FERUGLIOCLADACEAE, A NEW CONIFER FAMILY FROM THE PERMIAN OF GONDWANA
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About This Presentation
Comferous cones occurring m connection with leafy twigs are described from the early Permian of the La Rloja and
Chubut provinces of Argentina The female cones are terminal and compact, their bracts having subaxlllary ovules or
ovuhferous complexes d,sposed hehcally on the cone axm_ The cones are o...
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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/223764866
Ferugliocladaceae, a new conifer family from the Permian of
Gondwana
Article in Review of Palaeobotany and Palynology · January 1987
DOI: 10.1016/0034-6667(87)90016-9
CITATIONS
109
READS
353
2 authors, including:
Rubèn Cunéo
Museo Paleontológico Egidio Feruglio
192 PUBLICATIONS 6,397 CITATIONS
SEE PROFILE
All content following this page was uploaded by Rubèn Cunéo on 03 December 2018.
The user has requested enhancement of the downloaded file.
Ferugliocladaceae, a new conifer family from the Permian of
Gondwana
Article in Review of Palaeobotany and Palynology · January 1987
DOI: 10.1016/0034-6667(87)90016-9
CITATIONS
109
READS
353
2 authors, including:
Rubèn Cunéo
Museo Paleontológico Egidio Feruglio
192 PUBLICATIONS 6,397 CITATIONS
SEE PROFILE
All content following this page was uploaded by Rubèn Cunéo on 03 December 2018.
The user has requested enhancement of the downloaded file.
Rewew of Palaeobotany and Palynology, 51 (1987) 3 30 3
Elsewer Scmnce Pubhshers B V, Amsterdam Printed m The Netherlands
FERUGLIOCLADACEAE, A NEW CONIFER FAMILY FROM THE
PERMIAN OF GONDWANA
S ARCHANGELSKY and R. CUNEO
Natmnal Research Council of Argentzna (CONICET) and Palaeobotany Dw~smn, Museo Argent*no de Clene~as
Naturales "B Rwadawa", Av Angel Gallardo 470, Buenos A~res (1405) (Argentina)
(Rewsed manuscript accepted February 26, 1986)
Abstract
Archangelsky, S and Cflneo. R, 1987 Ferughocladaceae, a new conifer family from the Permmn of Gondwana Rev
Palaeobot Palynol, 51 3-30_
Comferous cones occurring m connection with leafy twigs are described from the early Permian of the La Rloja and
Chubut provinces of Argentina The female cones are terminal and compact, their bracts having subaxlllary ovules or
ovuhferous complexes d,sposed hehcally on the cone axm_ The cones are of two kinds_ One, m whmh the ovules have a
bffid apex, is made the basis of Ferughocladus gen nov, with two specms F rmjanum sp nov and F patagomcus
(Ferugho) comb nov The other has ovules or ovuhferous complexes with no specmhzed apical structure,
Ugartecladus gen nov, with a single specms U genoens~s sp nov The seeds of these genera had prevmusly been
referred to Eucerospermum Ferugho and the shoots to Paranocladus Florin Ferughocladus rmjanum has a wholly
cutlmzed nucellus with a long mlcropylar tube, enclosing a granular megaspore membrane and covered by an
integument The male cones of Ferughocladus are terminal and composed of a central axis bearing hehcally disposed
curved mlcrosporophylls The pollen was probably monosaccate, referable to Cannanoropolhs Potom6 and Sah
The phylogenetm lmphcatmns, reproductive bmlogy and palaeoecology of these conifers are discussed and a new
family, the Ferughocladaceae is proposed_ Comparisons with other conifers show slgmficant morphologmal
differences, mainly m the apparent lack of ovuhferous scales from the female cones This suggests that their ancestors
may have had either a simple bract and ovule orgamzatmn or that the ovule-bearing organ is strongly reduced, with
the ovuhferous scales and ovules fused together so strongly to form an ovuhferous complex as to be mdmtmgmshable
In this case these cones would be advanced m relatmn to those of other conifers of eqmvalent stratlgraphlc age from
other regmns
Introduction
Palaeozoic Gondwana comfers are known
mainly as vegetahve shoots such as Paranocla-
dus Florin, Walkomiella Florin and shoots of
BurzadLa Seward et Sahm which differ from
those typical of equatorial Walchiaceae and
~Contrlbutlon to IGCP Project 211 Late Palaeozom of
South Amemca
may well belong to different families (Pant,
1982). Because female cones have only rarely
been reported, however, relationships to com-
fers from elsewhere have remained obscure.
Permian conifers m particular are usually rare
m Gondwana except for some areas of South
America, parhcularly Argentina and Brazil
Several formations m the Paganzo and Patago-
man basins have ymlded abundant remains,
mlhally reported by Feruglio (1934, 1946, 1951)
who recogmzed several species of Paranocla-
Elsewer Scmnce Pubhshers B V, Amsterdam Printed m The Netherlands
FERUGLIOCLADACEAE, A NEW CONIFER FAMILY FROM THE
PERMIAN OF GONDWANA
S ARCHANGELSKY and R. CUNEO
Natmnal Research Council of Argentzna (CONICET) and Palaeobotany Dw~smn, Museo Argent*no de Clene~as
Naturales "B Rwadawa", Av Angel Gallardo 470, Buenos A~res (1405) (Argentina)
(Rewsed manuscript accepted February 26, 1986)
Abstract
Archangelsky, S and Cflneo. R, 1987 Ferughocladaceae, a new conifer family from the Permmn of Gondwana Rev
Palaeobot Palynol, 51 3-30_
Comferous cones occurring m connection with leafy twigs are described from the early Permian of the La Rloja and
Chubut provinces of Argentina The female cones are terminal and compact, their bracts having subaxlllary ovules or
ovuhferous complexes d,sposed hehcally on the cone axm_ The cones are of two kinds_ One, m whmh the ovules have a
bffid apex, is made the basis of Ferughocladus gen nov, with two specms F rmjanum sp nov and F patagomcus
(Ferugho) comb nov The other has ovules or ovuhferous complexes with no specmhzed apical structure,
Ugartecladus gen nov, with a single specms U genoens~s sp nov The seeds of these genera had prevmusly been
referred to Eucerospermum Ferugho and the shoots to Paranocladus Florin Ferughocladus rmjanum has a wholly
cutlmzed nucellus with a long mlcropylar tube, enclosing a granular megaspore membrane and covered by an
integument The male cones of Ferughocladus are terminal and composed of a central axis bearing hehcally disposed
curved mlcrosporophylls The pollen was probably monosaccate, referable to Cannanoropolhs Potom6 and Sah
The phylogenetm lmphcatmns, reproductive bmlogy and palaeoecology of these conifers are discussed and a new
family, the Ferughocladaceae is proposed_ Comparisons with other conifers show slgmficant morphologmal
differences, mainly m the apparent lack of ovuhferous scales from the female cones This suggests that their ancestors
may have had either a simple bract and ovule orgamzatmn or that the ovule-bearing organ is strongly reduced, with
the ovuhferous scales and ovules fused together so strongly to form an ovuhferous complex as to be mdmtmgmshable
In this case these cones would be advanced m relatmn to those of other conifers of eqmvalent stratlgraphlc age from
other regmns
Introduction
Palaeozoic Gondwana comfers are known
mainly as vegetahve shoots such as Paranocla-
dus Florin, Walkomiella Florin and shoots of
BurzadLa Seward et Sahm which differ from
those typical of equatorial Walchiaceae and
~Contrlbutlon to IGCP Project 211 Late Palaeozom of
South Amemca
may well belong to different families (Pant,
1982). Because female cones have only rarely
been reported, however, relationships to com-
fers from elsewhere have remained obscure.
Permian conifers m particular are usually rare
m Gondwana except for some areas of South
America, parhcularly Argentina and Brazil
Several formations m the Paganzo and Patago-
man basins have ymlded abundant remains,
mlhally reported by Feruglio (1934, 1946, 1951)
who recogmzed several species of Paranocla-
dus leafy shoots and also of Eucerospermum
Feruglio (1946), a genus based on isolated seeds
found m association with the shoots. The
object of this paper is to describe additional,
fertile, material discovered recently in Argen-
tina m organic connection to shoots, which
contributes to an understanding of the
"whole" plants and therefore to a better
understanding of aspects of gondwanan conifer
evolutmn and palaeobiology.
Material and methods
Intensive collecting by the present authors
in the formations just referred to shows that
comfers are numermally dominant in several
of the plant beds (Archangelsky, 1981; Cfineo,
1983). Abundant cones were found, both as
isolated specimens and in organic attachment
to shoots In the Arroyo Totoral Formation (La
Rloja Province), cuticles of seeds, leaves, and
pollen are preserved, whilst the Rio Genoa
Formatmn of Chubut Province has yielded
only impressions, although structural details
are available m these which have facilitated
reconstruction of the strobilar organization.
Although this paper concentrates on re-
cently collected material, a revision has also
been undertaken of previous collections: the
material of A. Piatnltzky, F. Ugarte and other
geologists in the palaeobotamca] collections of
the La Plata Natural History Museum (LP Pb),
and the original specimens of E Ferugho and
A. Platnitzky m the Buenos Aires Natural
History Museum (BA Pb). Other material is in
the Palaeobotanical Unit of the Geological
Research Center, CIRGEO, of Buenos Aires
(CIRGEO Pb).
The fossil localities are shown in Fig 1 For
stratlgraphic details and section numbers of
the Patagonian material see Archangelsky
(1981) and C6neo (1983). Other Patagoman
sections are now under study. The stratigraphy
of the La Rloja occurrences is described by
Andrels et al. (1984). The age of the maternal is
early Permian, perhaps latest Carbomferous m
some lower parts of the sections Locally it is
Fig 1 Map of the [ossll locahtms
,-"" "i' (' ~':~"-":"
AMERICA ,~.~
REFERENCES
(~ ARROYO TOTORAL
FORMATION
(~) GENOA GROUP RIO
Feruglio (1946), a genus based on isolated seeds
found m association with the shoots. The
object of this paper is to describe additional,
fertile, material discovered recently in Argen-
tina m organic connection to shoots, which
contributes to an understanding of the
"whole" plants and therefore to a better
understanding of aspects of gondwanan conifer
evolutmn and palaeobiology.
Material and methods
Intensive collecting by the present authors
in the formations just referred to shows that
comfers are numermally dominant in several
of the plant beds (Archangelsky, 1981; Cfineo,
1983). Abundant cones were found, both as
isolated specimens and in organic attachment
to shoots In the Arroyo Totoral Formation (La
Rloja Province), cuticles of seeds, leaves, and
pollen are preserved, whilst the Rio Genoa
Formatmn of Chubut Province has yielded
only impressions, although structural details
are available m these which have facilitated
reconstruction of the strobilar organization.
Although this paper concentrates on re-
cently collected material, a revision has also
been undertaken of previous collections: the
material of A. Piatnltzky, F. Ugarte and other
geologists in the palaeobotamca] collections of
the La Plata Natural History Museum (LP Pb),
and the original specimens of E Ferugho and
A. Platnitzky m the Buenos Aires Natural
History Museum (BA Pb). Other material is in
the Palaeobotanical Unit of the Geological
Research Center, CIRGEO, of Buenos Aires
(CIRGEO Pb).
The fossil localities are shown in Fig 1 For
stratlgraphic details and section numbers of
the Patagonian material see Archangelsky
(1981) and C6neo (1983). Other Patagoman
sections are now under study. The stratigraphy
of the La Rloja occurrences is described by
Andrels et al. (1984). The age of the maternal is
early Permian, perhaps latest Carbomferous m
some lower parts of the sections Locally it is
Fig 1 Map of the [ossll locahtms
,-"" "i' (' ~':~"-":"
AMERICA ,~.~
REFERENCES
(~ ARROYO TOTORAL
FORMATION
(~) GENOA GROUP RIO
referred to the Gangamoptens zone (La Rioja
Prownce) and
Ferughocladus superzone (Chu-
but Province) (Archangelsky and Cfineo, 1984).
The material was studied using standard
palaeobotanical techmques of hght mmroscopy
and maceratmn, including bulk maceratmn.
Systematic
description
Genus:
UgartecIadus gen. nov.
D~agnosis: Shoots branching at least up to
fourth order; branches attached radially and
~rregularly at acute angles, strmght to shghtly
curved, leafless or with hehcally disposed
leaves Leaves all of one kind, with a single
vein, lanceolate, acute Female cones compact,
ovoid, terminal; composed of rumple bracts and
either free ovules or ovuhferous complexes,
attached hehcally to cone axis. Presumed
ovules or ovuhferous complexes subaxillary to
bracts, subclrcular, ovules orthotropous with a
small and rumple nucellar apex. Presumed seeds
platyspermm w~th a single mucronate apex, a
strongly developed central body and with nar-
row wings.
Type species: Ugartecladus genoenszs sp nov_
Derwatmn of name: After F61ix Ugarte, who
contmbuted to the geology of Patagoma m the
area of this study
Comments There is some doubt about whether
the ovules are wholly free or are ovuhferous
complexes composed of ovule and ovuhferous
scale or scales The ovuhferous appendage
consists clearly of an ovule but there is no
ewdence for an independent ovuhferous scale
or ovuhferous branch scales. The ovule is
surrounded by a poorly developed tissue which
develops into a narrow wmg at the seed stage.
This may have a purely integumentary origin
or may represent m part a greatly reduced
ovuhferous scale or scales fused to the ovule,
m which case the term ovuliferous complex is
appropriate. Our reference to "ovules or ovuhf-
erous appendages" reflects this uncertainty
Compartson: The irregular branching pattern,
the lanceolate and acute, single-vetoed leaves
mostly of one kind, attached helically to the
branches and either appressed to the axis or
forming angles up to 50 L, are all features which
closely resemble those of
Paranocladus Florin
(1940). This is a genus for leafy shoots known
from the Late Palaeozmc of Argentina (Feru-
gho, 1951; Legulzamdn, 1972, Archangelsky
and Arrondo, 1973), Brazd (Florin, 1940a;
Mlllan, 1974; Flttlpaldl and Rosler, 1978) and
India (Surange and Lele, 1957). The work of
Florin (1940a) and Flttlpaldl and Rosler (1978)
reveals a wide range of cutmle structure,
probably mdmatmg that the natural affinitms
of
Paranocladus are heterogeneous as for
Brachyphyllum Lmdley et Hutton ex Brongni-
art and
Pagmphyllum Heer, discussed by
Harris (1979, pp 4, 24) This heterogeneity is
reinforced by the occurrence of the two differ-
ent kinds of female cones in attachment
descmbed here
(Ferughocladus, Ugartecladus)
We therefore regard Paranocladus as a form
genus which is approprmte for detached shoots
hke those known for the natural genera
Ferughocladus and Ugartecladus. Shoots of
Bras~locladus Yoshlda from the Permian of
Brazil (Bernardes de Ohvelra and Yoshlda,
1982) are also similar in their external mor-
phology and partly overlap
Paranocladus
Kr(~uselcladus
Yoshlda (1970) from the Per-
mlan of Brazil and Carboniferous of Argentina
(Archangelsky, 1978), like
Buriadta Seward et
Sahnl 1920 (Pant and Nauhyal, 1967; Ber-
nardes de Ohvelra and Yoshida, 1982, Pant,
1982) from the Permian of India and Brazil,
differs m its heteromorphm fohage. Isolated
female organs found m associatmn with
Krau-
selcladus
shoots are composed of loosely ar-
ranged, helically attached bracts with axdlary
ovules or seeds. The seeds are bdaterally
symmetrical and diwded by a "mmropylar
hne" (Yoshlda, 1970). The acutely pointed
micropyle appears to point outwards, suggest-
mg that the ovules were orthotropous Yoshida
(1970) compared these bodms, although
smaller, with
Corda~carpus seeds These female
organs also have some similarity to those of
Burmdm (Pant and Nautiyal, 1967) The seeds
are more like those of
Ugartecladus than
Ferugliocladus (described below)
Prownce) and
Ferughocladus superzone (Chu-
but Province) (Archangelsky and Cfineo, 1984).
