Fungal fruiting bodies in the Cullen Formation (Miocene) in Tierra del Fuego, Argentina

Abstract

Abstract. An abundant and diverse fungal assemblage was recovered from the Cullen Formation suggesting humid paleoenvironmental conditions. Thirty one different forms of fungal fruiting bodies, mostly belonging to the Microthyriales (Ascomycota), were recognized. Nineteen were assigned to fossil genera, including 6 known species. The others are presented as twelve different Types, and possibly represent new morphogenera. The genera/species Plochmopeltinites cooksonia Ramanujan and Rao, Microthyriella diporata Rao and Ramanujan, Microthyrites, Phragmothyrites eocaenicus Edwards, P. kiandrensis Selkirk and Perisporiacites are recorded for the first time in Argentina, enlarging their palaeogeographical distribution. A Miocene age is proposed for this formation in coincidence with that suggested for the fungal spore content and other palynological data.

Full text

Introduction
Fruiting bodies, particularly the Microthyriales,
are some of the best known fungal remains through-
out the geological record (Dilcher, 1965; Elsik, 1968).
Assignment to extant genera and species has been
based on morphological features, and the fact that
many have been found “in situ” associated with leaf
surfaces, stems and flowers of higher plants (Elsik,
1992).
In spite of the preliminary nature of the palaeo-
mycological studies carried out in Argentina, some
papers describing fruiting bodies of Microthyriales
suggest their potential as palaeoenviromental mark-
ers (Martínez, 1968; Durango de Cabrera and
Rodríguez de Sarmiento, 1995; Romero and Castro,
1986). In the Cullen Formation, spores and other
fungal remains, together with fruiting bodies, have
been noted in the literature, but these generally lack
morphological descriptions and/or illustrations
(Vergel and Durango de Cabrera, 1988; Durango de
Cabrera and Vergel, 1989; Zamaloa, 1999, 2000; Zet-
ter et al., 1999; García–Massini et al., 1999). Recently,
more detailed information about the Cullen For-
mation mycoflora has been provided by
García–Massini (2001).
In this paper, we report the presence of abundant
and diverse fruiting bodies recovered from the
Cullen Formation. Most of them conform to Micro-
thyriales (Ascomycetes) and others are of unknown
affinity.
The Cullen Formation is part of the Austral Basin
and crops out along the Atlantic coast in the north-
east of Isla Grande de Tierra del Fuego in southern
Argentina (Codignotto and Malumián, 1981). The
outcrops extend from near Punta Catalina to 2 km
south of the Tapera Sur Dell (figure 1).
AMEGHINIANA (Rev. Asoc. Paleontol. Argent.) - 41 (1): 83-90. Buenos Aires, 30-03-2004 ISSN 0002-7014
©Asociación Paleontológica Argentina AMGHB2-0002-7014/04$00.00+.50
1Department of Ecology and Evolutionary Biology, The
University of Kansas, Haworth Hall, 6022, Lawrence, Kansas
66045-2106 EE.UU. mass112@ku.edu
2Departamento de Ecología, Genética y Evolución, Universidad
de Buenos Aires, Intendente Güiraldes 2620, 1428 Buenos Aires,
Argentina. mzamaloa@bg.fcen.uba.ar
3Departamento de Ecología, Genética y Evolución, Universidad
de Buenos Aires, Intendente Güiraldes 2620, 1428 Buenos Aires,
Argentina. Museo Argentino de Ciencias Naturales B. Rivadavia,
A. Gallardo 470, 1405 Buenos Aires, Argentina.
director@macn.gov.ar
Fungal fruiting bodies in the Cullen Formation (Miocene)
in Tierra del Fuego, Argentina
Juan L. GARCÍA-MASSINI1, María del C. ZAMALOA2and Edgardo J. ROMERO3
Key words. Fungal fruiting bodies. Microthyriales. Miocene. Tierra del Fuego. Argentina.
Palabras clave. Cuerpos fructíferos fúngicos. Microthyriales. Mioceno. Tierra del Fuego. Argentina.
