Rafael Gioia MARTINS-NETO, Oscar Florencio GALLEGO

Acta zoologica cracoviensia, 46(suppl.– Fossil Insects): 229-256, Kraków, 15 Oct., 2003 

The Triassic insect fauna from South America (Argentina, Brazil 

and Chile): a checklist (except Blattoptera and Coleoptera) and 

descriptions of new taxa 

Rafael Gioia MARTINS-NETO, Oscar Florencio GALLEGO 

and Ricardo Nestor MELCHOR 

Received: 25 Feb., 2002 

Accepted for publication: 30 Sep., 2002 

MARTINS-NETO R. G., GALLEGO O. F., MELCHOR R. N. 2003. The Triassic insect fauna 

from South America (Argentina, Brazil and Chile): a checklist (except Blattoptera and 

Coleoptera) and descriptions of new taxa. Acta zoologica cracoviensia, 46(suppl.– Fossil 

Insects): 229-256. 

Abstract. The present paper describes new fossil insect taxa from the Los Rastros and Ischichuca 

Formations, La Rioja Province, Argentina; and from the Santa Juana Formation, 

Chile. These formations are dated as late Middle Triassic to early Late Triassic. Eight new 

genera and nine new species are described: Frenguelliphlebia labandeirai n. gen. et n. sp. 

(Odonatoptera); Argentinoperlidium rogersi n. gen. et n. sp. (Plecoptera: Gripopterigidae); 

Dysmorphoptiloides pulcherrima n. sp.; Bandelnielsenia chilena n. gen. et n. sp., 

Gallegomorphoptila breviptera n. gen. et n. sp. (all Auchenorhyncha: Dysmorphoptilidae), 

Gualoscytina mayae n. gen. et n. sp. (Auchenorrhyncha: Scytinopteridae), Argentinopheloscyta 

forsterae n. gen. et n. sp. (Auchenorrhyncha: Stenoviciidae); Yurigomicex 

popovi n. gen. et n. sp., and Popovigocimex yurii n. gen. et n. sp. (both Coleorrhyncha: 

Progonomicidae). Additionally a checklist of all known described species of South America 

Triassic is provided (except Blattoptera and Coleoptera, paper in preparation), as well 

as a discussion of dispersal paths of Triassic conchostracan and insects faunas of Gondwana. 

Key words: Triassic insects, new genus, new species, Argentina, Brazil, Chile, Gondwana. 

MARTINS-NETO R. G., Universidade Guarulhos / PPGEO-UNISINOS / Sociedade Brasileira 

de Paleoartropodologia - SBPr: Rua Arnaldo Vitaliano, 150, apto 81; 14091-220 – 

Ribeirão Preto – SP, Brasil. 

E-mail: martinsneto@terra.com.br 

GALLEGO O. F., Catedra de Paleontología (Facultad de Ciencias Exactas y Naturales y 

Agrimensura – Universidad Nacional del Nordeste) y Área Paleontología (Centro de 

Ecología Aplicada del Litoral – CONICET), Casilla de Correo 128, 3400 Corrientes – Argentina. 

E-mail: ofgallego@hotmail.com 

MELCHOR R. N., CONICET and Universidad Nacional de la Pampa, Av. Uruguay 151, 

6300 Santa Rosa, Argentina. 

E-mail: melchor_r@cpsarg.com

230 

R. G. MARTINS-NETO et al. 

I. INTRODUCTION 

This paper presents new data about the fossil record of Triassic insects from Argentina, Brazil 

and Chile, and a compendium of reported occurrences. The Triassic insect faunas from South 

America are sparse and poorly known 

from this time. Only six Triassic insects 

localities are presently known: Mendoza (Cacheuta hill), La Rioja (Gualo River and “Quebrada de 

Ischichuca Chica”) Provinces (western Argentina), São Paulo (Timbó river), Rio Grande do Sul 

(Passo das Tropas) states (Brazil), and Concepción (southern Chile). 

Previous records of these faunas were summarized by GALLEGO (1997), GALLEGO &MAR- 

TINS-NETO (1999) and MARTINS-NETO &GALLEGO (1999). The presence of fossil insects in Triassic 

sediments of South America was first reported by WIELAND (1925); this record has been significantly 

expanded by subsequent authors. 

Argentinian faunas are the richest in South America, with the most described species in an extensive 

literature. Only three species have been described from Brazil; Chilean occurrences have 

been mentioned only as undetermined insects (FUENZALIDA 1937; CECIONI &WESTERMANN 1968) 

from localities Punta Puquén, Los Lobos and Los Molles in Central Chile. 

In the present paper we describe new taxa from Argentina, the described species from Argentina 

and Brazil are listed, and new specimens from Chile are reported for the first time (Concepción locality, 

southern Chile). 

This paper contains the first overview of the Triassic insect fauna of Argentina (excluding Coleoptera 

and Blattoptera), including many recent findings. Coleoptera and Blattoptera (which include 

a total of about 1355 specimens) will be treated by us elsewhere. The material described herein includes 

23 genera and 28 species (33 specimens, Table I) from Cuyo and Ischigualasto – Villa Unión 

Basins of Argentina (Fig. 1) (most specimens) and from the Paraná basin of Brazil (two specimens). 

Terminology and classification adopted here conforms mainly to that of KUKALOVÁ-PECK 

(1991) and VILESOV &NOVOKSHONOV (1994). 

Abbreviations: 

CORD-PZ – Palaeozoological Collection, Universidad Nacional de Cordoba, Argentina 

MACN – Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina 

MLP – Museo de La Plata (Invertebrate Paleontology), Argentina 

MMHN – Museo Municipal de Historia Natural de San Rafael, Mendoza, Argentina 

PULR-I – Invertebrate Paleontological Collection, Universidad Nacional de La Rioja, Argentina 

PZ-CTES – Paleozoological Collection of the Facultad de Ciencias Exactas, Naturales 

y Agrimensura de la Universidad Nacional del Nordeste, Mendoza, Argentina. 

SGO-PI – Paleontology Section of the Museo Nacional de Historia Natural, Santiago, Chile. 

USP – Universidade de São Paulo, campus de Ribeirão Preto, SP, Brazil. 

Acknowledgements.Wethank Dr. Ana Maria ZAVATTIERI (Ianigla-Cricyt- 

CONICET, Mendoza, Argentina), Dr. Augusto MENÉNDEZ, Lic. Jorge CRUZ and Sr. Alberto 

ACOSTA for field assistance and collaboration in the Mendoza Province Triassic outcrops. We also 

acknowledge Lic. Andrea ARCUCCI (Universidad Nacional de San Luís, Argentina), Drs. Catherine 

FORSTER, Cathleen MAY and Raymond ROGERS from the project “El contexto del origen de los Dinosaurios: 

Paleontología y Paleoecología de la Formación Los Rastros, Triásico Medio de La 

Rioja”, National Geographic Society (Grant 5317-94). Special thanks to Dr. Klaus BANDEL and 

Geol. Sven NIELSEN (Universität Hamburg, Geologisch-Paläontologisches Institut and Museum) 

for providing us with material collected in the Chilean Triassic. Thanks are also due to our special 

friend, Dr. Yuri POPOV (Russian Academy of Sciences), for his help in the description of the Heteroptera 

specimen, and to our “computer magician”, Vinicius Moreno DE GODOI (FFCLRP/SBPr), 

for help in all of the informatics problems and solutions.

Triassic insect fauna from South America 

Table I 

231 

Summary of the insect taxa described in this paper, grouped by unit and locality 

Order Family Species 

Potrerillos Formation (Cerro Cacheuta, Mendoza, Cuyo basin) 

Odonatoptera Incertae sedis Triassothemis mendozensis CARPENTER 1960 

Odonatoptera Incertae sedis Frenguelliphlebia labandeirai MARTINS-NETO &GALLEGO n. sp. 

Plecoptera Gripopterygidae: Euxenoperlinae Gondwanoperlidium argentinarum PINTO &PURPER 1978 

Gondwanoperlidium mendozensis PINTO &PURPER 1978 

Miomoptera Incertae sedis Miomina mendozina MARTINS-NETO &GALLEGO 1999 

Grylloblattida Atactophlebiidae Triasseuryptilon acostai (MARQUAT) STOROZHENKO 1997 

Ensifera Haglidae Notopamphagopsis bolivari CABRERA 1928 

Auchenorrhyncha Dysmorphoptilidae Dysmorphoptiloides acostai MARTINS-NETO &GALLEGO 1999 (*) 

Auchenorrhyncha Chiliocyclidae Tipuloidea rhaetica WIELAND 1925 

Argentinocicada magna MARTINS-NETO &GALLEGO 1999 

Argentinocicada minima MARTINS-NETO &GALLEGO 1999 

Auchenorrhyncha Scytinopteridae Potrerillia nervosa MARTINS-NETO &GALLEGO 1999 

Trichoptera? / Tipulidites affinis WIELAND 1926 

Diptera? Necrotaulius affinis WIELAND 1926 

Los Rastros Formation (Río Gualo, La Rioja, Ischigualasto – Villa Unión basin) 

Glosselytrodea Polycytellidae Argentinoglosselytrina pulchella MARTINS-NETO &GALLEGO 2000 (*) 

Plecoptera Gripopterygidae: Euxenoperlinae Argentinoperlidium rogersi MARTINS-NETO &GALLEGO n. sp 

Ensifera Haglidae Notopamphagopsis? sp. 1 

Notopamphagopsis? sp. 2 


Auchenorrhyncha Dysmorphoptilidae Dysmorphoptiloides acostai MARTINS-NETO &GALLEGO 1999 (*) 

Dys morphoptiloides gigantea MARTINS-NETO &GALLEGO 2000 

Dysmorphoptiloides pulcherrima MARTINS-NETO &GALLEGO n. sp. 

Gallegomorphoptila breviptera MARTINS-NETO n. sp. 

Auchenorrhyncha Scytinopteridae Australocicada arcucciae MARTINS-NETO &GALLEGO 2000 (*) 

Gualoscytina mayae MARTINS-NETO &GALLEGO n. sp 

Auchenorrhyncha Stenoviciidae 

Argentinopheloscyta forsterae MARTINS-NETO &GALLEGO n. sp 

Coleorrhyncha Progonomicidae Yurigocimex popovi MARTINS-NETO &GALLEGO n. sp. 

Popovigocimex yurii MARTINS-NETO &GALLEGO n. sp. 

Ischichuca Formation (Quebrada Ischichuca, La Rioja, Ischigualasto – Villa Unión basin) 

Auchenorrhyncha Prosboloidea: Prosbolidae (2) Lariojaprosbole melchori MARTINS-NETO &GALLEGO 2000 (*) 

Rio Do Rastro Formation (near Poço Preto, Santa Catarina, Paraná basin) 

Auchenorrhyncha Prosboloidea: Prosbolidae Prosbolidinella riorastensis MARTINS-NETO &ROHN 1996 

Santa María Formation (Passo das Tropas, Rio Grande do Sul, Paraná basin) 

Trichoptera? 

Sanctipaulus mendesi PINTO1956 

(*) Indicates two specimens. The remaining occurrences are known from a single specimen.