The material was studied using standard
palaeobotanical techmques of hght mmroscopy
and maceratmn, including bulk maceratmn.
Systematic
description
Genus:
UgartecIadus gen. nov.
D~agnosis: Shoots branching at least up to
fourth order; branches attached radially and
~rregularly at acute angles, strmght to shghtly
curved, leafless or with hehcally disposed
leaves Leaves all of one kind, with a single
vein, lanceolate, acute Female cones compact,
ovoid, terminal; composed of rumple bracts and
either free ovules or ovuhferous complexes,
attached hehcally to cone axis. Presumed
ovules or ovuhferous complexes subaxillary to
bracts, subclrcular, ovules orthotropous with a
small and rumple nucellar apex. Presumed seeds
platyspermm w~th a single mucronate apex, a
strongly developed central body and with nar-
row wings.
Type species: Ugartecladus genoenszs sp nov_
Derwatmn of name: After F61ix Ugarte, who
contmbuted to the geology of Patagoma m the
area of this study
Comments There is some doubt about whether
the ovules are wholly free or are ovuhferous
complexes composed of ovule and ovuhferous
scale or scales The ovuhferous appendage
consists clearly of an ovule but there is no
ewdence for an independent ovuhferous scale
or ovuhferous branch scales. The ovule is
surrounded by a poorly developed tissue which
develops into a narrow wmg at the seed stage.
This may have a purely integumentary origin
or may represent m part a greatly reduced
ovuhferous scale or scales fused to the ovule,
m which case the term ovuliferous complex is
appropriate. Our reference to "ovules or ovuhf-
erous appendages" reflects this uncertainty
Compartson: The irregular branching pattern,
the lanceolate and acute, single-vetoed leaves
mostly of one kind, attached helically to the
branches and either appressed to the axis or
forming angles up to 50 L, are all features which
closely resemble those of
Paranocladus Florin
(1940). This is a genus for leafy shoots known
from the Late Palaeozmc of Argentina (Feru-
gho, 1951; Legulzamdn, 1972, Archangelsky
and Arrondo, 1973), Brazd (Florin, 1940a;
Mlllan, 1974; Flttlpaldl and Rosler, 1978) and
India (Surange and Lele, 1957). The work of
Florin (1940a) and Flttlpaldl and Rosler (1978)
reveals a wide range of cutmle structure,
probably mdmatmg that the natural affinitms
of
Paranocladus are heterogeneous as for
Brachyphyllum Lmdley et Hutton ex Brongni-
art and
Pagmphyllum Heer, discussed by
Harris (1979, pp 4, 24) This heterogeneity is
reinforced by the occurrence of the two differ-
ent kinds of female cones in attachment
descmbed here
(Ferughocladus, Ugartecladus)
We therefore regard Paranocladus as a form
genus which is approprmte for detached shoots
hke those known for the natural genera
Ferughocladus and Ugartecladus. Shoots of
Bras~locladus Yoshlda from the Permian of
Brazil (Bernardes de Ohvelra and Yoshlda,
1982) are also similar in their external mor-
phology and partly overlap
Paranocladus
Kr(~uselcladus
Yoshlda (1970) from the Per-
mlan of Brazil and Carboniferous of Argentina
(Archangelsky, 1978), like
Buriadta Seward et
Sahnl 1920 (Pant and Nauhyal, 1967; Ber-
nardes de Ohvelra and Yoshida, 1982, Pant,
1982) from the Permian of India and Brazil,
differs m its heteromorphm fohage. Isolated
female organs found m associatmn with
Krau-
selcladus
shoots are composed of loosely ar-
ranged, helically attached bracts with axdlary
ovules or seeds. The seeds are bdaterally
symmetrical and diwded by a "mmropylar
hne" (Yoshlda, 1970). The acutely pointed
micropyle appears to point outwards, suggest-
mg that the ovules were orthotropous Yoshida
(1970) compared these bodms, although
smaller, with
Corda~carpus seeds These female
organs also have some similarity to those of
Burmdm (Pant and Nautiyal, 1967) The seeds
are more like those of
Ugartecladus than
Ferugliocladus (described below)
The leaf morphology of Walkom~ella Florin
(1940b) is similar to that of Ugartecladus and
Paranocladus but differs m cutlcular struc-
ture The female cones (White, 1981) are
terminal (described as "tuft flowers"), com-
posed of compact cone-scales bearing one small
seed per scale No exphclt mention was made of
bracts. From Whlte's figure (1981, pl I, 3) it is
difficult to establish the cone orgamzation, but
from the shape, seed size and evident lack of
bracts it appears to differ from Ugartecladus.
Surange and Smgh (1951, 1953) described
another species, W. ind~ca, having ovules m
orgamc connectmn with a cone axis. These
ovules are oval but are somewhat hke those of
Ugartecladus which are usually more rounded.
Without more knowledge about Walkom~ella
cones, fully effective compamson with Ugarte-
cladus is precluded.
Palaeozom genera of the equatorial belt are
better known and can therefore be compared
more effectively All differ in their more
regular branching pattern of the vegetative
shoots and m hawng more or less distinct
ovuliferous scales, e.g Walch~a Sternberg
(= Lebach~a Florin), Ernest~odendron Florin,
Moyhostrobus Miller et Brown (1973), Pseudo-
voltz~a Florin, Ullmania Goeppert, Glyptolep~s
Florin, as do Triassic genera such as Voltzmp-
sts H. Potonl~ and Voltzla Brongmart (Florin,
1940a, 1951; Clement-Westerhof, 1984, Mapes
and Rothwell, 1984).
Ugartecladus genoensis sp. nov (Plates I and II,
Figs.2 and 3)
Diagnos~s: Branches of first (main) order
leafless, up to 12 mm wide, with transverse
lentmular scars 1-5 mm long placed closely or
more distantly, never as wide as branch width.
Branches of second order up to 17 cm
long × 5 mm wide, third (penultimate) order
7 cm × 3 5 mm, attached at an angle of 35-45 °,
branches of fourth (ultimate) order 3 cm ×
3mm, attached at about 30 c. Branches of
second, third and fourth orders bearing ap-
pressed, curved leaves, 8 mm long × 2 5 mm
wide; length-breadth ratio 4 or 5 1, greatest
width near middle of leaf; base of leaf subrhom-
bic, veto median, vlmble m distal portion.
Female cones up to 4 cm long × 1.2 cm wide,
length-breadth ratio up to 3 5:1, borne on
branches of last two orders Bract scales up to
1 cm long at base of cone, lanceolate, curved
towards cone apex, acute, with subrhombic
base, attached at right angles to the 1 mm wide
cone axis. Presumed ovules sessile, outhne m
plan view subclrcular, up to 5 mm in diameter
Presumed seeds oval to subclrcular, 5mm
long × 3 mm wide, length-breadth ratio 1.5"1
Shght subrectangular prolongation of seed
body protruding at base of seed Central body
elhptmal, 2.3 mm long × 0.9 mm wide, with
apical mmropylar tube culminating in mucro-
nate, cuspldate, apex 0 5 mm long; body sur-
rounded by 1 mm wide wing, covered from base
to apex by a thin film of dehcate tissue
Derivation of name: After the Genoa river,
Chubut Province, Argentina
Holotype: LP Pb 12003.
Paratypes LP Pb 12004a b, 12005a-b, CIRGEO
Pb 540
Type locahty Puesto Alambre, Rio Genoa,
Chubut Province, Argentina.
Type horizon Fine-grained sfltstones of the Rio
Genoa Group, Ferughocladus superzone, Lu-
beckense Stage, early Permian
Material stud~ed: Puesto Alambre section, Rio
Genoa: (SP numbers refer to plantlferous
levels), SPI-NF A, CIRGEO Pb 540: SP II-NF 3,
PLATEI
Ugartecladus genoens~s sp nov
1 Showing branches of different orders and a female cone, CIRGEO Pb 540, scale 1 cm
2 Mare branch with transverse scars, LP Pb 12010, scale 1 cm
3 Detail of leaves showing fine longitudinal striations, CIRGEO PB 526, × 12
4 Holotype, showing branch with terminal female cone, LP Pb 12003, scale 1 cm
(1940b) is similar to that of Ugartecladus and
Paranocladus but differs m cutlcular struc-
ture The female cones (White, 1981) are
terminal (described as "tuft flowers"), com-
posed of compact cone-scales bearing one small
seed per scale No exphclt mention was made of
bracts. From Whlte's figure (1981, pl I, 3) it is
difficult to establish the cone orgamzation, but
from the shape, seed size and evident lack of
bracts it appears to differ from Ugartecladus.
Surange and Smgh (1951, 1953) described
another species, W. ind~ca, having ovules m
orgamc connectmn with a cone axis. These
ovules are oval but are somewhat hke those of
Ugartecladus which are usually more rounded.
Without more knowledge about Walkom~ella
cones, fully effective compamson with Ugarte-
cladus is precluded.
Palaeozom genera of the equatorial belt are
better known and can therefore be compared
more effectively All differ in their more
regular branching pattern of the vegetative
shoots and m hawng more or less distinct
ovuliferous scales, e.g Walch~a Sternberg
(= Lebach~a Florin), Ernest~odendron Florin,
Moyhostrobus Miller et Brown (1973), Pseudo-
voltz~a Florin, Ullmania Goeppert, Glyptolep~s
Florin, as do Triassic genera such as Voltzmp-
sts H. Potonl~ and Voltzla Brongmart (Florin,
1940a, 1951; Clement-Westerhof, 1984, Mapes
and Rothwell, 1984).
Ugartecladus genoensis sp. nov (Plates I and II,
Figs.2 and 3)
Diagnos~s: Branches of first (main) order
leafless, up to 12 mm wide, with transverse
lentmular scars 1-5 mm long placed closely or
more distantly, never as wide as branch width.
Branches of second order up to 17 cm
long × 5 mm wide, third (penultimate) order
7 cm × 3 5 mm, attached at an angle of 35-45 °,
branches of fourth (ultimate) order 3 cm ×
3mm, attached at about 30 c. Branches of
second, third and fourth orders bearing ap-
pressed, curved leaves, 8 mm long × 2 5 mm
wide; length-breadth ratio 4 or 5 1, greatest
width near middle of leaf; base of leaf subrhom-
bic, veto median, vlmble m distal portion.
Female cones up to 4 cm long × 1.2 cm wide,
length-breadth ratio up to 3 5:1, borne on
branches of last two orders Bract scales up to
1 cm long at base of cone, lanceolate, curved
towards cone apex, acute, with subrhombic
base, attached at right angles to the 1 mm wide
cone axis. Presumed ovules sessile, outhne m
plan view subclrcular, up to 5 mm in diameter
Presumed seeds oval to subclrcular, 5mm
long × 3 mm wide, length-breadth ratio 1.5"1
Shght subrectangular prolongation of seed
body protruding at base of seed Central body
elhptmal, 2.3 mm long × 0.9 mm wide, with
apical mmropylar tube culminating in mucro-
nate, cuspldate, apex 0 5 mm long; body sur-
rounded by 1 mm wide wing, covered from base
to apex by a thin film of dehcate tissue
Derivation of name: After the Genoa river,
Chubut Province, Argentina
Holotype: LP Pb 12003.
Paratypes LP Pb 12004a b, 12005a-b, CIRGEO
Pb 540
Type locahty Puesto Alambre, Rio Genoa,
Chubut Province, Argentina.
Type horizon Fine-grained sfltstones of the Rio
Genoa Group, Ferughocladus superzone, Lu-
beckense Stage, early Permian
Material stud~ed: Puesto Alambre section, Rio
Genoa: (SP numbers refer to plantlferous
levels), SPI-NF A, CIRGEO Pb 540: SP II-NF 3,
PLATEI
Ugartecladus genoens~s sp nov
1 Showing branches of different orders and a female cone, CIRGEO Pb 540, scale 1 cm
2 Mare branch with transverse scars, LP Pb 12010, scale 1 cm
3 Detail of leaves showing fine longitudinal striations, CIRGEO PB 526, × 12
4 Holotype, showing branch with terminal female cone, LP Pb 12003, scale 1 cm
PLATEI
- o
/
~' B NB
J " 1 .'-"
It M
A B
B
N
M
A I B
M N
D
I
Fig 2 Ugartecladus genoenszs sp nov A. Reconstructed radial longitudinal and transverse sections of female cone. × 1 5
B Ideahzed homzontal longitudinal section of a seed, ×7 C Ideahzed adaxlal view of a bract and ovule, x 4 D
Ideahzed transverse sectzon of C, along hne
A--B, × 3 E Ideahzed vertmal longitudinal section of bract and ovule,
× 2 B = bract,
M= megaspore, N = nucellus, I = integument, Ch = chalaza, MC= mmropzlar canal, NB = nucellar beak,
W= wing, O = ovule
LP Pb 12006, SP IV-NF B, LP Pb 12005, 12007;
SP IV-NF C, LP Pb 12004, 12008, 12010; SP VI-
NF A, LP Pb 12011, SP VI-NF C, LP Pb 12012-
19, SP VI, between NF E and F, LP Pb 12020,
SP VI-NF F, LP Pb 12021
Estancla La Casilda Section, Rio Genoa
SPI-NF D, CIRGEO Pb 526a-b, 527, 528a-b,
536.
Mulangulfieo, Betancourt area LP Pb 3649.
Description: The main branches, although not
connected to second order branches, are
closely associated on the same blocks (Plate I,
2) In LP Pb 12011 a large round scar appears to
be the base of a second order axis The
branches of more ultimate orders (Plate I, 1)
have an irregular attachment in more than one
plane (CIRGEO Pb 527, 540, LP Pb 12006)The
phyllotaxis of the leaves is estimated at 3/8,
PLATE II
Ugartecladus genoensls sp nov
1 A seed with carbomzed soft external tissue and basal rectangular remnant of chalaza arrowed, CIRGEO Pb 528, × 12
2_ Vertically compressed cone, A = cone axzs, B = bracts, O= subclrcular ovules, P= stalks of ovules, LP Pb 12004, × 4 5
3 Cone compressed sideways with an attached ovule at straight arrow and detached seeds at curved arrow, LP Pb 12005
×25
4 Cone with shghtly curved bracts but no ovules, LP Pb 12020, scale 1 cm
/
~' B NB
J " 1 .'-"
It M
A B
B
N
M
A I B
M N
D
I
Fig 2 Ugartecladus genoenszs sp nov A. Reconstructed radial longitudinal and transverse sections of female cone. × 1 5
B Ideahzed homzontal longitudinal section of a seed, ×7 C Ideahzed adaxlal view of a bract and ovule, x 4 D
Ideahzed transverse sectzon of C, along hne
A--B, × 3 E Ideahzed vertmal longitudinal section of bract and ovule,
× 2 B = bract,
M= megaspore, N = nucellus, I = integument, Ch = chalaza, MC= mmropzlar canal, NB = nucellar beak,
W= wing, O = ovule
LP Pb 12006, SP IV-NF B, LP Pb 12005, 12007;
SP IV-NF C, LP Pb 12004, 12008, 12010; SP VI-
NF A, LP Pb 12011, SP VI-NF C, LP Pb 12012-
19, SP VI, between NF E and F, LP Pb 12020,
SP VI-NF F, LP Pb 12021
Estancla La Casilda Section, Rio Genoa
SPI-NF D, CIRGEO Pb 526a-b, 527, 528a-b,
536.