Abstract. An abundant and diverse fungal assemblage was recovered from the Cullen Formation sug-
gesting humid paleoenvironmental conditions. Thirty one different forms of fungal fruiting bodies, most-
ly belonging to the Microthyriales (Ascomycota), were recognized. Nineteen were assigned to fossil gen-
era, including 6 known species. The others are presented as twelve different Types, and possibly repre-
sent new morphogenera. The genera/species Plochmopeltinites cooksonia Ramanujan and Rao,
Microthyrielladiporata Rao and Ramanujan, Microthyrites, Phragmothyriteseocaenicus Edwards, P. kiandren-
sisSelkirk and Perisporiacites are recorded for the first time in Argentina, enlarging their palaeogeograph-
ical distribution. A Miocene age is proposed for this formation in coincidence with that suggested for the
fungal spore content and other palynological data.
Resumen. CUERPOS FRUCTÍFEROS FÚNGICOS DE LA FORMACIÓN CULLEN (MIOCENO) EN TIERRA DEL FUEGO,
ARGENTINA.Se estudia una abundante y diversa micoflora de la Formación Cullen que sugiere condiciones
paleoambientales de elevada humedad. Se reconocieron 31 diferentes cuerpos de fructificación, la mayoría
perteneciente a los Microthyriales (Ascomycota). Diecinueve de ellos fueron asignados a géneros fósiles
incluyendo 6 especies conocidas. Los restantes se describen como Tipos, y posiblemente representen
nuevos morfogéneros. Los géneros/especies PlochmopeltinitescooksoniaRamanujan y Rao, Microthyriella di-
porata Rao y Ramanujan, Microthyrites, Phragmothyrites eocaenicus Edwards, P. kiandrensis Selkirk y
Perisporiacites se registran por primera vez en Argentina, ampliando su distribución paleogeográfica. Se
propone una edad miocena para esta formación en coincidencia con la sugerida por el contenido de es-
poras fúngicas y por la evidencia de otros datos palinológicos.

J.L. García-Massini, M.del C. Zamaloa and E.J. Romero84
Materials and methods
Three palynological sections taken from the
Cullen Formation by Zamaloa (1999) have been ex-
amined for the presence of fungal fruiting bodies.
Standard palynological techniques were used to
process the samples (Traverse, 1988). Slides exam-
ined are housed in the Laboratorio de Paleobotánica,
Departamento de Ecología, Genética y Evolución,
Universidad de Buenos Aires, Argentina, as BAFCB
p.m. The coordinates given are those from the micro-
scope Dialux 20 N° 967412 in that department.
For the classification and identification of the fun-
gal material, we have followed Elsik (1992) and
Kalgutkar and Jansonius (2001). Primary diagnostic
characters used include: 1) Presence or absence of ra-
diate symmetry, 2) Presence or absence of an ostiole,
3) Nature of the margin of the ascoma, 4) Character
of the cells composing the ascoma, 5) Other special
features such as porate cells. Descriptions were based
on the terminology of Elsik et al. (1983).
Results
Of the three studied sections from the Cullen
Formation, 23 stratigraphic levels yielded fungal re-
mains. More than 500 different forms were recog-
nized. Those samples that contain the best preserved
and abundant spores also contain a large number of
fruiting bodies. The forms described and/or illus-
trated in this contribution were selected because of
their excellent preservation and/or similarity to mo-
dern fungi. None of the fruiting bodies were found in
association with mycelia, spores, or attached to the
leaf cuticles of any recognizable plant. Some form
genera and species could be assigned to modern taxa
based on information provided in the current litera-
ture (Dilcher, 1965; Romero and Castro, 1986;
Durango de Cabrera and Rodriguez de Sarmiento,
1995; among others). Nineteen different forms of
fruiting bodies could be accommodated within 9
known form genera, including 6 known form species,
12 forms could not be unequivocally assigned to any
known taxa and are described as Types.
Systematic paleontology
Selected fruiting bodies identified in the present
study are listed in Table 1. In addition, for the Types
we have assigned a number only reflecting a sequen-
tial ordering, and these are presented below quali-
fied by a brief description.
Type I (aff. Arnaudiella “andina” Butin and
Peredo 1986)
Figure 3.G
Description. Circular fruiting body formed of a single
layer of elongate–lobulate cells; more than one layer in
its central portion. Cells radially arranged. Circular os-
tiole present in a raised central position, surrounded
by a ring of thick–walled cells. The margin is lobulate.
