A COMPARATIVE STUDY OF THE TYPES OF THREE SPECIES
OF MYXOMYCETES: TRICHIA CRATERIFORMIS, T. FALLAX VAR.
OLIVACEA AND T. FERNBANKENSIS
G. MORENO & A. CASTILLO
Departamento de Biología Vegetal, Facultad de Biología, Universidad de Alcalá,
Alcalá de Henares, E-28871 Madrid
gabriel.moreno@uah.es; aurelio.castillo@uah.es
Summary. MORENO, G. & A. CASTILLO (2013). A comparative study of the types of three
species of Myxomycetes: Trichia crateriformis, T. fallax var. olivacea and T. fernbankensis. Bol.
Soc. Micol. Madrid 37: 85-98.
The type of Trichia crateriformis was examined with light and electron microscopy and compared
with a lectotype of Trichia fallax var. olivacea. The two taxa are here considered to be conspeciic.
Also, the type of Trichia fernbankensis was examined with light and electron microscopy and is
a synonym of Trichia decipiens.
Keywords: Amoebozoa, Myxomycota, scanning electron microscopy, taxonomy, type material.
Resumen. MORENO, G. & A. CASTILLO (2013). Un estudio comparativo de los tipos de tres
especies de Myxomycetes: Trichia crateriformis, T. fallax var. olivacea and T. fernbankensis. Bol.
Soc. Micol. Madrid 37: 85-98.
Se estudia el tipo de Trichia crateriformis con microscopía óptica y electrónica y se compara
con el lectotipo de Trichia fallax var. olivacea. Los dos taxones son considerados coespeciicos.
También, el tipo de Trichia fernbankensis fue examinado con microscopía óptica y electrónica y
es un sinónimo de Trichia decipiens.
Palabras clave: Amoebozoa, material tipo, microscopio electrónico de barrido, Myxomycota,
taxonomía.
INTRODUCTION
An examination of type material is fundamental for knowing any species well and this is
particulary true for the myxomycetes. However,
locating a type specimen may be a dificult task,
especially when it is very old material. Sometimes
the location of type material was not provided
in the original description, or the material may
subsequently have been moved. Furthermore, access to a type may be restricted due to its age or
Bol. Soc. Micol. Madrid 37. 2013
scarcity of the material.
Investigators therefore frequently rely on the
original species description or subsequent descriptions made by other investigators who may not
have actually examined the type specimen. Older
descriptions are often brief and their interpretation may be dificult and ambiguous. As a consequence, misinterpretations of taxa are passed on,
as in the case of Lamproderma cribrarioides (Fr.)
R.E. Fr., which had been misinterpreted since its
description in 1829. The holotype of this species
85
G. MORENO & A. CASTILLO
was examined by SINGER & al. (2003) found
that corresponds to a new species, L. retirugisporum G. Moreno, H. Singer, Illana & A. Sánchez,
that had previously not been properly described.
The solutions to various taxonomic problems
may depend on the careful study of the type material. Here we continue with this line of research
to review taxa in the genus Trichia. We compared
the original diagnoses with a reexamination of the
type material under SEM, using the technique of
critical point drying to examine spores and capillitium which has been essential in obtaining the
results presented below.
MATERIALS AND METHODS
The material is preserved in the herbarium
of the Dept. of Plant Biology, University of Alcalá, (AH) and in the herbarium of the National
Fungus Collections (BPI). Other specimens were
from the herbaria AH, MA-Fungi, and the herbarium of D.W. Mitchell. The collections studied were mounted in Hoyer’s medium. Spores
were measured including surface structures such
as spines or warts with an oil immersion lens.
Light microscopy (LM) was made with a Nikon
eclipse 80i microscope equipped with a digital
automatic photographic system Nikon DS-5M.
The SEM micrographs were made with a Zeiss
DSM-950 microscope. For ultramicroscopic studies, the material to be examined was rehydrated in
concentrated ammonium hydroxide (28–30%) for
30 minutes, dehydrated in aqueous ethanol (70%)
for 30 minutes, ixed for 2 hours in pure ethylene
glycol dimethyl ether (= 1,2–dimethoxymethane)
and inally immersed in pure acetone for at least
2 hours. This was followed by critical point drying and sputter-coating with gold-palladium. This
technique uses very little material (one sporocarp,
a part of it or only a small portion of spores). Terminology used to describe spore ornamentation
follows that of RAMMELOO (1974, 1975).