Mulangulfieo, Betancourt area LP Pb 3649.
Description: The main branches, although not
connected to second order branches, are
closely associated on the same blocks (Plate I,
2) In LP Pb 12011 a large round scar appears to
be the base of a second order axis The
branches of more ultimate orders (Plate I, 1)
have an irregular attachment in more than one
plane (CIRGEO Pb 527, 540, LP Pb 12006)The
phyllotaxis of the leaves is estimated at 3/8,
PLATE II
Ugartecladus genoensls sp nov
1 A seed with carbomzed soft external tissue and basal rectangular remnant of chalaza arrowed, CIRGEO Pb 528, × 12
2_ Vertically compressed cone, A = cone axzs, B = bracts, O= subclrcular ovules, P= stalks of ovules, LP Pb 12004, × 4 5
3 Cone compressed sideways with an attached ovule at straight arrow and detached seeds at curved arrow, LP Pb 12005
×25
4 Cone with shghtly curved bracts but no ovules, LP Pb 12020, scale 1 cm
PLATE II
3
2
4
3
2
4
L0
Fig 3
Ugartecladus genoens~s sp nov Reconstruction of branching system with female cones and vegetative buds, × 0 75
based on CIRGEO Pb 526 The leaf bases,
which are clearly vmlble in some specimens
(LP Pb 12008, 12017), are not decurrent and are
rhombm to typically subrhombm m section.
The mldvein of the leaf is only rarely visible,
usually m the apmal region. Free longitudinal
and parallel stmatlons converge towards the
leaf apices (Plate I, 3). The apical leaves of the
ultimate branches often sheath the bases of the
cones (LP Pb 12003, 12018, Plate I, 4), the size
and angle of attachment of the leaves gradually
increasing towards the cone base (ranging from
4 5mm to 8-9mm long), and the leaves
appearing to change gradually into bracts.
Most of the 15 female cones studmd are
orgamcally connected to branches of ultimate
order (LP Pb 12012, 12015), but isolated cones
have also been found (CIRGEO Pb 536).
The cone bracts are usually attached at 90 °
to the axis and commonly have a depression
near the base where the ovules are placed (LP
Pb 12003). Distally, the bracts may be slightly
curved towards the cone apex (Plate II, 4), or
may be perpendmular to the cone axis m
presumably more mature stages At the cone
apices of LP Pb 12003, 12005, 12009, 12018 and
12020 the bracts project distally (Plates I, 4 and
II, 3). Dispersed bracts (LP Pb 12007) are
typically lanceolate and wider at the base and
middle Their scars of attachment on the cone
axis are rhombic (LP Pb 12020) In some of the
bracts a midvein is vlmble (LP Pb 12009) and
fine parallel stmaUons converge towards the
apex, with minute pits probably representing
stomata. There are up to six bracts per cm m a
parastlchy (LP Pb 12018), the largest usually
occurring at the cone base
The ovules are usually subclrcular m adax-
Fig 3
Ugartecladus genoens~s sp nov Reconstruction of branching system with female cones and vegetative buds, × 0 75
based on CIRGEO Pb 526 The leaf bases,
which are clearly vmlble in some specimens
(LP Pb 12008, 12017), are not decurrent and are
rhombm to typically subrhombm m section.
The mldvein of the leaf is only rarely visible,
usually m the apmal region. Free longitudinal
and parallel stmatlons converge towards the
leaf apices (Plate I, 3). The apical leaves of the
ultimate branches often sheath the bases of the
cones (LP Pb 12003, 12018, Plate I, 4), the size
and angle of attachment of the leaves gradually
increasing towards the cone base (ranging from
4 5mm to 8-9mm long), and the leaves
appearing to change gradually into bracts.
Most of the 15 female cones studmd are
orgamcally connected to branches of ultimate
order (LP Pb 12012, 12015), but isolated cones
have also been found (CIRGEO Pb 536).
The cone bracts are usually attached at 90 °
to the axis and commonly have a depression
near the base where the ovules are placed (LP
Pb 12003). Distally, the bracts may be slightly
curved towards the cone apex (Plate II, 4), or
may be perpendmular to the cone axis m
presumably more mature stages At the cone
apices of LP Pb 12003, 12005, 12009, 12018 and
12020 the bracts project distally (Plates I, 4 and
II, 3). Dispersed bracts (LP Pb 12007) are
typically lanceolate and wider at the base and
middle Their scars of attachment on the cone
axis are rhombic (LP Pb 12020) In some of the
bracts a midvein is vlmble (LP Pb 12009) and
fine parallel stmaUons converge towards the
apex, with minute pits probably representing
stomata. There are up to six bracts per cm m a
parastlchy (LP Pb 12018), the largest usually
occurring at the cone base
The ovules are usually subclrcular m adax-
ial view (Plate II, 2), and are free from the
bracts Oval shaped ovules (LP Pb 12003)
probably result from sideways compression
Nucellar outlines and narrow wings (?integu-
ment) have been seen in LP Pb 12005 and ovule
stalks m LP Pb 12004 (Plate II, 2: P).
The presumed seeds, found dispersed in the
same slabs, (CIRGEO Pb 528a-b) are only
slightly bigger than the ovules. A delicate thin
film of soft tissue surrounding them is seen in
some specimens, as well as a rectangular basal
region presumed to represent the chalaza
(Plate II, 1)
A reconstruction of U genoensis branches
and female cones and seed is dlustrated m
Figs 2 and 3
Comparison. Elatocladus halle~ Ferugho (1934,
pl.II, figs 1, 4; pl III, figs 1, 8) from plant-rich
horizon '<II" at the Betancourt locahty, was
later recomblned as Paranocladus~ hallei
(Ferugho) Ferugho (1951, p.16, pl.III, fig5)
based mainly on new stratlgraphic evidence
which showed its age to be Permian rather
than 3urasslc This specms, known only from
vegetative remains, is very similar to ours but
the leaves are usually bigger (7.2-15mm
long × 2.2 4 2 mm wide) The material was not
numbered m these publicatmns; nothing was
stated about Its repository, and we have not
been able to find the illustrated specimens m
Ferugho's collections in the Buenos Aires and
La Plata museums. Full comparison with
U_ genoens~s is therefore precluded.
Assocmted with P.~ (Elatocladus) hallez,
Ferugho described fertile scales which he
attributed to it (1934, pl II, figs.3 7), although
later he estabhshed a new genus for this
material, Eucerospermum (Ferugho, 1946) The
species attributed to his P? hallez was E.
patagontcum (Ferugllo, 1946, p.213, pl.II,
figs.3 7, pl III, figs 1 7). As for the vegetative
material, these scales cannot be located.
Although the leaves of Paranocladus~ hallel
are often larger than those of Ugartecladus
genoensts, those of U genoenszs (8mm
long × 2 5 mm wMe) fall within the total range
of Its leaf size given above In wew of the size
differences we choose at present to distinguish
11
the new species from hallei, In a mmflar way to
Harris (1979, p.73) for distmgmshing Elattdes
thomasn Harris from Elatides wilhamsonzz
(Lmdley et Hutton) Nathorst Most probably
Feruglio's sterile material represents a specms
of Ugartecladus and is close to U. genoensts.
Ferugliocladus gen. nov.
Diagnos~s Shoots branching up to fifth order,
attached radially and irregularly at acute
angles, straight to slightly curved, leafless or
with helically disposed leaves. Leaves all of
one kind, lanceolate, acute, with a single
vein Female cones compact, ovoid to sub-
rounded, terminal; composed of simple bracts
and free ovules or ovuhferous complexes (see
p 5) attached hehcally to the cone axis. Pre-
sumed ovules or ovuhferous complexes axil-
lary to subaxfllary, ovules orthotropous with
bifid apex. Presumed seeds platyspermic, with
a bifid apex, a well-marked central body, a
micropylar tube, and wings Male cones borne
terminally, oblong to lanceolate, composed of
central axis and helically disposed mmrosporo-
phylls.
Type species Ferughocladus rio)anum sp. nov.
Derwatmn of name: After Egldlo Ferugllo,
pioneer of Patagonian stratigraphy and palae-
ontology.
Comparison. The vegetative and basra female
cone structure of Ugartecladus is similar The
main differences are in the ovule and seeds,
those of Ferughocladus constantly having a
bifurcate apex as opposed to the simple one
of Ugartecladus The ovules and the seeds of
Ferughocladus are also larger than those
of Ugartecladus, and have more strongly
developed wings.
The comparisons given on p.5 for Ugartecla-
dus also apply to Ferughocladus.
The male cones of F nojanum resemble
those of Krauselcladus canoznhensts Yoshida
(1970), who compared his cones with Walchi-
anthus (=Lebachta) Florin Male cones of
Walkom~ella australis (Fmstmantel) Florin,
described by White (1981) are different from
those found in Ferugliocladus, composed of
bracts Oval shaped ovules (LP Pb 12003)
probably result from sideways compression
Nucellar outlines and narrow wings (?integu-
ment) have been seen in LP Pb 12005 and ovule
stalks m LP Pb 12004 (Plate II, 2: P).
The presumed seeds, found dispersed in the
same slabs, (CIRGEO Pb 528a-b) are only
slightly bigger than the ovules. A delicate thin
film of soft tissue surrounding them is seen in
some specimens, as well as a rectangular basal
region presumed to represent the chalaza
(Plate II, 1)
A reconstruction of U genoensis branches
and female cones and seed is dlustrated m
Figs 2 and 3
Comparison. Elatocladus halle~ Ferugho (1934,
pl.II, figs 1, 4; pl III, figs 1, 8) from plant-rich
horizon '<II" at the Betancourt locahty, was
later recomblned as Paranocladus~ hallei
(Ferugho) Ferugho (1951, p.16, pl.III, fig5)
based mainly on new stratlgraphic evidence
which showed its age to be Permian rather
than 3urasslc This specms, known only from
vegetative remains, is very similar to ours but
the leaves are usually bigger (7.2-15mm
long × 2.2 4 2 mm wide) The material was not
numbered m these publicatmns; nothing was
stated about Its repository, and we have not
been able to find the illustrated specimens m
Ferugho's collections in the Buenos Aires and
La Plata museums. Full comparison with
U_ genoens~s is therefore precluded.
Assocmted with P.~ (Elatocladus) hallez,
Ferugho described fertile scales which he
attributed to it (1934, pl II, figs.3 7), although
later he estabhshed a new genus for this
material, Eucerospermum (Ferugho, 1946) The
species attributed to his P? hallez was E.
patagontcum (Ferugllo, 1946, p.213, pl.II,
figs.3 7, pl III, figs 1 7). As for the vegetative
material, these scales cannot be located.
Although the leaves of Paranocladus~ hallel
are often larger than those of Ugartecladus
genoensts, those of U genoenszs (8mm
long × 2 5 mm wMe) fall within the total range
of Its leaf size given above In wew of the size
differences we choose at present to distinguish
11
the new species from hallei, In a mmflar way to
Harris (1979, p.73) for distmgmshing Elattdes
thomasn Harris from Elatides wilhamsonzz
(Lmdley et Hutton) Nathorst Most probably
Feruglio's sterile material represents a specms
of Ugartecladus and is close to U. genoensts.
Ferugliocladus gen. nov.
Diagnos~s Shoots branching up to fifth order,
attached radially and irregularly at acute
angles, straight to slightly curved, leafless or
with helically disposed leaves. Leaves all of
one kind, lanceolate, acute, with a single
vein Female cones compact, ovoid to sub-
rounded, terminal; composed of simple bracts
and free ovules or ovuhferous complexes (see
p 5) attached hehcally to the cone axis. Pre-
sumed ovules or ovuhferous complexes axil-
lary to subaxfllary, ovules orthotropous with
bifid apex. Presumed seeds platyspermic, with
a bifid apex, a well-marked central body, a
micropylar tube, and wings Male cones borne
terminally, oblong to lanceolate, composed of
central axis and helically disposed mmrosporo-
phylls.
Type species Ferughocladus rio)anum sp. nov.
Derwatmn of name: After Egldlo Ferugllo,
pioneer of Patagonian stratigraphy and palae-
ontology.
Comparison. The vegetative and basra female
cone structure of Ugartecladus is similar The
main differences are in the ovule and seeds,
those of Ferughocladus constantly having a
bifurcate apex as opposed to the simple one
of Ugartecladus The ovules and the seeds of
Ferughocladus are also larger than those
of Ugartecladus, and have more strongly
developed wings.
The comparisons given on p.5 for Ugartecla-
dus also apply to Ferughocladus.
The male cones of F nojanum resemble
those of Krauselcladus canoznhensts Yoshida
(1970), who compared his cones with Walchi-
anthus (=Lebachta) Florin Male cones of
Walkom~ella australis (Fmstmantel) Florin,
described by White (1981) are different from
those found in Ferugliocladus, composed of
12
fertile scales with microsporangla Their re-
semblance is closer to Squamella White, which
is referred to the Glossoptendales
Ferughocladus riojanum sp nov (Plates
III V; see Flg.5.B, 6 B).
Dtagnosls Mare branch (of first order) un-
known, branching dense Branches of second
order more than 12 cm long × 6 mm wide ex-
cluding leaves, showing transverse, lentlcular
to hnear scars extending across at least half of
branch width Third order branches attached
at 40-45 ° at intervals of about 4 cm, more than
13 cm long × 3 mm wide (excluding leaves).
Branches of fourth order up to 8 cm × 2 mm
excluding leaves, attached at 4g' Ultimate
(fifth order) branches occasional, 2 cm × 15 mm
excluding leaves, disposed at very acute
angles. Branches of second to fifth orders
bearing linear, acute, curved leaves, attached
at 5": 1 cm long × 1.5 mm wide at base, length-
breadth ratm 6 8:1. Leaves with entire margin,
lacking trichomes
Female cones ovoid, borne on branches of
fourth or fifth order, largest 2 cm long x 1 cm
wide, length-breadth ratio 1.8-2:1 Cone axis
bearing curved lanceolate bracts up to 7 mm
long; ovules or ovuliferous complexes subcir-
cular, up to 6 mm m dmmeter.
Seeds subclrcular to cuneiform, up to 7 mm
long × 6 mm wide, length-breadth ratio 1 1:1,
with apex 3 mm long Central body subclrcu-
lar to oval, up to 3 mm long x 2 7 mm wide,
with mlcropyle 2 3 mm long reaching apex
Lateral wings up to 1.6 mm wide, surrounded
by delicate tissue up to 0.7 mm wide which
covers the seed evenly from its usually
rounded base to the apex Megaspore wall
3-4pm think, appearing granular in hght
microscope, apex prominent, external thick
cuticle, presumably of integument, showing
isodiametrlc cells.
Male cones borne on ulUmate to penulUmate
branches, the largest 2 cm long × 1 cm wide,
length-breadth ratio 2.1; cone axis up to 1.5 cm
wide. Mmrosporophylls 9mm long, slightly
reflexed, with perpendicular basal stalk, distal
region of sporophyll bent towards cone apex at
almost 90 °, overlapping sporophylls above.
Pollen sacs of uncertain number and position.