232 


Fig. 1. Map showing the Cuyo and Ischigualasto – Villa Unión basins and the locations of the three insect-bearing localities 

mentioned in the text. 

II. GEOLOGICAL ASPECTS 

Most of the known Triassic insect fauna from Argentina come from the Potrerillos and Los 

Rastros Formations, which belong to Cuyo and Ischigualasto Villa-Unión rift basins respectively 

(Fig. 1). Each of these units have yielded a single insect-bearing locality or area: Cerro Cacheuta, of 

the Potrerillos Formation (northern Mendoza Province) and the Río Gualo area of the Los Rastros 

Formation (Talampaya Park, southern La Rioja Province). The Potrerillos Formation is included in 

the Uspallata Group and represents the depositional record of the second sinrift phase of KOKOGIÁN 

et al. (1993). Occurrences of insect remains at Cerro Cacheuta were concentrated in a 3 km-long 

tract along the strike of the upper part of the Potrerillos Formation in the southern flank of the hill 

(approximately 33º 05’ S, 69º 07’ W; see MOREL 1994: Fig. 1). The stratigraphic interval where 

most insect remains were collected has been interpreted as floodplain, crevasse, and levee facies 

(MOREL 1994; KOKOGIÁN et al. 1999) in an intermediate to high sinuosity fluvial system (KOKO- 

GIÁN et al. 1999; BELLOSI et al., in press). At Cerro Cacheuta, these facies grade to distributary sys-


233 

tems (mouth bar, channel, and interdistributary bay) of lacustrine deltas (KOKOGIÁN et al. 1999). 

Associated plant remains have been attributed to the BNP biozone of SPALLETTI et al. (1999). 

Both the Los Rastros and Ischichuca Formations belong to the Agua de la Peña Group of the Ischigualasto 

– Villa Unión Basin. The insect fauna from the Los Rastros Formation was recovered 

from Río Gualo (about 29° 53’ S; 67° 46’ W), southern La Rioja Province, Talampaya Park (Fig. 1). 

In this area the exposures of the Los Rastros Formation are mostly composed of deep-lacustrine 

mudstones, storm and wave-influenced delta-front siltstones, and channel sandstones. Insect remains 

are preferentially preserved in olive-green, thinly laminated mudstones deposited in deep, 

well-oxygenated lakes. 

Ischichuca Formation is dominantly composed of shallow and deep lacustrine facies with progadational 

deltaic successions (MELCHOR 1998). In particular, insect remains come from dark 

brown to olive-green claystones with abundant plant remains from the lower part of the unit. This 

interval is interpreted as a shallow, partially saline lake. The Ischichuca Formation continental palynologic 

assemblage is closely comparable to those of the Ipswich Microflora of Australia (ZAVAT- 

TIERI &MELCHOR 1999). The Macroflora of this unit was included by SPALLETTI et al. (1999) in 

the CSD biozone. 

There is general agreement that the age of the Potrerillos Formation is late Middle Triassic to 

early Late Triassic (e.g., STIPANICIC 1983; MOREL 1994; ZAVATTIERI &BATTEN 1996; SPAL- 

LETTI et al. 1999; KOKOGIÁN et al. 1999). In contrast, disagreement exists on the age of the Ischichuca 

and Los Rastros Formations (mainly concerning the former). Most authors consider these 

units to be of approximately the same age as the Potrerillos Formation (STIPANICIC &BONAPARTE 

1979; STIPANICIC 1983; KOKOGIÁN et al., 1999). However, in the recent chronostratigraphic 

scheme of SPALLETTI et al. (1999) both the Ischichuca and Los Rastros Formations are restricted to 

the lower part of the Middle Triassic. Conversely, ZAVATTIERI &MELCHOR (1999) suggest that the 

age of the Ischichuca Formation, on the basis of its palynological content, is late Ladinian-nearly 

Carnian. This is in agreement with a previous palynological assessment of Carnian age for the overlying 

Los Rastros Formation (ZAVATTIERI &BATTEN 1996). 

The remaining occurrences of Triassic insects discussed in this paper (Table I) come from the 

Paraná basin in southeast Brazil, from Rio do Rasto, and from the Santa Maria Formation. The Rio 

do Rasto Formation (uppermost unit of the Passa Dois Group) have been considered of Late Permian 

age until recent findings of MARTINS-NETO &ROHN (1996), who suggest a maximum Early 

Triassic age for this unit based on its flora, Conchostracan fauna, and insect remains (Prosbolidinella 

riorastensis MARTINS-NETO &ROHN 1996). This formation is considered essentially of fluvial 

origin. The overlying Santa Maria Formation (Rosario do Sul Group) is a continental red-bed 

sequence from the central region of Rio Grande do Sul. Typical facies include mudstones intercalated 

with fine-grained sandstones, deposited in the floodplains of a braided fluvial system (HOLZ & 

BARBERENA 1994). The Santa Maria Formation is well known for its high diversity of fossil tetrapods 

(e.g., BONAPARTE 1982), which allowed its assignment to the Late Triassic (e.g., ANDERSON 

&ANDERSON 1993; BENTON 1983), specifically the Carnian (SHUBIN &SUES 1991). Its fauna is 

similar to that of the Ischigualasto Formation of the Ischigualasto – Villa Unión basin of Argentina, 

which may imply a slightly younger age than the Ischichuca and Los Rastros Formations. 

III. SYSTEMATIC PALEONTOLOGY 

ODONATOPTERA 

Family incertae sedis 

Triassothemis mendozensis CARPENTER, 1960 

Fig. 2G-H 

*1960 Triassothemis mendozensis CARPENTER: 71-73, Fig. 1

234 


1992 Triassothemis mendozensis CARPENTER: 78; 1997 Triassothemis mendozensis CARPENTER in GAL- 

LEGO: 512-513; 1999 Triassothemis mendozensis CARPENTER in GALLEGO &MARTINS-NETO: 87-88, Fig. 16. 

Holotype.MACN no. 18040, housed in MACN, Argentina. Type locality: Aguas de 

las Avispas, Cerro Cacheuta, Argentina. Type stratum : Potrerillos Formation. Age: late Middle Triassic 

to early Late Triassic. 

Frenguelliphlebia MARTINS-NETO &GALLEGO, n. gen. 

Diagnosis.Wing similar to Petrotypus ZALESSKY, 1950 in the general aspect of the 

venation, but with M and Cu moderately wavy. 

Type species. Frenguelliphlebia labandeirai MARTINS-NETO &GALLEGO n. sp., designated 

here. 

Etymology.Inhonor of Dr. Joaquin FRENGUELLI, the paleobotanist who first mentioned 

the insect fossils of this region, and phlebia, a common odonate suffix. Gender: feminine. 

Discussion. Frenguelliphlebia n. gen. is similar to Petrotypus ZALESSKY, 1950 

(Meganeuridae), from the Upper Permian of the Urals by having CuA and CuP multi-branched, differing 

however, by having M and Cu moderately wavy (notably wavy in Petrotypus). In this aspect 

Frenguelliphlebia n. gen. is similar to Triadotypus GRAUVOGEL &LAURENTIAUX, 1952 (Triadotypidae) 

from the Lower Triassic of France, differing, however, by having a wider cubital area and 

long and curved branches of Cu. The Meganeuridae as well as Triadotypidae are mostly consisting 

of giant species ( 90 mm in length), unlike this specimen, which is not more than 25 mm long. Additionally, 

Frenguelliphlebia n. gen. exhibits a combination of characters present separately in Meganeuridae 

and Triadotypidae. Because of this, correct family placement is uncertain, and 

Frenguelliphlebia n. gen. may belong to a new family. More complete material is necessary to confidently 

determine the family placement of this genus. 

Frenguelliphlebia labandeirai MARTINS-NETO &GALLEGO, n. sp. 

Fig. 2 I 

Diagnosis.Asforthegenus. 

Etymology.Inhonor of Dr. Conrad Christopher LABANDEIRA (Smithsonian Institution), 

in recognition of his great contribution to the paleoentomology. 

Holotype. PULR-I nº 269, housed in PULR-I, Argentina. Type locality: Gualo, La 

Rioja Province, Argentina. Type stratum: Los Rastros Formation. Age: late Middle Triassic to early 

Late Triassic. 

Description(Fig. 2I). Basal fragment of incomplete fore wing, 12 mm in length 

preserved. R+MA and Cu moderately wavy. Cu origin close to the wing base. CuA with at least five 

secondary branches with intercalary veins; CuP single, distally branched. Anal branches long and 

curved. 

GLOSSELYTRODEA 

Family: Polycytellidae MARTYNOVA, 1952 

Argentinoglosselytrina pulchella MARTINS-NETO &GALLEGO 2001 

Fig. 2A, B 

*2001 Argentinoglosselytrina pulchella MARTINS-NETO &GALLEGO. 



Late Triassic.


235 

Fig. 2.A–B:Argentinoglosselytrina pulchella MARTINS-NETO &GALLEGO 2001 (Odonatoptera: Policytellidae), holotype 

PULR-I n° 253 (A); Argentinoglosselytrina pulchella MARTINS-NETO &GALLEGO, 2001, specimen PULR-I n° 263 (B); 

C. Tryasseuryptilon acostai (MARQUAT) STOROZHENKO 1997 (Grylloblattida: Atactophlebiidae), holotype MMNH n° 

1518, reproduced from STOROZHENKO (1997: Fig. 1); D. Necrotaulius affinis WIELAND 1925 (Mecoptera?), holotype, 

redrawn from WIELAND (1925: Fig. 1); E. Argentinoperlidium rogersi n. sp. (Plecoptera: Gripopterigidae), holotype 

PULR-I, n° 258; F. Miomima mendozina MARTINS-NETO &GALLEGO 1999 (Miomoptera, fam. inc. sed.), holotype 

PZ-CTES n° 5731; G-H: Triassothemis mendozensis CARPENTER 1960 (Odonatoptera, fam. inc. sed.), holotype MACN 

n°18040; I. Frenguelliphlebia labanderai MARTINS-NETO &GALLEGO n. sp. (Odonatoptera, fam. inc. sed.), holotype 

PULR-I, n° 269; J. Gondwanoperlidium argentinarum PINTO &PURPER 1978 (Plecoptera: Gripopterigidae), holotype 

MLP n° 8908; K. Gondwanoperlidium mendozensis PINTO &PURPER 1978, holotype CORD-PZ n° 336. Scale bar 

1 mm. All drawings are original, if not otherwise stated.

236 


Supplementary material. (Fig. 2B); PULR-I nº 263. Same age, locality, stratum and destination 

as in Holotype. 

PLECOPTERA 

Family: Gripopterigidae sensu RIEK, 1976 

Subfamily: Euxenoperlinae RIEK, 1976 

Gondwanoperlidium argentinarum PINTO &PURPER, 1978 

Fig. 2J 

1921 Baiera argentinii KURTZ: Pl. 21, Fig. 336 

1926 Elcana? argentina COCKERELL in: WIELAND: 131. 

1927 ?Elcana sp. COCKERELL 

1939 Elcana? argentina HANDLIRSCH 

1941 Locustopsis? argentina ZEUNER 

*1978 Gondwanoperlidium argentinarum, PINTO &PURPER: 79-80, Pl. 1A, Fig. 1C. 

