the Santa Lucia Range are ssp. doudoroffi. 

Distribution: The subspecies is restricted to three distinct areas 

on the northern San Francisco peninsula: Montara Mountain; 

Milagra Ridge; and San Bruno Mountain. Each of these localities 

supports an array of highly local demographic units tied together 

by occasional adult migration. Populations probably once existed 

within San Francisco at Twin Peaks and Mount Davidson, but 

have disappeared with urbanisation (Emmel and Ferris 1972). 

Population Size: The San Bruno Elfin was never common, 

because of specialized habitat requirements (see below). The 

butterfly exists in local discrete populations of ten to several 

hundred adults at higher altitudes. A thousand or more adults 

may exist in about 15 total subpopulations on San Bruno 

Mountain in a good year. Montara Mountain supports about 10 

local populations, and Milagra Ridge supports about four. 

Virtually all of the existing habitat is now protected as parkland, 

and numerous populations have been qualitatively monitored 

since 1982. Colonies noted by Arnold (1984) occupied small 

areas (0.15–8.0ha) on steep, north-facing slopes. 

141 

Habitat and ecology: The distribution of the butterfly closely 

follows the narrow, fragmented distribution of its larval 

hostplant, stonecrop, Sedum spathulifolium (Brown 1969b). 

This succulent plant grows in abundance only on thin-soiled or 

rocky north-facing slopes within the coastal fog belt. Sedum 

occurs in both short-statured coastal scrub and grassland 

vegetation types, and is most common near the summits of 

coastal mountains and around rocky outcrops on lower slopes. 

Sedum readily invades roadcuts and old quarry faces provided 

the aspect is correct. Local populations of the Elfin correspond 

closely to these patches of the larval hostplant, which range 

from a hundred square meters to several hectares in extent. 

San Bruno Elfin adults fly from February into April, during 

the latter part of the rainy season in northern California, but 

before the onset of persistent summer fog. Adults usually 

appear after the first extended warm sunny period of the season, 

as early as the first week in February, or as late as April. The 

window of sunny, calm conditions during the flight season is 

highly variable from year to year, and adults run the risk of 

being grounded by inclement weather for weeks on end. 

Populations were greatly reduced during and after near record 

rainfall in 1983, but appear to be less affected by recent drought 

conditions (San Bruno Mountain Conservation Plan Monitoring 

Reports, 1982–1991). 

Habitat topography may be limiting for Elfin populations in 

certain cases. Because of low winter sun angles, the steepest 

habitat areas may be in deep shade for much of the day, limiting 

access by adults (Weiss and Murphy 1991). While natural 

contours are rarely shaded all day even in February, roadcuts 

and quarry faces may face severe shading limitations. Steep 

(>40°) north-facing slopes provide minimal solar exposure 

even in March, and are rarely occupied by Elfin. Equally steep 

northeast-facing slopes receive direct morning light when winds 

are calm, and provide excellent Elfin habitat. Elfin activity on 

steep northwest-facing slopes, however, may be limited by 

strong afternoon winds. 

Adults are highly sedentary, typically moving less than 100 

metres (Arnold 1983), with a maximum recorded movement of 

about 800m (Arnold 1984). Males perch on rocks and vegetation, 

and dart out at passing insects; females spend much of their time 

crawling among the foodplants (Emmel and Ferris 1972; Arnold

1983). Both sexes visit flowers (especially Lomatium 

utriculatum, but also other early blooming coastal species) 

when plants are available, but a number of local Elfin populations 

on Montara Mountain exist where nectar resources are 

astonishingly sparse. 

The butterfly is univoltine. Females oviposit on Sedum 

rosettes. Early instars feed on the fleshy leaves, but third and 

fourth instars feed on the flowers when they appear. Third and 

fourth instars are easily observed basking and feeding on 

Sedum flowerheads, and exhibit a continuous colour 

polymorphism ranging from deep red (the colour of Sedum 

foliage) through orange to bright yellow (the colours of Sedum 

flowers); larvae may change colour morph over a few days 

(Orsak and Whitman 1986). Larvae may be tended by ants of up 

to nine different species, but the relationship appears facultative 

(Arnold 1983). However, most larvae in the field are observed 

without ants (Emmel and Ferris 1972, S.B. Weiss pers. obs.). 

Larvae are parasitized by a tachinid, Aplomya theclarum that 

emerges from the fourth instar Elfin larvae. Parasitisation rates 

of reared larvae and collected pupae are high, of the order of 

50–80% (Arnold 1983). Given the high densities of larvae 

observed (one or more per square meter), such high mortality 

appears necessary to produce the typically low density of adults 

the following year. The fourth instar caterpillar pupates in the 

duff immediately below the hostplants, and diapause lasts 

through the summer, fall and early winter. 

Threats: Because the vast majority of San Bruno Elfin 

populations are on public land (including San Bruno Mountain 

County Park, Golden Gate National Recreation Area, and 

McNee Ranch State Park) further opportunities for habitat 

conversion are limited. A proposed six lane road would skirt a 

population on Montara Mountain, but that construction has 

been challenged on a host of environmental and development 

issues other than the San Bruno Elfin. Continued expansion of 

a quarry on San Bruno Mountain could destroy some habitat on 

that property; however, the quarry is scheduled to be shut down 

within a decade. The prohibitions of the Endangered Species 

Act and park rules provide a strong deterrent against 

overcollecting by both amateurs and scientists. Wildfires may 

threaten in more heavily vegetated areas, but the thin vegetation 

on rocky outcrops is relatively safe from fires. Vegetation 

succession also appears unimportant, given the thin soils and 

windswept conditions of the habitat. Invasive introduced species 

such as gorse (Ulex europeaus), brooms (Cytisus spp.), pampas 

grass, ice plant (Mesembryanthemum spp.), and blue gum 

(Eucalyptus globulus) are encroaching on some local 

populations. 

Conservation: Since most of the habitat is already protected, 

conservation prescriptions for the San Bruno Mountain fall 

under the jurisdiction of the San Bruno Mountain Habitat 

142 

Conservation Plan, 1982 (Bean et al. 1991), which includes 

yearly monitoring of adult numbers and management activities. 

Elements of a Recovery Plan (Arnold 1984) included: (1) 

protection of essential habitat (which was designated on each 

site), through a range of strategies including cooperative 

agreements, easements and others; (2) prevention of further 

habitat degradation, and habitat enhancement where possible, 

through minimising toxin use, removal of weeds, control of offroad 

vehicles and revegetating with native flora; (3) development 

and implementation of management plans for extant colonies 

by utilising annual surveys and fostering autecological studies; 

(4) re-establishment of the species in restored sites within its 

historical range; (5) increase in public awareness; and (6) 

enforcement and evaluation of protective laws and regulations 

at all levels. 

Control of invasive species is currently under way at Milagra 

Ridge and San Bruno Mountain. Large areas of Sedum along 

Wolf Ridge in Marin County (just outside the historical range 

of the Elfin) are presently unoccupied by the butterfly, raising 

the possibility of an introduction attempt there. Revegetating 

abandoned quarry faces on San Bruno Mountain would provide 

a great opportunity to increase the habitat of the San Bruno 

Elfin, because Sedum will rapidly invade bare rock surfaces. 

The quarry configuration would provide large areas of 

appropriate solar exposure (especially northeast-facing slopes 

and flat benches) if restoration were attempted (Weiss and 

Murphy 1991). 

References 


(Lepidoptera, Lycaenidae): island biogeography, patch dynamics, and 


ARNOLD, R.A. 1984. (Main author of) U.S. Fish and Wildlife Service. 1984. 

Recovery plan for the San Bruno Elfin and Mission Blue butterflies. U.S. 

Fish and Wildlife Service, Portland, Oregon. 

BEAN, M., FITZGERALD, S. and O'CONNOR, M. 1991. The San Bruno 

habitat conservation plan. In: Reconciling Conflicts under the Endangered 

Species Act. World Wildlife Fund, pp. 52–65. 

BROWN, R.M. 1969a. A new subspecies of Callophrys fotis from the San 

Francisco Bay Area. J. Lepid. Soc. 23: 95–96. 

BROWN, R.M. 1969b. Larva and habitat of Callophrys fotis bayensis.J. Res. 

Lepid. 8: 49–50. 

EMMEL, J.F. and FERRIS, CD. 1972. The biology of Callophrys fotis 

bayensis. J. Lepid. Soc. 26: 237–244. 

MACNEILL, CD. 1963. Callophrys fotis (Strecker) from the San Francisco 

Bay area. Pan-Pacific Entomologist 39: 60. 

ORSAK, L. and WHITMAN, D.W. 1986. Chromatic color polymorphism in 

Callophrys mossii bayensis larvae (Lycaenidae): spectral characteristics, 

short-term color shifts, and natural morph frequencies. J. Res. Lepid. 25: 

188–201. 

WEISS, S.B. and MURPHY, D.D. 1991. Thermal microenvironments and the 

restoration of rare butterfly habitat. In: J. Berger (Ed.) Environmental 

Restoration. Island Press, Covelo, California, pp. 50–60.


The Lotis Blue, Lycaeides idas lotis (Lintner) 

R.A. ARNOLD 

Entomological Consulting Services Limited, 104 Mountain View Court, Pleasant Hill, CA 94523, U.S.A. 

Status and Conservation Interest: Status – probably extinct; 

endangered (Red List). 

The Lotis Blue was regarded by Arnold (1985) as probably 

the rarest resident butterfly in the continental United States. 

Historically, it was probably restricted to just a few localised 

colonies in coastal northern California. Although it was listed 

as an endangered species in 1976, today it is feared to be extinct. 

Taxonomy and Description: The Lotis Blue was formerly 

considered to be a subspecies of Lycaeides argyrognomon 

(Bergstrasser), and was listed as endangered under the name L. 

argyrognomon lotis. However, European butterfly taxonomists 

have recently examined the types of L. argyrognomom and L. 

idas. They concluded that all North American taxa, which were 

formerly called L. argyrognomon, should now be called L. idas. 

Hence, the Lotis Blue is now referred to as L. idas lotis. Twelve 

subspecies of L. idas (Linneaus) have been described from 

North America (Downey 1975). The Lotis Blue is one of the 

larger subspecies of L. idas. It is also separable from other 

described species by its wing colours and markings. 

Distribution: Nearly all collections or sightings of the Lotis 

Blue since 1933 have been from a single location, near the town 

of Mendocino in Mendocino County, California (Arnold 1991). 

This location is a Sphagnum bog situated in the right-of-way for 

the Elk-Fort Bragg 60-kV transmission line, operated by the 

Pacific Gas and Electric Company (P.G. & E). The bog is about 

lha in size and is surrounded by Red Alder (Alnus rubra) 

riparian forest, Northern Bishop Pine (Pinus muricata) forest, 

and Mendocino Pygmy Cypress (Cupressus pygmaea) forest. 

Characteristic understory vegetation includes various ericaceous 

shrubs, sedges, and ferns. 

Historical records of the Lotis Blue reveal that it was known 

from only a few other locations, between Point Arena and Fort 

Bragg in coastal Mendocino County. Reports of the butterfly's 

probable occurrence in Sonoma and northern Marin counties by 

Tilden (1965) are unsubstantiated by any specimens. Because 

of substantial differences in the types of vegetation that occur 

in these areas, it is doubtful that the butterfly ever occurred 

143 

outside of coastal Mendocino County. Fewer than 75 specimens 

are housed in North American entomological collections (Arnold 

1991). 

In 1990, P.G. & E. sponsored an extensive survey of the 

butterfly and its suspected larval foodplant at 23 locations in 

coastal Mendocino County, the historical geographic range of 

the Lotis Blue. Unfortunately, no specimens of the Lotis Blue 

were found; indeed, none have been seen since 1983 and the 

negative results of this survey suggest that the butterfly may 

now be extinct (Arnold 1991). 

Population Size: On June 19th and 20th, 1953, J.W. Tilden 

collected at least 26 adults of the Lotis Blue from the bog 

population of the eventual P.G. & E. transmission line, which 

are preserved in North American entomological collections. 

During 1977–1989, Arnold saw only 26 adult butterflies during 

67 days of field work at the P.G. & E. location, and none after 

1983. Because of the small size of the butterfly's habitat, its 

population numbers probably were never greater than a few 

hundred individuals per season at the powerline bog. The 

butterfly's very rarity has precluded detailed investigations 

about its population size and structure, as have been obtained 

for other endangered lycaenids that occur in California (Arnold 

1983). 

Habitat and ecology: Because of the Lotis Blue's rarity, little 

is known about its specific habitat requirements and ecology. In 

northern California, other subspecific taxa of Lycaeides idas 

typically occur in wet meadows, bogs, seeps or springs, and 

along streamsides. Populations of these butterflies are typically 

associated with small, and often isolated, patches of their larval 

foodplants. Known larval foodplants include legumes that 

grow in these wet habitats, in particular Lotus oblongifolius, 

Lupinus polyphyllus and Astragalus whitneyi (A. Shapiro, pers. 

comm.). 

Four legume species have been observed growing in or very 

near the bog at the primary Lotis Blue population site along the 

transmission line. Of these legumes, Lotus formosissimus (Coast 

Trefoil) is the most likely candidate to be the butterfly's larval 

foodplant, since it grows in the bog where most specimens of 

the Lotis Blue have been observed. Also, a female was observed

(J.F. Emmel, cited in Arnold 1985) attempting to oviposit on 

this plant. 

A hypothetical life cycle and natural history of the Lotis 

Blue can be surmised based on circumstantial evidence from 

related taxa whose biologies are better known. Like other taxa 

of L. idas in northern California, the Lotis Blue is probably 

univoltine. Historical collection records indicate that adults 

may be present from mid-May through mid-July. Eggs are laid 

throughout the adult flight season and newly hatched larvae 

probably begin to feed immediately. Partially grown larvae, 

probably second instars, diapause until the following spring, 

when larval development is completed in about four to six 

weeks after feeding resumes. Presumably, the pupal stage lasts 

no more than a few weeks. 