Pollen evidently monosaccate
Denvatmn of name: From La Rloja Province,
Argentina.
Holotype. CIRGEO Pb 521a-b.
Paratypes: CIRGEO Pb 541, 548.
Type locahty- Arroyo Totoral, La Rloja Prov-
ince, Argentina.
Type horizon: Free-grained yellow mltstones of
the Arroyo Totoral Formation Gangamopterts
zone, early Permian.
Material studted Arroyo Totoral Section B,
Anzul6n SPI-NF 1, CIRGEO Pb 521-525, LP
Palynologmal collection 2099-2107 (shdes with
ovule cuticles) and 2108-2109 (shdes with leaf
cuticles), SPI-NF 2, CIRGEO Pb 548-555
Description: CIRGEO Pb 523 and 525 have
second order branch fragments which gradu-
ally narrow from base to apex where there is a
greater leaf density. The attachment of third
order branches at 45-50", 1.5 mm wide, is also
clearly visible (Plate III, 1). CIRGEO 524 shows
fourth order branches whmh distally are paral-
lel to the third order ones Vegetative buds
occur on some of the ultimate (fifth) order
branches (CIRGEO Pb 552-555) and may be
mistaken for cones (Plate III, 3) The apical
leaves of fertile branches sheath the bases of
the female cones as for Ugartecladus (CIRGEO
Pb 548, Plate III, 4)
The cuUcle of the leaves is much altered (LP
PLATE III
Ferughocladus rzojanum sp nov, scale 1 cm
1 Second order branch with branches of th,rd and fourth orders, CIRGEO Pb 523
2 Range of different branch orders, CIRGEO Pb 525
3 Vegetat,ve bud, CIRGEO Pb 552
4 Female cone with sheathing leaves of branch at base, CIRGEO Pb 548
fertile scales with microsporangla Their re-
semblance is closer to Squamella White, which
is referred to the Glossoptendales
Ferughocladus riojanum sp nov (Plates
III V; see Flg.5.B, 6 B).
Dtagnosls Mare branch (of first order) un-
known, branching dense Branches of second
order more than 12 cm long × 6 mm wide ex-
cluding leaves, showing transverse, lentlcular
to hnear scars extending across at least half of
branch width Third order branches attached
at 40-45 ° at intervals of about 4 cm, more than
13 cm long × 3 mm wide (excluding leaves).
Branches of fourth order up to 8 cm × 2 mm
excluding leaves, attached at 4g' Ultimate
(fifth order) branches occasional, 2 cm × 15 mm
excluding leaves, disposed at very acute
angles. Branches of second to fifth orders
bearing linear, acute, curved leaves, attached
at 5": 1 cm long × 1.5 mm wide at base, length-
breadth ratm 6 8:1. Leaves with entire margin,
lacking trichomes
Female cones ovoid, borne on branches of
fourth or fifth order, largest 2 cm long x 1 cm
wide, length-breadth ratio 1.8-2:1 Cone axis
bearing curved lanceolate bracts up to 7 mm
long; ovules or ovuliferous complexes subcir-
cular, up to 6 mm m dmmeter.
Seeds subclrcular to cuneiform, up to 7 mm
long × 6 mm wide, length-breadth ratio 1 1:1,
with apex 3 mm long Central body subclrcu-
lar to oval, up to 3 mm long x 2 7 mm wide,
with mlcropyle 2 3 mm long reaching apex
Lateral wings up to 1.6 mm wide, surrounded
by delicate tissue up to 0.7 mm wide which
covers the seed evenly from its usually
rounded base to the apex Megaspore wall
3-4pm think, appearing granular in hght
microscope, apex prominent, external thick
cuticle, presumably of integument, showing
isodiametrlc cells.
Male cones borne on ulUmate to penulUmate
branches, the largest 2 cm long × 1 cm wide,
length-breadth ratio 2.1; cone axis up to 1.5 cm
wide. Mmrosporophylls 9mm long, slightly
reflexed, with perpendicular basal stalk, distal
region of sporophyll bent towards cone apex at
almost 90 °, overlapping sporophylls above.
Pollen sacs of uncertain number and position.
Pollen evidently monosaccate
Denvatmn of name: From La Rloja Province,
Argentina.
Holotype. CIRGEO Pb 521a-b.
Paratypes: CIRGEO Pb 541, 548.
Type locahty- Arroyo Totoral, La Rloja Prov-
ince, Argentina.
Type horizon: Free-grained yellow mltstones of
the Arroyo Totoral Formation Gangamopterts
zone, early Permian.
Material studted Arroyo Totoral Section B,
Anzul6n SPI-NF 1, CIRGEO Pb 521-525, LP
Palynologmal collection 2099-2107 (shdes with
ovule cuticles) and 2108-2109 (shdes with leaf
cuticles), SPI-NF 2, CIRGEO Pb 548-555
Description: CIRGEO Pb 523 and 525 have
second order branch fragments which gradu-
ally narrow from base to apex where there is a
greater leaf density. The attachment of third
order branches at 45-50", 1.5 mm wide, is also
clearly visible (Plate III, 1). CIRGEO 524 shows
fourth order branches whmh distally are paral-
lel to the third order ones Vegetative buds
occur on some of the ultimate (fifth) order
branches (CIRGEO Pb 552-555) and may be
mistaken for cones (Plate III, 3) The apical
leaves of fertile branches sheath the bases of
the female cones as for Ugartecladus (CIRGEO
Pb 548, Plate III, 4)
The cuUcle of the leaves is much altered (LP
PLATE III
Ferughocladus rzojanum sp nov, scale 1 cm
1 Second order branch with branches of th,rd and fourth orders, CIRGEO Pb 523
2 Range of different branch orders, CIRGEO Pb 525
3 Vegetat,ve bud, CIRGEO Pb 552
4 Female cone with sheathing leaves of branch at base, CIRGEO Pb 548
PLATE III
13
1
2
3
4
13
1
2
3
4
14
palynologlcal collection 12108-9) and it was
impossible to separate upper from lower. Cell
outlines are not clear, except near the margins
where elongate cells form several longitudinal
rows. Small elliptical pits which represent
possible stomata are found on both cuticles,
indistinctly ormnted and scattered. The stoma-
tal apparatus appears to be oval, 60 pm long,
with six subsldmry cells (see Flg.5.B). The type
specimen (Plate IV, 1, 2) has dispersed seeds
and female cones as well as cones attached to
shoots. The seeds typically have a bffid apex
(Plate IV, 3-5), which is prominent and api-
cally curved. The base is generally smooth
though a small depressmn sometimes seen
probably corresponds to the original chalazal
region (CIRGEO Pb 549, Plate IV, 3, 4 at base
of seeds). Well preserved specimens have a
dehcate tissue which is better developed api-
cally, underneath the curved extrem~tms, and
apparently covers umformly the rest of the
seed (CIRGEO Pb 522, Plate IV, 5). Seed
cuticles have been obtained both directly from
the specimens and by bulk maceration The
innermost is the megaspore membrane (Plate
V, 1, lower half of 2) which totally occupies the
internal cavity of the nucellus except at its
apex. The nucellar membrane (Plate V, upper
half of 2) is adherent to the megaspore, but
after alkah treatment portions could be de-
tached. It is up to 1 #m thick and has numerous
longitudinal folds whmh may or may not be
original (Plate V, 1). The cell outhnes are only
rarely visible clearly, but are rectangular and
m more or less regular rows. They are about
50 70 pm long x 30-35 ttm wide, with strmght
or shghtly sinuous antichnal walls. A layer of
carbonized tissue covering the nucellar cutmle
dissolves readily with alkali and has numerous
minute perforations 3-5 pm in diameter which
may be original (Plate V, 3). The chalaza shows as
a subrectangular region, 130 × 170 #m (Plate V,
4, 5), including several carbonized thin bundles
without visible structure which spread in vari-
ous directions A possible canal or duct, 15 #m
wide, runs towards the nucellar apex (Plate V, 5 at
arrow, VI, 1 at arrow). Finally, a 9 #m thick
cuticle was recovered, possibly belonging to the
integument (Plate VI, 2, 3) It has lsodlametric-
polygonal, sometimes slightly elongated, cells,
20-30 #m wide, with anticlinal flanges about 6 pm
thick. No stomata were observed.
The male cones (CIRGEO Pb 541a-b, Plate
VI, 4) have sporophylls that markedly overlap
those attached above, nearer to the cone apex.
The fertile branches have well developed apical
leaves that may extend to as much as half the
length of the cone, sometimes more It has been
difficult to estabhsh the position and number of
the pollen sacs, but subcircular bodms up to
I mm In diameter occur on the adaxlal surface
of the sporophylls Pollen grams have not been
observed in the mmropyles However, we have
macerated samples of sediment exclusively
containing the seeds Apart from cuticles,
poorly preserved but identffiable pollen has
been recovered Counts of 200 identifiable
grains showed that 76% were monosaccates,
9.5% strmte blsaccates, 8°/0 spores and 6.5%
bisaccates. The monosaccates are dominated by
Cannanoropolhs Potom~ et Sah (54%) with
Poton~etspor~tes Bharadwaj (20°). Cannanoro-
polhs is the only species m whmh the internal
body is still preserved We suspect, therefore,
that Ferugliocladus riojanum probably pro-
duced pollen of this type.
PLATE IV
Ferughocladus rmTanum sp nov
1 Holotype, showing seed attached to the female cone. later removed for maceration, bffid apex arrowed, CIRGEO Pb 521a
×4
2 Counterpart of holotype showing several seeds m close association with the cone, seed arrowed has prominent blfid apex
and was used for preparation of nucellar cutmle and megaspore membrane, scale 1 cm
3 Holotype Seed showing blfid apex, basal scar, dehcate soft tmsue at arrows, and central oval nucellar body, × 12
4_ Basal scar of seed, central body detached, CIRGEO Pb 549, × 12
5 Seed apex with remnants of soft tmsue proximal to horns, CIRGEO Pb 522, × 12
palynologlcal collection 12108-9) and it was
impossible to separate upper from lower. Cell
outlines are not clear, except near the margins
where elongate cells form several longitudinal
rows. Small elliptical pits which represent
possible stomata are found on both cuticles,
indistinctly ormnted and scattered. The stoma-
tal apparatus appears to be oval, 60 pm long,
with six subsldmry cells (see Flg.5.B). The type
specimen (Plate IV, 1, 2) has dispersed seeds
and female cones as well as cones attached to
shoots. The seeds typically have a bffid apex
(Plate IV, 3-5), which is prominent and api-
cally curved. The base is generally smooth
though a small depressmn sometimes seen
probably corresponds to the original chalazal
region (CIRGEO Pb 549, Plate IV, 3, 4 at base
of seeds). Well preserved specimens have a
dehcate tissue which is better developed api-
cally, underneath the curved extrem~tms, and
apparently covers umformly the rest of the
seed (CIRGEO Pb 522, Plate IV, 5). Seed
cuticles have been obtained both directly from
the specimens and by bulk maceration The
innermost is the megaspore membrane (Plate
V, 1, lower half of 2) which totally occupies the
internal cavity of the nucellus except at its
apex. The nucellar membrane (Plate V, upper
half of 2) is adherent to the megaspore, but
after alkah treatment portions could be de-
tached. It is up to 1 #m thick and has numerous
longitudinal folds whmh may or may not be
original (Plate V, 1). The cell outhnes are only
rarely visible clearly, but are rectangular and
m more or less regular rows. They are about
50 70 pm long x 30-35 ttm wide, with strmght
or shghtly sinuous antichnal walls. A layer of
carbonized tissue covering the nucellar cutmle
dissolves readily with alkali and has numerous
minute perforations 3-5 pm in diameter which
may be original (Plate V, 3). The chalaza shows as
a subrectangular region, 130 × 170 #m (Plate V,
4, 5), including several carbonized thin bundles
without visible structure which spread in vari-
ous directions A possible canal or duct, 15 #m
wide, runs towards the nucellar apex (Plate V, 5 at
arrow, VI, 1 at arrow). Finally, a 9 #m thick
cuticle was recovered, possibly belonging to the
integument (Plate VI, 2, 3) It has lsodlametric-
polygonal, sometimes slightly elongated, cells,
20-30 #m wide, with anticlinal flanges about 6 pm
thick. No stomata were observed.
The male cones (CIRGEO Pb 541a-b, Plate
VI, 4) have sporophylls that markedly overlap
those attached above, nearer to the cone apex.
The fertile branches have well developed apical
leaves that may extend to as much as half the
length of the cone, sometimes more It has been
difficult to estabhsh the position and number of
the pollen sacs, but subcircular bodms up to
I mm In diameter occur on the adaxlal surface
of the sporophylls Pollen grams have not been
observed in the mmropyles However, we have
macerated samples of sediment exclusively
containing the seeds Apart from cuticles,
poorly preserved but identffiable pollen has
been recovered Counts of 200 identifiable
grains showed that 76% were monosaccates,
9.5% strmte blsaccates, 8°/0 spores and 6.5%
bisaccates. The monosaccates are dominated by
Cannanoropolhs Potom~ et Sah (54%) with
Poton~etspor~tes Bharadwaj (20°). Cannanoro-
polhs is the only species m whmh the internal
body is still preserved We suspect, therefore,
that Ferugliocladus riojanum probably pro-
duced pollen of this type.
PLATE IV
Ferughocladus rmTanum sp nov
1 Holotype, showing seed attached to the female cone. later removed for maceration, bffid apex arrowed, CIRGEO Pb 521a
×4
2 Counterpart of holotype showing several seeds m close association with the cone, seed arrowed has prominent blfid apex
and was used for preparation of nucellar cutmle and megaspore membrane, scale 1 cm
3 Holotype Seed showing blfid apex, basal scar, dehcate soft tmsue at arrows, and central oval nucellar body, × 12
4_ Basal scar of seed, central body detached, CIRGEO Pb 549, × 12
5 Seed apex with remnants of soft tmsue proximal to horns, CIRGEO Pb 522, × 12
PLATE IV
15
2
3
5
L-~,I ~
4
15
2
3
5
L-~,I ~
4
16
Comparison The leaves and female cones of this
specms are smaller, the seeds are more circular,
and the nucellar beak longer than m F.
patagonicus (Feruglio) comb. nov. described
below, which lacks sheathing leaves at the base
of the female cone The vegetative regmns are
similar to Paranocladus? halle~ (Feruglio) Feru-
gho (1934, 1951), but F. rm]anum has a much
greater leaf length breadth ratio. The seeds are
very hke Eucerospermum Ferugho but differ
from its three descmbed specms in their size,
their more rounded shape, and the absence of a
medmn longitudinal furrow. They would never-
theless constitute a fourth species of Eucero-
spermum if the isolated seeds were to be named.
Ferughocladus patagonzcus (Ferugho) nov,
comb (Plates VII-X, Figs.4, 5A, 6.A)
Basionym Elatocladus patagonicus Ferugho,
1934, pl I, 6. Specimens from Betancourt.
1934 Elatocladus scales, Ferugho, pl I, 7-9 Female cone
scales, molated
1946 Eucerospermum mtens Ferugho, pl II, 1, 2 Formal
name for [emale cone scales, good lllustratmns
1951 Paranocladus~ patagon~cus (Ferugho) Ferugho New
combmatmn, age shown to be Permmn rather than
Jurassic
1958 Cardmcarpus n~tens (Ferugho) Barbosa, pp 206, 213
Name only, see p 26 for dlscussmn
1972 CordaLcarpus n~tens (Ferugho) Mlllan, p 96, pl XII,
figs 1 7 Pro parte, see p 26
Diagnos~s. Mare (first order) branches leafless,
up to 1.5 cm wide, with transverse hnear scars
up to 1.2 cm long Second order branches more
than 20 cm long × 6-8 mm wide excluding
leaves; third order branches attached at 30 45 ,
1 cm or less apart, up to 17 cm long × 6 mm wide
(excluding leaves). Fourth order branches at-
tached at up to 50 ° , 7cm longx3mm wide
excluding leaves; fifth order branches occa-
sional, attached at acute angle of about 20",
4 cm long × 2 mm wide excluding leaves.