Overall size is 119 µm, ostiole 9.1 µm in diameter.
Comments. This specimen has most of the morpho-
logical features found in the extant Arnaudiella “andi-
na” (Ascomycota, Microthyriaceae), a common para-
sitic fungus of the Podocarpaceae of the Chilean
Andean region (Butin and Peredo, 1986).
Studied material. BAFCB p.m. 240, 48.8/111.6.
Type II (aff. Arnaudiella “andina”
Butin and Peredo 1986)
Figure 3.H
Description. Polygonal fruiting body, formed by one
layer of elongated cells. Two layers appear to deve-
lop toward the central part of the body. The body has
2 approximately circular slightly raised ostioles.
Around each ostiole the cell wall is markedly thicker
forming circular rings. The cells are approximately
AMEGHINIANA 41 (1), 2004
Figure 1. Location map. B) Austral Basin distribution, C) Cullen
Formation outcrop, Northeast of Tierra del Fuego / Mapa de ubi-
cación. B) Ubicación de la Cuenca Austral, C) Afloramientos de
Formación Cullen, Noreste de Tierra del Fuego.

Fungal fruiting bodies from the Miocene of Argentina 85
radially arranged. The margin is slightly lobulate to
fimbriate. Size 110.5 x 100.5 µm, ostioles 11 and 14
µm.
Comments. The shape and spatial arrangement of
cells in this specimen resembles Type I, only differ-
ing in the number of ostioles.
Studied material. BAFCB p.m. 213, 38.7/104.5.
Type III
Figures 3.I, J
Description. Fruiting body with an irregular to quad-
rangular shape, and formed of one to two layers of
large lobulate, thin-walled, radially arranged cells. The
body shows a large pseudo–ostiole of irregular to near
circular outline. The margin is lobulate. Size 67 x 59
µm, 35 x 29 µm, pseudo–ostiole 14 to 16 µm, 5 to 9 µm.
Comments. The main feature of this type is that the
cells, considered individually, resemble those desig-
nated as “germlings of Microthyriaceae” by Dilcher
(1965).
Studied material. BAFCB p.m. 240, 43.8/101.3,
44.1/102.
Type IV (“Parabrefeldiellites” Elsik 1992)
Figure 3.K
Description. Flattened fruiting body of polygonal
outline with 7 sides. Body formed of one layer of
cells, aparently two layers occur in the central part of
the body. Cells rectangular to quadrangular, with
thick walls, and arranged in parallel rows that deve-
lop in four directions. Margin smooth and entire.
Astomate body. Size 85 x 59 µm.
Comments.This specimen is closely similar to the in-
formal genus “Parabrefeldiellites” Elsik 1992, previ-
ously reported from sediments of Miocene to Recent
age (Elsik, 1992).
Studied material. BAFCB p.m. 193, 40.8/100.
Type V
Figure 3.L
Description. This specimen is represented by ap-
proximately half of a spherical body, and formed by
thin–walled, rectangular to circular cells. Extending
from the convex part of the body are septate process-
es, 3 to 5 cells long. These processes have a slightly
pyramidal shape and blunt apex. Diameter of the
body 40 µm; length of processes 25 to 30 µm cell wall
thickness 1 to 2 µm.
Comments. According to its shape and cell arrange-
ment, this fruiting body looks similar to Appen-
dicisporonites Saxena and Khare 1992, but differs in
the presence of septa in the processes that arise from
the central body. The named genus was originally
described as dispersed spores from the Tertiary of
India (Saxena and Khare, 1992) but later suggested as
a fruiting body either of Microthyriaceae or
Coelomycetes (Kalgutkar and Jansonius, 2001).
Studied material. BAFCB p.m. 240, 30.6/99.
Type VI
Figure 3.M
Description. Circular fruiting body of, at least, two
layers of cells of irregular shape, arranged in approx-
imately radial pattern. Four zones are distinguish-
able constituting triangular sectors of a circle; cells
inside are heavily pigmented. A pseudo–ostiole of ir-
regular shape is in the center of the body. The margin
is lobulate to irregular. Diameter of the body 58.5 µm,
pseudo–ostiole 9 µm.
Studied material. BAFCB p.m. 240, 44/98.