Threats: Because of the butterfly's extremely limited 

geographical range, and the small size of its only known habitat, 

the Lotis Blue is extremely vulnerable to any type of habitat loss 

or alteration. Collecting of specimens could also be detrimental. 

Arnold (1985) speculated that drought may have previously 

affected the butterfly's habitat by decreasing water levels in the 

bog. However, more recent information suggests that 

successional changes in the vegetation at the transmission line 

site, and at other potential population sites, are probably 

responsible for the recently observed decline of the butterfly. 

The leguminous foodplants of other northern California 

populations of L. idas generally grow in small patches in wet 

habitats that are at the early stages of vegetation succession. 

Circumstantial evidence from the 1990 P.G. & E. sponsored 

study suggests that Lotus formosissimus, the suspected larval 

foodplant of the Lotis Blue, also grows in greatest abundance in 

the early successional stages of wet areas, such as bogs, plus the 

headwaters and shorelines of streams (Arnold 1991). 

Furthermore, examination of field notes from cadastral surveys 

and maps prepared by the U.S. Coast and Geodetic Survey, 

during the late 1800s and early 1900s indicate that the 

transmission line site and much of the coastal area within the 

butterfly's historic range were logged then. Indeed, many areas, 

including the transmission line bog, which today support dense 

forest vegetation, were open fields a century ago. Thus, the 

forests surrounding the transmission line bog represent regrowth 

rather than natural habitat. Comparison of a series of aerial 

144 

photos covering the past several decades depict how rapidly the 

vegetation has changed from a more open area to a dense forest 

with a closed canopy in many places. Presumably, as the 

successional changes have proceeded, the increasing density of 

the vegetation gradually choked out the suspected larval 

foodplant of the butterfly, which prefers open areas. By 1990, 

only 15 specimens of L. formosissimus were observed at two 

locations in or near the transmission line bog (Arnold 1991), 

and only two specimens were observed growing in the bog 

during 1992 (Arnold, unpublished data). 

Conservation: Clearly, basic ecological and natural history 

information about the butterfly is needed to identify potential 

habitat sites within its historic range and to manage these areas 

to benefit the Lotis Blue. Confirmation of the butterfly's larval 

foodplant is essential to identify its breeding habitat. Because 

of the butterfly's dubious status, surveys should be undertaken 

to determine if it even still exists. 

Objectives of the butterfly's recovery plan (Arnold 1985) 

are to: (1) protect the butterfly and its habitat at the only known 

site; (2) establish three new viable populations at different sites; 

and (3) determine the extent of the population and size of secure 

habitats needed so the butterfly can be reclassified as 'threatened' 

rather than 'endangered'. However, until the basic biological 

information about the butterfly is obtained, the objectives of the 

recovery plan probably cannot be achieved. 

References 


(Lepidoptera: Lycaenidae): island biogeography, patch dynamics, and 

the design of habitat preserves. Univ. Calif. Publns Entomol. 99: 1–161. 

ARNOLD, R.A. 1985. (Main author of) U.S. Fish & Wildlife Service. Lotis 

Blue butterfly recovery plan. U.S. Fish & Wildlife Service. Portland, 

Oregon. 46pp. 

ARNOLD, R.A. 1991. Biological studies of the endangered Lotis Blue 

butterfly for P.G. & E. 's Elk-Fort 60kV transmission line. Pacific Gas & 

Electric Company. San Ramon, California. 46pp. and appendices. 

DOWNEY, J.C. 1975. Genus Plebejus Kluk. In: Howe, W.H. (Ed.). The 

Butterflies of North America. Doubleday. Garden City, New York. pp. 

337–350. 

TILDEN, J.W. 1965. Butterflies of the San Francisco Bay Region. Univ. of 

Calif. Press. Berkeley, California. 88pp.

Additional taxa of concern in southern California 

Three additional lycaenid butterflies of conservation interest 

occur in southern California: 

The Human Folly Blue, Philotes 

sonorensis extinctis Mattoni 

The subspecies occurred across a l000ha site in the upper San 

Gabriel river wash. It has been extinct since 1968, having been 

eliminated by the activities of the U.S. Army Corps of Engineers 

to provide a spreading basin for subsurface water recharge. 

Ironically the Corps is today charged with the responsibility of 

protecting wetlands and species. Details of this subspecies can 

be found in Mattoni (1991). 

Reference 

MATTONI, R.H.T. 1991. An unrecognised, now extinct, Los Angeles area 

butterfly (Lycaenidae). J. Res. Lepid. 28: 297–309 (1989). 

The Santa Monica Mountains Hairstreak, 

Satyrium auretorum fumosum Emmel & 

Mattoni 

This hairstreak is only known from a few localities within a 

circumscribed area of about 25,000ha in the western Santa 

Monica mountains. The range is completely surrounded by 

urban development and itself is being further fragmented by 

developments. Battlelines between environmentalists and 

developers have been drawn to define the future of the mountains. 

The compromises finally reached regarding habitat preservation 

of the area may serve as a model for the rest of the United States. 

With land values in the order of one million dollars per hectare, 

economic pressures even affect biologist consultants. An effort 

to federally list the species is underway. 



145 

The San Gabriel Blue, Plebejus saepiolus 

undescribed subspecies 

This is another extinct taxon. Known only from a few meadows 

in the Big Pine recreation area on the north slope of the San 

Gabriel mountains it was last seen in the mid 1980s. Its 

distribution was very limited, since its wet meadow habitat, 

necessary to support the clover foodplant, is very restricted 

across the dry north slope. Extinction was brought on by 

draining of the limited meadow habitats by the U.S. Forestry 

Service.

Selected Neotropical species 

K.S. BROWN, JR. 

Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109, Campinas, São Paulo, 

13.081, Brazil 

Styx infernalis Staudinger 

Country: Peru. 


This species is probably the most primitive of the Riodininae, 

confined to a very small region of high species diversity and 

endemicity. It is very rarely observed. 

Taxonomy and Description: A medium-sized, dirty transparent 

grey butterfly with narrow wings and a heavy black body, 

seeming rather like a Geometrid or Lymantriid moth. It has 

short antennae. 

Distribution: S. infernalis is known only from central and 

southern Peru, at elevations between 1000 and 1600m (Figure 1). 

Population Size: Not known. 

Habitat and Ecology: This species inhabits primeval cloud 

forest with steep slopes and torrents. It is active in sun patches 

near midday, with a weak, almost gliding flight, in small 

aggregations near rushing streams. The early stages are unknown. 

Threats: Habitat conversion for coffee or other plantations 

(very scattered colonies). 

Conservation: The main need is to locate colonies and secure 

large tracts of undisturbed cloud forest. 

Nirodia belphegor (Westwood) 

Country: Brazil. 


Nirodia is a monotypic genus (very close toRhetus, composed 

of common tropical species), whose only species is confined to 

high-altitude rockfields in a very ancient environment. It has 

146 

been observed fewer than ten times since its original description 

140 years ago in spite of extensive human activity in the region. 

Taxonomy and Description: N. belphegor is a medium-small, 

yellow-spotted, dark blue butterfly with pointed forewings and 

a short broad tail on the hindwings (Figure 1d of regional 

account), strongly reminiscent of Rhetus periander (Cramer). It 

seems quite close to a fossil riodinine from the Eocene (Durden 

and Rose 1978). 

Distribution: It is known from only four localities in the Serras 

do Espinhago and Cipó in central Minas Gerais, Brazil, at 

elevations above 1000m (Figure 1) where it is seen very 

sporadically. 


Habitat and Ecology: It inhabits 'campo rupestre', a xeric 

system on rocky soils at about 1000m elevation, characterised 

by strong endemism in plants (Velloziaceae, Eriocaulaceae, 

Xyridaceae) and animals (including amphibians), with ancient 

affinities to similar systems in Africa. Males are seen resting on 

the ground with wings outspread (like Rhetus) drinking water 

beside small creeks rushing down through rockfields. They 

make short sallies out from perching places on sunlit rocks and 

dart irregularly about as if defending a territory in that space 

(fide Ivan Sazima). Early stages are unknown, as are any further 

biological details on the species. 

Threats: The 'campo rupestre' system, restricted in area, is 

under very heavy pressure from removal and exportation of 

dried plants, and is subject to extensive burning and trampling. 

Conservation: The species should be preserved in the recently 

declared Serra do Cipó National Park, and the proposed Caraça 

Natural Park. 

Reference 

DURDEN, C.J. and ROSE, H. 1978. Butterflies from the middle Eocene: the 

earliest occurrence of Fossil Papilionoidea (Lepidoptera). Pearce-Sellards 

Series, Texas Memorial Museum 29: 1–25.

Figure 1. Distribution (A–E) of selected Neotropical species of Lycaenidae. 

Joiceya praeclarus Talbot 



Joiceya is a monotypic genus known only from the type 

series, from a small, heavily collected and increasingly converted 

region. 

147 

Taxonomy and Description: A 'Setabis alcmaeon (Lewis 

1973, plate 77: Figure 5) -like' pattern: small, with a pointed 

forewing and elongated hindwing; dull grey-brown underside 

with faint distal lines; strong blue upperside with a black baseto-margin 

line on the hindwing; and a black area between 

forewing base and submarginal spots. It is like no other species 

or genus.

Distribution: J. praeclarus is known only from an original 

collection in central Mato Grosso (Cuiabá and Tombador), 

Brazil (Figure 1). It has not been collected in intensive recent 

work in the region. 


Habitat and Ecology: The ecology of this species is not 

known, but it probably inhabits the understorey of isolated 

humid headwater or spring-fed forests in strongly scarped 

regions within the cerrado landscape (chapadas). 

Threats: The area is being used intensively for colonisation, 

ranching, hydroelectric projects, mines, and industrial farms, 

with removal of all original vegetation in parts. 

Conservation: The species may be present in the new Chapada 

dos Guimarães National Park and other small reserves in the 

region, but it needs to be relocated and colonies specifically 

saved. 

Reference 

LEWIS, H.L. 1973. Butterflies of the World. Follett, Chicago. 

Arcas, five rarer species 

Countries: Panama, Ecuador, Peru, Brazil. 

Status and Conservation Interest: Status – rare or vulnerable 

(with the exception of A. imperialis which is not considered 

threatened). 

These are the most exquisite of all neotropical Theclinae, 

typical of large areas of virgin wet forest and usually disappearing 

in disturbed areas. They are easy to find when present and are 

thus good indicators of undisturbed forest systems. 

Taxonomy and Description: These are largish, two-tailed 

species with an additional well-separated long anal lobe of the 

hindwing (thus flashing six mobile 'antennae' to the false 

head). They are brilliant striated green on the ventral surface 

(often with additional nuances of rose and chrome yellow) and 

iridescent blue or green above. The genus was revised by 

Nicolay (1971). Illustrations of at least A. imperialis appear in 

most popular butterfly books, and A. ducalis has been called the 

most beautiful small butterfly in the neotropics. 

Distribution: The non-threatened A. imperialis (Cramer) occurs 

from Mexico to southern Brazil, with northerly females 

sometimes showing much rose colour ventrally. A. cypria 

(Geyer) is infrequent from Mexico to western Colombia, and A. 

ducalis (Westwood) is very local in southeastern Brazilian 

mountains. The A. tuneta superspecies (delphia Nicolay in 

148 

Costa Rica, and tuneta (Hewitson) from Peru to southeastern 

Brazil) are very rarely encountered. A. jivaro Nicolay is known 

only from the two types in a single locality, in eastern Ecuador 

and A. splendor (Druce), not seen for 110 years after its original 

collection, is restricted to scattered cloud forest localities from 

Costa Rica to W. Ecuador (Figure 1). Further taxa may still be 

found. 


Habitat and Ecology: Arcas males are most often seen in early 

afternoon on forested hilltops or ridges, where they sit high 

(5–10m) in the trees in the sun, looking out from leaves over a 

green sunlit space. They vigorously defend this space against 

other males while waiting for a female to arrive. Mated females 

are found lower down, inspecting growing tips of the foodplants 

(Annonaceae: Rollinia, and Lauraceae for A. ducalis). Two or 

three species can be found on a single hilltop in Panama or 

southeastern Brazil, but such favoured sites are very rarely 

found. Flower visiting is infrequent, mostly before midday, but 

may be prolonged. Flight is very rapid, a brilliant green swirl 

accompanied by blue flashes, with the insect returning to the 

same or a nearby perch. Most populations occur throughout the 

year but the insects are commoner on sunny days in the rainier 

seasons. 

Threats: Moderate modification of the habitat may eliminate 

male perching sites and prevent the sexes from meeting and 

mating, in the sparse populations of these species. Two of the 

species are known from very few localities and may easily be 

eliminated. 

Conservation: For the rarest of these species, A. jivaro and A. 

splendor, further localities should be looked for and protected 

Arcas ducalis (photo by K.S. Brown, Jr.).

wherever possible. Three of the more widespread species 

(cypria, ducalis, and tuneta), though known from more areas, 

are still very rarely encountered. Along with the more frequently 

encountered A, imperialis, they should be reported and monitored 

as good indicators of the health of intact, rich tropical wet forest 

ecosystems. 

Reference 

NICOLA Y, S.S. 1971. A review of the genus Arcas with descriptions of new 

species (Lycaenidae, Strymonini). J. Lepid. Soc. 25: 87–108. 

Arawacus aethesa (Hewitson) 



This species is endemic to wet forests in a restricted area of 

the Atlantic lowlands of southeastern Brazil. Since the building 

of a major new highway twenty years ago, over 90% of the 

vegetation has been cut over, greatly disturbed or removed. 

Taxonomy and Description: A. aethesa is closely related to A. 

aetolus (= linus Auctt.), the classical 'false head' phenotype 

which is very widespread in the neotropics, with convergent 

black lines on the cubital spot of the hindwing margin, from the 

forewing costa. It is easily distinguished by its smoky brown 

undersurface, with a yellow submarginal band. 