1992 Gondwanoperlidium argentinarum PINTO &PURPER in CARPENTER: 97; 1997 Gondwanoperlidium 

argentinarum PINTO &PURPER, GALLEGO: 512-513; 1999 Gondwanoperlidium argentinarum PINTO & 

PURPER in GALLEGO &MARTINS-NETO: 87-88 

Holotype. MLPnº8908, housed in MLP, Argentina. Type locality: “Minas de Petróleo”, 

at Cacheuta, 14 km W of Mendoza city, Mendoza Province, Argentina. Type stratum: 

Potrerillos Formation. Age: late Middle Triassic to early Late Triassic. 

Gondwanoperlidium mendozensis PINTO &PURPER 1978 

Fig. 2K 

*1978. Gondwanoperlidium mendozensis PINTO &PURPER: 81, Pl. 1B, Fig. 1D. 

1995 Gondwanoperlidium mendozensis PINTO &PURPER in MAZZONI: 229, 230, Lám XXII, Fig. 22; 

Fig-text. 45; 1997 Gondwanoperlidium mendozensis PINTO &PURPER in GALLEGO: 512-513; 1999 Gondwanoperlidium 

mendozensis PINTO &PURPER in GALLEGO &MARTINS-NETO: 87-88, Fig. 17. 

Holotype. CORD-PZ nº 336, housed in CORD-PZ, Argentina. Type locality: “Minas 

de Petróleo”, at Cacheuta, 14 km W of Mendoza city, Mendoza Province, Argentina. Type stratum: 

Potrerillos Formation (Cacheuta Fm., sensu MAZZONI, 1995). Age: late Middle Triassic to early 


Argentinoperlidium MARTINS-NETO &GALLEGO, n. gen. 

Diagnosis. Wing similar to Euxenoperla RIEK, 1976 in general aspect of venation, 

but ScP fused distally with RA, and M strongly curved. 

Type species. Argentinoperlidium rogersi MARTINS-NETO &GALLEGO n. sp., designated here. 

Etymology.Argentina, from which the material originates, and perlidium, from Perlida. 

Gender: neutrum. 

Discussion.Argentinoperlidium n. gen. differs of all known Euxenoperlinae by having 

ScP distally fused with RA (connected by a cross-vein in Euxenoperla RIEK, 1976 as well as in 

Gondwanoperlidium PINTO &PURPER, 1978), and M strongly curved. Additionally, Argentinoperlidium 

n. gen. has a more complex cross-veins pattern.


237 

Argentinoperlidium rogersi MARTINS-NETO &GALLEGO, n. sp. 

Fig. 2E 


Etymology.Inhonor of Dr. Raymond ROGERS (Department of Geology, McAlester 

College, St. Paul, Minnesota), who collaborated in fieldwork, working in the collections, and did 

sedimentological analysis at the Gualo river site (La Rioja) for the NGS project (grant 5317-94). 

Holotype. nº258, housed in PULR-I. Type locality: Picos Gemelos, Gualo, La Rioja, 

Argentina. Type stratum: Los Rastros Formation. Age: late Middle Triassic to early Late Triassic. 

Description.Fore wing fragment 5 mm in length. Costal area very narrow where 

preserved, filled with pectinate cross-veins. ScP relatively long, distally fused with RA. RA sigmoid. 

M very curved towards apex. Complex pattern of cross-veins forming a heterogeneous mosaic 

of cells. 

MIOMOPTERA 

Family incertae sedis 

Miomina mendozina MARTINS-NETO &GALLEGO 1999 

Figs 2F 

*1999 Miomina mendozina MARTINS-NETO &GALLEGO: 198, Figs 1H, 5B. 

Holotype:nº5731, housed in PZ-CTES, Argentina. Type locality: 300 mWofQuebrada 

del Durazno. South of the Cerro Cacheuta. Mendoza, Argentina, levels EP I and EP II of 

MOREL (1994). Type stratum: Upper Section of the Potrerillos Formation. Age: late Middle Triassic 


GRYLLOBLATTIDA 

Family: Atactophlebiidae sensu STOROZHENKO, 1997 

Triasseuryptilon acostai (MARQUAT) STOROZHENKO, 1997 

Fig. 2C 

1991 Delopterum acostai MARQUAT: 6, Fig. 2 

1997 Paleomantis acostai (MARQUAT) GALLEGO: 513-514. 

*1997 Triasseuryptilon acostai (MARQUAT) STOROZHENKO: 8, Fig. 1. 

1999 Triasseuryptilon acostai (MARQUAT) STOROZHENKO in GALLEGO &MARTINS-NETO: 87; Fig. 15 

Holotype. MMHN nº 1518, housed in MMHN, Argentina. Type locality: Cerro 

Cacheuta, Mendoza Province, Argentina. Type stratum : Potrerillos Formation. Age: late Middle 

Triassic to early Late Triassic. 

ENSIFERA 

Family: Haglidae HANDLIRSCH, 1906 

Notopamphagopsis bolivari CABRERA 1928 

Fig. 3A 

*1928 Notopamphagopsis bolivari CABRERA: 372, Figs 1 and 2 

1939 Notopamphagopsis bolivari CABRERA in ZEUNER: 18; 1962 Notopamphagopsis bolivari CABRERA 

in ROHDENDORF: 201; 1968 Notopamphagopsis bolivari CABRERA in SHAROV: 51; 1991 Notopamphagopsis

238 


bolivari CABRERA in MARTINS-NETO: 26; 1992 Notopamphagopsis bolivari CABRERA in CARPENTER: 168; 

1995 Notopamphagopsis bolivari CABRERA in GOROCHOV: 137; 1997 Notopamphagopsis bolivari CABRERA 

in GALLEGO: 511; 1999 Notopamphagopsis bolivari CABRERA in GALLEGO &MARTINS-NETO: 87, Fig. 11; 

1999 Notopamphagopsis bolivari CABRERA in MARTINS-NETO &GALLEGO: 198-200, Fig. 3A. 

Fig. 3. Ensifera: Haglidae. A. Notopamphagopsis bolivari CABRERA 1928, holotype MLP n° 4354; B-F: Notopamphagopsis? 

sp. 1, specimen PULR-I n° 223 (B); unnumbered specimens fragments, possibly attributable to genus (C – F); G. Notopamphagopsis? 

sp. 2, specimen PULR I n° 259; H. Notopamphagopsis? sp. 3, specimen PULR I n° 270. Terminology: 

ScP, Subcosta posterior, RA, RP, anterior and posterior Radius, respectively; MA, MP, anterior and posterior Media, respectively. 

Scale bar 1 mm.


239 

Holotype:MLPnº4354, housed in MLP, Argentina. Type locality: Cerro Cacheuta, 

Mendoza. Argentina (possibly the same horizon EP I and EP II of MOREL, 1994). Type stratum: 

Potrerillos Formation. Age: late Middle Triassic to early Late Triassic. 

Notopamphagopsis? sp1. 

Fig. 3B 

1999 Notopamphagopsis? sp. MARTINS-NETO &GALLEGO, 2001: Figs 3B-F, 4D-E 

Material. PULR-I nº 223, housed in PULR-I, Argentina. Type locality: Gualo, La 




Fig. 3G 

Material. PULR-I nº 259, housed in PULR-I, Argentina. Type locality: Gualo, La 




Fig. 3H 

Material: PULR-I nº 270, housed in PULR-I, Argentina. Type locality: Gualo, La 



AUCHENORRHYNCHA 

CERCOPOIDEA 

Family: Dysmorphoptilidae HANDLIRSCH, 1906 

Dysmorphoptiloides EVANS, 1956 

Type species: Dysmorphoptiloides elongata EVANS 1956, by original designation. 

Dysmorphoptiloides acostai MARTINS-NETO &GALLEGO, 1999 

Fig. 4B 

*1999 Dysmorphoptiloides acostai MARTINS-NETO &GALLEGO: 192, 194, Fig. 2A. 

2001 Dysmorphoptiloides acostai MARTINS-NETO &GALLEGO 

Holotype.PZ-CTES nº 5772, housed in PZ-CTES, Argentina. Type locality: 300 m 

west of Quebrada del Durazno. South of the Cerro Cacheuta. Mendoza, Argentina, levels EP I and 

EP II of MOREL (1994). Type stratum: Upper Section of the Potrerillos Formation. Age: late Middle 


Supplementary material. PULR-I nº 254 (Fig. 4A). PULR-I nº 260 

(Fig. 4C): fore wing basal fragment with 6 mm of length preserved. Both housed in PULR-I; locality: 

Gualo, La Rioja Province, Argentina; Los Rastros Formation. PZ-CTES nº 7200, (Fig. 4D): fore 

wing fragment with 10 mm of length preserved; destination and other data as in Holotype. 

Dysmorphoptiloides gigantea MARTINS-NETO &GALLEGO 2001 

Fig. 4G 

*2001 Dysmorphoptiloides gigantea MARTINS-NETO &GALLEGO.

240 


Holotype: PULR-I nº 255, housed in PULR-I, Argentina. Type locality: Gualo, La 



Dysmorphoptiloides pulcherrima MARTINS-NETO &GALLEGO n. sp. 

Fig. 4E-F 

Diagnosis.Costal and Anal margin concave at the Apical margin. r-m close to the 

R+MA fusion; MP1 and MP2 two-branched. 

Etymology.From Latin pulcherrima, “the most beautiful”. 




Description.Fore wing 18 mm long. Costal and Anal margin sigmoid, concave at 

apical margin boundary. Apical margin rounded. Costal area broad. ScP+RA five-branched, with 

first branch near mid length of wing, second and third enclosing concave area of the Costal margin. 

R+MA distally fused at same level of concave area of Costal margin. MP two-branched: MP1 and 

MP2 two-branched and MP1a, MP1b, MP2a and MP2b of similar size. r-m close to fusion of 

R+MA. CuA two-branched: CuA1 curved and longer than CuA2. CuA1 and CuA2 encloses concave 

area of Anal margin. m-cu relatively short. Color pattern consists of circular spots placed in 

R+MA fusion, between MP1 and MP2, close to Anal margin, between CuA1 and CuA2, close to 

anal margin, in the middle of wing, and in base of wing. Entire surface of wing rugose, with homogeneously 

granular surface. 

Discussion.Similar to Dysmorphoptiloides acostai, described for the same sediments 

in the general aspects of the wing venation. The new species differs, however, by having both 

the Costal and the Anal margins concave at the apical margin (just in the Costal margin in D. 

acostai), MP1 two-branched (unbranched or at least not branching at the same level in D. acostai) 

and r-m close to the R+MA fusion (far from the R+MA fusion in D. acostai). Additionally, D. pulcherrima 

n. sp. has a different color pattern, formed by spots distributed in the fore wing. 

Gallegomorphoptila MARTINS-NETO n. gen. 

Diagnosis.Costal margin distally notably deflected toward RA+RP, dome-like. Apical 

area short. Anal margin slightly concave at Apical margin. R+MA distally fused. MP1 and MP2 

unbranched. 