Type VII
Figure 3.N
Description. Circular fruiting body constructed of
one to two (centrally located) layers of cells, of a
roughly polygonal shape. Cell walls located in the
periphery of the body are less thick compared to
those of the more centrally located cells. Astomate
body. The cells are radially arranged. Margin is not
continuous but formed by the wall projections of the
peripheral cells. Diameter of the body 87 µm; thick-
ness of the cell walls 1 to 2 µm.
Studied material. BAFCB p.m. 249, 32.8/112.8.
Type VIII
Figure 3.O
AMEGHINIANA 41 (1), 2004
Micropeltaceae
* Plochmopeltinites cooksonia Ramanujam & Rao 1973 Figure 2.A
Plochmopeltinites masonii Cookson 1947 Figure 2.B-E
* Plochmopeltinites sp. 1 Figure 2.F
* Plochmopeltinites sp. 2 Figure 2.G
* Plochmopeltinites sp. 3 Figure 2.H
* Plochmopeltinites sp. 4 Figure 2.I
Microthyriaceae
Asterothyrites (Cookson) Kalgutkar &
Jansonius 2001 Figure 2.J
Callimothallus persutus Dilcher 1965 Figure . 2. K
* Callimothallus sp. Figure 2.L
* Microthyriella diporataRao & Ramanujam 1976 Figure 2.M
* Microthyrites sp. Figure 2.N
* Phragmothyrites eocaenicus Edwards 1922 Figure 2.O-Q
* Phragmothyrites kiandrensis Selkirk 1975 Figure 3.A
* Phragmothyrites sp. Figure 3.B
Perisporiaceae
* Perisporiacites sp. Figure 3.C
Trichopeltaceae
Trichopeltinites sp. Figure 3.F
Trichothyriaceae
* Trichothyrites sp. 1 Figure 3. D
* Trichothyrites sp. 2 Figure 3.E
Incertae sedis
* Types I to XII Figure 3.G-S
Table 1. List of taxa of fungal fruiting bodies recognized in the
Cullen Formation. Those marked with * are recorded for the first
time in Argentina / Lista de taxones de cuerpos fructíferos reconocidos
en la Formación Cullen. Los indicados con * son citados por primera vez
en Argentina.

J.L. García-Massini, M.del C. Zamaloa and E.J. Romero86
Description. Circular to elongate fruiting body con-
structed of a layer of elongate and lobulate cells; this
feature is evident at the periphery. The cells are radi-
ally arranged and a large ostiole of irregular contour
is distinguished. The margin is lobulate. Diameter of
the body 115 µm, ostiole 20 µm.
Studied material. BAFCB p.m. 240, 26.5/98.8.
Type IX
Figure 3.P
Description. Circular fruiting body, constructed of
semicircular to ovoid cells, arranged in a radial pat-
tern. The cell walls are thin and no ostiole is distin-
guished. The margin appears irregular because the
cells in the periphery are not intact. Diameter of the
body 59 µm.
Studied material. BAFCB p.m. 249, 39.9/102.2.
Type X
Figure 3.Q
Description. Large circular fruiting body having sev-
eral layers of heavily pigmented cells. Cells appear to
be elongated, thick–walled and radially arranged.
There is a slightly raised circular central ostiole. The
margin is irregularly sinuate. Diameter 303.5 µm; os-
tiole 24 µm.
Studied material. BAFCB p.m. 235, 37.3/96.9.
Type XI
Figure 3.R
Description. Fruiting body of irregular to polygonal
shape. Body formed of large lobulate cells radially
arranged with respect to one or two central cells.
Astomate body. From most cells elongate ap-
pendages emerge perpendicularly showing a rather
circular swollen head, which is curved. In some cas-
es, more than one appendix protrudes from the same
cell. The margin of the body is slightly lobulate. Size
35 µm.
Studied material. BAFCB p.m. 240, 38.8/110.6.
Type XII
Figure 3.S
Description. Fruiting body of quadrangular shape,
formed of four central cells and eleven lobulate pe-
ripheral cells. Astomate body, lobulate margin.
Central cells with differentially thickened walls of up
to 2 µm in thickness. Size 33 x 30 µm.