Distribution: It is known only from southern Bahia, northern 

Espírito Santo, and eastern Minas Gerais, in the Atlantic forests 

of eastern Brazil. 


Habitat and Ecology: It has been observed in sunlit columns 

in the interior of wet lowland forest. The foodplant is almost 

surely Solarium leaves. 

Threats: Habitat reduction is the primary threat to this species 

and many others endemic to Atlantic lowland forest. 

Conservation: The species exists in several reserves within its 

range, but needs to be specially protected as it uses secondarysuccession 

plants in a forest biome. 

149 

Cyanophrys bertha (Jones) 



A very rarely seen (only the single type existed among 

collections in all American and European museums until very 

recently) and spectacular species typical of large, species-rich, 

high-elevation sites in the coastal mountains, where many other 

unusual and little-known species occur. 

Taxonomy and Description: Like other Cyanophrys, it is blue 

above and pea-green below, but in contrast to them it has a white 

stripe across both wings on the underside, bordered basally with 

diffuse blue-white blotches. 

Distribution: The species is known in Brazilian collections 

from over 1000m elevation in the mountains of Minas Gerais 

(Pocos de Caldas, Barbacena), Rio de Janeiro (Petr6polis), Sao 

Paulo (Serra do Japi, where quite frequent at times), and Parana 

from a total of fewer than 20 specimens. 


Habitat and Ecology: Males frequent hilltops in the early 

afternoon, where they perch in the crowns (not usually the 

highest points) of tall scraggly trees, changing perches every 

few minutes or when approached. Both sexes can be found on 

flowers in the morning, and females frequent sunny glades on 

hillsides. Early stages are still unknown. It is associated with C. 

herodotus (F.), C. amyntor (Cramer), C. acaste (Prittwitz), C. 

remus (Hewitson) and C. nr. pseudolongula (Clench), the last 

three often perching on the same high-elevation hilltop trees. 

Threats: Like many highly localised and rarely seen Theclinae, 

it can be eliminated from the few local colonies by minor habitat 

alteration. 

Conservation: The species should be preserved in a number of 

parks and inaccessible areas.

Neotropical Lycaenidae endemic to high elevations in SE Brazil 


Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109, São Paulo, 13. 081, Brazil 


Status and Conservation Interest: Status – threatened 

community. 

These are rarely observed, often very specialised species 

and genera, in very local habitats in a strongly heterogeneous 

landscape. The area is subject to intensive human activity and 

there is considerable modification of the vegetation. 

Taxonomy and Description: In the Riodininae the community 

includes: three monotypicgenera:Nirodia belphegor (discussed 

under 'Selected Neotropical Species'); Eucorna sanarita 

(Schaus), dark with a 'fuzzy' pattern; Petrocerus catiena 

(Hewitson), similar to a dark Calydna; isolated species of 

Mesosemia (?) (M.? acuta Hewitson, beige with a falcate 

forewing); Calydna (a still undescribed species, very small 

with orange spots and a falcate forewing); Crocozona (?) (C.? 

croceifasciata Zikán, small with a transverse orange band 

across both wings); a species of Mesenopsis (M. albivitta 

(Lathy), imitating common Dioptid moths with orange bars on 

each wing (see next account); Panara ovifera Seitz; Mycastor 

leucarpis (Stichel); Argyrogrammana caesarion Rebillard and 

a number of species of Napaea (all dark brown). 

Endemic theclines are many, though not equal to the highaltitude 

groups in the Andes; some are more 'furry' than their 

lower-altitude congeners (due to the cold?) and many have 

rather sombre patterns (but see Cyanophrys bertha, Brown, this 

volume). 

150 

Distribution: The above groups are found in the Serras do Mar 

and da Mantiqueira, from the interior of Bahia and Espírito 

Santo south to Santa Catarina and northern Rio Grande do Sul. 

They are most common in very high areas in Rio de Janeiro/ 

Minas Gerais/São Paulo. 

Population Sizes: Not known. 

Habitat and Ecology: The species fly rarely, but have been 

noted flying in sunny weather near midday, in variable seasons 

and habitats. Most have been found too sporadically to make 

any generalisations, but as a rule they are likely to be in wet 

habitats or bamboo forests, not rare when found, occasionally 

partial to hilltops, and probably tightly associated with special 

host plants or ants. 

Threats: The exceedingly heterogeneous habitats are full of 

small micro-islands of different vegetation (often no more than 

a few hundred square metres on a hilltop, hillside or gully) 

whose destruction leads to extinction of the populations. 

Conservation: Although many large, inaccessible areas still 

exist in these regions, some of them officially preserved, little 

information is available about the presence of these rare species, 

and possibly other Lycaenidae still undescribed. Much 

exploration is needed to establish the localities of colonies and 

preserve them.


Riodininae: Amazonian genera with most species 

very rare or local 


Departamento de Zoologia, Institute de Biologia, Universidade Estadual de Campinas, C.P. 6109, 

Säo Paulo, 13. 081, Brazil 


community. 

There are a number of forest species that are very rarely 

seen, concentrated in compact taxonomic groups, whose local 

colonies, often very far apart, may be eliminated easily by 

modest alteration of the habitat. 

Description: The group includes: 

• four species of Alesa (telephae Boisduval, fournierae 

Rebillard, neagra Röber, thelydryas Bates, known from 

very few specimens but not A. prema Godart or A. amesis 

Cramer), 

• Mimocastnia rothschildi Seitz (a hypertrophic representative 

of the same phenotype and lineage); 

• both species of Colaciticus; 

• three species of Mesenopsis (a fourth is montane in SE 

Brazil, see preceding account); 

• all species of Xenandra including X. pulcherrima (Herrich- 

Schäffer) (usually placed in Melanis); 

• most species of Esthemopsis and Symmachia; 

• Zelotaea phasma Bates; 

151 

• all species of Dysmathia. 

The species vary from uniformly dingy (the last genus) to 

uniformly white (penultimate), with many having strong colours 

of yellow, orange and red or green; all are small except for 

Mimocastnia. 


Habitat and Ecology: They are typically found as isolated 

males in very high-diversity hilltops or clearings, very 

occasionally on flowers or in small assemblages. These species 

all inhabit the deep forest; some may be canopy dwellers, but 

most just seem to be very rare and sporadic in occurrence, 

presumably due to excessively narrow ecological tolerances. 

Threats: The very rarefied distributions indicate that the 

conversion of small areas may eliminate local colonies which 

will not be re-established in nearby intact habitat. 

Conservation: Wherever colonies are known, they should be 

protected by moderate-sized reserves which can maintain natural 

processes and heterogeneity.

Theclinae endemic to the Cerrado vegetation (central Brazil) 



Departamento de Zoologia, Institute de Biologia, Universidade Estadual de Campinas, 

C.P. 6109, Sao Paulo, 13. 081, Brazil 


community. 

These are very local species that are indicators of healthy 

and diverse, well-watered cerrado habitats which contain the 

full range of microenvironments and successional vegetation 

typical of the region. 

Taxonomy and Description: Coming from many groups and 

genera in the Theclinae, most of the characteristic 'cerrado 

species' have large red markings or blotches on the underside, 

almost as if they formed an environment-mediated mimicry 

ring. Typical representatives are Arawacus tarania (Hewitson), 

Strymon tegaea (Hewitson), S. ohausi (Spitz), S. sp. (zibagroup), 

'Thecla' mantica Druce, Magnastigma julia Nicolay, 

'Thecla' socia Hewitson, and 'Thecla' bagrada Hewitson. All 

are distinctive and restricted to the cerrado domain and its 

peripheries. 

Distribution: Central Brazil Plateau in Goiás, Distrito Federal, 

Minas Gerais, Mato Grosso, Mato Grosso do Sul, and parts of 

Bahia, Sao Paulo and Parana, in the cerrado. 


152 

Habitat and Ecology: The habitats of the group are variable: 

S. tegaea prefers wet grasslands by headwater woods, while the 

minute S. ohausi is restricted to tiny grassy marshes in small 

sinkholes within the cerrado. A. tarania is more widespread in 

scrub with a grassy understorey, the juveniles feeding on small 

Leguminosae (K. Ebert, pers. comm.); 'Th.' mantica and M. 

julia prefer bushy cerrado, the larvae of the former living on 

Chrysobalanaceae (K. Ebert, pers. comm.). The other three 

'Thecla', are often found perched in medium-sized trees, near 

the end of the afternoon. Most are scarce. 

Threats: Large areas of cerrado are being occupied by intensive, 

mechanised and chemical agriculture, completely destroying 

and poisoning the diverse and complex natural systems. As 

several of these species are intensely localised and occupy rare, 

scattered and very specific habitats, their few local populations 

are especially vulnerable. 

Conservation: A number of preserved areas in the cerrado 

region probably include the commoner species of Theclinae 

endemic to this biome, but colonies of the rarer ones such as S. 

ohausi and M. julia need to be localised and specifically 

protected. The latter has not been seen since the 1969 collecting 

trip that led to its discovery and description, when it was found 

in only three localities, all of precarious preservation today.

Neotropical Riodininae endemic to the Chocó region of western 

Colombia 

Country: Colombia. 

C.J. CALLAGHAN 

Louis Berger International Inc., 100 Halsted Street, P.O. Box 270, East Orange, N.J. 07019, U.S.A. 


community. 

The area supports a high number of rare, specialised species 

of riodinine butterflies, all very sensitive to alterations in 

vegetation patterns. The area is currently under pressure from 

human activity, especially logging of pulp-wood for paper 

mills. 

Taxonomy and Description: The fauna is taxonomically poorly 

known, so that future study may well reveal additional species 

or genera which are endemic to the region. Preliminary work by 

Callaghan (1985) suggests that 33 (36%) of a total of 91 species 

of the known Riodininae are endemic. Among these areEuselasia 

rhodogyne (Godman), E. violacea Lathy, two undescribed 

Euselasia, Mesosemia asa iphigenia Stichel, M. sibyllina 

Staudinger, Eurybia juturna cyclopia Stichel, Lucillella sp. 

nov., Calospila rubrica (Stichel), C. asteria (Stichel), C. caligata 

(Stichel), Nymphidium balbinus Staudinger, and Stalachtis 

magdalenae cleove Staudinger. In addition to these endemics, 

the Chocó above 1000m also forms the last refuge for Mesosemia 

mehida Hewitson and M. bifasciata Hewitson, described from 

western Ecuador. 

Distribution: The Chocó region extends from north of Quibdó 

south along the western slopes of the Cordillera Occidental to 

northwestern Ecuador. The fauna shows strongest affinities 

with Panama/Costa Rica, with which it shares 58% of its 

Riodininae. Because of this shared influence from Panama, 

43% of the Chocó fauna is also found in the Cauca Valley to the 

east of the Cordillera; but from there eastwards, the faunal 

similarities drop rapidly. Only 17% of the taxa are also 

encountered on the eastern (Amazonian) slope of the Cordillera 

Oriental. 

153 


Habitat and Ecology: The characteristic of the Chocó is its 

high rainfall, as much as 13 metres a year east of Quibdó. The 

rain is nearly constant, though slightly less in January-February 

and July-August, and often mostly in the afternoon and evening. 

The butterflies take to the wing during short intervals of sun, 

usually in the morning and early afternoon. Most species 

concentrate on hilltops, although many are found in the forests 

along trails and streams, often deeply cut into the weathered 

soil. 

Threats: Nearly all species are very sensitive to habitat 

alteration, particularly cutting of the rain forest. A significant 

fall in the number of hilltopping species has been seen as a 

function of the cutting of larger trees on the slopes. 

Conservation: Reserves in the Chocó are few and only 

established with great difficulty due to the conflicting economic 

interests, particularly from the paper industry. However, due to 

the uniqueness of the habitat and its high endemism, combined 

with the rudimentary level of existing knowledge, every effort 

should be made to support the investigation and establishment 

of sustainable reserves in the Chocó. 

Reference 

CALLAGHAN, C.J. 1985. Notes on the zoogeographic distribution of 

butterflies in the subfamily Riodininae in Colombia. In: Proc. 2nd Symp. 

Neotropical Lepidoptera. J. Res. Lep. Supplement 1: 50–69.

Aloeides dentatis dentatis (Swierstra), Aloeides dentatis maseruna 

(Riley); Subfamily Theclinae, Tribe Aphnaeini 

Country: South Africa. 

S.F. HENNING 1 , G.A. HENNING 2 and M.J. SAMWAYS 3 

1 5 Alexander St., Florida 1709, South Africa 

2 17 Sonerend St., Helderdruin 1724, South Africa 

3 Department of Zoology and Entomology, University of Natal, Pietermaritzburg 3200, South Africa 

Status and Conservation Interest: Status – A. d. dentatis: 

rare; A. d. maseruna: indeterminate; rare (Red List). 

These two subspecies are highly localised endemics from 

the central grassland savannah of southern Africa. Earlier 

recorded sites for both butterflies have been damaged, with 

localised loss of the species. 

Taxonomy and Description. The upperside of A. d. dentatis is 

orange, with narrow black margins and apical patches (Murray 

1935, Pennington 1978). The basal half of the forewing costa is 

orange. The underside of the hindwing is crimson-red with 

silvery white and black markings. The diagnostic feature is a 

medial series of small dentate markings with black along their 

outer edge. There is also a form with a pale brown rather than 

crimson ground colour, and with indistinct markings. The 

female is similar to the male but has more rounded wings. 

Forewing lengths: male 14–18mm; female 15–19mm. 

A. d. maseruna has a pale tawny-orange upperside with a 

dark grey border. The underside of the hindwing is pale brown 

or pinkish-red with the silvery dentate band more extensive 

than in the nominate subspecies. Forewing lengths: male 

13.5–18.5mm; female 15–19mm. 

Distribution. A. d. dentatis occurs on the highveld of the 

Transvaal, at Waterval Onder, Ruimsig, Pretoria, Springs, 

Alberton and Suikerbosrand. 

