Postilla /Yale Peabody Museum of Natural History.

Peabody Museum

of Natural History

Yale University

New Haven, CT 06511

Postilla Number 180
30 November 1980

See

Devonian Vertebrates
from Australia

Emily B. Giffin

(Received 3 September 1979)
Abstract

A vertebrate microfauna from the Lower De-
vonian (Emsian) of Australia is described. It is
& taxonomically diverse assemblage, includ-
ING disarticulated dermal skeletal fragments
and/or teeth of Thelodonti, Placodermi,
Cladodontida, Acanthodii, Onychodontidae,
Rhipidistia, and Paleoniscida. Elements of
this fauna have been described previously
from geographically diverse localities of ap-
Proximately equivalent age, but this Aus-
tralian fauna is unique in possessing all of the
above taxonomic groups in a single as-
SeMblage. This occurrence reinforces previ-
US Suggestions that the Lower Devonian fish
fauna was widespread and uniform.

Introduction

The vertebrate material discussed below was
Collected by B. D. E. Chatterton from a series
Of micrites, biomicrites, and biosparites in the
lower 200 feet of the Receptaculites Lime-
Stone. The Receptaculites unit is one of a
Sequence of limestones of the Taemas For-
Mation (Murrumbidgee Group) that occurs
Near the locality site (Fig. 1) designated as
Locality P, Bloomfield Property, Portion 229,

“ Copyright 1980 by the Peabody Museum of

atural History, Yale University. All rights re-
Served. No part of this publication, except
brief Quotations for scholarly purposes, may
be produced without the written permission
of the Director, Peabody Museum of Natural!
\story.

Parish of Warroo, near Yass, New South
Wales, Australia (personal communication,
B. D. E. Chatterton to K. S. Thomson, 1968).
The age of the Murrumbidgee Group has
been assigned on the basis of its invertebrate
and conodont faunas (Philip and Pedder,
1964, 1967; Pedder, 1967; Savage, 1973) as
near the Siegenian/Emsian boundary. Chat-
terton (personal communication, 1978) now
places the Receptaculites unit itself in the
middle Emsian on the basis of conodont
occurrences.

The general area surrounding the collec-
tion site described here has yielded a large
number of vertebrate remains over a consid-
erable period of years. Etheridge (1906) de-
scribed the dipnoan Ganorhynchus (later as-
signed to Dipnorhynchus by Jaeckel, 1927)
from this locality, and a series of placoderms
were described by Woodward (1941) and
White (1952). More recently Schultze (1968)
described a microvertebrate fauna that in-
cluded some, but not all, of the forms de-
scribed here. Schultze’s fauna was also col-
lected from the Taemas Formation, but the
majority of it was from the Spirifer yassensis
limestone, a lithic unit some 800-1200 feet
beneath the base of the Receptaculites
Limestone (Chatterton, 1971). Schultze de-
scribed scales of the new paleoniscoid
Ligulalepis toombsi from his fauna, and men-
tioned the occurrence of fragmentary re-
mains of Ohiolepis, Ohioaspis, and Ony-
chodus with them.

In addition to the vertebrate material,
the Receptaculites Limestone contains a
diverse invertebrate fauna composed of
brachiopods, gastropods, tabulate and
rugose corals, trilobites, ostracods, and con-
odonts. The diversity of this assemblage and

Devonian Vertebrates
from Australia

Postilla 180

Queensland

New South Wales

“a,

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Victoria Ace

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1
1
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'
[e
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1
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fi
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Receptaculites Limestone
Bloomfield Limestone
= Currajong Limestone
Spirifer yassensis Limestone
f=] Majurgong Formation

ea Cavan Bluff Limestone
Mountain Creek Tuffs
Narrangullen Rhyolite fe)

Lng
~

Sy
WY

1}

0

Ke
SC
x

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i

Coarse tuffs ;

1/2 | Taemas i
mile |

Collection site of vertebrate fauna, near Yass, New
South Wales, Australia. Map prepared by Brian
D. E. Chatterton.

the presence of calcareous algae suggest
that the environment of deposition was a
warm shallow sea of low-to-moderate energy.
The regime was apparently strictly marine.

r Site of collection
—<— Fault Geology partly from Browne 1959 sy
Fig. 1

Description of Specimens

The recovery of vertebrate material from the
matrix was a byproduct of its preparation for
conodonts. This was accomplished by diges-
tion of the rock in 10% monochloracetic acid,
sieving, and heavy mineral separation. The
resulting vertebrate fossils are extremely
fragile and lack all microstructure, although
fine surface detail is retained. Unfortunately,
damage occurred to some of this very fragile
material during the processes required for
SEM photography.