Type species. Gallegomorphoptila breviptera MARTINS-NETO n. sp., by present designation. 

Etymology.Inhonor of Dr. Oscar Florencio GALLEGO, who collected all of the material 

described here, and from morphoptila, alluding to Dysmorphoptila, the most similar genus. 

Gender: feminine. 

Discussion.Gallegomorphoptila n. gen. is very similar in general appearance to the 

Jurassic genus Dysmorphoptila HANDLIRSCH, recorded from the German Triassic as well as from 

the Russian Triassic. Gallegomorphoptila differs, however, by R+MA distally fused (R is connected 

to MA by a vertical cross-vein in Dysmorphoptila and in Tennentsia RIEK, 1974, from the 

South Africa Triassic). Gallegomorphoptila n. gen. differs of Dysmorphoptiloides, recorded in the 

Australian Triassic as well as the Argentinean Triassic, by having a costal margin dome-like in 

shape, apical area very short, MP1 and MP2 unbranched, and reduced secondary venation. 

Gallegomorphoptila breviptera MARTINS-NETO n. sp. 


Fig. 4H


241 

Fig. 4. Auchenorrhyncha: Dysmorphoptilidae. A-D. Dysmorphoptiloides acostai MARTINS-NETO &GALLEGO 1999: 

specimen PULR-I n° 254 (A); holotype PZ-CTES n° 5772 (B); specimen PULR-I n° 260 (C); and specimen PULR-I n° 

264 (D). E-F: Dysmorphoptiloides pulcherrima MARTINS-NETO &GALLEGO n. sp., holotype PULR-I n° 261, part (E), 

and counterpart (F); G. Dysmorphoptiloides gigantea MARTINS-NETO &GALLEGO, 2001, holotype PULR-I n° 255; H. 

Gallegomorphoptila breviptera MARTINS-NETO n. sp., holotype PULR-I n° 265. Terminology: ScA, anterior subcosta; 

CuA, anterior cubitus; m-cu, medial-cubital cross-vein; r-m; radial-medial cross-vein. Other abbreviations as for Fig. 2. 

Scale bar 1 mm.

242 


Etymology.From Latin brevis for ‘short’, alludes to the small apical area of the species. 


Rioja Province, Argentina. Type stratum: Los Rastros Formation (second cycle). Age: late Middle 


Description.Fore wing 14 mm long. Costal margin conspicuously deflected distally, 

forming a dome-like structure. Costal area as wide as Radial and Medial areas together. Apical 

margin quite straight where preserved, Apical area short. R+MA distally fused, after level of costal 

margin deflection. MP two-branched: MP1 and MP2 unbranched. CuA two-branched: CuA1 relatively 

long, curved and longer than CuA2. CuA1 and CuA2 branch before of concave area of Anal 

margin. m-cu relatively long. Color pattern absent, or at least, not preserved. Entire surface of wing 

rugose, with homogeneously granular surface. 

Discussion.Similar to Dysmorphoptiloides acostai (described for the same stratum) 

in the general aspects and bearing similar wing venation, differing however by having both the 

Costal area relatively wider and the Apical area shorter. The anal margin is slightly concave at the 

Apical margin (straight in D. acostai, curvature more pronounced in D. pulcherrima n. sp.), MP1 

and MP2 unbranched (MP2 two-branched in D. acostai and D. pulcherrima n. sp.). Gallegomorphoptila 

breviptera n. sp. is similar to Dysmorphoptiloides gigantea, described from the same sediments, 

by MP1 and MP2 unbranched, and differing by its small size, Costal margin dome-like 

(slightly concave in D. gigantea), Anal margin slightly concave (more pronounced in D. gigantea). 

Additionally, G. breviptera n. sp. does not have a color pattern formed by spots distributed in the 

fore wing. 

Bandelnielsenia MARTINS-NETO &GALLEGO n. gen. 

Diagnosis.Fore wing with apical area short, rounded and wide; ScP+RA and MA+R 

distally fused; MP three-branched with MP3 as long as MP2. 

Type species. Bandelnielsenia chilena MARTINS-NETO &GALLEGO n. sp., by present designation. 

Etymology.Inhonor of Dr. Klaus BANDEL and Geol. Sven NIELSEN (Universität 

Hamburg, Geologisch-Paläontologisches Institut and Museum), who collected the material described 

here. 

Discussion.Bandelnielsenia n. gen. differs of all known Dysmorphoptilidae by having 

a short and wide apical area, characteristically elongated and narrow in the other known genera. 

It differs from Dysmorphoptiloides, Dysmorphoptila HANDLIRSCH, Tennentsia RIEK and Gallegomorphoptila 

n. gen. by MP three-branched (four-branched in Dysmorphoptiloides, two-branched in 

Dysmorphoptila and Gallegomorphoptila, and five-branched in Tennentsia). Bandelnielsenia n. 

gen. is similar to Dysmorphoptiloides and Gallegomorphoptila by having SCP+RA and MA+R distally 

fused (unfused in Dysmorphoptila as well as in Tennentsia). Additionally, Bandelnielsenia n. 

gen. differs from all of these genera by CuA branches long (short in the others and CuA1 longer than 

CuA2). 

Bandelnielsenia chilena MARTINS-NETO &GALLEGO n. sp. 

Fig. 6G 

Diagnosis.Asforthegenus. Fore wing 4 mm long. Concave distal region in Costal 

margin incipient, absent in Anal area. 

Etymology.Named for the nation of Chile. 

Holotype:SGO.PI.5989, housed in the Paleontology Section of SGO, Chile. Type locality: 

outcrop at South of Santa Juana city, Southeast Concepción, southern Chile. Type stratum: 

Santa Juana Formation. Age: Upper Triassic.


243 

Description.Fore wing 4 mm long. Costal margin slightly deflected distally, forming 

an incipient dome-like structure. Costal area wider at Apical margin boundary. Apical margin 

wide and rounded, relatively short. R+MA distally fused, after level of Costal margin deflection. 

MP three-branched; MP1+MP2 origin before fusion of ScP+RA with MA+R, MP3 slightly longer 

than MP1+MP2. CuA two-branched, CuA1 long, as long as CuA2. CuA1+CuA2 origin before origin 

of MP3. m-cu relatively long, placed between both MP3 and CuA1+CuA2 origin. No color patterning 

detected. Entire surface of wing rugose, but not consisting of homogeneously granular 

surface. 

Discussion.Allknown Dysmorphoptilidae have a typical constriction in the distal 

part of the costal area, incipient in Bandelnielsenia chilena n. sp. and absent in the anal margin (conspicuous 

or at least incipient in the other known species). Additionally, the fore wing of Bandelnielsenia 

chilena n. sp. is very small when compared with the other known species, in which the fore 

wing is longer than 10 mm. 

SCYTINOPTEROIDEA 

Family: Chiliocyclidae EVANS, 1956 

Tipuloidea rhaetica WIELAND 1925 

Fig. 5F 

*1925 Tipuloidea rhaetica WIELAND: Figs 2, 3. 

1925 Wielandia rhaetica (WIELAND) inTILLYARD: Fig. 1 

1956 Tipuloidea rhaetica WIELAND in EVANS: 209, Fig. 12a; 1992 Tipuloidea rhaetica WIELAND in CAR- 

PENTER: 258, Fig. 162. 2; 1997 Tipuloidea rhaetica WIELAND in GALLEGO: 511-516; 1999 Tipuloidea rhaetica 

WIELAND in GALLEGO & MARTINS-NETO: 87, Fig. 3; 1999 Tipuloidea rhaetica WIELAND in 

MARTINS-NETO &GALLEGO: 194: Fig. 2D. 

Holotype. Unknown. Type locality: Minas de Petroleo (= Puesto Miguez), Cerro 

Cacheuta, southwest of Mendoza, Argentina. Type stratum : Potrerillos Formation ? Age: late Middle 


Remarks.TILLYARD (1925) discussed this material and placed it in the family Scytinopteridae 

HANDLIRSCH, 1926, which was followed by MARTINS-NETO &GALLEGO (1999). However, 

EVANS (1956) created the family Chiliocyclidae, which have a small discal cell in the fore 

wing, and removedTipuloidea to this new family, which is now adopted here, also including the genus 

Argentinocicada MARTINS-NETO &GALLEGO 1999. 

Argentinocicada magna MARTINS-NETO &GALLEGO 1999 

Fig. 5C 

*1999 Argentinocicada magna MARTINS-NETO &GALLEGO: 194, 196; Figs 2C, 4A 

1999 Argentinocicada magna GALLEGO &MARTINS-NETO: 87; Fig. 4 

Holotype.PZ-CTES nº 5729, housed at PZ-CTES, Argentina. Type locality: 300 m 

west of Quebrada del Durazno. South of the Cerro Cacheuta. Mendoza, Argentina, levels EP I and 



Argentinocicada minima MARTINS-NETO &GALLEGO 1999 

Fig. 5D 

*1999 Argentinocicada minima MARTINS-NETO &GALLEGO: Figs 2G, 4B-C 

1999 Argentinocicada minima GALLEGO &MARTINS-NETO: 87; Fig. 5 

Holotype.PZ-CTES nº 5776, housed at PZ-CTES, Argentina. Type locality: 300 m 

west of Quebrada del Durazno. South of the Cerro Cacheuta. Mendoza, Argentina, levels EP I and

244 


Fig. 5. A. Prosbolidinella riorastensis MARTINS-NETO &ROHN 1996 (Prosbolidae), holotype RGMN-T007; B. Potrerillia 

nervosa MARTINS-NETO &GALLEGO 1999 (Scynopteridae), holotype PZ-CTES n° 5728; C. Argentinocicada magna 

MARTINS-NETO &GALLEGO 1999 (Chiliocyclidae), holotype PZ-CTES n° 5729; D. Argentinocicada minima MAR- 

TINS-NETO & GALLEGO 1999 (Chiliocyclidae), holotype PZ-CTES n° 5776; F. Tipuloidea rhaetica WIELAND 

1925(Chiliocyclidae), redrawn from WIELAND (1925: Fig. 2); E, G: Lariojaprosbole melchori MARTINS-NETO &GAL- 

LEGO 2001 (Prosbolidae), holotype PULR-I n° 257, basal part (E), possible apical part (G), respectively. Scale bar 1 mm. 



Supplementary material. PZ-CTES nº 5733, same data as in Holotype. 

Family: Scytinopteridae HANDLIRSCH, 1906 

Potrerillia nervosa MARTINS-NETO &GALLEGO 1999 

Fig. 5B 

*1999 Potrerillia nervosa MARTINS-NETO &GALLEGO: 196, 198: Figs 2B, 5A 

1999 Potrerillia nervosa GALLEGO &MARTINS-NETO: 87, Figs 2;


245 

Holotype.PZ-CTES nº 5728, housed at PZ-CTES, Argentina. Type locality: 300 m west 

of Quebrada del Durazno, south of the Cerro Cacheuta. Mendoza, Argentina, levels EP I and EP II 

of MOREL (1994). Type stratum: Upper Section of the Potrerillos Formation. Age: late Middle Triassic 


Australocicada arcucciae MARTINS-NETO &GALLEGO 2001 

Fig. 6D 

*2001 Australocicada arcucciae MARTINS-NETO &GALLEGO: Fig. 1C. 