TAXONOMY
Trichia crateriformis G.W. Martin, Mycologia
55: 131 (1963) Figs. 1–10
≡ T. craterioides G.W. Martin, Brittonia 14: 183
86
(1962) nom. illeg., non T. craterioides Corda,
Icon. Fung. 2: 21 (1838)
Specimens examined: NEW ZEALAND. Riccarton Bush, South Island, on dead wood, June 1957,
P.S. Evans 47 (CANTY, type; portion in IA, now
in BPI). Coll. Peter S. Evans, leg. W.R. Philipson
47, BPI 835163 type (isotype; this material is a
portion sent to IA and currently deposited in BPI).
Two ixed slide preparations indicate the typus,
BPI 835156.
Observations: The type (BPI 835163) is on
a woody substrate attached to the lid of a cardboard box and consists of a group of approx. 12
sporocarps with stipes of 1–1.5 mm total height,
some of which are in poor condition. Sporotheca
green olive and 0.75–1 mm diam. (Fig. 3). Dehiscence is well marked by means of a circular
area in almost all of the sporocarps (Fig. 4). Stipe
cylindrical, 0.5–0.75 mm in height, dark to almost black, with the interior filled with cell-like
bodies, 13–20 µm diam. Capillitium formed of
pale yellow elaters 7–8 µm in diam. and 3–4
broad and smooth, long-tipped spirals (Figs. 5-7).
Spores 11–12.5 µm diam., globose, without refractive bodies, pale yellow, spinulose to crested,
with the ornamentation evenly distributed, wellmarked and approx. 1 µm high. In phase-contrast
the crests are easily observed. Under SEM the
spore ornamentation is cristate and is formed by
broad-based ridges that join to form short crests
with irregular and sinuous morphology (Figs.
8-10). Sometimes they join towards the apex to
assume a stellate or reticulate appearance. The
capillitial elaters under SEM are smooth and with
tightly-packed spirals (Figs. 5-7).
In addition, two slides remain in a separate
box (BPI 835156), one with a circular coverslip,
the other with a square coverslip and labelled as
Trichia craterioides, the epithet chosen by MARTIN (1962) subsequently called T. crateriformis
(MARTIN, 1963). On both it is indicated that the
specimen consists of type material. In the abovementioned slide preparations and with the effect
of the years passed, LM observations show the
elaters to be more than 250 µm in length and
5–6 µm diam., ornamented with broad spirals and
Bol. Soc. Micol. Madrid 37. 2013
A COMPARATIVE STUDY OF THE TYPES OF THREE SPECIES OF MYXOMYCETES: TRICHIA CRATERIFORMIS,
T. FALLAX VAR. OLIVACEA AND T. FERNBANKENSIS
Figs. 1-10 Trichia crateriformis type. 1. Label on the box in which the collection was conserved. 2. Two ixed preparations which indicate
the type deposited in BPI. 3. Sporocarps. 4. Detail of the circumscissile dehiscence of the sporocarps. 5-6. Endings of the elaters. 7. Smooth
spirals of the elaters. 8-9. Spores as viewed by SEM. 10. Detail of spore ornamentation as viewed by SEM.
Scale bars: 3-4 = 1 mm, 5-7 = 5 µm, 8-9 = 2 µm, 10 = 1 µm.
Bol. Soc. Micol. Madrid 37. 2013
87
G. MORENO & A. CASTILLO
with tapered tips more than 25 µm long. Spores
are collapsed and approximately 10–11 µm in
diam., pale yellow, spinulose and with short wellmarked crests.
There are few records of Trichia crateriformis.
The type locality is in New Zealand and there are
other records from Tierra del Fuego in Argentina
(ARAMBARRI, 1975), Austria (SCHUSSLER,
1975) and India (LAKHANPAL & MUKERJI,
1981).
NEUBERT & al. (1993: 197-198) and WANG &
LI (2006, plate 48).
Trichia decipiens var. decipiens (Pers.) T. Macbr.,
N. Amer. Slime-Moulds: 218 (1899) Figs. 11-22.
≡ Arcyria decipiens Pers., Ann. Bot. (Usteri) 15:
35 (1795)
= Trichia fallax Pers., Observ. Mycol. 1: 59
(1796)
Observations: The type material is well preserved with numerous clustered, pyriform sporocarps, on remains of wood that contain abundant
capillitium and spores. The sporocarps are short
stalked. Sporotheca 0.6-1 mm in diam., subglobose to ovoid with greenish yellow to greenish
brown shades (Fig. 24). Peridium single, thin,
translucent, with apical circumcissile dehiscence
that breaks to leave the sporocarp in the shape of
a deep calyculus with regular edges. Operculum
membranous, similar in structure to that of the
calyculus (Fig. 25). Stalk short, darker than the
sporotheca and tapered towards the base. Capillitium 7-8 µm diam., formed of yellowish elaters with thick, unbranched, smooth spirals and
long-tapered tips (Figs. 26-27). Spores 11-12 µm
diam., globose to subglobose, pale yellowish,
verruculose to shortly crested and which are seen
well in phase contrast. By SEM the spore ornamentation is cristate and the crests have an irregular morphology, formed by being linked to each
other by bacula with a broader base (Figs. 28-30).