Branches of second to fifth orders bearing
leaves. Leaves up to 1 8 cm long × 3 mm wide,
greatest width near the leaf base, length
breadth ratio 6 1, appressed or diverging at an
angle of up to 30°; when spreading, fastlgiate,
with rhombic base m section: median vein seen
only m distal half of leaf.
Female cones on branches of ultimate or
penultimate order, the largest 4 cm long × 3 cm
wide, length-breadth ratio 1.3:1. Cone axis
bearing lanceolate, acute, almost straight
bracts at a wide angle, up to 1 cm long.
Presumed ovules or ovuliferous complexes
subaxillary, cuneiform, up to 7.5 mm
long × 7 mm wide Presumed seeds trmngular to
cuneiform, up to 8 mm long × 7 mm wide, width
greatest in apical regmn, length-breadth ratio
1.1:1. Apex curved, up to 1.5 mm long Central
body cuneiform, up to 6 mm long × 4 mm wide.
Lateral wings up to 1.6 mm wide, covered by soft
tissue markedly developed beneath the apex,
composed of large cells with their long dlrectmn
parallel to seed margin. Medmn furrow (canal 9)
running from chalaza to base of blfid apex
Male cones borne on branches of ultimate to
penultimate order, largest 2 5 cm long × 1 5 cm
wide. Central axis up to 1.2 mm wide, bearing
mmrosporophylls up to 1 5 cm long × 1.5 mm
wide: sporophylls lanceolate, deflexed proxl-
PLATE V
Ferughocladus rlolanum sp nov
1 Cutmle of nucellar apex and megaspore membrane (dark), LP Palynologmal Collections, shde 2105, × 60
2 Megaspore membrane (dark) and dehcate nucellar cutmle, LP Palynologlcal Collections, shde 2101, × 120
3 Layer of carbomzed tissue whmh covers the nucellus cutmle, showing minute holes, LP Palynologlcal Collections, shde
2099, × 300
4 Cutmle at base of seed shghtly macerated, showing remnants of carbomzed tissue at its base, LP Palynologmal
Collections, shde 2099, × 50_
5 Same specimen after stronger maceration, showing basal plate of chalaza with presumed remnants of strands or bundles
and one shghtly larger duct arrowed, × 120
Comparison The leaves and female cones of this
specms are smaller, the seeds are more circular,
and the nucellar beak longer than m F.
patagonicus (Feruglio) comb. nov. described
below, which lacks sheathing leaves at the base
of the female cone The vegetative regmns are
similar to Paranocladus? halle~ (Feruglio) Feru-
gho (1934, 1951), but F. rm]anum has a much
greater leaf length breadth ratio. The seeds are
very hke Eucerospermum Ferugho but differ
from its three descmbed specms in their size,
their more rounded shape, and the absence of a
medmn longitudinal furrow. They would never-
theless constitute a fourth species of Eucero-
spermum if the isolated seeds were to be named.
Ferughocladus patagonzcus (Ferugho) nov,
comb (Plates VII-X, Figs.4, 5A, 6.A)
Basionym Elatocladus patagonicus Ferugho,
1934, pl I, 6. Specimens from Betancourt.
1934 Elatocladus scales, Ferugho, pl I, 7-9 Female cone
scales, molated
1946 Eucerospermum mtens Ferugho, pl II, 1, 2 Formal
name for [emale cone scales, good lllustratmns
1951 Paranocladus~ patagon~cus (Ferugho) Ferugho New
combmatmn, age shown to be Permmn rather than
Jurassic
1958 Cardmcarpus n~tens (Ferugho) Barbosa, pp 206, 213
Name only, see p 26 for dlscussmn
1972 CordaLcarpus n~tens (Ferugho) Mlllan, p 96, pl XII,
figs 1 7 Pro parte, see p 26
Diagnos~s. Mare (first order) branches leafless,
up to 1.5 cm wide, with transverse hnear scars
up to 1.2 cm long Second order branches more
than 20 cm long × 6-8 mm wide excluding
leaves; third order branches attached at 30 45 ,
1 cm or less apart, up to 17 cm long × 6 mm wide
(excluding leaves). Fourth order branches at-
tached at up to 50 ° , 7cm longx3mm wide
excluding leaves; fifth order branches occa-
sional, attached at acute angle of about 20",
4 cm long × 2 mm wide excluding leaves.
Branches of second to fifth orders bearing
leaves. Leaves up to 1 8 cm long × 3 mm wide,
greatest width near the leaf base, length
breadth ratio 6 1, appressed or diverging at an
angle of up to 30°; when spreading, fastlgiate,
with rhombic base m section: median vein seen
only m distal half of leaf.
Female cones on branches of ultimate or
penultimate order, the largest 4 cm long × 3 cm
wide, length-breadth ratio 1.3:1. Cone axis
bearing lanceolate, acute, almost straight
bracts at a wide angle, up to 1 cm long.
Presumed ovules or ovuliferous complexes
subaxillary, cuneiform, up to 7.5 mm
long × 7 mm wide Presumed seeds trmngular to
cuneiform, up to 8 mm long × 7 mm wide, width
greatest in apical regmn, length-breadth ratio
1.1:1. Apex curved, up to 1.5 mm long Central
body cuneiform, up to 6 mm long × 4 mm wide.
Lateral wings up to 1.6 mm wide, covered by soft
tissue markedly developed beneath the apex,
composed of large cells with their long dlrectmn
parallel to seed margin. Medmn furrow (canal 9)
running from chalaza to base of blfid apex
Male cones borne on branches of ultimate to
penultimate order, largest 2 5 cm long × 1 5 cm
wide. Central axis up to 1.2 mm wide, bearing
mmrosporophylls up to 1 5 cm long × 1.5 mm
wide: sporophylls lanceolate, deflexed proxl-
PLATE V
Ferughocladus rlolanum sp nov
1 Cutmle of nucellar apex and megaspore membrane (dark), LP Palynologmal Collections, shde 2105, × 60
2 Megaspore membrane (dark) and dehcate nucellar cutmle, LP Palynologlcal Collections, shde 2101, × 120
3 Layer of carbomzed tissue whmh covers the nucellus cutmle, showing minute holes, LP Palynologlcal Collections, shde
2099, × 300
4 Cutmle at base of seed shghtly macerated, showing remnants of carbomzed tissue at its base, LP Palynologmal
Collections, shde 2099, × 50_
5 Same specimen after stronger maceration, showing basal plate of chalaza with presumed remnants of strands or bundles
and one shghtly larger duct arrowed, × 120
PLATE V
o.
2
17
3
5
4
o.
2
17
3
5
4
18
A
/4
) B
bl
A--
/?
/ B
/
Fig 4 Ferughocladus patagonLcus (Ferugho) comb nov A. Reconstructmn of radml longitudinal and transverse sectmns of
female cone, x 1 B Ideahzed vertical longitudinal sectmn of bract and ovule, × 4 C Ideahzed tranverse section of D, along
hne A B, × 4 D Ideahzed adaxml wew of a bract and ovule, × 3 B = bract, M = megaspore, N= nucellus, I= integument,
Ch = chalaza, MC= mmropylar canal, NB = nucellar beak, O = ovule, C = canal
mally, overlapping one or more sporophylls
nearer cone apex Pollen sacs of uncertain
number and position
Neotype CIRGEO Pb 513a b.
Other specimens: CIRGEO Pb 462, 514, LP Pb
12002
Type locahty Betancourt, Chubut Province.
Type horizon: Orbiculo~dea level ( = homzon I of
Ferugho, 1934, 1951), Permian.
Materral studied Estancla La Camlda, Rio
Genoa SP III IV, CIRGEO Pb 462,513a b, 514,
529 533, 535a-b, 539, 542 547, 557
Puesto Alambre Section, Rio Genoa: SPIII-
NF D, LP Pb 12002; SPIV-NF B, LP Pb 12007;
SPIV-NF C, LP Pb 12008
Lomas Chatas Section, Rio Genoa: SPI-NF A,
CIRGEO Pb 537a-b, 538
Mulangmfieo, borehole NL 5, NF
Orbiculo~-
dea:
LP Pb 3001, 3009
Aguada de las Mulas, under plant inch
horizon 2 of Ferugho: LP Pb 3618, 8994.
Piedra Shoteh LP Pb 3090, 3125, 3303.
Descriptmn" The main, leafless, branches of the
first order have not been found m orgamc
connection with second order branches (CIR-
GEO Pb 539; Plate VII, 2). However they are so
closely associated that there is little doubt
about their relationship The leaf scars are
rhomboidal with a think margin (CIRGEO Pb
533; Plate VII, 3), their diameter varying
PLATE VI
Ferughocladus rmTanum sp nov
1 Same specimen as in Plate V, 5, showing detail of a duct at arrow and its hollow centre, × 300_
2, 3 Outer cuticle of seed showing modlametmc cells, LP Palynologlcal Collections, shde 2100, × 120 and × 300
4 Male cone attached to a branch, stemle leafy shoot at left, CIRGEO Pb 541, scale 1 cm
A
/4
) B
bl
A--
/?
/ B
/
Fig 4 Ferughocladus patagonLcus (Ferugho) comb nov A. Reconstructmn of radml longitudinal and transverse sectmns of
female cone, x 1 B Ideahzed vertical longitudinal sectmn of bract and ovule, × 4 C Ideahzed tranverse section of D, along
hne A B, × 4 D Ideahzed adaxml wew of a bract and ovule, × 3 B = bract, M = megaspore, N= nucellus, I= integument,
Ch = chalaza, MC= mmropylar canal, NB = nucellar beak, O = ovule, C = canal
mally, overlapping one or more sporophylls
nearer cone apex Pollen sacs of uncertain
number and position
Neotype CIRGEO Pb 513a b.
Other specimens: CIRGEO Pb 462, 514, LP Pb
12002
Type locahty Betancourt, Chubut Province.
Type horizon: Orbiculo~dea level ( = homzon I of
Ferugho, 1934, 1951), Permian.
Materral studied Estancla La Camlda, Rio
Genoa SP III IV, CIRGEO Pb 462,513a b, 514,
529 533, 535a-b, 539, 542 547, 557
Puesto Alambre Section, Rio Genoa: SPIII-
NF D, LP Pb 12002; SPIV-NF B, LP Pb 12007;
SPIV-NF C, LP Pb 12008
Lomas Chatas Section, Rio Genoa: SPI-NF A,
CIRGEO Pb 537a-b, 538
Mulangmfieo, borehole NL 5, NF
Orbiculo~-
dea:
LP Pb 3001, 3009
Aguada de las Mulas, under plant inch
horizon 2 of Ferugho: LP Pb 3618, 8994.
Piedra Shoteh LP Pb 3090, 3125, 3303.
Descriptmn" The main, leafless, branches of the
first order have not been found m orgamc
connection with second order branches (CIR-
GEO Pb 539; Plate VII, 2). However they are so
closely associated that there is little doubt
about their relationship The leaf scars are
rhomboidal with a think margin (CIRGEO Pb
533; Plate VII, 3), their diameter varying
PLATE VI
Ferughocladus rmTanum sp nov
1 Same specimen as in Plate V, 5, showing detail of a duct at arrow and its hollow centre, × 300_
2, 3 Outer cuticle of seed showing modlametmc cells, LP Palynologlcal Collections, shde 2100, × 120 and × 300
4 Male cone attached to a branch, stemle leafy shoot at left, CIRGEO Pb 541, scale 1 cm
19
PLATE VI
1
4 2
PLATE VI
1
4 2
20
EC
1 ~ ."k, % .<--
A
,',*x %
v!7'o :~,
,/
I
Fig 5 A Reconstruction of the branching system of Ferughocladus patagon~cus (Ferugho) comb nov, with female cones,
× 1 B Reconstruction of a stomatal apparatus of
F rmTanum sp nov, × 500 M= stomatal mouth, SC= subsidiary cells,
EC = enmrchng cells
cyclically at intervals of 2 1 2 cm along the
branches, suggesting that differential, probably
seasonal, growth occurred. The phyllotaxls is
estimated as 3/8 Leaves at the tips of the
branches tend to be spreading and are typically
fastlglate (CIRGEO Pb 534, 542, 545-6). Vegeta-
tive buds occur as apmal expansions of some
branches (CIRGEO Pb 462). The leaves nor-
mally show no midvein though fine longitudinal
striations are commonly seen converging
towards the apex The leaves at the female cone
base do not form a sheath, quite unhke
F. rmTanum.
The number of cones studmd, both male and
female, was ten. The female cone bracts in the
neotype show the impressions of posmbly two
cuticles having elongated cells m longitudinal
rows, one (upper?) has no stomata whereas the
other (lower?) has pits which possibly represent
stomata. They are scattered, mdmtinctly ormn-
ted, usually separate but sometimes abutting.
Dispersed seeds and bracts are common
PLATE VII
Ferughocladus patagomcus (Ferugho) comb nov , scale 1 cm
1 Branches of vamous orders and two fragmentary female cones at arrow. CIRGEO Pb 462
2 Leafless branch of first order, CIRGEO Pb 539
3 Rhomboidal leaf base scars, CIRGEO Pb 533
EC
1 ~ ."k, % .<--
A
,',*x %
v!7'o :~,
,/
I
Fig 5 A Reconstruction of the branching system of Ferughocladus patagon~cus (Ferugho) comb nov, with female cones,
× 1 B Reconstruction of a stomatal apparatus of
F rmTanum sp nov, × 500 M= stomatal mouth, SC= subsidiary cells,
EC = enmrchng cells
cyclically at intervals of 2 1 2 cm along the
branches, suggesting that differential, probably
seasonal, growth occurred. The phyllotaxls is
estimated as 3/8 Leaves at the tips of the
branches tend to be spreading and are typically
fastlglate (CIRGEO Pb 534, 542, 545-6). Vegeta-
tive buds occur as apmal expansions of some
branches (CIRGEO Pb 462). The leaves nor-
mally show no midvein though fine longitudinal
striations are commonly seen converging
towards the apex The leaves at the female cone
base do not form a sheath, quite unhke
F. rmTanum.
The number of cones studmd, both male and
female, was ten. The female cone bracts in the
neotype show the impressions of posmbly two
cuticles having elongated cells m longitudinal
rows, one (upper?) has no stomata whereas the
other (lower?) has pits which possibly represent
stomata. They are scattered, mdmtinctly ormn-
ted, usually separate but sometimes abutting.