The vertebrate fauna, like the invertebrate
fauna, is remarkable for its diversity. Al-
though lack of microstructure makes even
generic identification difficult, fish remains
belonging to a number of the larger tax-
onomic groups can be recognized. These
include Thelodonti, Placodermi, Cladodon-
tida, Acanthodii, Onychodontidae, Rhipidis-
tia, and Paleoniscida, as well as various inde-
terminate fish fragments.

By far the greatest bulk of the residue con-
sists of acanthodian scales and onychodon-
tid teeth. All of the remaining taxa are rep-
resented by only a few specimens. All of the
material is now part of the Australian National
University collections, numbers 35606 and
35607.

3 Devonian Vertebrates
from Australia

Postilla 180

Fig. 2A, B

Skamolepis fragilis, thelodont agnathan scales
from the Receptaculites Limestone. Scale =
0.1 mm.

Thelodonti

A very small number of scales presumed to
be those of the thelodont agnathans were
found in the Receptaculites Limestone res-
idue. Although at first glance the two Speci-
mens figured (Fig. 2A,B) appear to be quite
different, they may be considered end forms
Of the same morphological pattern. This pat-
tern includes a wide open, central pulp cavity
and a nearly flat basal plate. The neck is con-
Stricted and short. The crown bears a series
Of linear ornaments and shows suggestions
Of tripartite structure.

The central pulp cavities of these scales
demand their placement in the Thelodonti.
Despite lack of histological information, they
can be placed in the species Skamolepis
fragilis because of their remarkable external
similarities to this species, which was re-
cently described by Karatajute-Talimaa
(1978). S. fragilis has been reported previ-
ously only from the late Emsian of Latvia
and from the Grey Hoek (?Eifelian) of
Spitsbergen.

Thelodonts have been reported previously
in assemblages similar in age to that found in
the Receptaculites Limestone. Orvig (1957,

4 Devonian Vertebrates
from Australia

Postilla 180

Fig. 3
Ohioaspis tumulosa, placoderm tessera from the
Receptaculites Limestone. Scale = 0.1 mm.

1969a, 1969b) described the thelodont Amal-
theolepis winsnesi from the Grey Hoek of
Spitsbergen, part of a fauna that also in-
cludes fragments of a petalichthyid,
Arctolepis, a struniiform, and acanthodians.
Orvig particularly noted (1969b) that this
fauna is unlike the characteristic Emsian/
Eifelian fauna that contains ptyctoconts,
Ohioaspis, Ohiolepis, and undetermined
Struniiformes from a variety of localities.
Schultze (1968) noted the presence of
“horn-like thelodontid scales” from this same
unit in association with the paleoniscoid Or-
vikuina and Porolepis-like scales.

Placodermi

The placoderm material of the Receptacu-
lites Limestone consists of dermal tesserae of
varying size and generally polygonal shape
(Fig. 3). A base and sculptured crown sur-
face can be differentiated, but there is no
discrete neck. The base is usually flat or
slightly convex, and is often penetrated by
several vascular canals. The sculpture con-

sists of star-shaped tubercles which vary
in micro-ornament, shape, and number.
The tesserae range from 0.5 to 1.2 mm
in greatest diameter.

These tesserae closely resemble those
placed in the genus Ohioaspis by Wells
(1944) from the Middle Devonian bonebeds
of Ohio, Indiana, and Kentucky. Although
Wells recognized three species of Ohioaspis,
one with four “forms,” Gross (1973) has
grouped them all into the species O. tum-
ulosa. Gross also suggested that Ohioaspis
and the rhenanid placoderm genus Asteros-
teus are identical, but retained the genus
Ohioaspis until confirmation of the identity.