When the bacula are joined at the apex they take
a stellate or subreticulate appearance, sometimes
leaving small complete circles (Figs. 32-33). The
elaters of the capillitium have smooth, closely
spaced spirals with long, sinuous tips (Figs. 2627). The inner surface of the peridium is smooth
(Fig. 31).
Specimens examined: MEXICO. on woody debris, MEXU 11292 in AH 31771. JAPAN, Mt.
Tskura, Namischo, Fukushima Pref., coll. M.
Hario, 22.10.1996, YY 16456 with duplicate in
DWM 5315.
Observations: It is characterized by its pyriform or globose form and the brown to greenish
brown colour of the sporotheca (Fig. 11). Stalk
shorter and darker than the sporotheca (Fig. 12).
Peridium membranous, iridescent and translucent, with irregular dehiscence at the apex that
persists at the base as a calyculus with irregular
edges (Fig. 13). Capillitium formed of elaters with
smooth spirals and of equal diam. that are tapered
towards the ends (Figs. 14-15). We emphasize this
particular spore ornamentation by LM, formed by
a complete reticulum of small meshes per hemisphere and with warts and crests in the remaining surface. SEM studies confirm the existence
of this contrasting type of spore ornamentation
between hemispheres (simple reticulate and cristate) (Figs. 16-22). In addition, we observe that
the walls of the meshes are of uniform width and
height, without having hollows or perforations.
Sometimes the interior of the mesh has small free
projections that are not anastomosed. The meshes
have a variable morphology with a width up to 2
µm. This type of ornamentation has also been observed using SEM by HATANO (1986, plate 21),
88
Trichia fallax var. olivacea Meyl., Bull. Soc.
Vaud. Sci. Nat. 44: 300 (1908) Figs. 23-33.
Specimen examined: SWITZERLAND. Deneriaz-Dessus, Le Chasseron, Canton Vaud, on dead
wood, 1300 m, leg. C. Meylan, X-1908, lectotype
myxo 069 herbarium C. Meylan in LAU.
Trichia decipiens var. olivacea (Meyl.) Meyl.,
Bull. Soc. Vaud. Sci. Nat. 55: 244 (1924)
Specimens examined: CUBA. on logs, 1857, leg.
C. Wright, ex folder 193, nº 546 Curtis Fungus
Herbarium, FH, as T. varia. SPAIN. Badajoz,
Azuaga, Arroyo Argallón, on dead wood of
Bol. Soc. Micol. Madrid 37. 2013
A COMPARATIVE STUDY OF THE TYPES OF THREE SPECIES OF MYXOMYCETES: TRICHIA CRATERIFORMIS,
T. FALLAX VAR. OLIVACEA AND T. FERNBANKENSIS
Figs. 11-22 Trichia decipiens var. decipiens DWM 5315. 11-12. Sporocarps. 13. Detail of the irregular dehiscence of the sporocarp. 14.
Smooth spirals of the elaters. 15. Endings of the elaters. 16-17 and 19-22. Spores as viewed by SEM forming a complete reticulum of
small meshes in one hemisphere and with warts and crests on the remaining surface. 18. Detail of the complete reticulum ornamentation
of spore as viewed by SEM.
Scale bars: 11-13 = 0.5 mm, 14-15 = 5 µm, 16-17 and 19-22 = 2 µm, 18 = 1 µm.
Bol. Soc. Micol. Madrid 37. 2013
89
G. MORENO & A. CASTILLO
Figs. 23-33 Trichia fallax var. olivacea lectotype. 23. Label on the box in which the collection was conserved. 24. Sporocarps. 25. Detail
of the circumscissile dehiscence of the peridium. 26. Elater tip. 27. Smooth spirals of the elater. 28-29. Spores as viewed by SEM formed by
short crests. 30. Detail of the crested ornamentation of a spore viewed by SEM. 31. Detail of the inner peridium AH 14899. 32-33. Spores
as viewed by SEM formed by short crests AH 14899.
Scale bars: 24-25 = 1 mm, 26-27 = 5 µm, 28-29 and 32-33 = 2 µm, 30 = 1 µm. 31 = 50 µm.