A. dentatis maseruna was originally recorded from Maseru 

in Lesotho. Other localities in the Orange Free State and the 

Western Transvaal are now also known. 

Population Size: From mark-recapture data collected at the 

Ruimsig Reserve in 1989/1990, it was estimated that the 

population size of A. d. dentatis was 400 specimens on the wing. 

The population size of other populations of A. d. dentatis is not 

known. There is no information available on the sizes of the 

known colonies of A. dentatis maseruna. 

Habitat and Ecology: A. d. dentatis occurs in highveld 

grassland. The adults do not range far from the host plants and 

154 

ants. The males maintain small territories, on sandy patches 

among the foodplants, in which they can be found throughout 

most of the day. The females fly randomly throughout the area, 

and are as common as the males. During partly cloudy days, A. 

d. dentatis has been seen to bask on a rock by lying sideways. 

It feeds on a variety of flowers, and its foodplant at Ruimsig 

is Hermannia depressa (Sterculiaceae) at an elevation of 1500m. 

The eggs are laid in pairs on the underside of the leaves of the 

foodplant. In the Suikerbosrand Nature Reserve, at 1900m, the 

foodplant is Lotononis erianthe (Fabaceae), and the eggs are 

laid on the stems. At Ruimsig, the larvae shelter during the day 

in the nest of the widespread ant Acantholepis capensis Mayr. 

The larvae apparently release a pheromone which appears to 

mimic the brood pheromone of the ant, causing the ant to treat 

the larvae as its brood. At night, the larvae emerge from the nest 

to feed on the foodplant. During these journeys, they apparently 

release a pheromone which imitates the alarm pheromone of the 

ant. This excites the attendant ants and partially protects the 

larvae while they feed. The larvae pupate within the ants' nests. 

When they emerge from the pupae, the adults run along the 

tunnels in the nest with wings still unexpanded, only expanding 

them once outside the nest. 

The eggs of A. d. dentatis are 0.8mm in diameter and 0.5 mm 

high. They are bun-shaped, with a bold network of ridges. The 

colour is creamy-white at first, becoming purple-brown with 

development. All six larval instars are similar in appearance. 

The head is dark brown with light brown setae. The broad neckshield 

and small rounded anal shield are dark brown. The body 

is grey with longitudinal streaks and markings of reddishbrown. 

The tubercle cases on the eighth segment are black and 

bear the characteristic protective spines. The retractile tubercles 

are white. The honey gland is absent. Maximum length of the 

final instar larva is 18–19mm. The pupa is 12–13mm long, 

golden-brown and rounded (Henning 1983a,b, 1984a,b, 1987; 

Tite and Dickson 1968a,b). 

Adults are on the wing from August to April with a peak 

from October to December (Henningl983a,b, 1984a, 1987; 

Tite and Dickson 1968b). 

A. d. maseruna inhabits flat grassveld, usually near water or 

marshy areas. The habitats are sparsely grassed, the ground 

being sandy with gravel. The adults sit on sand or gravel

patches, the males establishing territories from which to chase 

intraspecific intruders and court females. The males fly low and 

fast, and in wide circles, generally returning to their original 

spots. The females fly slowly about in the same areas. When a 

female enters the territory, the male approaches from the rear 

and 'shimmies' just behind her. When unresponsive, she flies 

off. Alternatively, when responsive, she immediately settles. 

The male settles beside her and sidles up, but if she is not ready 

to mate, she turns away from him in a 'rejection posture', 

causing him to fly off. The actual mating has not yet been 

recorded. The fertilised female then spends her time searching 

for the foodplant Hermannia jacobifolia (Sterculiaceae) on 

which to lay eggs. The female alights on the plant and 

immediately begins searching with her antennae for ant 

pheromone trails. The ant species has not yet been recorded. 

The female will search the 200mm plant from top to bottom and 

even the ground around the base. When she is appropriately 

stimulated, she usually lays two eggs, side by side, on the stem 

of the foodplant. 

The egg of A. d. maseruna is also bun-shaped with a 

pronounced network of ridges. Initially, the egg is greyishwhite, 

becoming purple-brown. The remainder of the life history 

is unrecorded. 

A. d. maseruna is on the wing from November to February 

(Henning and Henning 1989). 

Threats. The establishment of the butterfly reserve at Ruimsig, 

specifically for A. d. dentatis, and its presence in the 

Suikerbosrand Nature Reserve suggests that this butterfly is no 

longer under immediate threat. However, it is essential to 

continue monitoring this subspecies, especially as the Ruimsig 

population is so small (Henning and Henning 1985, 1989). 

Towards this end, over a two week period in December 1989 

and January 1990, an estimation was made of the population 

size at the Ruimsig Reserve. By using a mark-recapture method 

it was estimated that the population size was about 400 specimens 

on the wing. This was far greater than was expected. This study 

will be continued on an annual basis to determine whether the 

management methods at the reserve are successful or not. 

Recent records for A. d. maseruna from Maseru are sparse. 

The most suitable habitats have been used for farming, or 

155 

washed away by erosion. Another recorded locality is Ladybrand, 

apparently along the banks of a river where it has not been seen 

in recent times. A new locality is at Heilbron, where it has been 

found next to a dam in the town and also 10km further north on 

a slight slope above a stream. The locality next to the dam is in 

the grounds of a recently erected old age home, and as this 

development continues, the colony may well disappear. The 

other locality north of Heilbron is apparently safe, as is the 

Boons locality, which is in a large, flat, marshy area. 

Conservation: In 1985 the Roodepoort City Council established 

a 12ha reserve at Ruimsig for A. d. dentatis. The butterfly is also 

found in the Suikerbosrand Nature Reserve. 

No conservation measures are currently in force for A. d. 

maseruna. 

References 

HENNING, S.F. 1983a. Biological groups within the Lycaenidae (Lepidoptera). 

J. ent. Soc. S. Afr. 46: 65–85. 

HENNING, S.F. 1983b. Chemical communication between lycaenid larvae 

(Lepidoptera: Lycaenidae) and ants (Hymenoptera: Formicidae). J. ent. 

Soc. S. Afr. 46: 341–366. 

HENNING, S.F. 1984a. The effect of ant association on lycaenid larval 

duration (Lepidoptera: Lycaenidae). Ent. Rec. J. Variation 96: 99–102. 

HENNING, S.F. 1984b. Southern African Butterflies, with illustrations by 

Clare Abbott. Macmillan South Africa, Johannesburg. 

HENNING, S.F. 1987. Myrmecophily in lycaenid butterflies (Lepidoptera: 

Lycaenidae). Ent. Rec. J. Variation 99: 215–222. 

HENNING, S.F. and HENNING, G.A. 1985. South Africa's endangered 

butterflies. Quagga 10: 16–17. 

HENNING, S.F. and HENNING, G.A. 1989. South African Red Data Book– 


158, Council for Scientific and Industrial Research, Pretoria, 175 pp. 

MURRAY, D.P. 1935. South African Butterflies: A Monograph of the Family 

Lycaenidae. John Bale, Sons and Danielsson, London. 

PENNINGTON, K.M. 1978. Pennington's Butterflies of Southern Africa. Ed. 

by C.G.C. Dickson, with the collaboration of D.M. Kroon. Ad. Donker, 

Johannesburg. 

TITE, G.E. and DICKSON, C.G.C. 1968a. The Aloeides thyra complex 

(Lepidoptera: Lycaenidae). Bull. Brit. Mus. (Nat. Hist.)Ent. 21:367–388. 

TITE, G.E. and DICKSON, C.G.C. 1968b. The genus Aloeides and allied 

genera (Lepidoptera: Lycaenidae). Bull. Brit. Mus. (Nat. Hist.) Ent. 29: 

225–280.

Erikssonia acraeina Trimen; Subfamily Theclinae, Tribe Aphnaeini 





Countries: South Africa, Namibia, Zaire, Zambia. 

Status and Conservation Interest: Status – vulnerable; rare 

(Red List). 

There are no recent reports of known colonies except for the 

South African colony, which is thriving on a private farm. This 

colony is presently being investigated by the Transvaal Nature 

Conservation Department, with a view to circumventing the 

threat of agricultural development destroying the colony. No 

conservation measures are being taken to preserve the other 

colonies. 

Taxonomy and Description: The upperside is orange with a 

narrow, black marginal border, a black, subcostal, discal patch 

on the forewing and a thin, black, post-discal band on the 

hindwing (Pennington 1978). The underside is orange with 

thin, post-discal, black lines and scattered black spots. The 

female is similar to the male but with a more rounded wing 

shape. Forewing lengths: male 15–18mm; female 16–21mm. 

Distribution: This species has a scattered distribution, occurring 

in Ovamboland in Namibia, and the Waterberg Mountains 

(1700m) west of Ny lstroom in the western Transvaal. It has also 

been recorded from Mongu (Barotse Province, Zambia) and in 

Zaire. 

Population Size: The Waterberg colony in South Africa is 

thought to be strong although no figures on population size are 

available. There are no recent reports from the Zambian colony, 

and it has not been recorded from Ovamboland, Namibia since 

the type was collected. 

Habitat and Ecology: E. acraeina flies in open, grassy savannah 

with sandy soil, in occasional localities where its foodplant 

Gnidia kraussiana (Thymelaeaceae) and host ant (Acantholepis 

sp.) occur together. It does not range far from its host plant and 

ant. The males establish small territories among the foodplants 

in which they can be found throughout most of the day. The 

females fly randomly within the area, and are as common as the 

males. When basking on a sandy patch in the sunshine, 

individuals have been seen to lie down on their side maximising 

156 

exposure. They settle on the ground, on small plants or on grass. 

Individuals are often seen feeding on flowers. E. acraeina flies 

slowly and weakly, the bright colour and slow flight indicate 

that the species is probably unpalatable through sequestering 

toxins from its foodplant. It does not mimic an Acraea as the 

name implies but has developed aposematic colouring 

independently. The most closely related genus is probably 

Aloeides, of which most species have tawny-orange colouring. 

The eggs are laid in coarse sand at the base of the foodplant 

near the entrance to the ants' nest. The egg is dome-shaped with 

irregular raised convolutions, giving a truffle-like appearance. 

Convolutions are absent at the micropyle, which is large, round 

and deeply indented. When first laid it is yellowish-ochre, later 

darkening to grey or greyish brown. 

The larva shelters in the nest during the day, emerging at 

night to feed on the foodplant. All instars appear similar. The 

body is pinkish-grey with a maroon longitudinal line down the 

centre of the dorsal surface, flanked on either side by a bluishgreen 

area. Laterally, the larvae are marked with regular reddishbrown 

markings. The honey gland on the seventh segment is 

well developed and the retractile tubercles on the eighth are 

white. The sixth (final) instar reaches a maximum length of 

35mm. 

Pupation takes place within the ants' nest. The pupa is at 

first bright yellow darkening to a deep ochre with a brownish 

dorsal line within 48 hours. Pupal length is about 15mm 

(Henning 1984). 

Adults are on the wing from November to February (Henning 

1984; Henning and Henning 1989). 

Threats: The Waterberg colony, although strong, is on a 

private farm, and therefore may be susceptible to agricultural 

development in the long term. There are no recent reports from 

the colonies in Zaire or Namibia. Similarly, threats to the 

Zairean colony are unknown (Henning and Henning 1989). 

Conservation: The status of this species is currently being 

investigated by the Transvaal Nature Conservation Department 

with assistance from the Lepidopterists' Society of Southern 

Africa. No conservation measures are being taken to conserve 

the species at the other localities.

References 

HENNING, S.F. 1984. Life history and behaviour of Erikssonia acraeina 

Trimen (Lepidoptera: Lycaenidae). J. ent. Soc. S. Afr. 47: 337–342. 

HENNING, S.F. and HENNING, G.A. 1989. South African Red Data Book- 

157 





Johannesburg.

Alaena margaritacea Eltringham; Subfamily Lipteninae, Tribe 

Pentilini 






Status and Conservation Interest: Status – vulnerable (Red 

List). 

This liptenine is endemic to one locality in the northeastern 

Transvaal. Two colonies are known high in the Strydpoort 

Mountains where the butterfly inhabits the steep grassy slopes 

and lichen-covered rocks 400m from the summit. Planting of 

stands of pine trees in the area poses a threat, although the 

Transvaal Nature Conservation Department is aware of the 

situation. 

Taxonomy and Description: This is a small species (Clark and 

Dickson 1971; Murray 1935; Pennington 1978) with elongated 

forewings. The upperside is black with a broad orange band, 

which is very broad in the female, almost reaching the base. The 

underside of the hindwing is creamy white with an intricate, 

lace-like pattern of thin black lines. Forewing lengths: male 

12–13.5mm; female 14–15mm. 

Distribution: A. margaritacea is endemic to South Africa, and 

is known only from one locality near Haenertsburg in the 

northeastern Transvaal (Henning and Henning 1989). 

Population Size: Although the two colonies near Haenertsburg 

only cover an area of about lha, there can be numerous 

individuals. 

Habitat and Ecology: Two secluded colonies are known from 

the slopes of the Strydpoort Mountains about 400m below the 

peaks. A. margaritacea flies on the steep grassy slopes with 

large lichen-covered rocks (Swanepoel 1953). Near midday, 

the males have been recorded ascending almost to the mountain 

tops, where they establish small territories at the base of rocky 

ridges just below the peaks. A. margaritacea has a weak 

fluttering flight, and when disturbed, it flies a few metres before 

settling again on a grass stem. If repeatedly disturbed, it may fly 

for a hundred metres before settling again. The males normally 

158 

establish territories in the breeding area, perching on grass 

stalks and fluttering around the grass. When another male 

enters its territory, it sometimes chases the intruder away. The 

female flutters slowly throughout the breeding area, searching 

for suitable lichen on which to lay her eggs. 