Studied material. BAFCB p.m. 240, 46.6/101.9.
Discussion
It has been suggested that with the origin of the
angiosperms a major diversification within the
Ascomycetes occurred giving rise to epiphytic and
saprophytic groups such as the Microthyriales
(Kalgutkar and Jansonius, 2001). Elsik (1978, 1992)
hypothesized that from the Early Cretaceous on
Microthyriales and Angiosperms coexisted in the
same habitats promoting a continuous enrichment of
the community and creating additional microhabi-
tats occupied by other fungi. As a consequence, it is
not unusual to find a high diversity of fossil fungi as-
sociated to the presence of angiosperms in sediments
spanning the Cretaceous, Paleogene and Neogene.
The Cullen Formation is a representative example of
such situation.
The sediments studied here contain numerous
morphologically diverse fungal fruiting bodies. Based
on the affinities with known extant and fossil forms,
some ideas may be presented regarding previously
suggested palaeoenvironmental conditions at the time
of deposition (Zamaloa, 1999, 2000). Other specimens
found in this formation could not be assigned to ex-
tant taxa or known fossil forms; however, some of
these display characteristic morphological features,
which facilitate their identification and may be useful
as palaeoenvironmental and/or perhaps stratigraphic
markers in future works. Particularly, the specimens
designated as Types I and II result similar to extant
Arnaudiella “andina” (Ascomycota, Microthyriaceae), a
common parasitic fungus of the Chilean Andean re-
gion (Butin and Peredo, 1986).
During the time of deposition of the sediments
comprising the Cullen Formation, it is postulated
that a temperate humid forest, mostly composed of
angiosperms of the Nothofagaceae and gym-
nosperms of the Podocarpaceae and Araucariaceae,
formed the environment where an abundant my-
coflora developed. Among angiosperms, representa-
tives of about 30 different families were recorded.
These include Apiaceae, Asteraceae, Chenopodia-
ceae, Cunoniaceae, Cyperaceae, Gunneraceae, Mal-
vaceae, Menyanthaceae, Myrtaceae, Onagraceae,
Poaceae, Proteaceae, Rosaceae, Rubiaceae, Sapinda-
ceae, Sparganiaceae/Typhaceae and Winteraceae. A
AMEGHINIANA 41 (1), 2004
Figure 2. Fossil fungal fruiting bodies from the Cullen Formation. Graphic scale = 20 µm /Cuerpos fructíferos fúngicos de la Formación
Cullen. Escala gráfica = 20 µm. A, Plochmopeltinites cooksonia Ramanujan & Rao BAFCB p.m. 240: 39.7/103.4; B-E, Plochmopeltinites
masonii Cookson, B, BAFCB p.m. 210: 43.5/109.4; C, BAFCB p.m. 240: 46.7/108.1; D, BAFCB p.m. 240: 37.3/95; E, BAFCB p.m. 252:
31.6/97.2; F, Plochmopeltinites sp. 1 BAFCB p.m. 235: 31.7/93.2; G, Plochmopeltinites sp. 2 BAFCB p.m. 240: 38//103; H,
Plochmopetinites sp. 3 BAFCB p.m. 240: 46.3/103.2; I, Plochmopetinites sp. 4 BAFCB p.m. 210: 29.6/91.7; J, Asterothyrites sp. BAFCB
p.m. 210: 42.6/92.6; K, Callimothallus persutusDilcher BAFCB p.m. 242: 47.1/102.4; L, Callimothallus sp. BAFCB p.m. 240: 48.7/110.7;
M, Microthyriella diporata Rao & Ramanujan BAFCB p.m. 193: 39.5/105.7; N, Microthyritessp. BAFCB p.m. 240: 27/98; O-Q,
Phragmothyrites eocaenicus Edwards, O, BAFCB p.m. 240: 32.4/98.5; P,BAFCB p.m. 240: 35.3/99; Q, BAFCB p.m. 227: 50.5/95.1.

Fungal fruiting bodies from the Miocene of Argentina 87
AMEGHINIANA 41 (1), 2004

J.L. García-Massini, M.del C. Zamaloa and E.J. Romero88
diverse flora of bryophytes and pteridophytes was
also documented (Zamaloa, 2000). It is suggested
that these bioclimatic conditions promoted the large
fungal diversity found in this formation.