Supplementary material.PULR-I nº 266 (Fig. 6C), same data as in 

Holotype. 

Gualoscytina MARTINS-NETO &GALLEGO n. gen. 

Diagnosis.Similar to Saaloscytina BRAUCKMANN and SCHLÜTER, 1993 in general aspect 

of venation, differing, however, by ScP long, ending close to apex (short, reaching costal margin 

before mid length of wing in Saaloscytina); RP origin close to the wing base; and CuA forking 

close to wing base (close to apex in Saaloscytina). 

Type species. Gualoscytina mayae MARTINS-NETO &GALLEGO n. sp., by present designation. 

Etymology.Named for the Gualo locality and scytina, common suffix for Scytinopteridae. 


Discussion.Gualoscytina n. gen. is very similar in general aspects of venation to 

Saaloscytina BRAUCKMANN &SCHLÜTER, 1993, described from the Upper Triassic of Germany, 

differing however, mainly by ScP morphology: short in Saaloscytina and notably long in Gualoscytina 

n. gen. (perhaps a plesiomorphic condition). Also, CuA forks differently: close to the wing 

base in Gualoscytina n. gen. (a plesiomorphic condition), with CuA1 and CuA2 long; close to the 

apex in Saaloscytina (an apomorphic condition), with CuA1 and CuA2 short. 

Gualoscytina mayae MARTINS-NETO &GALLEGO n. sp. 

Fig. 6B 

Diagnosis.Asforthegenus. Fore wing 9 mm long, foliaceous in shape. All principal 

veins with color pattern, contrasting with wing surface. 

Etymology.Inhonor of Dr. Cathleen MAY (Newkirk Engler & May Foundation, 

Denver, Colorado), who collected some of the material described here and coordinated the NGS 

project (grant 5317-94). 

Holotype.PULR(I) nº 267, housed in PULR-I, Argentina. Type locality: Gualo, La 

Rioja Province, Argentina. Type stratum: Los Rastros Formation (second cycle). Age: late Middle 


Description.Fore wing 9 mm long, foliaceous in shape. Wing base very narrow. 

Apex acuminate. ScP long, reaching Costal margin close to apex. RA long, reaching Apical margin 

at apex, with at least one secondary branch. RP origin close to wing base, unbranched. MP long and 

sigmoid, originating shortly after origin of RP, also unbranched. CuA forks at same level of MP origin, 

two-branched: CuA1 long and sigmoid; CuA2 quite straight. Color pattern consists of darker 

color covering all longitudinal veins. Entire surface of the wing rugose, with homogeneously granular 

surface. 

Discussion.Gualoscytina mayae n. sp. considerably differs from Saaloscytina perreticulata 

from the Triassic of Germany in wing shape: foliaceous acuminate in G. mayae n. sp.,

246 


petular in Saaloscytina perreticulata. Additionally G. mayae n. sp. is larger than Saaloscytina perreticulata 

(fore wing 9 mm in G. mayae n. sp., 4 mm in S. perreticulata) and G. mayae n. sp. has 

color patterning in the longitudinal veins, absent in S. Perreticulata. 

Family: Stenoviciidae EVANS, 1956 

Argentinopheloscyta MARTINS-NETO &GALLEGO, n. gen. 

Diagnosis.Similar to Apheloscyta TILLYARD, 1922 in general aspect of wing, differing 

by MP three-branched. 

Type species. Argentinopheloscyta fosterae MARTINS-NETO and GALLEGO n. sp., by present 

designation. 

Etymology.Named for Argentina and pheloscyta, from Apheloscyta, a similar genus. 


Discussion.Argentinopheloscyta n. gen. differs from all known Stenoviciidae 

genera by having MP three-branched. 

Argentinopheloscyta forsterae MARTINS-NETO &GALLEGO n. sp. 

Fig. 6A 


Etymology. Inhonor of Dr. Catherine FORSTER (Department of Anatomical Sciences, 

State University of New York, Stony Brook), who collected some of the material described 

here, and coordinated the NGS project (grant 5317-94). 


Rioja Province, Argentina. Type stratum: Los Rastros Formation (second circle). Age: late Middle 


Description.Fore wing 9 mm long. Costal margin slightly curved. Apical margin 

slightly acuminate, long. Costal area as wide as radial area. RA+RP parallel to Costal margin. RA 

short, reaching Costal margin before Apical margin. RP unbranched. M long and sigmoid, notably 

deflecting after mid length of wing, deflecting again toward Apical margin, with three short 

branches of similar size: MP1 reaching apex, MP2 reaching Apical margin at its mid length and 

MP3 reaching Apical margin close to Anal margin; r-m quite vertical; mp1-mp2 oblique to r-m; 

m-cu perpendicular to MP3, reaching it after MP2+MP3 fork. CuA long and straight, distally twobranched: 

CuA1 longer than CuA2, CuA2 oblique to the anal margin. CuP long, parallel to CuA, 

distally fused with CuA2 at anal margin. 

PROSBOLOIDEA 

Family: Prosbolidae HANDLIRSCH, 1904 

Prosbolidinella riorastensis MARTINS-NETO &ROHN 1996 

Fig. 5A 

*1996 Prosbolidinella riorastensis MARTINS-NETO &ROHN: 246-247, Figs 2, 3. 

Holotype. RGMN-T007, MARTINS-NETO Collection, housed in the Paleontological 

Collection, USP, Brazil. Type locality: 200 m East from Timbó river, in the road Canoinhas-Porto 

União (BR-280), Santa Catarina State, near Poço Preto locality. Type stratum: Morro Pelado Member, 

Rio do Rasto Formation, Paraná Basin. Age: Lower Triassic.


247 

Fig. 6. A. Argentinopheloscyta forsterae MARTINS-NETO &GALLEGO n. sp., (Stenoviciinae), holotype PULR-I n° 268; B. 

Gualoscytina mayae &GALLEGO n. sp. (Scynopteridae), holotype PULR-I n° 267; C-D. Australocicada arcucciae MAR- 

TINS-NETO &GALLEGO 2001 (Chiliocyclidae), specimen PULR-I n° 266 (C), and holotype PULR-I n° 266 (D); E. 

Yurigocimex popovi MARTINS-NETO &GALLEGO n. sp., holotype PULR-I n° 262; F. Popovigocimex yurii MAR- 

TINS-NETO & n. s., holotype PULR-I n° 271; G. Bandelnielsenia chilena MARTINS-NETO &GALLEGO n. sp. (Dysmorphoptilidae), 

holotype SGO.PI.5989; H. Sanctipaulus mendesi PINTO 1956 (Derbidae), redrawn from PINTO (1956: Fig. 

2); H. Auchenorrhyncha indet (probably attributable to Australocicada MARTINS-NETO &GALLEGO), specimen PULR-I 

n° 264.

248 


Lariojaprosbole melchori MARTINS-NETO &GALLEGO, 2001 

Fig. 5E 

*2001 Lariojaprosbole melchori MARTINS-NETO &GALLEGO: Pl. IB, Fig. 1E. 

Holotype.PULR-I nº 257a, housed in PULR-I, Argentina. Type locality and type stratum: 

Quebrada de Ischichuca Chica, La Rioja Province, Argentina. Ischichuca Formation. Age: 

late Middle Triassic (or early Late Triassic sensu ZAVATTIERI &MELCHOR, 1999). 

Supplementary specimen possibly attributable to Lariojaprosbole 

melchori MARTINS-NETO &GALLEGO, PULR-I nº 257b, preserved in the same slab as above (Fig. 5G). 

TRICHOPTERA 

Family: incertae sedis 

Sanctipaulus mendesi PINTO 1956 

Fig. 6H 

*1956 Sanctipaulus mendesi PINTO: 82-83, Fig. 2. 

Holotype.MP/241, Museu de Paleontologia of the ICN of the Universidade Federal 

do Rio Grande do Sul, Brazil. Type locality: the Passo das Tropas locality, Santa Maria, Rio Grande 

do Sul, Brazil. Type stratum: associated with the Thinnfeldia Flora, in the Santa Maria Formation. 

Age: Upper Triassic. 

Remarks.Originally this species was described as Auchenorrhyncha Derbidae, being removed 

here for Trichoptera incertae family. 

COLEORRHYNCHA 

Family: Progonocimicidae HANDLIRSCH, 1906 

Yurigocimex MARTINS-NETO &GALLEGO n. gen. 

Diagnosis.Similar to Archegocimex HANDLIRSCH, 1906 by fore wing with small basal 

cell and CuA zigzag-like. ScP far from RA. M connected to CuA by three cross-veins. 

Type species. Yurigocimex popovi MARTINS-NETO &GALLEGO n. sp., by present designation. 

Etymology.Inhonor of Dr. Yuri POPOV (Russian Academy of Sciences) and gocimex, 

from Archegocimex, the most similar genus. Gender: neutrum. 

Discussion.Yurigocimex n. gen. is similar to Archegocimex HANDLIRSCH, 1906, by 

having a small basal cell and CuA zigzag-like, differing however, by the shape of the basal cell (triangular 

in Archegocimex, pentagonal in Yurigocimex n. gen.). Yurigocimex n. gen. differs of all 

known Jurassic Progonomicidae genera (reviewed by POPOV &WOOTTON 1977), by having ScP far 

from RA. In this aspect, Yurigocimex n. gen. is similar to the Triassic genus Heteronella EVANS, 

1961, differing, however, by having a small basal cell (absent in Heteronella as figured, or if existing, 

is longer than in Yurigocimex n. gen.). Yurigocimex n. gen. differs from all known Jurassic Progonomicidae 

genera by having three cross-veins connecting M to CuA (two or less in the Jurassic 

genus). Three cross-veins connecting M to CuA are also found in Heteronella, differing however 

from those of Yurigocimex n. gen. by m-cua2 close to m-cu3, forming a small square cell (far from 

m-cua3, forming a great triangular cell in Yurigocimex n. General.). 

Yurigocimex popovi MARTINS-NETO &GALLEGO n. sp. 

Fig. 6E 

Diagnosis.Asforthegenus. MP2 distally branched; CuA two-branched with small 

distal dichotomy.


249 

Etymology.Inhonor of Dr. Yuri POPOV (Russian Academy of Sciences), specialist 

on Hemiptera. 




Description.Fore wing 5 mm long. Costal margin curved, Apical margin rounded. 

Costal area very wide, broad at wing apex, narrowing slightly towards wing base. ScP origin at MP 

origin, long, oblique to Costal margin. RA relatively far from ScP, connected to RP by small crossvein. 

r-m at origin of RP+MP1. MP zigzag-like, three-branched: MP1 origin at RP origin; MP2 distally 

branched, connected to RP+MP1 by cross-vein, forming distal part of basal cell. Basal cell 

pentagonal, relatively small. Three cross-veins connect MP to CuA: m-cu1, the longer, at MP origin; 

m-cu2 continuous to r-m; and m-cu3 in mid length of CuA1. CuA two-branched: CuA1 distally 

branched and CuA2 unbranched. CuA2 distally connected to CuA1 through small cross-vein. 