The foodplant is probably rock lichen, although the female 

sometimes lays on rocks and stones supporting only a little 

lichen. The eggs are laid singly or in small clusters. The eggs are 

0.9mm in diameter, 0.4mm high, and purple-brown. There are 

four rings of 14, round indentations on each egg. Those at the 

micropyle are narrow and elongated. Nothing is known of 

larval behaviour. The first instar is purple-brown with a pale 

yellow neck-shield and white humps which bear the outer 

dorsal and lateral setae. The head is purple-brown. The 

subsequent instars and pupa are unrecorded. 

The adult is on the wing in December and January, with a 

peak towards the end of December. 

Threats: The planting of plantation pine trees is a serious threat 

to this species. 

Conservation: The Transvaal Nature Conservation 

Department is aware of the threats to this species, and is 

currently monitoring it. 

References 

CLARK, G.C. and DICKSON, C.G.C. 1971. Life Histories of the South 

African Lycaenid Butterflies. Purnell and Sons, Cape Town. 

HENNING, S.F. and HENNING, G.A. 1989. South African Red Data Book– 



MURRAY, D.P. 1935. South African Butterflies: A Monograph of the Family 

Lycaenidae. John Bale, Sons and Danielsson, London. 



Johannesburg. 

SWANEPOEL,D.A. 1953. Butterflies of South Africa: Where, When and How 

they Fly. Maskew Miller, Cape Town.

Orachrysops (Lepidochrysops) ariadne (Butler); Subfamily 

Polyommatinae, Tribe Polyommatini 



' 5 Alexander St., Florida 1709, South Africa 



Status and Conservation Interest: Status – rare; endangered 

(Red List). 

O. ariadne is a highly localised Natal endemic. One of the 

two recorded colonies has become extinct, and the other occupies 

1 hectare on private land. Neglectful management of the site is 

posing a serious threat. Much more biological information on 

the species is required, as is a management plan for its protection. 

Taxonomy and Description: This species has rounded wings. 

The males are dull violaceous-blue on the upperside with 

narrow dark brown margins. The underside is dark greyishbrown 

with black spots and a distinct line of clearly-defined 

white post-discal marks. The female is brown on the upperside 

with blue areas reduced to the basal half of the wings; the 

underside is similar to that of the male. Forewing lengths: male 

13–17 mm; female 13–19 mm. 

Distribution: O. ariadne is endemic to Natal (Henning and 

Henning 1989). It has only been found in the Karkloof District, 

although previously it was recorded nearby at Balgowan. A few 

specimens which could represent this species have been recorded 

from Nkandla near Eshowe. 

Population Size: The population size of the only confirmed 

colony (at Karkloof Falls) is not known. 

Habitat and Ecology: O. ariadne inhabits a one hectare area of 

steep grassy slopes adjacent to forests (Pennington 1978). It 

occurs in tall grass on the north side of the stream running down 

to the top of the Karkloof Falls. On the south-facing steep slope, 

among the tall Hyparrhenia spp. grasses, the foodplant 

Indigofera astragalina (Fabaceae) is found (Swanepoel 1953). 

159 

O. ariadne is a fast, low flier, but may fly to a height of two 

metres to clear the tall grassheads. The males patrol up and 

down the valley, dodging in and out of the tall grass, sometimes 

venturing into the adjacent valley across the stream, but always 

returning to where the foodplant grows. They do not appear to 

be strongly territorial. 

This species is on the wing from l000h to 1430h. The 

females spend much time looking for the foodplant on which to 

lay their eggs. They fly more slowly than the males, and are 

always found in the vicinity. Little is known of the life-history, 

although early instars have been seen to feed on the foodplant 

before going down, as third instars, into an unidentified ant nest 

to feed on the brood. 

Threats: The habitat is being overgrown. In the past, vertebrate 

herbivores and natural fires kept the site in a more open, earlier 

successional stage suitable for the foodplant and ant host. 

Conservation: Careful monitoring of the site and active 

management is advisable to prevent the colony disappearing. 

No formal conservation measures are currently in force, aside 

from some interest by the owners of the land. 

References 

HENNING, S.F and HENNING, G.A. 1989. South African Red Data Book - 


158, Council for Scientific and Industrial Research, Pretoria. 175 pp. 



Johannesburg. 

SWANEPOEL,D.A. 1953. Butterflies of South Africa: Where, When and How 

they Fly. Maskew Miller, Cape Town.

Country: Australia 

Hypochrysops C. and R. Felder 

D.P.A. SANDS 

Division of Entomology, CSIRO, Meiers Road, Indooroopilly, Queensland 4068, Australia 

Status and Conservation Interest: Status – several species 

endangered or rare. 

Habitat destruction has affected seven species whose survival 

is now considered to be threatened in four Australian States 

(Table 1, last column). The most threatened species in the 

genus, H. piceatus, is only known from two localities in 

southern Queensland, one formerly at Millmerran but now 

cleared for farming and the other consisting of a few kilometres 

of roadside savannah near Leyburn. Together with Paralucia 

spinifera Edwards & Common, these two butterflies are the 

most 'at risk' of all the Australian butterflies. It is possible that 

other localities will be found for H. piceatus but none has yet 

been discovered despite searches of suitable intact open forest 

containing the foodplant, Casuarina luehmannii. Fortunately, 

the butterflies are not uncommon in some years and have 

maintained their abundance despite heavy collecting at times 

by amateurs. 

Taxonomy and Description: Species of Hypochrysops are 

renowned among lepidopterists for the distinctive and beautiful 

patterns on the undersides of their wings. Unlike the closely 

related genus Philiris Rober, most species of Hypochrysops are 

easily distinguished and were described before the turn of the 

century. All except one (H. piceatus Kerr, Macqueen & Sands) 

of the species occurring in Australia were described by the time 

Waterhouse's 'WhatButterfly is That?' (1932) was published. 

However, the specific and subspecific status of some populations 

has been reviewed recently (Sands 1986) and some taxonomic 

identities changed. 

Distribution: The majority of the 57 species recognised (Sands 

1986), occur on the neighbouring islands north of Australia and 

only one (H. coelisparsus [Butler]) occurs west of Wallace's 

line. Six of the 18 species of Hypochrysops in Australia are 

endemic while the remainder are also found on mainland New 

Guinea. 


160 

Habitat and Ecology: The life history of H. ignitus ignitus 

(Leach) was described by Waterhouse (1932) and it differs only 

slightly for subspecies erythrinus. Adults oviposit on small (up 

to 2m) plants of Eucalyptus confertiflora or Acacia spp. and 

their larvae shelter by day in byres of the attendant ant, 

Iridomyrmex nitidus, constructed on the stems and trunk of the 

foodplant. At the end of the wet season populations normally 

stabilise after the dry season stress – a time when much of their 

habitat is now burnt intentionally as a precaution against wild 

fires. 

Threats: Not all species 'hilltop' but for those that do, human 

activities such as clearing and installation for radio and television 

equipment, forestry observation towers and mining of rock 

outcrops often result in destruction of the hilltop habitats and 

local extinctions. The most susceptible to these activities are the 

subspecies of H. delicia Hewitson and H. ignitus (Leach). 

It is likely that the intentional annual burning of savannah in 

the Northern Territory near Darwin contributes to the extreme 

rarity of H, ignitus erythrinus (Waterhouse & Lyell). The firing 

is carried out by conservation authorities in the belief that the 

original inhabitants did so as a method for hunting wildlife. 

However, these fires now started at the end of every wet season 

are quite devastating to several insect species and particularly 

to H. ignitus erythrinus. 

At the end of the wet season populations of H, ignitus ignitus 

normally stabilise after the dry season stress. The abundance of 

this subspecies, dependant on suckers and low regrowth of the 

larval foodplants, has been dramatically reduced since routine 

fuel reduction burning began. It is said that vertebrates are not 

affected by these 'slow burns' since they are sufficiently mobile 

to escape from the advancing flames. Immature stages of 

arthropods, on the other hand, are incinerated when surrounded 

by combustible materials close to the ground. There is an urgent 

need to review these burn policies and to take account of sessile 

organisms, perhaps by providing fire-free areas where dry 

country habitats of rare insect fauna are allowed to stabilise. 

The Australian species of Hypochrysops are local, 

uncommon or rare, occurring in undisturbed habitats and very 

few if any are able to adapt to encroaching urbanisation. H.

pythias (Felder & Felder) occurs at times in hundreds in Papua 

New Guinea, especially where its foodplant Commersonia 

bartramia has regrown after human disturbance, but the 

Australian subspecies euclides has never been observed in such 

numbers even though the food plant is the same and often a 

predominant regrowth plant. It is an unusual species as adults 

are known to aggregate in large numbers at night in Papua New 

Guinea. 

For those species that congregate on undisturbed hilltops 

males are sometimes seen in numbers but the females are 

always uncommon. Hilltops are selected as locations where 

unmated female adults can, with a degree of certainty, find a 

mate among the competing males. 

Australian tropical and subtropical rainforests have been 

depleted to a fraction of their original area, and conservation of 

the remainder is essential if the survival of several species of 

Hypochrysops is not to be threatened. The fringing rainforests 

along the Rocky River have suffered from gold mining activities 

placing at risk survival of the unique H. theon cretatus Sands. 

Nowhere else is this recently-described subspecies known to 

occur. It is the most southern population of a species found 

throughout mainland New Guinea and is the most distinctive of 

all of the seven subspecies recognised (Sands 1986). Fortunately, 

much of the habitat for another subspecies, H. theon medocus 

(Fruhstorfer), and the related H. hippuris nebulosis Sands is not 

at risk. However, the rainforest margin species, H. miskini 

miskini (Waterhouse) is seriously threatened by habitat 

destruction in southeastern Queensland. Formerly common 

near Burleigh Heads, on the Coomera River and near Rainbow 

Beach, the species has disappeared from most localities and is 

becoming rare at the remainder. Although slightly different in 

morphology from the southern populations, populations of H. 

miskini from the northern localities are not so threatened. 

At the edge of its range in northern NSW, H. digglesii 

(Hewitson) is now confined mainly to moist savannah near 

Broken Head. Unfortunately, although most of the nearby 

rainforest is included in a National Park, the breeding sites for 

this species are at present threatened by local development 

proposals. 

It is important to note that this is the only locality remaining 

intact for this species in NSW and that it differs quite considerably 

in appearance from the more abundant populations in 

Queensland. 

Coastal paperbark (Melaleuca spp.) swamps are habitats for 

several species destroyed on a large scale in recent years. H. 

apollo apollo M iskin was once quite abundant at a few paperbark 

localities including Cardwell, the southern edge of its range and 

at Port Douglas, northern Queensland. However, forestry 

activities, mainly clearing and planting with Pinus sp. and earth 

works for a canal development at the first locality and clearing 

for a golflinks at the second, have seriously damaged the major 

breeding sites for this unusual butterfly. H. apollo apollo has 

suffered from clearing of the paperbark swamps that support 

the epiphytic plant, Myrmecodia beccari, the larval foodplant. 

Table 1. Hypochrysops spp. currently known to survive at five or fewer localities in States where they were formerly widely distributed. 

Taxon 

H. apelles apelles 

H. apollo apollo 

H. byzos byzos 

H. cleon 

H. digglesii 

H. epicurus 

H. hippuris nebulosus 

H. ignitus ignitus 

H. ignitus erythrinus 

H. piceatus 

H. theon cretatus 

Key 

NSW 

Q 

Q 

Q 

NSW 

NSW 

Q 

SA 

NT 

Q 

Q 

State† with nos 

localities 

intact 

† NSW New South Wales * MG mangroves 

0 Queensland SV savannah 

NT Northern Territory RF rainforest 

SA South Australia 

V Victoria 

WA Western Australia 

c. 4 

c. 5 

c.3 

1 

1 

c.5 

1 

1 

2 

1 

1 

161 

Present 

elsewhere 

(state*) 

Q, NT 

– 

NSW 

– 

Q 

Q 

– 

Q, V, NSW 

WA 

– 

– 

Type* 

MG 

SV 

SV 

RF 

SV 

MG 

RF 

SV 

SV 

SV 

RF 

Habitats 

At risk 

+ 

+ 

– 

– 

+ 

+ 

– 

+ 

+ 

+ 

+

Another factor that is decreasing the abundance of the butterfly 

is removal of the bulbous foodplants by collectors and who cut 

them up in search of pupae. Colonisation of the bulbs by the 

exotic ant Pheidole megacephala may also affect survival of H. 

apollo since it displaces the native Iridomyrmex cordatus, the 

natural bulb inhabitant. Indeed, P. megacephala may eventually 

threaten the survival of ant plants, as it is known to destroy the 

developing seeds. This is probably one of the very few examples 

of a conservation issue for Australian butterflies involving 

threat to the breeding sites by collectors. The same habitat is 

shared with the much more abundant H. narcissus narcissus 

(F.) which breeds on mistletoes, a species also present in and 

breeding on mangroves in northern Queensland. 

Mangroves are important breeding sites for a number of 

interesting lycaenid butterflies. In southeastern Queensland 

and northern NSW, the coppery H. apelles apelles (F.) only 

occurs near saltwater swamps where the larvae feed on several 

species of mangroves. The species was once very common near 

Southport and on the Tweed River but its numbers have declined 

162 

in recent years and it has disappeared from several localities. 

Housing developments, clearing of mangroves and possibly, 

spraying with insecticides have resulted in the species becoming 

scarce in NSW and uncommon in southern Queensland. Further 

north in Queensland a wide range of plants are food for larvae 

and the species is not at risk. Another species, H. epicurus 

Miskin, has suffered from similar destruction of its only 

mangrove (Avicennia marina) habitat in southeastern 

Queensland and NSW. This species has become quite rare at 

localities where it was formerly very common due to habitat 

interference. 

References 


(Lepidoplera: Lycaenidae). Entomonograph No. 7. E.J. Brill, Leiden. 

WATERHOUSE, G.A. 1932. What Butterfly is That? Angus & Robertson, 

Sydney.