Additionally, taphonomic conditions, particularly
the high rate of sedimentation, in part, as a conse-
quence of the periodic overflow of the meandriform
rivers, created an anoxygenic microenvironment that
enabled the excellent preservation of many of the
fungal remains.
Most of the recovered fruiting bodies are mem-
bers of the Ascomycota representing the families
Micropeltaceae (Plochmopeltinites cooksonia, P. maso-
nii, Plochmopeltinites sp. 1, Plochmopeltinites sp. 2,
Plochmopeltinites sp. 3, Plochmopeltinites sp. 4), Micro-
thyriaceae (Asterothyrites sp., Callimothallus persutus,
Callimothallus sp., Microthyriella diporata, Microthyrites
sp., Phragmotyrites eocaenicus, P. kiandrensis, Phrag-
mothyrites sp.), Trichopeltaceae (Trichopeltinites sp.),
Trichothyriaceae (Trichothyrites sp. 1, Trichothyrites
sp. 2), and Perisporiaceae (Perisporiacites sp.), the first
four families belonging to the Microthyriales and the
Perisporiaceae belonging to the Dothidiales (Tiffney
and Barghoorn, 1974; Kalgutkar and Jansonius,
2001).
Dilcher (1965) described fruiting bodies of Calli-
mothallus and Microthyriella from the Eocene of
Tennessee epiphytically associated to cuticles of
Sapindaceae and Rosaceae. These latter angiosperm
families were also part of the flora of the Cullen
Formation and pollen grains with these affinities
were found in stratigraphic levels coincident with
those where fruiting bodies of Callimothallus and
Microthyriella were found, further establishing a pos-
itive correlation between both observations.
Several authors have pointed out the value of
Microthyriaceous fungi as indicators of moist and
humid climates and/or subtropical temperatures
(Dilcher, 1965; Lange, 1976; Kalgutkar and Jansonius,
2001). Cookson (1947) suggested that the
Microthyriaceae are more abundant in regions of
warm temperate climates characterised by a high rel-
ative ambient moisture. Moreover, Elsik (1992) noted
that the single most important external factor re-
sponsible for their distribution was not temperature,
but rather the mean rainfall, and that this feature pro-
vided a positive correlation between their presence
and rainfall values higher than 1000 mm/year. As an
example, Trichothyrites was found in humid zones of
high temperature (Cookson, 1947) and in temperate,
non–tropical and high precipitation regions
(Rosendahl, 1943), showing once more that their
presence was more dependent on the availability of
humid conditions rather than on the temperature.
Finally, the presence of these fungi in conjunction
with spores and pollen belonging to water demand-
ing taxa (Zamaloa, 2000) reinforces the hypothesis of
prevailing humid climate conditions at the time of
deposition of the sediments of the Cullen Formation.
Many of the fruiting bodies described and illus-
trated here (i.e. Plochmopeltinites cooksonia, P. masonii,
Callimothallus persutus, Microthyriella diporata, Phrag-
mothyrites eocenicus, P. kiandrensis, “Parabrefeldiel-
lites”, Appendicisporonites) are similar to those found
in other basins (Cookson, 1947; Dilcher, 1965;
Edwards, 1922; Elsik, 1992; Ramanujam and Rao,
1973; Rao and Ramanujam, 1976; Saxena and Khare,
1992; Selkirk, 1975). Some of these forms are cited for
the first time for Argentina (Table 1) thus enlarging
their palaeogeographical and/or stratigraphic distri-
bution.
The presence of some selected taxa allows us to
hypothesize on the age of the Cullen Formation.
Phragmothyrites kiandrensis (Early Miocene), Plochmo-
peltinites masonii (Oligocene to Miocene), Plochmo-
peltinites cooksonia (Miocene), Microthyriella diporata
(Miocene) (age ranges taken from Kalgutkar and
Jansonius, 2001) and “Parabrefeldiellites” (Miocene to
Recent) (Elsik, 1992) indicate a Miocene age.