90
Bol. Soc. Micol. Madrid 37. 2013
A COMPARATIVE STUDY OF THE TYPES OF THREE SPECIES OF MYXOMYCETES: TRICHIA CRATERIFORMIS,
T. FALLAX VAR. OLIVACEA AND T. FERNBANKENSIS
Populus alba, leg. J.R. García, 10.4.1991, AH
13200. Ibidem on woody debris of Nerium oleander, 27.12.1991, AH 16317. Ibidem 2.2.1992,
AH 16314. Ibidem on dead wood of Quercus sp.,
31.5.1993, AH 16483. Barcelona, Santa Fé del
Montseny, on woody debris of deciduous, leg.
G. Moreno and C. Illana, 17.10.1989, AH 12187.
Burgos, Villafranca Montes de Oca, on wood of
Fagus sylvatica, leg. A. Castillo, 6.10.1991, AH
14700. Ibidem leg. A. Castillo and I. Sevilla,
25.7.1993, AH 16224. San Miguel de Pedroso,
on wood of Quercus faginea, leg. A. Castillo,
15.8.1992, AH 14812. Fresneda de la Sierra,
on wood of Fagus sylvatica, leg. A. Castillo,
7.11.1992, AH 15352. Cáceres, Sierra de Bernabé del Piornal de Tormantos, on woody debris
of Castanea sativa, leg. R. Galán, 11.11.1987,
AH 12348. Ibidem leg. G. Moreno and C. Illana,
28.11.1988, AH 11415. Cantabria, Parque Natural
del Saja-Besaya, Ucieda de Arriba, on wood of
Fagus sylvatica, 28.8.1996, 315 m, leg. A. Sánchez, AH 30309. Mirador Cruz de Cabezuela, Polaciones, on wood of Fagus sylvatica, 15.9.1999,
1150 m, leg. A. Sánchez, AH 30310. Córdoba,
Argallón, Finca Las Canalejas, on woody debris of deciduous, leg. J.R. García, 8.12.1990,
AH 12699. Guadalajara, Aldeanueva de Atienza,
river Pelagallinas, on wood of Pinus sylvestris,
leg. A. Castillo, 17.11.1993, AH 16393. La Rioja,
Ezcaray, near to resort of Vadezcaray, on wood of
Fagus sylvatica, leg. A. Castillo, 15.3.1992, AH
14899. Lérida, Sorpe, Pto. de la Bonaigua, on
wood of Abies alba, leg. F. Esteve-Raventós, A.
Altés, V. González and C. Illana, 3.10.1991, AH
13844. Madrid, Montejo de la Sierra, on wood
of Rosaceae, leg. G. Moreno and A. Castillo,
30.4.1992, AH 14807. Ibidem on wood of Fagus
sylvatica, AH 14808. Ibidem on wood of Quercus
pyrenaica, AH 14809. Puerto de Cotos, on wood
of Pinus sylvestris, 4.7.1998, 1900 m, leg. A. Sánchez, AH 19160. Málaga, Cortes de la Frontera,
Pto. de Galis, La Sauceda, on wood of Quercus
suber, leg. M.T. Vizoso and C. Illana, 11.12.1990,
AH 13078, AH 13080. Segovia: Puerto de la
Quesera, Riofrío de Riaza, on wood of Fagus
sylvatica, 13.12.1997, 1550 m, leg. A. Sánchez,
AH 19161. Ídem, 1.11.1999, 1500 m, AH 28866.
Ídem, 1.11.2000, 1600 m, AH 28863. Valsaín, on
Bol. Soc. Micol. Madrid 37. 2013
lumber, 23.5.1998, 1500 m, leg. A. Sánchez, AH
30327. Garden of Palacio, San Ildefonso de La
Granja, on lumber, 8.2.1999, 1500 m, leg. A. Sánchez, AH 30328. Caballar, on wood of Populus
sp., 17.2.2001, 1030 m, leg. A. Sánchez, AH
28835. Puerto de la Quesera, Riofrío de Riaza, on
wood of Fagus sylvatica, 9.7.2000, 1500 m, leg.
A. Sánchez, AH 30330. Ídem, 1.11.2000, 1650
m, AH 28883. Tenerife (Canary Islands): Monte
de los Silos, Teno, on broadleaf stick, 28.3.78,
DWM 3199. Las Mercedes, on rotting wood, leg.
C.L. Champion, 16.3.1975, DWM 1918. Monte
de los Silos, Teno, on dead wood; 28.3.78, DWM
3201. Los Organos, on dead wood of Pinus canariensis, 29.3.78. DWM 3224. Zaragoza, Moncayo, 1110 m, on wood of Quercus pyrenaica,
leg. M. Lizárraga and D. Cuadrado, 22.1.2005,
AH 34168, AH 34169, AH 34170, AH 34172, AH
34173. UNITED KINGDOM. Lavender Platt,
Ashdown Forest, East Sussex, Vice-county 14,
on dead wood of Fagus sylvatica; National Grid
Reference TQ3933, 20.8.1970, DWM 678. Paternoster Wood, Coleman’s Hatch, East Sussex, Vc.