Popovigocimex MARTINS-NETO &GALLEGO n. gen. 

Diagnosis.Similar to Archegocimex HANDLIRSCH, 1906, and Yurigocimex n. gen. by 

fore wing with small basal cell, and CuA zigzag-like. Similar to Yurigocimex n. gen. by ScP far 

from RA, differing however, by having two extra branches of RA. M connected to CuA by two 

cross-veins. 

Type species. Popovigocimex yurii MARTINS-NETO &GALLEGO n. sp., by present designation. 

Etymology.Inhonor of Dr. Yuri POPOV (Russian Academy of Sciences) and gocimex, 

from Archegocimex, the genus most similar. Gender: neutrum. 

Discussion.Popovigocimex n. gen. is similar to Archegocimex and Yurigocimex n. 

gen. by having a small basal cell and CuA zigzag-like, differing, however, by the shape of the basal 

cell (triangular in Archegocimex, pentagonal in Yurigocimex n. gen. and rectangular in Popovigocimex 

n. gen.). Popovigocimex n. gen. differs of all known Jurassic Progonomicidae genera (reviewed 

by POPOV &WOOTTON, 1977), by having ScP far from RA. In this aspect, Popovigocimex is similar 

to the Triassic genus Heteronella, differing however by having a small basal cell (absent in Heteronella 

as figured, or if existing, it is longer than in Popovigocimex). Popovigocimex n. gen. 

differs of Yurigocimex n. gen. by having two extra branches of RA (just one in Yurigocimex n. gen.), 

two cross-veins joining MP to CuA (three in Yurigocimex n. gen.),and CuA branches are longer, 

and not connected by a cross-vein. 

Popovigocimex yurii MARTINS-NETO &GALLEGO n. sp. 

Fig. 6F 

Diagnosis.Asforthegenus. MP2 distally unbranched. 

Etymology.Inhonor of Dr. Yuri POPOV (Russian Academy of Sciences), specialist on 

Hemiptera. 

Holotype.PULR-I nº 271, housed in PULR-I, Argentina. Type locality: Gualo, La 



Description.Fore wing 4 mm long. Costal margin curved, Apical margin rounded. 

Costal area very wide, broad at wing apex, narrowing slightly towards wing base. ScP origin anterior 

to MP origin, long, oblique to Costal margin. RA three-branched. MP two-branched: MP1 origin 

at the RP origin, MP2 unbranched, connected to RP+MP1 by cross-vein, forming distal part of 

basal cell. Basal cell rectangular, relatively small. Two cross-veins connecting MP to CuA: m-cu1, 

smaller, at MP1+RP origin, and m-cu2 close to apex. CuA two-branched: CuA1 and CuA2 unbranched, 

as preserved. Two cross-veins preserved, probable connected to CuP (not preserved).

250 


MECOPTERA? 

Tipulidites affinis WIELAND 1926 

Necrotaulius (Tipulidites) affinis WIELAND 1926 

?Necrotaulius (Tipulidites) affinis WIELAND 1926 

Fig. 2D 

Holotype. Unknown. Type locality: Minas de Petroleo (= Puesto Miguez), Cerro 

Cacheuta, southwest of Mendoza, Argentina. Type stratum. Potrerillos Formation? Age: late Middle 


Remarks. Inprevious papers this species was reported as Diptera, Trichoptera, and 

more recently as Mecoptera as proposed by W. KRZEMIÑSKI (pers. obs). 

IV. TAXONOMIC COMPOSITION AND DISPERSAL PATHS OF THE TRIASSIC 

CONCHOSTRACAN AND INSECT FAUNAS FROM ARGENTINA 

Triassic insect faunas from Argentina occur mostly in association with conchostracan faunas. 

For this reason it is appropriate to include a brief consideration of conchostracan assemblages, and 

their relation to dispersal and migration of insect faunas. 

Conchostracan fossils from Argentina and Chile have been recorded since the beginning of the 

last century, however, detailed taxonomic and stratigraphic studies are more recent. Studies carried 

out by one of the authors (GALLEGO 1992, 1999; GALLEGO &COVACEVICH 1998; GALLEGO & 

MELCHOR 2000) yielded new information on the taxonomic composition of these faunas. Due to the 

frequent occurrence of conchostracans in the Triassic successions of Argentina it is envisaged that 

this taxonomic list will be enlarged in the near future. For this reason the following comments are 

preliminary. 

Work on migration and dispersal paths of fossil Gondwanan conchostracan faunas is scarce. 

TASCH (1970, 1987) described Late Paleozoic – Middle Mesozoic migration routes displaying a 

general trend from the southern to the northern hemisphere, which were named as “Trans-Antarctic 

fresh-brackish water routes”. Taxa considered by TASCH range from the genus level (Paleolimnadia 

sp.), sub-generic (“lioestheriids”, sensu TASCH 1969), to the family level (Leaiidae) for the 

Permian-Jurassic interval. 

TASCH ´s (1970) proposal imposes serious problems for the taxonomy of the group, because the 

“lumping” scheme of TASCH (1969) implied a drastic reduction in the number of taxa. Furthermore, 

the information available at the time of his proposal was scarce, he did not also consider previous 

work, and he included secondary data in his conclusions. At present, there is new and abundant information 

from Argentina, Chile, and Brazil for the Triassic. TASCH (1970) made some inferences 

about the dispersal of “lioestheriids” in Gondwana using data from the Triassic of Argentina, where 

the record is fairly meager and dubious at the time of his proposal. Triassic Argentine fauna shows a 

marked diversity and higher frequency of species belonging to Euestheriidae (first recorded by RUS- 

CONI 1948), which is a family with a poor record in other regions of Gondwana (GALLEGO 1999). 

TASCH (1970) mentioned migratory routes in the southern hemisphere through Antarctica joining 

Australia, South Africa and South America. This author did not consider Triassic conchostracan 

faunas from South America, South Africa and Antarctica, as the available information was very 

scarce or non-existent. The Australian fauna was all referred to the genus Paleolimnadia, which was 

not recorded from the Triassic of Argentina and Chile, and is now known from the Triassic of Brazil 

and Bolivia and the Upper Permian of Brazil and Chile. 

New information produced in the last decade (GALLEGO 1992, 1999; GALLEGO &COVACEVICH 

1998; GALLEGO &MELCHOR 2000) allows some preliminary comments to be made on relationships 

with other conchostracan faunas and on probable migration paths of some taxa. It is clear that


251 

new information on this and other Gondwanan faunas (e.g., recent studies on Triassic faunas from 

Australia; P. JONES, pers. comm., 2001), might modify these conclusions. 

Six of the seven conchostracan families recorded from the Triassic of Argentina show close relationships 

with faunas from Laurasia and Gondwana. The families Palaeolimnadiopseidae (genus 

Endolimnadiopsis SHEN, 1994), Pemphilimnadiopseidae, Ulugkemiidae, Loxomegaglyptidae and 

Fushunograptidae (genus Liograpta GALLEGO &COVACEVICH 1998) recorded in the Triassic of 

Argentina have Devonian to Cretaceous representatives in Laurasia. In particular, Palaeolimnadiopseidae 

is documented by the genus Endolimnadiopsis, also recorded from the Upper Permian of 

Russia. Challaolimnadiopsis SHEN & GALLEGO, 2001 belongs to Pemphilimnadiopseidae, a family 

recorded from the Upper Devonian of China, Upper Carboniferous of North America, and the Upper 

Permian-Lower Triassic of Germany, Italy, Russia, China and Africa (SHEN et al. 2001). Ulugkemiidae 

are represented by the genus Triassulugkemia GALLEGO, 1999, which is also recorded 

from Middle Devonian and Upper Permian of Russia (GALLEGO &MELCHOR 2000). The Loxomegaglyptidae 

(documented in Argentina and the south of Brazil by the genus Triasoglypta GALLEGO, 

1999) have representatives from the Carboniferous to the Tertiary of Laurasia, Lower Carboniferous 

of England, Permian through Triassic of Russia, Triassic through Tertiary of China, and Triassic 

and Tertiary of South America. The Fushunograptidae (documented in Argentina and Chile by 

the genus Liograpta) was recorded in Russia (Upper Permian and Jurassic), Korea (Triassic), China 

(Triassic and Jurassic), Europe (Upper Triassic and Jurassic), and Africa (Triassic). The abovementioned 

evidence, along with the record from the Triassic of Argentina allows for the suggestion 

that it is highly probable that members of these families migrated from Asia either via India to Africa 

and South America or via India to Antarctica and South America through the southern point of 

Patagonia. 

The oldest records of Eosestheriidae are from the Triassic of Argentina and Chile (documented 

by the genus Menucoestheria GALLEGO &COVACEVICH, 1998). Prior to these findings its biochron 

was restricted to the Middle Jurassic to Lower Cretaceous of Asia, Europe and Antarctica. SHEN 

(1994) described the genus Carapacestheria (Eosestheriidae) from the Ferrar Group (Middle Jurassic 

of Victoria Land, Antarctica) and also noted that some of the characters of this genus are also 

present in specimens from the Middle Jurassic of Argentina, namely Cyzicus (Lioestheria) patagoniensis 

TASCH 1970 (Cañadón Asfalto Formation, Chubut Province) and in Cyzicus (Lioestheria) 

malacaraensis TASCH 1970 (La Matilde Formation, Santa Cruz Province). Cyzicus (Euestheria) 

taschi VALLATI and Cyzicus (Euestheria) sp. 1 (also from the Cañadón Asfalto Formation, Chubut) 

described by VALLATI (1986) might be added to this list of related forms, also belonging to 

Eosestheriidae. VALLATI (1986) already mentioned the similarities between Cyzicus (Euestheria) 

sp. 1 and Yanjiestheria xiaxiaensis WANG (Eosestheriidae), the last described from the Upper 

Jurassic-Lower Cretaceous of China. These new records of this family allow a hypothesis on its dispersal 

from Argentina or the southernmost end of South America through Antarctica to Asia and 

Europe (see Fig. 7). 

The family Polycytellidae (Glosselytrodea) has been restricted until now to the Upper Triassic. 

They are very rare; only 6 species are now known in the geological record. These include: Mesojurina 

sogjutensis MARTYNOVA (1952), Mongolojurina altaica PONOMARENKO (1988) and 

Karatajurina unica VILESOV & NOVOKSHONOV 1994, from the Russian Upper Triassic, 

Polycytella triassica TILLYARD 1922, and Permoberothella perplexa RIEK 1974, both from the Upper 

Triassic of Australia, and Argentinoglosselytrina pulchella MARTINS-NETO &GALLEGO 2001, 

from the Upper Triassic of Argentina. The paleogeographic distribution of the group is presented in 

Fig. 7. Although containing a relatively small number of well-known species, this is an important 

record, as Polycytellidae make excellent index-fossils, being restricted to the Upper Triassic, and 

with an exclusively Gondwanan distribution. In paleobiogeographic terms, this group possesses 

wide distribution within Gondwana, with records in Argentina, Australia and in the South Asia. It is 

very probable that Polycytellidae could be found in Antarctica in the future.