In addition to Wells’ localities, specimens
of Ohioaspis have been reported previously
from the Murrumbidgee area of Australia by
Schultze (1968) and from New York State
(Onondaga) and Spitsbergen (Lower/Middle
Devonian) by Orvig (1969b).

Cladodontida

Among the vertebrate remains of the Recep-

5 Devonian Vertebrates Postilla 180
from Australia

Fig. 4A, B

Ohiolepis sp., cladodont chondrichthyan scales
from the Receptaculites Limestone. Scale =

0.1 mm.

6 Devonian Vertebrates
from Australia

Postilla 180

Fig. 5
Cheiracanthoides comptus, acanthodian scale
from the Receptaculites Limestone. Scale =

0.1 mm

taculites Limestone, scales of the cladodont
chondrichthyan genus Ohiolepis are readily
identified by their crown sculpture (Fig. 4A,
B). This ornamentation consists of numerous
short, posteriorly directed thorn-shaped pro-
jections. Although arrangement and size of
the projections are variable, they are gener-
ally symmetrical in distribution. This pattern
and the low, flat profile of the crown are re-
flections of the scale’s growth pattern.
Cladodont scales grow by the addition of
successive, overlapping, and marginal in-
crements rather than by complete encircling
layers as in acanthodians. The ontogenet-
ically oldest part of the scale crown is central
in position (Gross, 1973).

The basal outline of the Ohiolepis scale is
round to rhombic, and forms the largest part
of the scale. On some specimens it bears
one or more vascular canal openings. The
base extends further anteriorly than the
crown but is overlapped by crown ornament

posteriorly. There is no discrete neck.

Other reported occurrences of Ohiolepis
include the Middle Devonian of Ohio, Indi-
ana, and Kentucky (Wells, 1944), the Heis-
dorf beds of Germany (Schmidt, 1961; Orvig,
1969b), the Murrumbidgee Group of Aus-
tralia (Schultze, 1968), and the Onodaga of
New York State (Orvig, 1969b).

Acanthodii

Acanthodian scales are a large component
of the vertebrate residue of the Receptacu-
lites Limestone. All acanthodian scales
present are variants of a single morphologi-
cal pattern (Fig. 5). The scale base is spheri-
cal, and like all acanthodian scales lacks a
pulp cavity opening. It is typically broader
than the scale crown, and extends further
forward than the anterior end of the crown.
The neck is low and bears the opening of the
vascular system that supplied the crown. The

7 Devonian Vertebrates
from Australia

Postilla 180

Fig. 6

Indet. acanthodian spine from the Receptaculites

Limestone. Scale = 0.1 mm.

Crown is nearly flat, rhombic in shape, and
possesses a pointed posterior tip. The
Characteristic crown sculpture consists of
more or less radial ribs that are broadest at
the anterior end of the crown. The ribs narrow
and often disappear near the center of the
Scale. Although microstructure is not pre-
served in this material, this structural pattern
iS Consistent with acanthodian scale genus
Cheiracanthoides comptus. Cheiracan-
thoides is known to possess a Nostolepis-
type histology. Scales range from 0.3 to 0.8
mm in length.

Wells (1944) first described the genus
Cheiracanthoides from the Middle Devonian
bonebeds of Ohio, Indiana, and Kentucky.
He recognized a total of two genera and six
Species on the basis of slight differences in
Crown sculpture and relative size of base and
Crown. Gross (1973) grouped all of these var-
lants into the single species C. comptus. He
reported the genus from various localities
near the Ostsee in Germany.

A single specimen of an acanthodian spine
was found in the fish residue (Fig 6). This

short (0.8 mm) fragment is flattened and very
slightly tapered. Its surface is smooth except
for a series of four conical projections in a
single row. These projections rise perpen-
dicular to the spine shaft without curving.

Acanthodian spine fragments were re-
ported by Wells (1944) from the Middle De-
vonian bonebeds noted above. He assigned
them provisionally to the organ genus
Gyracanthus because of their similarity to G.
primaevus, described by Eastman (1908)
from the Marcellus of New York State.