Illidge's Ant-Blue, Acrodipsas illidgei (Waterhouse and Lyell) 

Country: Australia. 

P.R. SAMSON 

Bureau of Sugar Experiment Stations, P.O. Box 651, Bundaberg, Queensland 4650, Australia 

Status and Conservation Interest: Status – rare (Common 

and Waterhouse 1981). 

Its life cycle is unusual, larvae being obligate 

myrmecophages. It has a restricted distribution and occurs 

almost exclusively among mangroves, a habitat which is under 

threat from development in the populous area where the species 

is known. A proposal for a canal estate and residential subdivision 

was rejected in a local government court decision in Brisbane 

in 1989, based in part on the need to preserve colonies of A. 

illidgei occurring in the area. In July 1990, A. illidgei became 

the first butterfly to be designated as 'Permanently Protected 

Fauna' in Queensland. 

Taxonomy and Description: A. illidgei was originally described 

as a subspecies of A. myrmecophylla (Waterhouse & Lyell), 

from which it differs by its larger size, the broader markings 

beneath the wings and the male genitalia (Kerr, Macqueen and 

Sands 1968). 

Acrodipsas Sands contains seven described species, all 

endemic to Australia (Common and Waterhouse 1981). They 

were previously referred to as Pseudodipsas C. and R. Felder. 

Four species have been described since 1965. Two more probable 

species await description (D. Sands, pers. comm.). All species 

except A. illidgei are known mostly from males collected on 

hilltops in open Eucalyptus forest. The life histories are poorly 

understood, and only A. illidgei and two others have been reared 

from the immature stages. 

Distribution: A. illidgei is restricted to coastal areas, mostly in 

mangrove habitats. For many years the species had only been 

recorded between Brisbane and Burleigh Heads in southern 

Queensland, but there are recent records from Mary River Heads 

near Maryborough, Queensland (Manskie and Manskie 1989) 

and Brunswick Heads, New South Wales (G. Miller pers. comm.). 

Population Size: Almost all recent specimens have been 

collected at Burleigh Heads or at Redland Bay near Brisbane. 

There are no estimates of the population sizes at these sites. 

Newly-emerged female of Acrodipsas illidgei (photo by P.R. Samson). Final instar larva of Acrodipsas illidgei with associated ants and brood 

(photo by P.R. Samson). 

163

Habitat and Ecology: The colonies at Burleigh and Redland 

Bay occur in communities dominated by the grey mangrove 

Avicennia marina. Other records from Mary River Heads, 

Hay's Inlet near Brisbane, and Brunswick Heads are also from 

mangrove areas. 

Adults of A. illidgei are small and inconspicuous, and are 

not easy to see among the dense mangrove vegetation. My 

observations at Redland Bay suggest that breeding occurs in 

patches of trees. I have found 25 eggs laid on a single mangrove 

tree with smaller numbers on adjacent trees. On that occasion 

the vast majority of trees apparently carried no eggs, despite the 

presence of ant colonies in some of them. Whether these 

patches remain stable over time is unknown. 

Of the immature stages, only the eggs are in exposed 

positions, and they are difficult to locate. Eggs are laid on 

branches or under loose bark of trees colonised by the associated 

ants. Eggs are aggregated on the particular trees used for 

breeding. They hatch in about one week during summer. 

The first instar larvae are carried into ant nests by the small 

ant, Crematogaster sp. (laeviceps group) (Samson 1987,1989). 

The nests occur in cavities (mostly in borer holes) in the wood 

of living trees. There, the larvae prey on the ant brood throughout 

their development. Pupae also occur inside the nest and several 

larvae or pupae may be found together in a hollow branch. 

Larvae and pupae occurring in these nests can only be found by 

destroying the branches and ant colonies in which they live. 

Almost all have been found inside nests in A. marina. However, 

several have been found beneath bark of Eucalyptus (Smales 

and Ledward 1942), indicating that there is no special 

relationship with Avicennia. 

Collecting records suggest that there are at least two 

generations each year, with maximum frequency of capture in 

September and December to February: no adults have been 

taken during the colder months of May through July. 

Adults have been seen feeding at flowers (Hagan 1980). 

Like some other species of Acrodipsas, females possess many 

fully developed eggs at emergence from the pupa (Sands 1979). 

Another lycaenid, Hypochrysops apelles (Fabricius), has an 

obligate relationship with Crematogaster sp. (laeviceps group) 

in southern Queensland. That butterfly occurs continuously 

along the coast to northern Queensland (Common and 

Waterhouse 1981), probably attended by the same ant. It is 

surprising, therefore, that A. illidgei has such a restricted 

distribution. 

Threats: Survival of A. illidgei is threatened by the loss of 

mangroves to residential development. The species' distribution 

coincides with the most populous region of Queensland, where 

coastal development is proceeding at a rapid rate. Fogging with 

insecticides to control biting insects that breed in mangroves 

may also threaten A. illidgei. 

Collecting poses little threat to the species. Only a small 

number of collectors have visited the breeding sites. The 

butterflies do not have a sharply defined flight period and are 

very difficult to find and capture in the habitat. Collecting of 

immature stages destroys the ant galleries that are searched. 

164 

However, it is only the galleries in the smaller, dead branches 

of the mangroves that are readily opened up by collectors: the 

greater part of the ant nests in the large branches and trunks of 

the mangroves are inaccessible. 

Conservation: Part of the habitat of A. illidgei at Burleigh 

Heads is protected within an environmental park established to 

preserve what little remains of mangroves in the area. Whether 

A. illidgei still breeds within the park is not known. 

Recently, approval for a canal estate proposal at Point 

Halloran, Redland Bay was rejected because of deleterious 

effects on the environment. The proposed development was to 

have included 610 residential allotments of which 385 were to 

have canal frontages on 106ha. The presence of A. illidgei in the 

area was one factor in the court's decision to reject the proposal 

(The Courier Mail, 17 June 1989). A small part of this area has 

since been rezoned for residential development, but the 

remainder is likely to become an environmental park. 

The colony at Redland Bay that is most often visited by 

lepidopterists is near to, but not contained within, the 

development proposal. There is at present no habitat protection 

afforded to this site. 

The mangrove habitat of A. illidgei at Mary River Heads is 

part of an area nominated for world heritage listing. A request 

from local residents for insecticidal fogging to control biting 

midges has been denied because of possible harm to the 

butterflies (The Courier Mail, 2 May 1992). 

The designation of A. illidgei as 'Permanently Protected 

Fauna' under the Queensland Fauna Conservation Act 1974–79 

on July 21, 1990 gives it an unusually high and controversial 

status (Monteith 1990). This status has otherwise been accorded 

to a few high-profile vertebrates, such as the koala and platypus. 

Permits are needed to keep specimens in private collections, 

and a separate permit is required in order to move specimens to 

different premises, or interstate. Permits may be obtained for 

scientific study of the species but a fear is that the degree of 

formality imposed is likely to deter the interest of enthusiastic 

amateurs who have contributed so much to knowledge of 

Australian Lycaenidae. 

References 


(2nd edn.) Angus and Robertson, Sydney. 

HAGAN, C.E. 1980. Recent records of Acrodipsas illidgei (Waterhouse and 

Lyell) (Lepidoptera: Lycaenidae) from the Brisbane area, Queensland. 

Aust. ent. Mag. 7: 39. 

KERR, J.F.R., MACQUEEN, J. and SANDS, D.P. 1968. The specific status 

of Pseudodipsas illidgei Waterhouse and Lyell stat. n. (Lepidoptera: 

Lycaenidae). J. Aust. ent. Soc. 7: 28. 

MANSKIE, R.C. and MANSKIE, N. 1989. New distribution records for four 

Queensland Lycaenidae (Lepidoptera). Aust. ent. Mag. 16: 98. 

MONTEITH, G. 1990. Another butterfly protected in Queensland. Myrmecia 

August 1990: 153–154. 

SAMSON, P.R. 1987. The blue connection: butterflies, ants and mangroves. 

Aust. nat. Hist. 22: 177–181.

SAMSON, P.R. 1989. Morphology and biology of Acrodipsas illidgei 

(Waterhouse and Lyell), a myrmecophagous lycaenid (Lepidoptera: 

Lycaenidae: Theclinae). J. Aust. ent. Soc. 28: 161–168. 

SANDS, D.P.A. 1979. A new genus, Acrodipsas, for a group of Lycaenidae 

(Lepidoptera) previously referred to Pseudodipsas C. & R. Felder, with 

165 

descriptions of two new species from northern Queensland. J. Aust. ent. 

Soc. 18: 251–265. 

SMALES, M. and LEDWARD, C.P. 1942. Notes on the life-histories of some 

lycaenid butterflies – Part I. Qd. Nat. 12: 14–18.


The Elthani Copper, Paralucia pyrodiscus lucida Crosby 

T.R. NEW 


Status and Conservation Interest: Status – rare, vulnerable. 

This local subspecies was feared to be extinct in the 

Melbourne area of southeastern Australia before a thriving 

colony was found in January 1987 on land already subdivided 

for imminent housing development. A major campaign was 

undertaken to (a) acquire and reserve the habitat, (b) search for 

other colonies and (c) design a management plan for the 

butterfly. This case broke new ground in increasing invertebrate 

conservation awareness in Victoria. 

Taxonomy and Description: The subspecies was described 

(Crosby 1951) from the Eltham (37°43'S, 149°09'E) / 

Greensborough area of outer northeastern Melbourne, and 

differs in the amount of copper scaling from the nominate 

subspecies, P. p. pyrodiscus (Doubleday) which occurs in parts 

of eastern Australia as far north as southern Queensland. 

Colonies around Kiata (36°22'S, 141°48'E) and Castlemaine 

(37°04'S, 144°13'E) are also at present referred to this 

subspecies. The extra-Victorian limits of the subspecies are not 

wholly clear: at present no other populations are referred to 

lucida formally. 

Paralucia Waterhouse and Turner contains three species, 

all endemic to Australia. One, P. aurifera (Blanchard), is 

widespread but local and the other two are rare. 

Distribution: The subspecies is very restricted in isolated parts 

of Victoria. Eight discrete colonies are known around Eltham; 

the subspecies otherwise occurs only at Kiata (six colonies 

within 3km of each other; one at nearby Salisbury) and 

Castlemaine (a single colony only) (Braby et al. 1992). 

Population Size: P. p. lucida was discovered near Melbourne 

in 1938. Accumulated collector wisdom shows that it was taken 

more or less regularly over the following decade or so but 

declined markedly from about 1950 onwards and thereafter 

became extremely rare. It was believed to be extinct in recent 

years until the discovery of a thriving colony near Melbourne 

in 1987. 

166 

All presently known colonies are very isolated, and the 

population structure is essentially closed. There is very little 

possibility of interchange of adults between most nearby 

suburban colonies, and none over the broader scale of 

distribution. Most of the Eltham colonies are small and seem 

unlikely to be viable in the long term. Population sizes have 

been estimated both by counts of adult butterflies during the 

flight season and nocturnal counts of feeding caterpillars 

(Vaughan 1987, 1988) and as the largest (presumed viable) 

colonies occupy only a few hundred square metres, these counts 

are likely to be reasonably accurate. The major Eltham colony, 

on the subdivision land, contained an estimated 300–500 larvae. 

At the other extreme only six butterflies were observed in a 

colony on private land. Several intermediate sized colonies 

each had populations estimated at 100–150 individuals. In 

contrast, the Kiata colonies contained 'many hundreds' of 

individuals, and there are 100–200 at Castlemaine. In 1988, the 

State population of the Eltham Copper was about 2600 

individuals (Braby and Crosby in prep.). 

Habitat and Ecology: Some colonies are confined to open 

forest areas, generally on north to west-facing slopes, and 

others occur on highly degraded areas such as roadsides. The 

life cycle appears to be limited obligatorily to a spiny dwarfed 

form of Sweet Bursaria (Bursaria spinosa: Pittosporaceae) as 

the sole larval foodplant, and to plants associated with nest 

chambers of the ant genus Notoncus. There is one major 

generation each year, with adults present from late November 

to early February, and a possible small, partial second generation 

represented by a few adults in March (Braby 1990). 

Eggs are laid singly or in small groups on or near the larval 

foodplant. Caterpillars hatch after about two weeks and are 

nocturnal feeders: they retreat to Notoncus chambers at the base 

of the plants during the daytime and are regularly tended by the 

ants, these being N. enormis Szabo at Eltham but N. 

ectatommoides (Forel) at Kiata. Caterpillars pupate in the ant 

chambers after five instars, and the pupal stage of the major 

generation lasts about a month. 

Adult P. p. lucida take nectar from various flowers, including 

Bursaria. They seem to be aggressive to other butterflies, and 

males actively pursue females.

Both the stunted Bursaria and Notoncus ants are widespread 

in Victoria, and there is seemingly no shortage of sites suitable 

for P. p. lucida. Its restricted distribution is at present difficult 

to explain. 

Threats: Recent declines appear to be attributable solely to 

urbanisation, with associated removal of native vegetation, 

fragmentation and destruction. The habitats of some former 

colonies near Eltham are now housing estates and this was 

clearly the major threat to the newly discovered colonies. The 

largest colony was reduced substantially during 1988–89. The 

very small colony noted above was in a suburban garden, but 

many such gardens are rapidly converted to exotic plant species 

rather than fostering native flora. Four of the Kiata colonies are 

within an 80ha Native Plant and Wildlife Reserve. Encroachment 

of exotic weeds on the site of the sole Castlemaine colony, 

which is not protected, may be a concern (Braby and Crosby in 

prep.). Sheep grazing has apparently reduced the number of 

host plants in one degraded Kiata colony. 