Dispersed seeds and bracts are common
PLATE VII
Ferughocladus patagomcus (Ferugho) comb nov , scale 1 cm
1 Branches of vamous orders and two fragmentary female cones at arrow. CIRGEO Pb 462
2 Leafless branch of first order, CIRGEO Pb 539
3 Rhomboidal leaf base scars, CIRGEO Pb 533
PLATE VII
'2
'2
22
At
B Ch "
Fig 6 Ideahzed views of ovule/seed m Ferughocladus A. Adaxml vmw of F patagon~cus (Ferugho) comb nov × 6 B
Horizontal long, tudmal sectmn of ovule off
rm.mnum n sp, × 6_ Ch = chalaza, M= megaspore, N = nucellus, I= integument,
PC = pollen chamber, NB = nucellar beak, AE = apical expansion, MF = medmn furrow, ST= soft tissue, SL = spongy layer,
VS = vascular strands, C = duct
(CIRGEO Pb 529-532). In the neotype the
presumed ovules occur in sltu (Plate VIII, 1-4)
and are similar to the seed
Eucerospermum
nitens
Ferugho (1946) The bifid and curved
apices protrude out from the cones (just visible
in Plates VIII, 4 and IX, 5). The bracts are
typmally lanceolate (CIRGEO Pb 532), with a
widened base and a shght concavity near the
axis which marks the position of the ovule.
LP Pb 12002 (Plate IX, 3 5) shows three
female cones, one of them (at right) borne on a
branch of ultimate order. The ovules in this
specimen are somewhat detached from their
original position because of the effects of
compression (Plate IX, 5)
Female cones borne on the ultimate branches
have also been found m other specimens
(CIRGEO Pb 537-8). In these the strobili are
probably mature, with opened out, straight
bracts (Plate IX, 1, 2), and poorly preserved
remnants of presumed seeds (one with typmal
expanded apex)
Seeds similar to those m sltu have been found
closely associated with the cones The median
furrow or channel is usually present and
sometimes widens near the seed apex, forming a
small chamber or cavity (CIRGEO Pb 532, Plate
X, 1, 3). This channel and the chamber are filled
with coal. The original position although
median is not central, but is peripheral and
probably adaxlal, as is revealed by ovules seen
m situ (CIRGEO Pb 513, Plate VIII, 3, 4, and LP
Pb 12002; Plate IX, 5) Feruglio (1946, pl.II, fig.l)
considered the channel to be a crest, located on
both sides of the seed The small upper cavity,
related to the channel, is situated at the top of
the mmropylar tube at the base of the b~fid apex
(Plate X, 1, 3, arrowed at A). CIRGEO Pb 532
PLATE VIII
Ferughocladus patagomcus (Ferugho) comb nov, Neotype, CIRGEO Pb 513
1 Arrows showing ovules m sltu, scale 1 cm
2 Distal region of female cone, arrows showing the same ovules as m 1 but with different hghtmg, × 4
3, 4 Ovule number 1 of 1, 2, illuminated differently m each photo Median furrow (canal ~) and bffid apex enclosing soft tissue
are visible Counterpart, ×6
At
B Ch "
Fig 6 Ideahzed views of ovule/seed m Ferughocladus A. Adaxml vmw of F patagon~cus (Ferugho) comb nov × 6 B
Horizontal long, tudmal sectmn of ovule off
rm.mnum n sp, × 6_ Ch = chalaza, M= megaspore, N = nucellus, I= integument,
PC = pollen chamber, NB = nucellar beak, AE = apical expansion, MF = medmn furrow, ST= soft tissue, SL = spongy layer,
VS = vascular strands, C = duct
(CIRGEO Pb 529-532). In the neotype the
presumed ovules occur in sltu (Plate VIII, 1-4)
and are similar to the seed
Eucerospermum
nitens
Ferugho (1946) The bifid and curved
apices protrude out from the cones (just visible
in Plates VIII, 4 and IX, 5). The bracts are
typmally lanceolate (CIRGEO Pb 532), with a
widened base and a shght concavity near the
axis which marks the position of the ovule.
LP Pb 12002 (Plate IX, 3 5) shows three
female cones, one of them (at right) borne on a
branch of ultimate order. The ovules in this
specimen are somewhat detached from their
original position because of the effects of
compression (Plate IX, 5)
Female cones borne on the ultimate branches
have also been found m other specimens
(CIRGEO Pb 537-8). In these the strobili are
probably mature, with opened out, straight
bracts (Plate IX, 1, 2), and poorly preserved
remnants of presumed seeds (one with typmal
expanded apex)
Seeds similar to those m sltu have been found
closely associated with the cones The median
furrow or channel is usually present and
sometimes widens near the seed apex, forming a
small chamber or cavity (CIRGEO Pb 532, Plate
X, 1, 3). This channel and the chamber are filled
with coal. The original position although
median is not central, but is peripheral and
probably adaxlal, as is revealed by ovules seen
m situ (CIRGEO Pb 513, Plate VIII, 3, 4, and LP
Pb 12002; Plate IX, 5) Feruglio (1946, pl.II, fig.l)
considered the channel to be a crest, located on
both sides of the seed The small upper cavity,
related to the channel, is situated at the top of
the mmropylar tube at the base of the b~fid apex
(Plate X, 1, 3, arrowed at A). CIRGEO Pb 532
PLATE VIII
Ferughocladus patagomcus (Ferugho) comb nov, Neotype, CIRGEO Pb 513
1 Arrows showing ovules m sltu, scale 1 cm
2 Distal region of female cone, arrows showing the same ovules as m 1 but with different hghtmg, × 4
3, 4 Ovule number 1 of 1, 2, illuminated differently m each photo Median furrow (canal ~) and bffid apex enclosing soft tissue
are visible Counterpart, ×6
PLATE VIII
3
4
23
3
4
23
24
shows, beneath the apex, a thick layer of tissue
composed of large cells forming a fleshy
structure, which has a markedly different cell
orientation from the rest. It is wslble clearly in
many specimens but in others may be absent
perhaps as a result of decay or poor preserva-
tion LP Pb 3598 is of interest in showing a
probable seedhng at an early stage of develop-
ment (Plate X, 4).
Some of the male cones (e.g. CIRGEO Pb 514)
show a transitmn between apical branch leaves
and the proximally de flexed basal sporophylls
of the cone The dmposition and number of
pollen sacs is uncertain because of their poor
preservation In some specimens, e.g. CIRGEO
Pb 529, subclrcular bodies 1.5 mm in diameter,
which seem to be abaxial, occupy the space
between the reflexed portmns of the mlcrospo-
rophylls (Plate X, 5).
Compartson and nomenclature Ferughocladus
patagonicus is compared with F. rmjanum sp.
nov. on p 16.
Paranocladus? patagon~cus (Feruglio) Feru-
gho (1934, 1951) has ultimate and penultimate
branches ~dentmal to ours m size, attachment
angle, shape of the leaves and their fastlgiate
character. As discussed on p 11, his illustrated
material cannot be found. Hence, we retain his
specific epithet and designate a neotype for the
species Ferugho also described "scales" associ-
ated with the foliage -shoots which he named
Eucerospermum n~tens (Ferugho, 1934, 1946).
They too are identical m shape, size and the
apex with a medmn channel, with the seeds
found in F. patagonzcus cones. Therefore, we
also include E. nztens in the synonymy of F
patagomcus. Since the name Eucerospermum
patagomcum Feruglio (1946) already exists, for
seeds of larger size, we suggest that dispersed
seeds referable to F patagonicus should be
referred to as '~F. patagonicus seeds", m a
similar manner to the cones of Elat~des Heer
(Harris, 1979). This will avoid difficultms with
homonymy.
Discussion
Notes on the genus Eucerospermum Ferugho
Ferugho (1946) described three specms, E.
n~tens, E. patagon~cum and E oplmum, and
referred them to the comfers (Abletineae),
recognizing that they are elements of female
cones. He interpreted the seeds as having two
apmal horns (euceros. Lat, true horns) and a
basal chalaza, based on comparison with Pma-
ceae Mlllan (1972) found similar seeds m Monte
Mor, Brazil (Itarar~ Subgroup, late Carbonifer-
ous to early Permian), but his interpretation
was different, suggesting that the seed horns
were m fact basal and related to the attachment
to the cone axis The outer structure surround-
ing the seed was interpreted as a sarcotesta by
Ferugho but as a sclerotesta by Mlllan. Mtllan
transferred n~tens and patagonzcum to Corda~-
carpus and combined them as C mtens. Cardm-
carpus nitens was used earlier by Barbosa (1958)
for Brazilian material, though without illustra-
tion of the specimens. Mlllan preferred the use
of Corda~carpus for lmpressions/compressmns,
retaining Cardmcarpus for petrifactions How-
ever, he retained the name Eucerospermum for
the third specms E opimum
The discovery of seeds identical to E. nttens m
the female cones of Ferughocladus patagontcus
shows clearly that the blfid apex points out-
wards from the cone axm and that Ferugho's
mterpretatmn is the correct one
The female cone appendages and their relevance
to conifer phylogeny: The vegetative and repro-
ductlve characters of Ugartecladus and Feru-
PLATE IX
Ferughocladus patagon~cus (Ferugho) comb nov
1, 2 Female cone with remains of elongate bracts (B) and ovules (O) with build apex just visible, CIRGEO Pb 537a, × 2 5 and
×7
3 5 Three female cones, that m the centre of 3 (enlarged m 4, 5), showing several superimposed ovules m distal region of cone,
LPPb 12002 3, scale=lcm:4, ×15,5, ×4
shows, beneath the apex, a thick layer of tissue
composed of large cells forming a fleshy
structure, which has a markedly different cell
orientation from the rest. It is wslble clearly in
many specimens but in others may be absent
perhaps as a result of decay or poor preserva-
tion LP Pb 3598 is of interest in showing a
probable seedhng at an early stage of develop-
ment (Plate X, 4).
Some of the male cones (e.g. CIRGEO Pb 514)
show a transitmn between apical branch leaves
and the proximally de flexed basal sporophylls
of the cone The dmposition and number of
pollen sacs is uncertain because of their poor
preservation In some specimens, e.g. CIRGEO
Pb 529, subclrcular bodies 1.5 mm in diameter,
which seem to be abaxial, occupy the space
between the reflexed portmns of the mlcrospo-
rophylls (Plate X, 5).
Compartson and nomenclature Ferughocladus
patagonicus is compared with F. rmjanum sp.
nov. on p 16.
Paranocladus? patagon~cus (Feruglio) Feru-
gho (1934, 1951) has ultimate and penultimate
branches ~dentmal to ours m size, attachment
angle, shape of the leaves and their fastlgiate
character. As discussed on p 11, his illustrated
material cannot be found. Hence, we retain his
specific epithet and designate a neotype for the
species Ferugho also described "scales" associ-
ated with the foliage -shoots which he named
Eucerospermum n~tens (Ferugho, 1934, 1946).
They too are identical m shape, size and the
apex with a medmn channel, with the seeds
found in F. patagonzcus cones. Therefore, we
also include E. nztens in the synonymy of F
patagomcus. Since the name Eucerospermum
patagomcum Feruglio (1946) already exists, for
seeds of larger size, we suggest that dispersed
seeds referable to F patagonicus should be
referred to as '~F. patagonicus seeds", m a
similar manner to the cones of Elat~des Heer
(Harris, 1979). This will avoid difficultms with
homonymy.
Discussion
Notes on the genus Eucerospermum Ferugho
Ferugho (1946) described three specms, E.
n~tens, E. patagon~cum and E oplmum, and
referred them to the comfers (Abletineae),
recognizing that they are elements of female
cones. He interpreted the seeds as having two
apmal horns (euceros. Lat, true horns) and a
basal chalaza, based on comparison with Pma-
ceae Mlllan (1972) found similar seeds m Monte
Mor, Brazil (Itarar~ Subgroup, late Carbonifer-
ous to early Permian), but his interpretation
was different, suggesting that the seed horns
were m fact basal and related to the attachment
to the cone axis The outer structure surround-
ing the seed was interpreted as a sarcotesta by
Ferugho but as a sclerotesta by Mlllan. Mtllan
transferred n~tens and patagonzcum to Corda~-
carpus and combined them as C mtens. Cardm-
carpus nitens was used earlier by Barbosa (1958)
for Brazilian material, though without illustra-
tion of the specimens. Mlllan preferred the use
of Corda~carpus for lmpressions/compressmns,
retaining Cardmcarpus for petrifactions How-
ever, he retained the name Eucerospermum for
the third specms E opimum
The discovery of seeds identical to E. nttens m
the female cones of Ferughocladus patagontcus
shows clearly that the blfid apex points out-
wards from the cone axm and that Ferugho's
mterpretatmn is the correct one
The female cone appendages and their relevance
to conifer phylogeny: The vegetative and repro-
ductlve characters of Ugartecladus and Feru-
PLATE IX
Ferughocladus patagon~cus (Ferugho) comb nov
1, 2 Female cone with remains of elongate bracts (B) and ovules (O) with build apex just visible, CIRGEO Pb 537a, × 2 5 and
×7
3 5 Three female cones, that m the centre of 3 (enlarged m 4, 5), showing several superimposed ovules m distal region of cone,
LPPb 12002 3, scale=lcm:4, ×15,5, ×4
PLATE IX
1 2
25
3
4
h4
1 2
25
3
4
h4
26
J
~US
Fig 7 Hypothetical evolutmnary steps from ancestor to Ugartecladus and Ferughocladus, mterpretatmn
mtermedmte stage shows a reduction of the dwar[ shoot, slmphficatlon of the bract, and an orthotropous ovule
(x)_ The
ghocladus show almost all the basic require-
ments for them to be regarded as true comfers
However, the female cone appendage is unusual
since there is no evidence for dwarf shoots
bearing both ovules and ovuliferous scales
(whmh are the norm in Palaeozom conifers).
Two possible interpretations were advanced for
this on p.5, here demgnated X and Y
(X) The ovules are entirely fused to an
ovuliferous scale or scales and may therefore be
regarded as ovuliferous complexes (Fig 7)
(Y) The ovules st&nd alone, without an
ovuliferous scale or scales, and the tissue
covering the nucellus is wholly integumentary
(Fig 8)
To accept either interpretation firmly is
difficult since direct evidence is lacking. It is
clear, however, that both
Ugartecladus and
Ferughocladus show no ovuliferous scale or
scales as independent or semi-independent
Ancestor
B
Burladla
~
ar tecladus
erug I,ocladus
Fig 8 Hypothetical common ancestor for Burmdta, Ugar-
tecladus
and Ferughocladus, interpretation (Y) From a
simple lax strobllus with orthotropous ovules, hne A leads
to
Bunadza, with lax cones and anatropous ovules, while
hne B leads to compact cones with stmphfied bracts
PLATE X
Ferughocladus patagonzcus (Ferugho) comb_ nov
1 3 Three seeds with longitudinal median canal (C) and small chamber at A, CIRGEO Pb 532, x 5
4 Probable germinating seed, LP Pb 3598, × 3
5 Male cone with strongly curved mlcrosporophylls, the small circular bodms arrowed at left may represent pollen sacs,
CIRGEO Pb 529, × 4
6 Basal region of a male cone showing curve of mlcrosporophylls, CIRGEO Pb 514, × 7
J
~US
Fig 7 Hypothetical evolutmnary steps from ancestor to Ugartecladus and Ferughocladus, mterpretatmn
mtermedmte stage shows a reduction of the dwar[ shoot, slmphficatlon of the bract, and an orthotropous ovule
(x)_ The
ghocladus show almost all the basic require-
ments for them to be regarded as true comfers
However, the female cone appendage is unusual
since there is no evidence for dwarf shoots
bearing both ovules and ovuliferous scales
(whmh are the norm in Palaeozom conifers).