14, on dead wood of Quercus robur; TQ4534,
2.10.1971, DWM 1018. Pam’s Wood, Horsted
Keynes, East Sussex, Vc. 14, on dead wood of
Quercus robur; TQ3728, 26.3.1972, DWM 1185.
Great Yews, South Wiltshire, Vc. 8, SU1122, on
dead wood, 16.3.1975. DWM 1870. Near Dores
by Loch Ness, Easterness-with-Nairn, Scotland,
Vc. 96, on dead wood, NH5934, 1.8.1975, DWM
2045. Winkhurst Green, West Kent, Vc. 16, on
dead wood, TQ4950, 20.2.1977, DWM 2890.
Alexandra Park, Hastings, East Sussex, Vc. 14,
on dead Ulmus, TQ8010, 10.2.80, DWM 3447.
Bouldner Plantation, Isle of White, Vc. 10, oak
branch on the ground; SZ3789, 25.8.1980, DWM
3508. Near Rushar’s Cross, Mayfield, East Sussex, Vc. 14, on dead wood, TQ5928, 15.3.1997,
DWM 5227. Hartfield, East Sussex, Vc. 14, on
dead wood; TQ4835, I.7.1997, DWM 5292.
Observations: Trichia decipiens var. olivacea
is characterized by its brown to olive-brown, often globose sporothecae. Peridium membranous,
with well-defined circumscissile operculate apical dehiscence which persists as a calyculus with
smooth edges. Operculum membranous, similar
91
G. MORENO & A. CASTILLO
to the structure of peridium of the calyculus,
sometimes with indentations on its inner surface
by pressure of spores. By LM the spore ornamentation is formed by warts, papillae or short irregular ridges across its surface. By SEM the spore
ornamentation is similar to that of Trichia crateriformis and consists of bacula that join to form
short ridges with irregular and sinuous morphology, the base appearing to be broadly columnar,
irregular and sometimes attached to the reticulate
apex and then taking a stellate appearance. The
elaters of capillitium have smooth, tightly packed
spirals. This type of spore ornamentation has also
been observed by SEM HATANO 1986 (plate 21)
and NEUBERT & al. (1993: 198).
Trichia fernbankensis Frederick, R. Simons &
I.L. Roth, Trans. Brit. Mycol. Soc. 83(2): 369
(1984) Figs. 34-43
Specimens examined: USA. Atlanta, Georgia,
Fernbank Science Center, DeKalb Country, mixed
hardwood forest, on rotting wood, 22.11.1976,
leg. R. Simons, det. M.L. Farr as Trichia crateriformis, BPI 71875 (835155) Holotype. Idem
(2 sporocarps metallic by SEM) BPI 835157 Isotype.
Observations: The type material is well preserved. On the remains of wood are two groups
of sporocarps, one with 11 sporocarps and the
other with 4 sporocarps. Sporocarps stalked,
1.5-2 mm total height (Fig. 36). Sporotheca approximately 1 mm in diam., brown to brownish
yellow in colour, with apical dehiscence more
or less circumcissile to irregular but not always
clearly circumsissile as indicated FREDERICK
& al. (1984). Operculum membranous, similar
to the structure of the calyculus, sometimes with
indentations on its inner surface caused by pressure of spores (Fig. 36). Stipe cylindrical, 1-1.5
mm of height, strongly ridged, dark brown (Fig.
36). Hypothallus dark-brown, thin, common to
groups of sporocarps. Capillitium abundant, 4-5
µm diam., formed by free elaters, pale yellow,
smooth spirals and long-tapered ends (Figs. 3738). Spores 8-10(-12) µm diam., globose, pale
yellow in LM, formed by a complete reticulum
92
of small meshes per hemisphere and with warts
and crests in the remaining surface (Figs. 3943). SEM studies conirm the existence of this
contrasting type of spore ornamentation between
hemispheres (simple reticulate and cristate).
DISCUSSION
PERSOON (1795), in his description of the
macroscopic characters of Arcyria decipiens, hesitated to assign this taxon to the genus Trichia,
possibly because the elaters present in Trichia
were not recognized as a character to differentiate the genus from the capillitium of Arcyria.
The following year, PERSOON (1796) described
Trichia fallax having doubts regarding the similarity of this species with Arcyria. He also used
the same iconography in both publications to describe the two species (A. decipiens and T. fallax). At this time the characters to delimit Trichia
and Arcyria were very subjective and undeined.