Conservation: The need for conservation of P. p. lucida 

became both apparent and urgent because of the imminent 

subdivision of the major site on which the butterfly was 

fortuitously discovered. A briefing paper to the State Minister 

for Conservation, Forests and Land led to negotiations with the 

developers, who agreed to a moratorium on development until 

the feasibility of purchasing the site as a 'Butterfly Reserve' 

could be investigated. The total cost of this was projected at 

A$l million. The State Government contributed A$250,000 

and the Eltham Shire Council committed A$125,000. A highly 

organized public appeal during the next year raised a further 

$56,000 and during that time the 'Eltham Copper' became an 

important local emblem. The total of some A$426,000 is by far 

the largest sum ever committed to conservation of an invertebrate 

species/subspecies in Australia and a major part of the prime 

colony site (0.7ha) was purchased in early 1989. This was 

augmented substantially by the State Government transferring 

to the butterfly reserve an area (c. 2.6ha) of Education Department 

land adjacent to this which supported a further important 

colony. 

This case did much to increase public awareness of butterfly 

conservation and the general importance of invertebrates in the 

environment. Following initial assessment of conservation 

status (Crosby 1987), a management plan for P. p. lucida has 

been prepared (Vaughan 1987, 1988). The habitats reserved, 

though only a few hectares in extent, should be sufficient to 

sustain the populations at Eltham with adequate management. 

Management recommendations for these remnant urban 

populations include: 

i) protection of all existing colonies from threatening processes 

associated with urbanisation including: human activity 

(trampling, slashing or burning vegetation, dumping of 

garbage, sullage overflow and changes to drainage patterns, 

potential overcollection); weed invasion; overgrowth of 

food plants; and activities of other species of animals; 

167 

ii) provision for expansion of the habitat by prompting natural 

regeneration of Bursaria and propagation from seeds or 

cuttings, or transplanting plants from any sites to be destroyed 

by development; 

iii) provision for a ranger to foster practical management 

activities and monitor the effects of these. 

Listing of the subspecies under the Victorian Flora and 

Fauna Guarantee Act is controversial, although full provision 

for invertebrates is included in this pioneering legislation. A 

requirement of the Flora and Fauna Guarantee Act listing is the 

preparation of an Action Statement from the earlier management 

plan. This statement, detailing steps needed to ensure the 

butterfly's long-term survival, is at present in draft form (Webster 

1993). A further leaflet on the Eltham Copper has been issued 

recently to encourage its well being by habitat protection 

(Ahern 1993), and a Coordinating Group of scientists is 

overseeing the developing management plan. The Entomological 

Society of Victoria has placed P. p. lucida on its list of 

butterflies to which a Voluntary Restricted Collecting Code 

applies. 

Current concerns include destruction of some of the small 

(unreserved) Eltham colonies and the coordination of effective 

measures to conserve the butterfly in other parts of Victoria. 

Attempts are being made, using the facilities of the 'Butterfly 

House' at the Royal Melbourne Zoological Gardens, to establish 

captive stock for future reintroduction to the wild. 


M. Braby, D. Crosby and P. Vaughan are thanked for information 

additional to that included in the published reports. 

References 

AHERN, L. 1993. Eltham Copper Butterfly 'The People's choice'. Land for 

Wildlife Note No. 21. Department of Conservation and Natural Resources, 

Victoria. 

BRABY, M.F. 1990. The life history and biology of Paralucia pyrodiscus 

lucida Crosby (Lepidoptera: Lycaenidae).J. Aust. ent. Soc. 29: 41–51. 

BRABY, M.F. and CROSBY, D.F. In prep. The status and conservation of the 

Eltham Copper Butterfly, Paralucia pyrodiscus lucida Crosby, in Victoria, 

Australia. 

BRABY, M.F., CROSBY, D.F. and VAUGHAN, P.J. 1992. Distribution and 

range reduction in Victoria of the Eltham Copper butterfly Paralucia 

pyrodiscus lucida Crosby. Viet. Nat. 109: 154–161. 

CROSBY, D.F. 1951. A new geographical race of an Australian butterfly. 

Viet. Nat. 67: 225–227. 

CROSBY, D.F. 1987. The conservation status of the Eltham Copper Butterfly 

(Paralucia pyrodiscus lucida Crosby) (Lepidoptera: Lycaenidae). Arthur 

Rylah Institute for Environmental Research. Tech. Rpt. No. 8, Melbourne. 

VAUGHAN, P.J. 1987. (1988). The Eltham Copper Butterfly draft management 

plan. Arthur Rylah Institute for Environmental Research. Tech. Rpt. No. 

57, Melbourne. 

VAUGHAN, P.J. 1988. Management plan for the Eltham Copper Butterfly 

(Paralucia pyrodiscus lucida Crosby) (Lepidoptera: Lycaenidae). Arthur 

Rylah Institute for Environmental Research. Tech. Rpt. No. 79, Melbourne. 

WEBSTER, A. 1993. Eltham Copper Butterfly, Paralucia pyrodiscus lucida. 

Action Statement No. 39. (draft). Department of Conservation and Natural 

Resources, Victoria.

The Bathurst Copper, Paralucia spinifera Edwards and Common 

Country: Australia 

E.M. DEXTER and R.L. KITCHING 

Department of Ecosystem Management, University of New England, Armidale, NSW 2351, Australia 

Status and Conservation Interest: Status – endangered 

(Common and Waterhouse 1981). 

The species, endemic to Australia, is known only from three 

localities all within 50km of each other. The populations are 

highly fragmented and are subject to various threats including 

grazing, pasture improvements and establishment of forestry 

plantations. 

Taxonomy and Description: The Australian endemic genus 

Paralucia Waterhouse and Turner contains three species, P. 

spinifera, P. aurifera and P. pyrodiscus. 

The first specimen of P. spinifera was collected in October 

1964 at Yetholme (33°27'S 149°48'E) New South Wales, by 

I.F.B. Common and M.S. Upton. Despite subsequent searches, 

the species was not found again until October 1977 and it was 

officially described and named by Edwards and Common 

(1978). Apart from the difference in size, shape and colour of 

the wing, Edwards and Common (1978) distinguished P. 

spinifera from its congeners P. aurifera (Blanchard) and P. 

pyrodiscus (Rosenstock) by its spine-like process that extends 

over the base of the tarsus on the tip of each fore tibia. Both 

sexes have this feature. 

Distribution: P. spinifera is known to occur at three main 

localities which are all within 50km of each other. The 

populations are highly fragmented both naturally, by occurring 

on ridges above 900m in altitude, and artificially, by pine 

plantations and areas of improved pasture. 

Population Size: Site A (see below) has the highest population 

of P. spinifera amounting to approximately 1120 individuals 

sighted over an eight-week period. Population sizes at the other 

two localities are smaller. 

Habitat and ecology: The three known localities for the 

species are remarkably uniform in geography, which suggests 

that the species has very specific microhabitat requirements. 

This has been confirmed by later studies (Dexter and Kitching 

unpublished). 

168 

All sites are on the edge of open eucalypt woodland (Specht 

1981) and usually on west to north-west facing slopes. The 

density of P. spinifera differs markedly in the three areas and is 

correlated with a spatial pattern of the larval food plant, native 

blackthorn Bursaria spinosa Cav. (Pittosporaceae). 

The butterfly has a mutualistic relationship with the ant 

Anonychomyrma itinerans. Ant surveys of known P. spinifera 

localities repeatedly yielded other ant species which may be 

important to the butterfly's life history. These were Iridomyrmex 

sp. 1,2,3, I. purpureus, I. rufoniger group 1 and 2, and 

Ochetellus sp. 

Ant surveys were also done in nearby areas, some only 

metres away, which appeared to be suitable habitat but lacked 

P. spinifera, and on every occasion the attendant ant was absent. 

One population which was high in numbers in 1989 crashed in 

1991. A follow-up survey showed that A. itinerans was not 

present. 

Athough A. itinerans has a wider distribution than P. 

spinifera, it too is restricted to regions above 900m in altitude. 

Apart from the central tablelands, A. itinerans is also found at 

Piccadilly Circus (35°22'S 148°49'E) (Australian Capital 

Territory), Barrington Tops (31°59'S 151°27'E) and Brown 

Mountain (36°36'S 149°23'E) in New South Wales. 

P. spinifera is univoltine and has a very early flight season 

which starts in the last week of September and finishes in mid- 

November. During this period the adult emergence is scattered 

and matings occur throughout the eight-week period. 

After mating, the female oviposits on bushes with ants, 

presumably detected using olfactory cues, and lays singletons 

or groups of up to four white eggs which darken to green with 

maturity. The egg group size of P. spinifera is considerably 

smaller than that of the closely related P. aurifera and P. 

pyrodiscus which are 15 and 12 eggs respectively (Braby 

1990). Eggs are laid on the lower third of the bush and are 

positioned on either the underside of leaves on the main trunk 

or on debris at the base of the plant. Eggs take approximately 15 

days to hatch. During the egg phase, the attendant ants are 

constantly searching the host plant, possibly seeking newly 

hatched larvae. 

After hatching, first instar larvae are immediately attended 

by one ant, which is curious as the early instar larvae do not have

ant-attracting organs. From instars one to three the larvae and 

ants are diurnal, feeding in both morning and afternoon and 

retreating to the nest at midday and dusk. After the third instar, 

the larvae and ants become nocturnal. The larvae of P. spinifera 

can have up to eight instars and can take between 60 and 70 days 

to pupate in laboratory conditions without ants. 

Preliminary laboratory studies (Dexter and Kitching, 

unpublished) have shown that larvae without ants remain on the 

bush permanently whilst larvae with ants return to the base of 

the plant during daylight hours. Larvae reared without ants are 

also considerably smaller at pupation than larvae which have 

been reared with ants. 

Threats: The three main localities, which for the purposes of 

this paper are termed sites A, B and C, differ greatly in the 

degree of habitat degradation and thus level of threat, which 

appears to be a reflection of the land tenure. 

Site A, which has the highest population of P. spinifera, is 

closest to the 'natural state' with a high level of plant diversity 

and a large community of ants present. This land is owned by 

the Department of Defence with restricted access, has not been 

burnt for 17 years, and has never been grazed by livestock. Site 

A also includes a nearby smaller site which is similar in 

condition to the main block described above. 

Site B supports five fragmented subpopulations and, although 

together the populations occupy an area of 5km 2 , a markrecapture 

study (Dexter and Kitching, unpublished) showed 

that the subpopulations do not intermingle and that the vagility 

of the adults is low. These sites are all on private land and have 

been or are still subject to grazing. 

Site C occurs within a nature reserve where it is fully 

protected by law. However, the population at this site is probably 

the most threatened of all. The site is heavily infested with 

blackberry, Rubus sp., is disturbed by feral pigs and threatened 

by fire control burns. Unfortunately the managing body is 

required to burn the area to reduce the risk of wildfire spreading 

from within the reserve to the neighbouring pine plantation. 

Unlike some lycaenids, for which grazing seems to be 

beneficial to the butterfly, grazing is detrimental to this species, 

as livestock trample and eat the seedlings of B. spinosa. This in 

turn inhibits juvenile recruitment of B. spinosa and can cause 

changes in the spatial pattern of the plant, leading to plant 

isolation. Larvae which occur on isolated bushes usually deplete 

their food supply so heavily that they starve or pupate 

prematurely. If larvae pupate at a smaller body size, the emerging 

adult will also be small and this can affect reproductive fitness 

because fecundity in butterflies is positively correlated with 

body size (Gilbert 1984). 

One sympathetic landholder is actively involved in managing 

her P. spinifera populations and provides hungry larvae with 

new bushes which ants and larvae colonise. It should be noted 

that for this management strategy to be successful the branches 

need to be entwined, as larvae do not seem to move across the 

ground. 

Overall the main threats to the survival of this species are: 

• ignorance of its existence and habitat degradation. The type 

169 

locality narrowly avoided being cleared during installation 

of a major power line, with no local appreciation of the 

butterfly's existence whatsoever. 

• grazing by cattle, sheep and goats, either by direct impact on 

the food plants or by use of pasture fertilisation to improve 

fodder quality. 

• clearing for establishment of forestry plantations. Along 

with pasture improvement, this process has probably 

produced the current highly fragmented distribution of the 

species in the area, although it must always have been a very 

local species. Clearing and habitat change were of course 

carried out in ignorance of the existence of the species. 

• all sites are under some threat from exotic weeds, the most 

significant being blackberry (Rubus spp.) and scotch broom 

(Cytisus scoparius). At one site, the Bursaria is in danger of 

being completely overgrown by the broom and at the only 

protected site (Winburndale Nature Reserve) uncontrolled 

spread of blackberries is a threat to part of the site at least. 

Conservation: Paralucia spinifera encapsulates many of the 

problems associated with the conservation of lycaenids. The 

scatter of highly restricted sites each with a different set of 

actual and potential problems and the relatively small changes 

that would be required to actually eliminate whole populations 

make generalisations difficult. Crucial to any management of 

the species is local awareness and sympathy and that has been 

created over the last few years. Only those who have regular, 

albeit casual, contact with the sites concerned can monitor day 

to day changes. 

Some general threats, particularly overgrowth by noxious 

weeds and damage to hostplants by feral animals, should be 

controlled by application of existing programmmes in National 

Parks and other public lands. Sites on private land need to be 

monitored for these impacts and the impact of stock, particularly 

goats, which may browse upon the host plants, and cattle which 

can produce soil impaction and other disturbance inimical to ant 

populations. 

Major land use changes involving clearing for conifer 

plantations and fertilisation for pasture 'improvement' are 

probably responsible for the present limited distribution of the 

species and obviously any further changes present additional 

threats. The occurrence of some butterfly populations on 

protected land is encouraging although other sites need to be 

monitored in this connection. One site is being proposed for 

listing under National Estate regulations because of the presence 

of this butterfly. 

There is circumstantial evidence for one site that 

overcollecting has severely reduced the population. The short 

flight period, very restricted areas in which populations occur, 

and a tendency by collectors to take long series for no justifiable 

reason may have contributed to making this species more 

sensitive than most to collecting pressures. Lepidopterists have 

argued vigorously for a self-regulatory approach to butterfly 

conservation rather than one driven by species-specific 

legislation. Collectors need to be very aware of the sensitivity 

of species like Paralucia spinifera for which self-regulation

does not appear to have worked. The case could certainly 

be made for total protection of the species. 