Two possible interpretations were advanced for
this on p.5, here demgnated X and Y
(X) The ovules are entirely fused to an
ovuliferous scale or scales and may therefore be
regarded as ovuliferous complexes (Fig 7)
(Y) The ovules st&nd alone, without an
ovuliferous scale or scales, and the tissue
covering the nucellus is wholly integumentary
(Fig 8)
To accept either interpretation firmly is
difficult since direct evidence is lacking. It is
clear, however, that both
Ugartecladus and
Ferughocladus show no ovuliferous scale or
scales as independent or semi-independent
Ancestor
B
Burladla
~
ar tecladus
erug I,ocladus
Fig 8 Hypothetical common ancestor for Burmdta, Ugar-
tecladus
and Ferughocladus, interpretation (Y) From a
simple lax strobllus with orthotropous ovules, hne A leads
to
Bunadza, with lax cones and anatropous ovules, while
hne B leads to compact cones with stmphfied bracts
PLATE X
Ferughocladus patagonzcus (Ferugho) comb_ nov
1 3 Three seeds with longitudinal median canal (C) and small chamber at A, CIRGEO Pb 532, x 5
4 Probable germinating seed, LP Pb 3598, × 3
5 Male cone with strongly curved mlcrosporophylls, the small circular bodms arrowed at left may represent pollen sacs,
CIRGEO Pb 529, × 4
6 Basal region of a male cone showing curve of mlcrosporophylls, CIRGEO Pb 514, × 7
PLATE X
2
1 2
4
3 5
6
2
1 2
4
3 5
6
28
units from the megasporophyll and ovule The
specialized blfid apices in Ferughocladus seem
to be of integumentary origin, rather than
modified scales, and the lack of this structure in
Ugartecladus is consistent with this assump-
tion If interpretation X is correct, reduction of
the scale or fusion of ovule and scale in one
complex, must be regarded as total or nearly
total, suggesting a highly advanced stage of
cone appendage evolution (Fig.7) The relative
age of our material is younger, although only
slightly so, than the first known stratigraphlc
occurrence of conifers which have ovuliferous
scales together with the ovules (Hill and Crane,
1982: Hill and Camus, 1986). This is consistent
with the accepted concepts of comferalean
phylogeny which are represented in our inter-
pretatlon (X) (Florin, 1951). In this context,
these genera are of interest in extending the
earliest stratlgraphIc occurrence of highly
specialized (1 e. fused/reduced) female cone-
scales backwards in time from the Trias to the
early Permian, showing that conifer evolution
was more rapid than has been thought, at any
rate in South America
Interpretation (Y), on the other hand (Fig 8)
represents a less orthodox but interesting view
of conifer evolution which would suggest that
the female cone m some conifers or conifer-like
plants did not have a compound origin.
(Whether such plants should then be accepted
as true conifers is an important question). On
this interpretation, Ugartecladus and Feru-
ghocladus would be considered to have simple
strobih (i e of ovules and bracts) such as are
known already to characterize the Permian
Gondwanan genus Buriadta Seward et Sahni
(Pant and Nautiyal, 1967). This is considered to
represent a distinct order within the conifers,
Burladlales, by Pant (1982) It has small,
stalked, anatropous ovules located between
leaves that do not form cones (Pant and
Nautiyal, 1967) Interestingly, the seeds have
blfid apices as in Ferugliocladus It is therefore
not unreasonable to suggest an ancestor com-
mon to both our genera and Buriadia (Fig.8, see
also Archangelsky, 1985). In evolutionary hne
A of Fig 8, the ancestor changes only in ~ts
ovule, which becomes anatropous m Burtadla.
whilst in line B, leading to Ugartecladus and
Ferughocladus the ovules lose their stalks, the
bracts become simple and the whole structure
forms a compact cone. If the Buriadlales are
indeed to be considered true conifers (Krassl-
lov, 1971; Meyen, 1978) they, hke Ugartecladus
and Ferugliocladus, would clearly represent on
this interpretation a parallel hne of evolunon
to that which led to Voltziales
Proposal of a new faintly The distractive
characters of Ugartecladus and Ferughocladus
preclude their inclusion in the most similar
conifer families Walchlaceae, Buriadlaceae
and Araucarlaceae These new genera, further-
more, are restricted m time and space to the
Permian of Gondwanan South America. There-
fore we propose the following definmon for a
new family, the Ferughocladaceae, of which
Ferughocladus is selected as the type genus.
Members of this family are conifers (sensu
lato or sensu stricto), with Irregular radial
branching, bearing terminal compact male and
female cones_ The leaves are small, helically
disposed, all of one kind, lanceolate and single
vetoed. The female cones are composed of a
central axis with bracts and apparently free or
actually free, orthotropous ovules which are
disposed helically The seeds are platyspermlc,
reaching their greatest size while still in the
cones. The male cones are composed of an axis
bearing helically arranged, proximally deflexed
mlcrosporophylls. The pollen is monosaccate.
Other genera Ugartecladus gen nov (male
cones unknown) Eucerospermum Ferugho (dis-
persed seeds) Addmonal genera, at least m
part, are Paranocladus Florin (branches and
leaves with cuticles) and Brastlocladus Yoshida
(branches and leaves which have not yet yielded
cuticles) The status of these genera is probably
similar to that of Brachyphyllum Lmdley et
Hutton ex Brongniart and Pagiophyllum Heer
(Harris, 1979).
Reproductwe biology. The external structures
of the platyspermlc ovules and seeds are weakly
developed in Ugartecladus though specialized
in Ferughocladus, which has a strongly devel-
oped mlcropylar apex and a delicate sur-
units from the megasporophyll and ovule The
specialized blfid apices in Ferughocladus seem
to be of integumentary origin, rather than
modified scales, and the lack of this structure in
Ugartecladus is consistent with this assump-
tion If interpretation X is correct, reduction of
the scale or fusion of ovule and scale in one
complex, must be regarded as total or nearly
total, suggesting a highly advanced stage of
cone appendage evolution (Fig.7) The relative
age of our material is younger, although only
slightly so, than the first known stratigraphlc
occurrence of conifers which have ovuliferous
scales together with the ovules (Hill and Crane,
1982: Hill and Camus, 1986). This is consistent
with the accepted concepts of comferalean
phylogeny which are represented in our inter-
pretatlon (X) (Florin, 1951). In this context,
these genera are of interest in extending the
earliest stratlgraphIc occurrence of highly
specialized (1 e. fused/reduced) female cone-
scales backwards in time from the Trias to the
early Permian, showing that conifer evolution
was more rapid than has been thought, at any
rate in South America
Interpretation (Y), on the other hand (Fig 8)
represents a less orthodox but interesting view
of conifer evolution which would suggest that
the female cone m some conifers or conifer-like
plants did not have a compound origin.
(Whether such plants should then be accepted
as true conifers is an important question). On
this interpretation, Ugartecladus and Feru-
ghocladus would be considered to have simple
strobih (i e of ovules and bracts) such as are
known already to characterize the Permian
Gondwanan genus Buriadta Seward et Sahni
(Pant and Nautiyal, 1967). This is considered to
represent a distinct order within the conifers,
Burladlales, by Pant (1982) It has small,
stalked, anatropous ovules located between
leaves that do not form cones (Pant and
Nautiyal, 1967) Interestingly, the seeds have
blfid apices as in Ferugliocladus It is therefore
not unreasonable to suggest an ancestor com-
mon to both our genera and Buriadia (Fig.8, see
also Archangelsky, 1985). In evolutionary hne
A of Fig 8, the ancestor changes only in ~ts
ovule, which becomes anatropous m Burtadla.
whilst in line B, leading to Ugartecladus and
Ferughocladus the ovules lose their stalks, the
bracts become simple and the whole structure
forms a compact cone. If the Buriadlales are
indeed to be considered true conifers (Krassl-
lov, 1971; Meyen, 1978) they, hke Ugartecladus
and Ferugliocladus, would clearly represent on
this interpretation a parallel hne of evolunon
to that which led to Voltziales
Proposal of a new faintly The distractive
characters of Ugartecladus and Ferughocladus
preclude their inclusion in the most similar
conifer families Walchlaceae, Buriadlaceae
and Araucarlaceae These new genera, further-
more, are restricted m time and space to the
Permian of Gondwanan South America. There-
fore we propose the following definmon for a
new family, the Ferughocladaceae, of which
Ferughocladus is selected as the type genus.
Members of this family are conifers (sensu
lato or sensu stricto), with Irregular radial
branching, bearing terminal compact male and
female cones_ The leaves are small, helically
disposed, all of one kind, lanceolate and single
vetoed. The female cones are composed of a
central axis with bracts and apparently free or
actually free, orthotropous ovules which are
disposed helically The seeds are platyspermlc,
reaching their greatest size while still in the
cones. The male cones are composed of an axis
bearing helically arranged, proximally deflexed
mlcrosporophylls. The pollen is monosaccate.
Other genera Ugartecladus gen nov (male
cones unknown) Eucerospermum Ferugho (dis-
persed seeds) Addmonal genera, at least m
part, are Paranocladus Florin (branches and
leaves with cuticles) and Brastlocladus Yoshida
(branches and leaves which have not yet yielded
cuticles) The status of these genera is probably
similar to that of Brachyphyllum Lmdley et
Hutton ex Brongniart and Pagiophyllum Heer
(Harris, 1979).
Reproductwe biology. The external structures
of the platyspermlc ovules and seeds are weakly
developed in Ugartecladus though specialized
in Ferughocladus, which has a strongly devel-
oped mlcropylar apex and a delicate sur-
rounding tissue The large "cells" m the apical
regmn of the ovule of F. patagon~cus may have
decomposed to form a hqmd, and the channel
whmh runs from base to apex, where there is a
small chamber or depression, may have served
for storage of some kind of liquid. These
structures may mdmate that Ferughocladus
was insect polhnated, unhke other conifers,
and are interesting m their similarities to the
apical structures of Bur~ad~a and of several
pteridosperms The small size difference be-
tween the m sltu ovules and dispersed seeds, in
both genera, suggests that pollinatmn occurred
on the plant, where the seed subsequently
developed When mature they were shed and
germinated
With the destructmn of the external soft
tissue, the apical hooks remained and may have
asmsted in dispersal of the seeds by animals. It is
of interest that blattold insects have been found
m the same homzons as Ferughocladus (Pinto,
1972).
Palaeoecology: Quanhtahve fieldwork has been
undertaken m several plant-inch sectmns yield-
mg conifers, m Chubut (Lomas Chatas, Puesto
Alambre, La Casllda) and Rloja (five localities).
In Patagoma, conifers were deposited in
extensive plains with meandering rivers
(Archangelsky, 1981; Cflneo, 1983), whmh
were suitable for the development of different
ecological inches hawng varied vegetation
Interbeddmg with horizons having marine
invertebrates mdmates proximity to the sea. In
Patagonlan assemblages, conifers have an
abundance of about 30% of the total plant
remains, ranging from 10 to 40~/o. Their habitat
was probably mesophllous to hygro-mesophi-
lous sensu Remy and Remy (1977). Moreover,
they are assocmted with ferns, sphenophytes
(sphenophylls and Equisetales) and lycophytes
in coa]-bearmg sequences, suggesting that they
grew m swampy enwronments close to the
water table. The fossils are represented by large
fragments of branches bearing cones, and by
isolated cones and abundant detached seeds.
The large fragments with branches of different
orders shll connected suggest that the plants
were shoot-dropping conifers and that trans-
29
port was mimmal Isolated leaves are less
common. Petrographm analyses of the coals
(H. Villar, pers. commun., 1983) and the
presence of large Aphleblae suggest a humid
climate The conifers are often associated with
Glossoptemdales, whmh probably had similar
ecologmal reqmrements There is no marked
field relationship with groups such as Cordal-
tales, Ginkgoales and pteridosperms
In La Rloja Province, palaeoenvlronments
were similar though smaller in area and flanked
by hills m a continental mountain system.
Consequently the assemblages are less diverse.
Conifers represent 24% of the total megafosslls
and are commonly associated with Equlsetales
(Phyllotheca) and more randomly with ferns,
Cordaltales, Gmkgoales and ptemdosperms
This suggests that a wider range of habitats
were occupmd here by the comfers than m
Chubut. Ferughocladus r~ojanum was probably
hygro-mesophllous, occurring associated with
Phyllotheca or with Cordaltes or Glossopterls. It
occurs as shoots with attached cones.
It appears that Ferughocladaceae had a more
vamed ecological range than those comfers
found m the equatorial belt, whmh are mainly
thought to have been xerophilous, and that they
lived in temperate humid conditions. This may
have been important m leading to their more
rapid evolutmn than the equatomal conifers, m
relation to present-day cone morphology of
conifers. Similar ecological conditmns have
been suggested for the Angara regmn by Meyen
(1978).
Acknowledgements
Suggestions by Dr K.L. Alvin, London, Dr
C N Miller, Mlssoula, Montana are gratefully
acknowledged. We also thank Drs B Petriella
(La Plata Museum), W Volkhelmer (Buenos
Aires Museum) and R LeguizamSn (CIRGEO)
for loan of specimens, and Mr. M. Archangel-
sky for photographm and field assistance. The
work was partly supported by grants from
the National Scmnce Foundation (USA) BSR
8313786 and CONICET (Argentina) to T.N
Taylor and S Archangelsky
regmn of the ovule of F. patagon~cus may have
decomposed to form a hqmd, and the channel
whmh runs from base to apex, where there is a
small chamber or depression, may have served
for storage of some kind of liquid. These
structures may mdmate that Ferughocladus
was insect polhnated, unhke other conifers,
and are interesting m their similarities to the
apical structures of Bur~ad~a and of several
pteridosperms The small size difference be-
tween the m sltu ovules and dispersed seeds, in
both genera, suggests that pollinatmn occurred
on the plant, where the seed subsequently
developed When mature they were shed and
germinated
With the destructmn of the external soft
tissue, the apical hooks remained and may have
asmsted in dispersal of the seeds by animals. It is
of interest that blattold insects have been found
m the same homzons as Ferughocladus (Pinto,
1972).
Palaeoecology: Quanhtahve fieldwork has been
undertaken m several plant-inch sectmns yield-
mg conifers, m Chubut (Lomas Chatas, Puesto
Alambre, La Casllda) and Rloja (five localities).
In Patagoma, conifers were deposited in
extensive plains with meandering rivers
(Archangelsky, 1981; Cflneo, 1983), whmh
were suitable for the development of different
ecological inches hawng varied vegetation
Interbeddmg with horizons having marine
invertebrates mdmates proximity to the sea. In
Patagonlan assemblages, conifers have an
abundance of about 30% of the total plant
remains, ranging from 10 to 40~/o. Their habitat
was probably mesophllous to hygro-mesophi-
lous sensu Remy and Remy (1977). Moreover,
they are assocmted with ferns, sphenophytes
(sphenophylls and Equisetales) and lycophytes
in coa]-bearmg sequences, suggesting that they
grew m swampy enwronments close to the
water table. The fossils are represented by large
fragments of branches bearing cones, and by
isolated cones and abundant detached seeds.
The large fragments with branches of different
orders shll connected suggest that the plants
were shoot-dropping conifers and that trans-
29
port was mimmal Isolated leaves are less
common. Petrographm analyses of the coals
(H. Villar, pers. commun., 1983) and the
presence of large Aphleblae suggest a humid
climate The conifers are often associated with
Glossoptemdales, whmh probably had similar
ecologmal reqmrements There is no marked
field relationship with groups such as Cordal-
tales, Ginkgoales and pteridosperms
In La Rloja Province, palaeoenvlronments
were similar though smaller in area and flanked
by hills m a continental mountain system.
Consequently the assemblages are less diverse.