LISTER (1894), in the irst edition of his book,
synonymised Arcyria decipiens with Trichia fallax, using the latter epithet to refer to this species
although it had priority over A. decipiens.
MACBRIDE (1899) later prioritized Trichia
decipiens over T. fallax by the relevant combination and, for this species, described the spores as
“delicatelly minutely reticulate, 10-12 μm” and
stating that “this is, of course, our familar T. fallax of all authors from Persoon down”. But this
author failed to mention the type of dehiscence
of the peridium in his description (either irregular
or preformed by a line).
From this date onwards, most authors such as
LISTER (1925), MARTIN & ALEXOPOULOS
(1969), HAGELSTEIN (1944) and NANNENGA-BREMEKAMP (1991) used this combination.
Trichia decipiens s. lato is a common taxon
with cosmopolitan distribution (MARTIN &
ALEXOPOULOS, 1969) and fruiting mainly on
woody debris. Within this concept we can include variations that are manifested in the morphology and colour of the sporotheca, peridium
type, size, capillitium morphology and spore
ornamentation. Hence some authors such as
NANNENGA-BREMEKAMP (1991), LADO &
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A COMPARATIVE STUDY OF THE TYPES OF THREE SPECIES OF MYXOMYCETES: TRICHIA CRATERIFORMIS,
T. FALLAX VAR. OLIVACEA AND T. FERNBANKENSIS
Figs. 34-43 Trichia fernbankensis type. 34. Label on the box in which the holotype was conserved. 35. Label on the box in which the
isotype was conserved. 36. Sporocarps. 37. Smooth spirals of the elater. 39. Ending of the elater. 39-40. Spores as viewed by SEM formed
by a complete reticulum of small meshes in a hemisphere. 41. Spores as viewed by SEM formed by a complete reticulum of small meshes
in one hemisphere and with warts and crests on the remaining surface. 42-43. Spores as viewed by SEM formed by short crests.
Scale bars: 36 = 1 mm, 37 = 5 µm, 38 = 10 µm, 39-43 = 2 µm.
Bol. Soc. Micol. Madrid 37. 2013
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G. MORENO & A. CASTILLO
PANDO (1997) and ING (1999) have recognized
at least two varieties T. decipiens var. decipiens
and T. decipiens var. olivacea, while others such
as MARTIN & ALEXOPOULOS (1969) and
FARR (1976) regarded them as a single species
with variable morphology. Tables I and II show
the differences indicated in the descriptions of
previous authors.
Introducir tabla 1 y tabla 2
If one looks closely at the differences in the
two tables, it can be seen that regarding the morphology of the sporotheca the authors agree to
separate T. decipiens var. olivacea by its morphology being almost spherical, subglobose to
globose, while T. decipiens var. decipiens has a
turbinate morphology, ovate to pyriform. However, LADO & PANDO (1997) also describe the
latter as being subglobose. Regarding the dehiscence of the peridium, that of T. decipiens var.
olivacea is clearly circumcissile and leaving a
calyculus, while in T. decipiens var. decipiens it
is irregular but also leaves a calyculus. Regarding the colour of the sporotheca of T. decipiens
var. olivacea, authors have included the olive-coloured, less yellowish or ochraceous appearance
which is unlike that of T. decipiens var. decipiens.
Regarding the capillitium, those authors who cite
the var. olivacea, do not provide measurements or
data to indicate differences. In T. decipiens var.
decipiens there is a large variation in the width
of capillitium according to the authors listed,
varying from 4.5-5.5 μm diam. (LISTER, 1925;
NANNENGA-BREMEKAMP, 1991) to 6-7 μm
in diam. (LADO & PANDO, 1997). The spores
do show differences in size but, nevertheless in T.
decipiens var. olivacea, the spore ornamentation
is warted while in T. decipiens var. decipiens it
is delicately reticulate over part of the surface.
Some authors have mentioned the two varieties as appearing to represent forms rather than important varieties (LISTER, 1925) or to be of little
taxonomic value (LADO & PANDO, 1997).
In our view the main differences used to separate the two taxa are the smaller size of the sporocarps of T. decipiens var. olivacea, dehiscence
of the peridium being clearly circumcissile and
remaining as a calyculus with a regular margin
and having non-reticulate spore ornamentation.
94
On the other hand, T. decipiens var. decipiens is
characterized by its larger sporocarps, irregular
dehiscence and persisting in the lower part as a
cup or funnel with an irregular edge. Also the
spore ornamentation is very different with half
of the spore surface having a delicate complete
reticulum and the other with a crested ornamentation similar to that of T. decipiens var. olivacea.
Thus we observed spores with different spore
ornamentation, a reticulate ornamentation and
another completely crested, depending on the
spore orientation. This feature of T. decipiens
var. decipiens spores is difficult to see by LM
and is the character that has probably confused
those who have not recognised the differences
or have simply treated the variations as forms or
varieties.