252 


Fig. 7. Dispersion tracks of Triassic Conchostraca and selected Triassic insect groups herein proposed. Conchostraca associations: 

C1 – Ulugkemiidae, Pemphilimnadiopseidae, Paleolimnadiopseidae, Fushunograptidae, Loxomegaglyptidae 

and Eosestheriidae; C2 – Loxomegaglyptidae; C3 – Pemphilimnadiopseidae; C4 – Fushunograptidae; C5 – Eosestheriidae; 

C6 – Loxomegaglyptidae, Eosestheriidae, Pemphilimnadiopseidae, and Fushunograptidae; C7 – Loxomegaglyptidae, 

Fushunograptidae, Ulugkemiidae, Pemphilimnadiopseidae and Paleolimnadiopseidae; C8 – Pemphilimnadiopseidae, 

Paleolimnadiopseidae, Fushunograptidae, Loxomegaglyptidae and Eosestheriidae;C9 – Eosestheriidae. Insect associations: 

I1 – Dysmorphoptilidae, Scytinopteridae, Grosselytrodea and Ademosynidae; I2 – Dysmorphoptilidae, Scytinopteroidea, 

Grosselytrodea, Locustopsidae and Ademosynidae; I3 – Locustopsidae; I4 – Scytinopteroidea and Ademosynidae; 

I5, Dysmorphoptilidae, Scytinopteroidea, Grosselytrodea, Locustopsidae, and Ademosynidae. 

Another group of the Auchenorrhyncha, the Dysmorphoptilidae, is more abundant in the Upper 

Triassic, and is extinct in the Lower Jurassic. Ten species are presently known: Dysmorphoptiloides 

elongata EVANS 1956 and Dysmorphoptiloides parva EVANS 1956 (both from the Australian Upper 

Triassic), Dysmorphoptiloides acostai MARTINS-NETO &GALLEGO 1999, Dysmorphoptiloides gigantea 

MARTINS-NETO &GALLEGO 2001, Dysmorphoptiloides pulchra MARTINS-NETO &GAL- 

LEGO 2001, Gallegomorphoptila breviptera MARTINS-NETO n. sp. (all from the Upper Triassic of 

Argentina), Bandelnielsenia chilena MARTINS-NETO &GALLEGO n. sp., from the Upper Triassic of 

Chile, Tennentsia protuberans RIEK 1974, from the Upper Triassic of South Africa, Dysmorphoptila 

notodon SHCHERBAKOV 1988 (Lower Jurassic, Mongolia), and Dysmorphoptila liasina 

HANDLIRSCH 1906 (Lower Jurassic, Germany). The known record of these species is supplied in 

Fig. 7. Species of Polycitellidae are also excellent index fossils. This family has few species, but 

those have wide geographic distribution. Species of the genus Dysmorphoptiloides, represented in 

both the Australian and the Argentinean Triassic, are an excellent index fossils for the Upper Triassic 

of Gondwana, and the genus Dysmorphoptila is a useful index fossil for the Lower Jurassic of 

Laurasia. 

The informal group Scytinopteromorpha is discussed here as including the families Scytinopteridae, 

Serpentiveniidae, Cicadoprosbolidae, Chiliocyclidae and Hylicellidae among others: all with


253 

restricted distribution in the Upper Triassic of Gondwana to the Lower Jurassic Inferior of Laurasia. 

Twenty three species are known for the Australian Upper Triassic: Chiliocycla scolopoides TIL- 

LYARD 1920; Mesoscytina australis TILLYARD 1920; Mesoscytina affinis TILLYARD 1920; Mesodiphthera 

subcostalis TILLYARD 1920; Mesodiphthera prosboloides TILLYARD 1922; 

Mesodiphthera dunstani TILLYARD 1922; Triassojassus proavitus TILLYARD 1920; Triassocarta 

subcostalis TILLYARD 1920; Mesocixius triassicus TILLYARD 1920; Mesocixiodes terminoneura 

TILLYARD 1920; Triassoscytina incompleta EVANS 1956; Triassoscytinopsis stenuata EVANS 

1956; Triassoscytinopsis aberrans EVANS 1956; Triassoscytinopsis paranotalis EVANS 1956; 

Mesothymbris perkinsi EVANS 1956; Mesothymbris woodwardi EVANS 1956; Mesonirvana 

abrupta EVANS 1956; Triassoscelis anomala EVANS 1956; Mesocicadella venosa EVANS 1956; 

Triassocotis australis EVANS 1956; Apheloscysta mesocampia TILLYARD 1922; Crosbella elongata 

EVANS 1956; and Crosbella alata EVANS 1956. The species Scytinoptera distorta RIEK 1976 is 

known from the South Africa Upper Triassic. From the Argentina Upper Triassic, eight species are 

known: Tipuloidea rhaetica WIELAND 1925; Argentinocicada magna MARTINS-NETO &GAL- 

LEGO, 1999; Argentinocicada minima MARTINS-NETO &GALLEGO 1999; Potrerillia nervosa MAR- 

TINS-NETO &GALLEGO 1999; Australocicada arcucciae MARTINS-NETO &GALLEGO 2001, 

Gualoscytina mayae MARTINS-NETO &GALLEGO n. sp., a possible new genus et species (Fig. 6 I), 

and Argentinopheloscyta forsterae MARTINS-NETO &GALLEGO n. sp. The only species known 

from the Upper Triassic of Germany is Saaloscytina perreticulata BRAUCKMANN &SCHLÜTER 

1993. The record of this group in the Lower Jurassic consists of seven species: Mesocixiella asiatica 

MARTYNOV 1937; Mesocixiella furcata MARTYNOV 1937; Mesocixiella extensa MARTYNOV 1937 

and Mesocixiella major MARTYNOV, 1937. All of these are from the Lower Jurassic of Russia. Finally, 

Archijassus vicinus HANDLIRSCH 1906; Liojassus affinis HANDLIRSCH 1906; and Mesojassus 

pachynavus HANDLIRSCH 1906, are known from the Lower Jurassic of Germany. 

As in the previously discussed group, the Scytinopteromorpha origin is clearly gondwanan, well 

represented in Argentina and Australia, index fossils for the Upper Triassic of Gondwana, and for 

the Lower Jurassic of Laurasia through Asia, reaching Germany, westward of which they are unknown 

on the continent. The paleogeography of Scytinopteromorpha is virtually identical of the 

previously discussed groups. 

Two important zones based on the insect taxa discussed above may be proposed: 

(a) the association zone Dysmorphoptila/Mesocixiella, marking the Lower Jurassic (Liassic) of 

Laurasia, correlating the deposits of central Asia and of Europe; and 

(b) association zone Polycitellidae/Dysmorphoptiloides, marking the upper part of the Middle Triassic 

to the beginning of the Upper Triassic in Gondwana, correlating the deposits of Argentina 

(Potrerillos and Los Rastros formations), of the south of Africa (Molteno Formation), of Antarctica 

(Graham Land Series), of Australia (Ipswich Series) and of the south of Asia. 

The Triassic of Gondwana has very representative insect fossils that allow, through their distribution, 

correlation to be established between several deposits. The Triassic insect assemblage of 

Argentina (Potrerillos and Los Rastros formations) is virtually identical with that of the Ipswich Series 

in Australia. The Ipswich Series in Australia is probably the region of origin for these insect 

groups. 

The dispersion routes for insects are very well defined from Australia through Antarctica to Argentina, 

and are not very different to the conchostracans migration routes proposed above. The faunas 

of the Triassic of South Africa and of South Brazil are not correlated to each other (in the case of 

Brazil, this is likely a result of few records reported), but correlation of the South Africa deposits 

with those of the Argentina + Australia Triassic is very low. On the other hand, Triassic conchostracan 

faunas from Argentina are correlated with those of Chile and Brazil, and these differ greatly 

with the faunas of South Africa and Australia. 

Different migration trackway of insects and conchostracans would be due to their distinct dispersion 

forms. Conchostracans migrate by wind transport, and insects mainly by powered flight. 

GALLEGO &MELCHOR (2000) mentioned that the presence of some conchostracan taxa in the

254 


northern and southern hemispheres during the Triassic would be due to the presence of monsoonal 

(transecuatorial) circulation of winds. New expectable insects finds, mainly from Brazilian Triassic, 

could give us a more accurate picture of the distribution of Gondwanan paleoentomofauna. 

REFERENCES 

ANDERSON J. M., ANDERSON H. M. 1993. Terrestrial flora and fauna of the Gondwana Triassic: part 1- Occurrences. 

[In:] S. G. LUCAS,M.MORALES (eds) – The non-marine Triassic. Bulletin of the New Mexico Museum 

of Natural History and Sciences, 3: 3-12. 

BELLOSI E. S., JALFIN G. G., BOSSI G. E., BOGGETTI D., CHEBLI P., MURUAGA C. Facies y sedimentación. 

[In:] A. ARTABE,E.MOREL,A.ZAMUNER (eds) – El Sistema Triásico en Argentina. Fundación Museo de 

La Plata “Francisco P. Moreno”, La Plata (in press). 

BENTON M. J. 1983. Dinosaur success in the Late Triassic, a noncompetitive ecological model. The Quarterly 

Review of Biology, 58:29-55. 

BONAPARTE J. F. 1982. Faunal replacement in the Triassic of South America. Journal of Vertebrate Paleontology, 

2: 362-371. 

BRAUCKMANN C., SCHLÜTER T. 1993. Neue Insekten aus der Trias von Unter-Franker. Geologica et Palaeontologica, 

27: 181-199. 

CARPENTER F. M. 1960. A Triassic Odonate from Argentina. Psyche, 76: 418-425. 

CECIONI G., WESTERMANN G. E. G. 1968. The Triassic/Jurassic marine transition of Coastal Chile. Pacific 

Geology, 1: 41-75. 

COCKERELL T. D. A. 1927. A Journey in South America. Natural History, 27(1): 77-92. 

EVANS J. W. 1956. Paleozoic and Mesozoic Hemiptera (Insecta). Australian Journal of Zoology, 4: 165-268. 

EVANS J. W. 1961. Some Upper Triassic Hemiptera from Queensland. Memoirs of the Queensland Museum, 

14: 13-33. 

FUENZALIDA H. 1937. El Triásico en la costa de Chile Central. Ministerio de Fomento, Departamento de Minas 

y Petróleo, 6 (65): 739-747. 

GALLEGO O. F. 1992. Conchóstracos Triásicos de Mendoza y San Juan, Argentina. Ameghiniana, 29(2): 159-175. 

GALLEGO O. F. 1994. Conchóstracos (Cyzicidae) del Jurásico de Santa Cruz y Chubut, Argentina. Ameghiniana, 

30(4): 333-345. 

GALLEGO O. F. 1996. Revisión de algunos conchóstracos de la Formación Santa María (Triásico Medio a Superior) 

del Estado de Rio Grande del Sur (Brasil). Acta Geológica Leopoldensia, 19(43): 59-76. 

GALLEGO O. F. 1997. Hallazgos de insectos triásicos en la Argentina. Ameghiniana, 34 (4): 511-516. 