Conifers represent 24% of the total megafosslls
and are commonly associated with Equlsetales
(Phyllotheca) and more randomly with ferns,
Cordaltales, Gmkgoales and ptemdosperms
This suggests that a wider range of habitats
were occupmd here by the comfers than m
Chubut. Ferughocladus r~ojanum was probably
hygro-mesophllous, occurring associated with
Phyllotheca or with Cordaltes or Glossopterls. It
occurs as shoots with attached cones.
It appears that Ferughocladaceae had a more
vamed ecological range than those comfers
found m the equatorial belt, whmh are mainly
thought to have been xerophilous, and that they
lived in temperate humid conditions. This may
have been important m leading to their more
rapid evolutmn than the equatomal conifers, m
relation to present-day cone morphology of
conifers. Similar ecological conditmns have
been suggested for the Angara regmn by Meyen
(1978).
Acknowledgements
Suggestions by Dr K.L. Alvin, London, Dr
C N Miller, Mlssoula, Montana are gratefully
acknowledged. We also thank Drs B Petriella
(La Plata Museum), W Volkhelmer (Buenos
Aires Museum) and R LeguizamSn (CIRGEO)
for loan of specimens, and Mr. M. Archangel-
sky for photographm and field assistance. The
work was partly supported by grants from
the National Scmnce Foundation (USA) BSR
8313786 and CONICET (Argentina) to T.N
Taylor and S Archangelsky
30
References
Andrels, R.R_, Cflneo, R and Rol6n, A D, 1984 Defimc16n
formal de los "Estratos de Arroyo Totoral", P~rmmo
referrer de Smrra de los Llanos, Prowncm de La Rmja,
Argentina Actas 9th Congr Geol Arg. Barfloche, 5
209 229
Archangelsky. S, 1978 Nuevos elementos paleoflomstmos
de la Formac16n Trampeadero (Carb6nlco), Smrra de
Ambato, Provlncm de la Rmja_ Ameghmmna, 15 429 439
Archangelsky, S. 1981 Estudms fitopaleoecol6glcos en el
Paleozmco Supermr del Oeste de Chubut, Argentina An
2nd Congr Latmoam Paleontol, Porto Alegre, 1
141 156
Archangelsky, S, 1985 Aspectos evolutlvos de las coniferas
gondwAmcas del Paleozolco Bull Sect Scl Cam Trav
Hlst Scl,8 115 124
Archangelsky, S and Arrondo, O G, 1973 Palaeophyto-
logm Kurtmana III 10 La tafoflora permma de Sierra de
los Llanos, Provmcm de La Rm]a Ameghmlana, 10
201 228
Archangelsky, S and Cdneo, R, 1984_ Zonac16n del P~rmlco
continental de Argentina sabre la base de sus plantas
fdsfles Mem 3rd Congr Latmoam Paleontol. M~xlco,
1984, pp 143 153
Barbosa, O, 1958 On the age of the Lower Gondwana floras
m Brazil and abroad In 20th Int_ Geol Congr_, M~xlco,
1956, Comls16n para la correlacldn del Slstema Karroo
(Gondwana). pp 205 236
Bernardes de Ohvelra. M E and Yoshlda, R, 1982
Comferdfitas da "Tafofl6rula Irapuh", Formacfio Rm
Bomto, Grupo Tubarfio, em Santa Catarlna Bol_ Asoc
Latlnoam Paleobot Pahnol_, (8) 39 55
Clement-Westerhof, J A, 1984 Aspects of Permmn palaeo-
botany and palynology IV The conifer Ortlse~a Florin
from the Val Gardena Formatmn of the Dolomites and the
Vlcentlman Alps (Italy) with specml reference to a
rewsed concept of the Walchlaceae (Goppert) Schlmper
Rev Palaeobot_ Palynol, 41 51 166
Cdneo, R, 1983 Paleoecologla de mlcrosecuenclas plantl-
feras del Grupo Rm Genoa. P~rmlco de Chubut, Argen-
tina Ameghmmna. 20 111 131
Ferugho, E, 1934 Fossil1 hasslcl della Valle del Rm Genoa
(Patagoma) G Geol 9 1 64
Ferugllo, E. 1946 La flora hgs~ca del valle del Rm Genoa
(Patagoma) Semma mcertae sed~s Rev Sac Geol
Argent, 1 209 218
Ferugho, E , 1951 Su alcune pmnte del Gondwana mferlore
della Patagoma Publ Ist Geol_ Umv Tormo, 1 1 34
F~ttlpaldL F C and Rosler. O, 1978 Paranocladus~ fallax
(Comfera) Estudos cut~culares Balm IG Silo Paulo, 9
109 113
Florin, R, 1940a Dm Komferen des Oberkarbons und des
Unteren Perms. 5 Palaeontographlca B, 85 243 363
Florin, R, 1940b On Walkom~a n gen_ A genus of upper
Palaeozoic conifers from Gondwanaland K Sven Vet-
ensk Akad Handl (ser 3), 18{5) 1 23
Florin, R. 1951 Evolutmn m Cordmtes and Conifers Acta
Hart1 Bergmm, 15 285 388
Harms. T M, 1979 The Yorkshire Jurassic flora, V
Comferales Brutish Museum (Natural History). London,
166 pp
HH1, C R and Camus. J M, 1986 Evolutionary cladlstlcs of
marattlalean ferns Bull Br Mus Nat Hlst (Bat), 14
219 300
Hill, C R and Crane, P R, 1982 Evoluhonary cladlshcs and
the omgm of Angiosperms In KA Joysey and A E
Fmday (Editors). Problems of phylogenetw reconstruc-
ban Syst Assoc Spec Vol, 21 269 361
KrassHov, V A, 1971 Evolution and systematlcs of Conl-
ferae (a critical review) Paleontol Zh, (1) 7 20 (in
Rusman)
Legmzam6n, R_, 1972 Estudlo paleobot~mco de la Forma-
c16n Tasa Cuna, P6rmlco inferior de la Provmcla de
Cdrdoba Ameghmlana, 9 305 342
Mapes, G and Rothwell, G W, 1984 Permmerahzed
ovulate cones of Lebach~a from late Paleozolc hmestones
of Kansas Palaeontology, 27 69 94
Meyen, S V, 1978 Permian comfers of West Angaraland
and new puzzles m the comferalean phylogeny Palaeobo-
tamst, 25 298 313
Mfllan. J E, 1972 Macrofl6rula carbomfera de Monte Mar.
Estado de Silo Paul() Thesis Unlv $3.o Paulo, Instlt
Geoclenc, 165 pp
Mfllan, J E, 1974 Sabre as Conlferopslda da fl6rula
gondw~mca de Monte Mar, Estado de S~o Paulo, Brasfl
Ameghlmana, 11 124 134
Miller, C N and Brown, J T, 1973_ A new voltzlalean cone
bearing seeds with embryos from the Permmn of Texas
Am J Bat, 60 561 569
Pant. D D, 1982 The Lower Gondwana gymnosperms and
their relationships Rev Palaeobot Palynol. 37 55 70
Pant, D D and Naut~yal. D D, 1967 On the structure
of BurtadLa heterophylla (Felstmantel) Seward and
Sahm, and its fructlficatmn Phflos Trans R Sac, B
252 27 48
Pinto. I D, 1972 A new msecta, Archangelskyblatta
vzshmakova~ Pinto, gen nov, spec nov, a Permmn blattold
fl'om Patagoma, Argentina Ameghmmna, 9 7~89
Remy, W and Remy, R, 1977 Die Floren des Erdaltertums
Gluckauf, Essen, 468 pp
Seward, A_C and Sabra, B, 1920 Indian Gondwana plants
a revision Mem Geol Surv Indm. n ser, 7(1) 1 41
Surange, K R and Lele, K M, 1957 Studms in the
Glossopterls flora of India, 6 Plant fossils from Talchlr
beds of South Rewa Gondwana basra Palaeobotamst,
5(2) 82 90
Surange K R and Smgh, P, 1951 Walkomtella Lndtca, a
new conifer from the Lower Gondwanas of Indm J
Indian Bat Sac, 30 143 147
Surange. K R and Smgh, P. 1953 The female dwarf shoot of
Walkormella znd~ca a conifer from the Lower
Gondwanas of India Palaeobotamst, 2 5 8
White. M E, 1981 The cones of Walkomtella austrahs
(Felst) Florin Palaeobotanlst 28 29 75 80
Yoshlda, R, 1970 Nova genera de comfera da Formac~o
Estrada Nova, norte do Estado de Santa Catarma Bol
Dlv Geol Miner Braz, 249 1 17
View publication stats
References
Andrels, R.R_, Cflneo, R and Rol6n, A D, 1984 Defimc16n
formal de los "Estratos de Arroyo Totoral", P~rmmo
referrer de Smrra de los Llanos, Prowncm de La Rmja,
Argentina Actas 9th Congr Geol Arg. Barfloche, 5
209 229
Archangelsky. S, 1978 Nuevos elementos paleoflomstmos
de la Formac16n Trampeadero (Carb6nlco), Smrra de
Ambato, Provlncm de la Rmja_ Ameghmmna, 15 429 439
Archangelsky, S. 1981 Estudms fitopaleoecol6glcos en el
Paleozmco Supermr del Oeste de Chubut, Argentina An
2nd Congr Latmoam Paleontol, Porto Alegre, 1
141 156
Archangelsky, S, 1985 Aspectos evolutlvos de las coniferas
gondwAmcas del Paleozolco Bull Sect Scl Cam Trav
Hlst Scl,8 115 124
Archangelsky, S and Arrondo, O G, 1973 Palaeophyto-
logm Kurtmana III 10 La tafoflora permma de Sierra de
los Llanos, Provmcm de La Rm]a Ameghmlana, 10
201 228
Archangelsky, S and Cdneo, R, 1984_ Zonac16n del P~rmlco
continental de Argentina sabre la base de sus plantas
fdsfles Mem 3rd Congr Latmoam Paleontol. M~xlco,
1984, pp 143 153
Barbosa, O, 1958 On the age of the Lower Gondwana floras
m Brazil and abroad In 20th Int_ Geol Congr_, M~xlco,
1956, Comls16n para la correlacldn del Slstema Karroo
(Gondwana). pp 205 236
Bernardes de Ohvelra. M E and Yoshlda, R, 1982
Comferdfitas da "Tafofl6rula Irapuh", Formacfio Rm
Bomto, Grupo Tubarfio, em Santa Catarlna Bol_ Asoc
Latlnoam Paleobot Pahnol_, (8) 39 55
Clement-Westerhof, J A, 1984 Aspects of Permmn palaeo-
botany and palynology IV The conifer Ortlse~a Florin
from the Val Gardena Formatmn of the Dolomites and the
Vlcentlman Alps (Italy) with specml reference to a
rewsed concept of the Walchlaceae (Goppert) Schlmper
Rev Palaeobot_ Palynol, 41 51 166
Cdneo, R, 1983 Paleoecologla de mlcrosecuenclas plantl-
feras del Grupo Rm Genoa. P~rmlco de Chubut, Argen-
tina Ameghmmna. 20 111 131
Ferugho, E, 1934 Fossil1 hasslcl della Valle del Rm Genoa
(Patagoma) G Geol 9 1 64
Ferugllo, E. 1946 La flora hgs~ca del valle del Rm Genoa
(Patagoma) Semma mcertae sed~s Rev Sac Geol
Argent, 1 209 218
Ferugho, E , 1951 Su alcune pmnte del Gondwana mferlore
della Patagoma Publ Ist Geol_ Umv Tormo, 1 1 34
F~ttlpaldL F C and Rosler. O, 1978 Paranocladus~ fallax
(Comfera) Estudos cut~culares Balm IG Silo Paulo, 9
109 113
Florin, R, 1940a Dm Komferen des Oberkarbons und des
Unteren Perms. 5 Palaeontographlca B, 85 243 363
Florin, R, 1940b On Walkom~a n gen_ A genus of upper
Palaeozoic conifers from Gondwanaland K Sven Vet-
ensk Akad Handl (ser 3), 18{5) 1 23
Florin, R. 1951 Evolutmn m Cordmtes and Conifers Acta
Hart1 Bergmm, 15 285 388
Harms. T M, 1979 The Yorkshire Jurassic flora, V
Comferales Brutish Museum (Natural History). London,
166 pp
HH1, C R and Camus. J M, 1986 Evolutionary cladlstlcs of
marattlalean ferns Bull Br Mus Nat Hlst (Bat), 14
219 300
Hill, C R and Crane, P R, 1982 Evoluhonary cladlshcs and
the omgm of Angiosperms In KA Joysey and A E
Fmday (Editors). Problems of phylogenetw reconstruc-
ban Syst Assoc Spec Vol, 21 269 361
KrassHov, V A, 1971 Evolution and systematlcs of Conl-
ferae (a critical review) Paleontol Zh, (1) 7 20 (in
Rusman)
Legmzam6n, R_, 1972 Estudlo paleobot~mco de la Forma-
c16n Tasa Cuna, P6rmlco inferior de la Provmcla de
Cdrdoba Ameghmlana, 9 305 342
Mapes, G and Rothwell, G W, 1984 Permmerahzed
ovulate cones of Lebach~a from late Paleozolc hmestones
of Kansas Palaeontology, 27 69 94
Meyen, S V, 1978 Permian comfers of West Angaraland
and new puzzles m the comferalean phylogeny Palaeobo-
tamst, 25 298 313
Mfllan. J E, 1972 Macrofl6rula carbomfera de Monte Mar.
Estado de Silo Paul() Thesis Unlv $3.o Paulo, Instlt
Geoclenc, 165 pp
Mfllan, J E, 1974 Sabre as Conlferopslda da fl6rula
gondw~mca de Monte Mar, Estado de S~o Paulo, Brasfl
Ameghlmana, 11 124 134
Miller, C N and Brown, J T, 1973_ A new voltzlalean cone
bearing seeds with embryos from the Permmn of Texas
Am J Bat, 60 561 569
Pant. D D, 1982 The Lower Gondwana gymnosperms and
their relationships Rev Palaeobot Palynol. 37 55 70
Pant, D D and Naut~yal. D D, 1967 On the structure
of BurtadLa heterophylla (Felstmantel) Seward and
Sahm, and its fructlficatmn Phflos Trans R Sac, B
252 27 48
Pinto. I D, 1972 A new msecta, Archangelskyblatta
vzshmakova~ Pinto, gen nov, spec nov, a Permmn blattold
fl'om Patagoma, Argentina Ameghmmna, 9 7~89
Remy, W and Remy, R, 1977 Die Floren des Erdaltertums
Gluckauf, Essen, 468 pp
Seward, A_C and Sabra, B, 1920 Indian Gondwana plants
a revision Mem Geol Surv Indm. n ser, 7(1) 1 41
Surange, K R and Lele, K M, 1957 Studms in the
Glossopterls flora of India, 6 Plant fossils from Talchlr
beds of South Rewa Gondwana basra Palaeobotamst,
5(2) 82 90
Surange K R and Smgh, P, 1951 Walkomtella Lndtca, a
new conifer from the Lower Gondwanas of Indm J
Indian Bat Sac, 30 143 147
Surange. K R and Smgh, P. 1953 The female dwarf shoot of
Walkormella znd~ca a conifer from the Lower
Gondwanas of India Palaeobotamst, 2 5 8
White. M E, 1981 The cones of Walkomtella austrahs
(Felst) Florin Palaeobotanlst 28 29 75 80
Yoshlda, R, 1970 Nova genera de comfera da Formac~o
Estrada Nova, norte do Estado de Santa Catarma Bol
Dlv Geol Miner Braz, 249 1 17
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