KOWALSKI (1975) reviewed material described by Meylan as Trichia fallax var. olivacea
and created a lectotype. His observations agree
with those of Meylan and he came to the conclusion that “T. fallax var. olivacea is a distinct taxon
and should be recognized because T. decipiens
var. decipiens is much yellower in color, longer
stalked, and it has a peridial apex that tears irregularly, leaving at most a highly roughened opening”. While for T. fallax var. olivacea he stated
that “by being olivaceous instead of tawny yellow, smaller in size, and by having a dull, evanescent, apical cap to the sporangium that left a
perfectly smooth, round opening upon disappearance”. However, neither Meylan nor Kowalski
mention differences in spore ornamentation.
HATANO (1986) and NEUBERT & al. (1993)
presented SEM images of the spore of Trichia
decipiens var. decipiens and T. decipiens var.
olivacea but did not show the different spore
ornamentation present in each hemisphere of T.
decipiens var. decipiens.
Trichia decipiens may be confused with Hemitrichia clavata (Pers.) Rostaf. by the shape of its
sporocarps, hemitrichioid capillitium consisting
of a few, very long unbranched elaters with almost no free ends and different spore ornamentation.
FREDERICK & al. 1983, stated that “the
operculum of Trichia crateriformis consists of a
single layer of spore-like bodies that have the cell
Bol. Soc. Micol. Madrid 37. 2013
A COMPARATIVE STUDY OF THE TYPES OF THREE SPECIES OF MYXOMYCETES: TRICHIA CRATERIFORMIS,
T. FALLAX VAR. OLIVACEA AND T. FERNBANKENSIS
Table I.- Main features of Trichia decipiens var. decipiens according to different authors.
* Only describe T. decipiens s. lato
T. decipiens
var. decipiens
Morphology
sporotheca
Peridium
Colour
sporotheca
Macbride (1899)
Turbinate
No mention to peridial
features or dehiscence
Shining olive or
olivaceous brown
Lister (1925)
Turbinate
Breaks up leaving a circular
opening
Shining olive or
yellow-brown
Martin &
Alexopoulos
(1969)*
Turbinate
Persisting below as a deep
or sometimes rather shallow
cup. Irregular dehiscence
Shining olive or
olivaceous brown
Farr (1976)*
Turbinate
to pyriform,
occasionally
goblet-shaped
Thicker below, persisting as
a deep to moderately shallow
calyculus. No mention
to dehiscence system of
sporotheca
NannengaBremekamp
(1991)
Ovate or pyriform
Lado & Pando
(1997)
Capillitium
Tapering gradually
to the long, 20-40
µm, slender apices.
4.5-5.5 µm,
gradually tapering
into long slender
points
Spores
Minutely, delicately
reticulate, 10-12 µm
Either minutely
warted or very
closely and often
irregularly reticulated
on one side, 9-12 µm
5-6 µm, slender tips
10-13 µm, bearing a
delicate reticulation
Shining olive to
olivaceous yellow
or brown
5-6 µm, tapering
gradually to the
long, slender tips
10-13 µm, faintly
and delicately
reticulate over part or
most of the surface
Dehiscing irregularly at the
apex leaving a basal cup
Ochraceous to
orchraceous
brown
4.5-5.5 µm,
gradually attenuate
into very long points
10-13 µm, with
short irregular ridges
and an interrupted
reticulum with small
meshes
Subglobose to
pyriform
Usually remaining as a basal
deep funnel- or trumpetshaped calyculus, border
usually rolled backwards
and/or torn. Irregular or
circumscissile dehiscence
Olive brown or
strong yellowish
brown or strong
orange yellow
6-7 µm, free ends
are pointed, 70100µm long
11-13 µm,
faintly reticulate,
fragmented and finemeshed
Ing (1999)
Top-shaped
Dehiscing at about midheight, either by a performed
fissure, leaving a cup below
and a lid above
Shining olive or
yellow- brown
5-6 µm, tapering to
long thin points
10-13 µm, marked
with a delicate
reticulation over
much of the surface,
the rest minutely
warted.
As Trichia
fernbankensis
(Frederick et al.
(1984)
Turbinate
Operculum appearing smooth
mooth
separating irregularly or
circumscissile at a performed
dehiscence line
Honey-coloured
4-5 µm diam. with
2-3 smooth to finely
roughened spirals
and long tapering
ends
8-10 (-12) µm,
distinctly reticulate
with ridges low and
narrow
Bol. Soc. Micol. Madrid 37. 2013
95
G. MORENO & A. CASTILLO
Table II.- Main features of Trichia decipiens var. olivacea according to different authors.