GALLEGO O. F. 1999. Estudio Sistemático de las Faunas de Conchóstracos Triásicos de la República Argentina. 

Córdoba, Facultad de Ciencias Exactas, Físicas y Naturales, Universidad Nacional de Córdoba. Thesis 

Doctoral. Inédita, 210pp. 

GALLEGO O. F., BREITKREUZ C. 1994. Conchóstracos (Crustacea-Conchostraca) paleozoicos de la Región de 

Antofagasta, norte de Chile. Revista Geológica de Chile, 21(1): 31-53. 

GALLEGO O. F., COVACEVICH V. 1998. Conchóstracos Triásicos de las Regiones de Antofagasta, Atacama y 

Coquimbo, Chile. Revista Geológica de Chile, 25(2): 115-137. 

GALLEGO O. F., MARTINS-NETO R. G. 1999. La entomofauna mesozoica de la Argentina: Estado actual del 

conocimiento. Revista Sociedad Entomológica Argentina, 58 (1-2): 86-94. 

GALLEGO O. F., MELCHOR R. N. 2000. La Familia Ulugkemiidae Novozhilov, 1958 (Conchostraca) en el 

Triásico de Argentina: Implicancias paleogeográficas. Ameghiniana, 37(1): 47-58. 

GOROCHOV A. V. 1995. System and evolution of the suborder Ensifera (Orthoptera). Parts 1 and 2., Trudy Paleontologicheskogo 

Instituta AN SSSR, 260. Sankt Petersburg, 224 and 213 pp. (In Russian). 

HANDLIRSCH A. 1906-1908. Die Fossilen Insekten und die Phylogenie der Rezenten Formen. Ein Handbuch 

für Paläontologie und Zoology. ENGELMANN (ed.), Leipzig, 640pp. 

HOLZ M., BARBERENA M. C. 1994. Taphonomy of the south Brazilian Triassic paleoherpetofauna: patterns of 

death, transport and burial. Palaeogeography, Palaeoclimatology, Palaeoecology, 107: 179-197. 

KOKOGIÁN D. A., FERNANDEZ SEVESO F., MOSQUERA A. 1993. Las secuencias sedimentarias triásicas. [In:] 

RAMOS V. A. (ed.). Geología y Recursos Naturales de Mendoza. 12° Congreso Geológico Argentino & 2º 

Congreso de Exploración de Hidrocarburos, p. 65-78. 

KOKOGIÁN D. A., SPALLETTI L. A., MOREL E., ARTABE A., MARTÍNEZ R. N., ALCOBER O. A., MILANA J. P., 

ZAVATTIERI A. M., PAPÚ O. H. 1999. Los depósitos continentals triásicos. [In:] R. CAMINOS – Geología 

Argentina. Anales Instituto de Geología y Recursos Minerales, 29: 377-398. 

KUKALOVÁ-PECK J. 1991. Fossil History and the Evolution of Hexapod Structures. [In:] I. D. NAUMANN, P. 

B. CARNE,J.F.LAWRENCE,E.S.NIELSEN,J.P.SPRADBERY,R.W.TAYLOR,M.J.WHITTEN,M.J.LIT- 

TLEJOHN (eds) –The Insects of Australia. Melbourne, University Press, 1: 141-179.


255 

MARQUAT F. J. 1991. Ninfa de Miomóptero (Insecta) del Triásico de Mendoza, República Argentina. Revista 

Museo Historia Natural San Rafael, Mendoza, 11(1): 3-13. 

MARTINS-NETO R. G., GALLEGO O. F. 1999. The Triassic Insect Fauna from Argentina. I. Auchenorrhyncha, 

Miomoptera and Ensifera. Revista Española de Paleontología, 14(2): 191-202. 

MARTINS-NETO R. G., GALLEGO O. F. 2001. The Triassic Insect Fauna from Argentina. III. Auchenorrhyncha 

and Glosselytrodea. Acta Geologica Leopoldensia, 24(52/53): 105-114. 

MARTINS-NETO R. G., ROHN R. 1996. Primeiro registro de inseto na Formação Rio do Rasto, Bacia do Paraná, 

com descrição de novo táxon. Geociências (São Paulo), 15: 243-251. 

MARTYNOV A. V. 1937. Liasovye nasekomye Shuraba i Kizil-kii. Trudy Paleontologicheskogo Instituta NA 

SSSR, 7(1): 5-232. 

MARTYNOVA O. 1952. Order Glosselytrodea from the Permian beds of Kemerov region. Tr. Paleont. in-ta AN 

SSSR, 40: 187-196. (In Russian). 

MAZZONI A. F. 1995. Los insectos fósiles. [In:] P. N. STIPANICIC,M.A.HÜNICKEN (eds) – Contribuciones a 

la paleophytologia Argentina. Actas Acad. Nac. Ci., 1995: 227-231. 

MELCHOR R. N. 1998. Icnología de los depósitos fluviales y lacustres de la Fm. Ischichuca (Triásico), Cuenca 

Ischigualasto Villa Unión, Quebrada Ischichuca Chica, La Rioja, Argentina. Tercera Reunión Argentina de 

Icnología y Primera Reunión de Icnología del Mercosur, Resúmenes: 19-20. Mar del Plata. 

MOREL E. M. 1994. El Triásico del Cerro Cacheuta, Mendoza (Argentina). Parte I. Geología, contenido paleoflorístico 

y cronoestratigrafía. Ameghiniana, 31(2): 161-176. 

PINTO I. D. 1956. Artrópodes da Formação Santa Maria (Triássico Superior) do Rio Grande do Sul, com 

notícias sobre alguns restos vegetais. Bol. Sociedad Brasil Geologia, São Paulo, 5(1): 76-87. 

PINTO I. D. PURPER I. 1978. A new genus and two new species of Plecopteran Insects, from the Triassic of Argentina. 

Pesquisas, 10: 77-86. 

PONOMARENKO A. G. 1988. New Mesozoic Insects. [In:] New species of Fossil Invertebrates of Mongolia. 

Transactions of the joint Soviet-Mongolian Paleontological Expedition, Nauka, 33: 71-80. [in Russian] 

POPOV Yu. A., WOOTTON R. J. 1977. The Upper Liassic Heteroptera of Mecklenburg and Saxony. Systematic 

Entomology, 2: 333-351. 

RIEK E. F. 1976. A new collection of insects from the Upper Triassic of South Africa. Annals of the Natal Museum, 

Pietermaritzburg, 22(3): 791-820. 

RUSCONI C. 1948. Algunas especies de Estherias del Triásico en Mendoza. Revista del Museo de Historia 

Natural, 2: 99-202. 

SHEN Y.-B. 1994. Jurassic conchostracans from Carapace Nunatak, southern Victoria Land, Antarctica. Antarctic 

Science, 6(1): 105-113. 

SHEN Y.-B., GALLEGO O. F., ZAVATTIERI A. M. 2001. A new conchostracan genus from the Triassic Potrerillos 

Formation, Argentina. Acta Geologica Leopoldensia, 24 (52/53): 227-236. 

SHUBIN N. H., SUES H.-D. 1991. Biogeography of early Mesozoic continental tetrapods: patterns and implications. 

Paleobiology, 17: 214-230. 

SPALLETTI L. A., ARTABE A., MOREL E., BREA M. 1999. Biozonación paleoflorística y cronoestratigrafía del 

Triásico Argentino. Ameghiniana, 36: 419-451. 

STIPANICIC P. N., 1983. The Triassic of Argentina and Chile. [In:] M. MOULLADE, A.E.M.NAIRN (eds) – 

The Phanerozoic Geology of the World. II. The Mesozoic, B: 181-199, Elsevier, Amsterdam. 

STIPANICIC P. N., BONAPARTE J. F. 1979. Cuenca triásica de Ischigualasto-Villa Unión (provincias de San 

Juan y La Rioja). [In:] J. C. M. TURNER (ed.) – Segundo Simpósio de Geología Regional Argentina. Academia 

Nacional de Ciências, 1: 523-575. 

TASCH P. 1969. Branchiopoda. [In:] R. C. MOORE (ed.) – Treatise on Invertebrate Paleontology. Geological 

Society of America and University of Kansas, Part R., Arthropoda, 4(1): R128-R191. 

TASCH P. 1970. Antarctic and other gondwana conchostracans and insects: new data; significance for the Drift 

theory. Proceedings and Papers Second Gondwana Symposium. Pp. 589-592. 

TASCH P. 1987. Fossil Conchostraca of the Southern Hemisphere and Continental Drift. Paleontology, Biostratigraphy 

and Dispersal. Geological Society of America, Memoir, 165. 290 p. 

TILLYARD R. J. 1919. Mesozoic Insects of Queensland. N° 5. Mecoptera, the new order Paratrichoptera, and 

additions to Planipennia. Proceedings of the Linnean Society, New South Wales, 44(1): 194-212. 

TILLYARD R. J. 1920. Mesozoic Insects of Queensland. N° 7. Hemiptera Homoptera; with a note on the Phylogeny 

of the Suborder. Proceedings of the Linnean Society, New South Wales, 44(4): 857-896. 

TILLYARD R. J. 1922. Mesozoic Insects of Queensland. N° 9. Orthoptera and additions to the Protorthoptera, 

Odonata, Hemiptera, and Planipennia. Proceedings of the Linnean Society, New South Wales, 47: 447-470. 

TILLYARD R. J. 1925. Alleged Rhaetic Crane Flies. Nature, 116 (2923): 676-677. 

VALLATI P. 1986. Conchostracos Jurásicos de la Provincia de Chubut, Argentina., 1986. Atas del V Congreso 

Argentino de Paleontologia y Bioestratigrafia, Mendoza 1986, 4: 29-38. 

VILESOV A. P., NOVOKSHONOV V. G. 1994. New fossil insects (Myrmeleontida, Jurinida) from the Upper 

Permian of Eastern Kazakhstan. Paleontological Journal, 2: 66-74. (In Russian).

256 


WIELAND G. R. 1925. Rhaetic crane flies from South American. American Journal of Sciences, 9: 21-28. 

ZALESSKY Yu. M. 1950. New representatives of fossil insects of order Protodonata. Byulleten Moskovskogo 

Obshchestva Ispytateley Prirody, otdel. Geol., 25(4): 98-108. 

ZAVATTIERI A. M., BATTEN D. J. 1996. Miospores from Argentinian Triassic deposits and their potential for 

intercontinental correlation. [In:] J. JANSONIUS,D.C.McGREGOR (eds) – Palynology: principles and applications; 

American Association of Stratigraphic Palynologists Foundation, vol. 2, p. 767-778. 

ZAVATTIERI A. M., MELCHOR R. N. 1999. Estudio palinológico preliminar de la Fm. Ischichuca (Triásico), en 

su localidad tipo (Quebrada de Ischichuca Chica), provincia de La Rioja, Argentina. Asociación Paleontológica 

Argentina, Publicación Especial 6, 33-38. 

ZEUNER F. E., 1939. Fossil Orthoptera Ensifera. London, Monograph British Museum Natural History, 321p.

Rafael Gioia MARTINS-NETO, Oscar Florencio GALLEGO

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?