Additionally, the presence of fungal spores referred
to Dyadosporites bhardwaji (Varma and Rawat) Kal-
gutkar and Jansonius 2001 (Eocene to Miocene), D.
cannanorensis Ramanujan and Rao 1978 (Miocene),
Pucciniasporites arcotensis Ramanujan and Ramachar
1980 (Miocene) and Frasnacritetrus siwalikus Saxena,
Singh and Rao 1987 (Miocene to Pliocene) (age
ranges taken from Kalgutkar and Jansonius, 2001) in
the Cullen Formation assemblage supports a
Miocene age. This age is coincident with that ob-
tained by analyzing the overall pollen/spore con-
tents (Zamaloa, 2000). According to these results, this
is the first record of Perisporiacites, Callimothallus per-
sutus and Phragmothyrites eocaenicus in sediments
younger than Eocene.
AMEGHINIANA 41 (1), 2004
Figure 3. Fossil fungal fruiting bodies from the Cullen Formation. Graphic scale = 20 µm / Cuerpos fructíferos fúngicos de la Formación
Cullen. Escal gráfica = 20 µm. A, Phragmothyrites kiandrensis Selkirk BAFCB p.m. 240: 37.9/99.1; B, Phragmothyrites sp. BAFCB p.m.
240: 38.1/110.1; C, Perisporiacites sp. BAFCB p.m. 240: 31.2/104.8; D, Trichothyrites sp. 1 BAFCB p.m. 187: 32.1/92.2; E, Trichothyrites
sp. 2 BAFCB p.m. 240: 47.6/101; F, Trichopeltinites sp. BAFCB p.m. 235: 36.7/100.5; G, Type I (aff. Arnaudiella “andina” Butin & Peredo)
BAFCB p.m. 240: 48.8/111.6; H, Type II (aff. Arnaudiella “andina” Butin & Peredo) BAFCB p.m. 213: 38.7/104.5; I-J, Type III, I BAFCB
p.m. 240: 43.8/101.3; J, BAFCB p.m. 240: 44.1/102; K, Type IV (“Parabrefeldiellites” Elsik) BAFCB p.m. 193: 40.8/100; L, Type V BAFCB
p.m. 240: 30.6/99; M, Type VI BAFCB p.m. 240: 44/98; N,Type VII BAFCB p.m. 249: 32.8/112.8; O, Type VIII BAFCB p.m. 240: 26.5/98.8;
P, Type IX BAFCB p.m. 249: 39.8/102.2; Q, Type X BAFCB p.m. 235: 37.3/96.9; R, Type XI BAFCB p.m. 240: 38.8/110.6; S, Type XII
BAFCB p.m. 240: 46.6/101.9.

Fungal fruiting bodies from the Miocene of Argentina 89
AMEGHINIANA 41 (1), 2004

J.L. García-Massini, M.del C. Zamaloa and E.J. Romero90
Conclusions
The Cullen Formation sediments have yielded a
great diversity of fungal fruiting bodies, which are
abundant in several samples. Of the hundreds of
specimens observed, 31 different forms are presented
in this contribution. Of these, 19 were assigned to
known fossil genera, including 6 known fossil
species. The others correspond to 12 different Types
and possibly represent new morphogenera.
The genera and/or species Plochmopeltinites cookso-
nia, Microthyriella diporata, Microthyrites, Phragmothy-
rites eocaenicus, P. kiandrensis and Perisporiacites are
recorded for the first time in Argentina, thus enlar-
ging their palaeogeographical distribution. The gen-
era and/or species Perisporiacites, Callimothallus persu-
tusand Phragmothyrites eocaenicus are recorded for the
first time in sediments younger than Eocene. A posi-
tive correlation appears to exist between the diversity
of fungal fruiting bodies and the diversity of spores
and pollen, especially to that of angiosperm source.
The fungal palaeoflora flourished under rather
humid climate conditions. A Miocene age is pro-
posed for this formation coinciding with that sug-
gested for the fungal spore content, as well as other
palynological data.
Acknowledgements
The authors thank Professor T. N. Taylor for critically reading
the manuscript. One of the authors (JLGM) wishes to acknowledge
the permanent scientific and personal support provided by J.E.
Wright, Universidad de Buenos Aires, and hereby dedicates him
this manuscript as a token of his thankfulness.
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Recibido: 15 de abril de 2002.
Aceptado: 21 de agosto de 2003.
AMEGHINIANA 41 (1), 2004