T. decipiens
var. olivacea
Morphology
sporotheca
Peridium
Colour
sporotheca
Lister (1925)
No mention
morphology of
sporotheca
Clean circular opening
No mention
colour of
sporotheca
NannengaBremekamp
(1991)
almost spherical
clearly marked, dull,
regular lid at the top
olive or olivebrown
Lado & Pando subglobose
(1997)
With calyculus and
dehiscence clearly
circumscissile
Olivaceous
Ing (1999)
Circumscissile
dehiscence, as if by a
performed lid
Olive and more or
less yellow
Globose
wall ornamented with warty excresences similar
to those of the spores”, noting that this ornamentation could be unique for the Myxomycetes. In
our opinion, and in view of the SEM photographs
of the operculum published by them and after
examination of type material of T. crateriformis,
we conclude that this unusual structure of the
operculum consists of tightly collapsed spores,
especially as the type material is in poor condition. The technique used for SEM in that paper
did not use the critical point method: Fruiting
body structures of Myxomycetes are not formed
by cells, except for the spores.
Trichia fernbankensis as we see in the igures (34-43) is a synonym of T. decipiens. The
holotype is deposited in BPI with the two numbers 71875 (835155) which must be retained
only by the number BPI 71875 as indicated by
FREDERICK & al. (1984) in their publication.
The sample’s label (ig. 34) reads “Trichia crateriformis”, while it should be “T. fernbankensis
holotype”.
In conclusion, regarding the dehiscence of the
sporotheca, we are able to establish two groups
among the species studied of the genus Trichia:
Group 1 with irregular dehiscence: Trichia
fernbankensis and Trichia decipiens var. decipiens.
Group 2 with regular dehiscence: Trichia crateriformis is well marked by means of a circular
area, T. fallax var. olivacea with apical circum-
96
Capillitium
No mention of
capillitium or
characteristic
Spores
No mention of spore
characteristics
Rather widely
dispersed, irregular,
low warts
Warted, with no trace
of reticulum
cissile dehiscence and T. decipiens var. olivacea
has well-defined circumcissile operculate apical
dehiscence.
Regarding the spore ornamentation, we conclude that there are two distinct types in the genus
Trichia studied. The ornamentation formed by a
reticulum on only half of the spore surface, while
the other hemisphere has crested warts and another type of spore ornamentation consisting only of
crested warts without forming a reticulum. There
is no variation in spore ornamentation between
the two groups.
First group: Trichia decipiens and T. fernbankensis.
Second group: Trichia crateriformis, T. fallax
var. olivacea and T. decipiens var. olivacea.
TAXONOMIC CONCLUSIONS
After detailed studies and analysis of the material we propose the synonymy between T. crateriformis and T. decipiens var. olivacea, and also
between Trichia decipiens and T. fernbankensis.
1.- Trichia decipiens (Pers.) T. Macbr., N. Amer.
Slime-Moulds: 218 (1899)
≡ Arcyria decipiens Pers., Ann. Bot. (Usteri) 15:
35 (1795)
= T. fernbankensis Frederick, R. Simons &
I.L. Roth, Trans. Brit. Mycol. Soc. 83(2): 369
(1984)
Bol. Soc. Micol. Madrid 37. 2013
A COMPARATIVE STUDY OF THE TYPES OF THREE SPECIES OF MYXOMYCETES: TRICHIA CRATERIFORMIS,
T. FALLAX VAR. OLIVACEA AND T. FERNBANKENSIS
2.- Trichia crateriformis G.W. Martin, Mycologia 55 (1):131 (1963)
= T. fallax var. olivacea Meyl., Bull. Soc. Vaud.
Sci. Nat. 44: 300 (1908)
= T. decipiens (Pers.) T. Macbr. var. olivacea
(Meyl.) Meyl., Bull. Soc. Vaud. Sci. Nat. 55: 244
(1924)
= T
. decipiens f. olivacea (Meyl.) Y. Yamam.,
The Myxomycete Biota of Japan (Tokyo): 237
(1998)
ACKNOWLEDGEMENTS
We wish to express our gratitude to Mr. A. Priego
and Mr. J.A. Pérez of the Electron Microscopy
Service of the University of Alcalá de Henares
for their invaluable help with the SEM. We also
thank Luis Monje of the Department of Drawing
and Scientiic Photography at the Alcalá University for his help in the digital preparation of the
photographs and we are grateful to Dr. J. Rejos,
curator of the AH herbarium, for his assistance
with the specimens examined in the present study.
We want to express our gratitude to D.W. Mitchell for sending his herbarium material and for
reviewing the manuscript.
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