References 

BRABY, M.F. 1990. The life history and biology of Paralucia 

pyrodiscuslucida Crosby (Lepidoptera: Lycaenidae).J. Aust. ent. 

Soc. 29:41–51. 

170 


Angus and Robertson, Sydney. 

EDWARDS, E.D. and COMMON, I.F.B. 1978. A new species of Paralucia 

Waterhouse and Turner from New South Wales (Lepidoptera: Lycaenidae). 

Aust. ent. Mag. 5 (4): 65–70. 

GILBERT, N. 1984. Control of fecundity in Pieris rapae I. The problem. J. 

Anim. Ecol. 53: 581–588. 

SPECHT, R.L. 1981. Foliage projective cover and standing biomass. In: 

Gillison, A.N. and Anderson, D.J. (Eds). Vegetation classification in 

Australia. CSIRO and ANU Press, Canberra.

The Australian Hairstreak, Pseudalmenus chlorinda (Blanchard) 

G.B. PRINCE 

Department of Parks, Wildlife and Heritage, 134 Mrs. Macquarie's Road, Hobart, Tasmania 7001, Australia 


Status and Conservation Interest: Status – Mainland 

subspecies: P. c. chloris, rare; P. c. fisheri and P.c. 

barringtonensis, vulnerable; P. c. zephyrus, not threatened. 

Tasmanian subspecies: P. c. conara and P. c. chlorinda, 

vulnerable (both indeterminate: Red List); P. c. myrsilus and 

the un-named P. c. zephyrus-likc form, endangered. 

Several of the subspecies of P. chlorinda are extremely 

localised, particularly in Tasmania where most conservation 

attention to this species has been paid. Several forms of the 

Hairstreak in Tasmania were stated by Couchman and Couchman 

(1977) 'to have suffered more than any local species of butterfly', 

and much suitable habitat has been destroyed. The Couchmans 

expressed grave concern for the butterfly's future in Tasmania, 

and a more recent survey (Prince 1988) also stressed the need 

for conservation measures. Many colonies have become extinct, 

and most of the remainder face threats to their continued well 

being. The species as a whole in Tasmania is considered 

vulnerable. 

Taxonomy and Description: P. chlorinda is the sole species of 

the endemic Australian thecline genus Pseudalmenus Druce, 

and is known from southeastern mainland Australia and 

Tasmania. It shows substantial geographical variation (Figure 

1) and seven named subspecies are widely recognised (Common 

and Waterhouse 1981), together with at least one other 

Tasmanian form. Couchman and Couchman (1977) noted 

another two Tasmanian forms which were by then extinct. 

Collectively, these forms constitute one of the most diverse 

polytypic Lycaenidae in Australia, probably representing 

incipient speciation, and of considerable evolutionary interest. 

Distribution: All eight subspecies have highly circumscribed 

distributions (see Figure 1) and only one, P. c. zephyrus, can be 

considered to be reasonably widespread. P. c. chloris is regarded 

as rare and local in New South Wales and the other mainland 

subspecies, P. c. fisheri and P. c. barringtonensis , are highly 

localised in the Grampians Mountains, Victoria and Barrington 

Tops, New South Wales, respectively. Of the Tasmanian taxa, 

P. c. chlorinda is found from Hobart to north of Swansea and 

171 

westward to the South Esk and Upper Tamar Valleys, P. c. 

conara is known from the midlands and P. c. myrsilus from the 

Tasman and Forestier Peninsulas and a small part of the east 

coast. The un-namedP. c. zephyrus-like form is known from the 

northeast of Tasmania. The recent discovery of the species in 

western Tasmania suggests the likelihood that other colonies 

may exist in that remote area: that race has been referred 

tentatively to P. c. chlorinda, but specimens are not available. 

Population Size: Most subspecies are clearly very localised 

but little information is available on population size, especially 

for the mainland taxa. Many of the Tasmanian sites surveyed by 

Prince (1988) had very low 'occupation rates' of suitable 

eucalypt trees, with few hairstreak pupae recorded from them. 

Many populations may, indeed, be confined to single eucalypt 

trees in very small isolated patches of habitat, from which 

recolonisation is probably unlikely. Detailed information on 

Pseudalmenus dispersal is not available, but it is believed to be 

largely sedentary in habit. 

Habitat and Ecology: Eggs are laid singly or in small groups 

on young twigs of a larval foodplant, Acacia. In Tasmania, the 

usual foodplant is the bipinnate A. dealbata (80% of Prince's 

records), with the closely related A. mearnsii (9%) also utilised. 

Other acacias, particularly the phyllodinous A. melanoxylon, 

have also been recorded, more especially for the mainland 

subspecies. Larvae feed on the Acacia foliage and are attended 

by small, black ants (Iridomyrmex foetans). The role of the ants 

is not clear, but they appear to be essential, and probably protect 

the caterpillars from parasitoids. Caterpillars pupate under the 

loose bark of nearby eucalypts, predominantly Eucalyptus 

viminalis (92% of pupae) in Tasmania, growing within a few 

metres of the acacias. Most (77%) grew within 2m, and few 

pupae were found on eucalypts only 10m from larval foodplants. 

The most frequently selected eucalypts were 15–18m tall, with 

diameters of 75–150cm. Pupae occurred up to around 2m from 

the ground, and most were on the northern half of the tree. 

Adults fly in spring and early summer, with slight differences 

in flight period between subspecies, and most forms are 

univoltine. A partial second generation has been suggested for 

P. c. fisheri (Common and Waterhouse 1981) but otherwise the

Figure 1. Distribution of subspecies of Pseudalmenus chlorinda in 

southeastern Australia, with details of Tasmanian subspecies. Mainland 

data after Common and Waterhouse (1981). 

period from December to at least August or September is passed 

as pupae. This stage therefore constitutes a suitable one for 

monitoring population sizes of Pseudalmenus. Pupae are often 

on trees occupied by I. foetans, and the scent of the ants might 

influence selection of host trees (Prince 1988). The ant can also 

be abundant on trees lacking Pseudalmenus. 

Threats: At many sites in Tasmania where P. chlorinda has 

become extinct, the habitat has been destroyed or severely 

disturbed. Pastoral clearing (52%) and fire (direct implication 

in loss of 24% of sites, contributory to the loss of a further 20%) 

were the predominant factors involved, and grazing, forestry 

(clearing for timber and woodchips) and housing subdivisions 

172 

accounted for the remainder. For extant colonies, Prince (1988) 

assessed fire as the most common threat (31 %) though clearing 

and subdivision are also significant. 

Pupae were recorded in the mid-1980s at only 15 of the 43 

sites noted by earlier workers, but some new sites were also 

found. Of 66 locations recorded for P. chlorinda in Tasmania, 

only 12 of the 40 which still support the hairstreak were 

considered to be reasonably secure (Prince 1988). P. c. myrsilus 

is especially restricted, and P. c. near zephyrus was found in 

only four areas. Most known sites are small and threatened. At 

many, pupae were found only on a single tree. Couchman and 

Couchman (in Prince 1988) observed instances where a single 

tree had supported a population of P. chlorinda for many years, 

and that population had disappeared once the tree had been cut 

down. 

Conservation: Couchman and Couchman (1977) emphasised 

the threefold needs of P. chlorinda: the close associations of a 

suitable Eucalyptus; a suitable Acacia; and the specific ant. 

They emphasised that the destruction of any of these might lead 

to the butterfly's local extermination. Protection of these 

resources in suitable habitats is clearly the major conservation 

need for P. chlorinda, both in terms of reservation of habitat and 

management within reserves and on private land. Many of the 

populations are in reserved sites, but some of these have been 

subject to clearing and fire. Protection of habitats on private 

land may be feasible in some instances, but sites close to urban 

centres are under considerable pressure from urban expansion. 

Increased community awareness of the butterfly is also needed. 

Active management, for example guarding against clearing 

of acacias near eucalypts, merits promotion, and a trial reported 

by Prince (1988) suggests that translocation of the butterfly to 

new suitable habitats might be feasible as a management tool. 

Studies of the status of mainland subspecies are also a priority 

for this species, together with clarification of the status of 

populations in western Tasmania. 

References 


(2nd edn.) Angus & Robertson, Sydney. 

COUCHMAN, L.E. and COUCHMAN, R. 1977. The butterflies of Tasmania. 

Tasmanian Year Book (1977): 66–96. 

PRINCE, G.B. 1988. The conservation status of the Hairstreak Butterfly 

Pseudalmenus chlorinda Blanchard in Tasmania (report to Department of 

Lands, Parks and Wildlife, Tasmania).

APPENDIX 1 

IUCN Red Data Book categories 

Many of the species discussed in this book have been allocated, 

sometimes tentatively, to the traditional IUCN Red Data Book 

categories. They are subject to revision of status as new 

information is incorporated. The categories are defined as 

follows: 

Extinct (Ex) 

Species not definitely located in the wild during the past 50 

years (criterion as used in the Convention on International 

Trade in Endangered Species of Wild Fauna and Flora). (For 

some butterflies, the status is applied to taxa which are known 

to have become extinct by, for example, destruction of the last 

or only known colony, without regard to the 50 year period: 

TRN.) 

Endangered (E) 

Taxa in danger of extinction and whose survival is unlikely if 

the causal factors continue to operate. These include species 

whose habitats or numbers have been reduced drastically and 

may be in danger of imminent extinction. The category also 

includes taxa which are probably already extinct, but which 

have been seen during the last 50 years (see comment in 

parentheses, above: 'Endangered' implies some uncertainty 

over whether or not the species is extinct: TRN). 

Vulnerable (V) 

Taxa believed likely to become Endangered in the near future 

if causal factors continue to operate. A wide range of threats is 

173 

included, from habitat destruction to overexploitation, and 

other environmental disturbances. The taxa need not, necessarily, 

be rare if threats operate over a large range. 

Rare (R) 

Taxa with small world populations which are not at present 

Endangered or Vulnerable, but are at risk. They are usually in 

limited geographical areas or habitats, or in low numbers over 

a more extensive range. 

Indeterminate (I) 

Taxa known to be Endangered, Vulnerable, or Rare, but for 

which there is insufficient information to determine which of 

these is the appropriate category. 

Out of Danger (O) 

Taxa formerly in one of the above categories but which are now 

considered relatively secure because of effective conservation 

measures and/or removal of previous threats to their survival. 

'Threatened' is a general term to denote species which are 

Endangered, Vulnerable, Rare, or Indeterminate. 

A 'Threatened Community' is a group of ecologically linked 

taxa occurring within a defined area, which are all under the 

same threat and require similar conservation measures.

Other Occasional Papers of the IUCN Species Survival Commission 

1. Species Conservation Priorities in the Tropical Forests of Southeast Asia. Edited by R.A. Mittermeier and W.R. 

Constant, 1985, 58pp, £7.50, US$15.00. (Out of print) 

2. Priorités en matière de conservation des espèces à Madagascar. Edited by R.A. Mittermeier, L.H. Rakotovao, V. 

Randrianasolo, E.J. Sterling and D. Devitre, 1987, 167pp, £7.50, US$15.00. 

3. Biology and Conservation of River Dolphins. Edited by W.F. Perrin, R.K. Brownell, Zhou Kaiya and Liu Jiankang, 

1989, 173pp, £10.00, US$20.00. 

4. Rodents. A World Survey of Species of Conservation Concern. Edited by W.Z. Lidicker, Jr., 1989, 60pp, £7.50, 

US$15.00. 

5. The Conservation Biology of Tortoises. Edited by I.R. Swingland and M.W. Klemens, 1989, 202pp, £12.50, 

US$25,00. 

6. Biodiversity in Sub-Saharan Africa and its Islands, 1991, £12.50, US$25.00. 

7. Polar Bears: Proceedings of the Tenth Working Meeting of the IUCN/SSC Polar Bear Specialist Group, 1991, 

£10.00, US$20.00. 

Where to order: 

IUCN Publications Services Unit, 219 Huntingdon Road, Cambridge, CB3 0DL, U.K. Please pay by cheque/ 

international money order to IUCN. Add 15% for packing and surface mail costs. A catalogue of IUCN publications can 

be obtained from the above address.

IUCN Species Survival Commission 

The Species Survival Commission (SSC) is one of six volunteer commissions of IUCN – The World Conservation Union, 

a union of sovereign states, government agencies and non-governmental organizations. IUCN has three basic 

conservation objectives: to secure the conservation of nature, and especially of biological diversity, as an essential 

foundation for the future; to ensure that where the earth's natural resources are used this is done in a wise, equitable and 

sustainable way; and to guide the development of human communities towards ways of life that are both of good quality 

and in enduring harmony with other components of the biosphere. 

The SSC's mission is to conserve biological diversity by developing and executing programs to save, restore and wisely 

manage species and their habitats. A volunteer network comprised of 4,800 scientists, field researchers, government 

officials and conservation leaders from 169 countries, the SSC membership is an unmatched source of information about 

biological diversity and its conservation. As such, SSC members provide technical and scientific counsel for conservation 

projects throughout the world and serve as resources to governments, international conventions and conservation 

organizations. 

The IUCN/SSC Occasional Paper series focuses on a variety of conservation topics including conservation overviews on 

a regional or taxonomic basis and proceedings of important meetings. 

IUCN/SSC also publishes an Action Plan series that assesses the conservation status of species and their habitats, and 

specifies conservation priorities. The series is one of the world's most authoritative sources of species conservation 

information available to nature resource managers, conservationists and government officials around the world. 

Published by IUCN 

This book is part of The IUCN Conservation Library 

For a free copy of the complete catalogue please write to 

IUCN Publications Services Unit, 

219 Huntingdon Road, Cambridge CB3 0DL, U.K.

Conservation Biology of Lycaenidae (Butterflies) - IUCN

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