The specimen described here differs from
G. primaevus in the straightness of its conical
projections, which are more typical of those
of G. sherwood as illustrated by Newberry
(1889). However, the specimen is much
smaller than either of these species, and
cannot be certainly assigned to either.

Onychodontidae

The onychodont material present in the Re-
ceptaculites residue consists of both dermal

8 Devonian Vertebrates
from Australia

Postilla 180

Fig. 7
Dermal ornament, possibly of Onychodus sig-
moides, from the Receptaculites Limestone.

Scale = 1.0mm.

ornament and teeth. The fragments of
dermal ornament are small and rare, but
the teeth form the most common element in
the residue.

The dermal ornament (Fig. 7) is recognized
by its peculiar pattern of raised, horseshoe-
shaped tubercles, which occur in neatly or-
dered rows. The tubercles are about 0.2 mm
in diameter, and the dermal fragments range
up to 1.5mm in greatest dimension. They are
most probably the ‘first generation” tuber-
cles on the dorsal surface of early crossop-
terygian scales. In these forms, radiating
rows Of first generation tubercles ornament
an area between the unornamented and
overlapped anterior portion and the ridged
and exposed posterior portion of the scales.
In detail they most closely resemble those
described as Onychodus sigmoides by
Newberry (1873), Wells (1944), and Orvig
(1957). However they are similar in general
form to those identified as Litoptychus (Deni-
son, 1951) and Glyptolepis (Gross, 1930;
Orvig 1957). The small amount of the material

and the variability of dermal ornament within
all of these genera makes definitive identifi-
cation difficult.

The teeth (Fig. 8) are narrow, pointed, and
hollow, without any identifiable ornamenta-
tion. The shaft tapers and is distinctly curved.
The base is typically constricted, and bears a
flange. They average about 1.1 mm in length.

Wells (1944) identified very similar teeth as
Onychodus interlaniaries. However, Gross
(1969) described teeth of almost identical
morphology among Lophosteus material
from the Beyrichia Limestone of northern
Germany. His study included a histological
examination of these teeth and comparable
material of known actinopterygian and cros-
sopterygian origin. It revealed no histological
characteristics suitable for distinguishing be-
tween isolated teeth of these groups. This
tooth type is possessed by the earliest known
representatives of the Osteichthyes, An-
dreolepis and Lophosteus, and is presuma-
bly primitive to the group as a whole.

The teeth found in the Receptaculites

9 Devonian Vertebrates
from Australia

Postilla 180

Fig. 8
?Onychodus sp. teeth from the Receptaculites
Limestone. Scale = 1.0 mm.

fauna are tentatively identified as Onychodus
because other Onychodus remains are pre-
Sumed present in the residue. However, no
Structural characteristic was found that would
Prohibit their assignment to Lophosteiformes
Or to several other Devonian taxa.

Rhipidistia

The Receptaculites fauna contains a single
isolated cosmoid scale (Fig. 9) and various
Ccosmine-covered dermal fragments. The
Scale is distinguished by a continuous
enamel (cosmine) layer that covers the crown
and a series of pores that puncture this layer.
The enamel surface of the crown is slightly
grooved near the pores, but is otherwise
completely smooth. The crown is irregularly
polygonal in outline and nearly flat, with its
Margins turned slightly downward. There is
no evidence of a crown area that was over-
lapped by neighboring scales, nor of the

groove that commonly separates overlapped
and exposed areas.

The neck of the scale is essentially indis-
tinguishable from the high, straight-walled
base. Pores can be seen to open on its sur-
face. On the inner surface of the base two
ridges parallel the long axis of the scale, with
a broad groove between them. There is a dis-
tinct peg and socket for articulation with
neighboring scales.

The complete cosmine covering of this
scale suggests that it be classified as
rhipidistian. Further, the thick base and polyg-
onal shape are reminiscent of scales of
early members of this group, either Osteo-
lepidae or Porolepidae. With some excep-
tions (Jarvik, 1950), these two groups may be
distinguished by the absence (osteolepid)
or presence (porolepid) of scale ornament
(Jarvik, 1950; Orvig, 1951).

Despite similarities to osteolepid and
porolepid scales, the Receptaculites cos-

Devonian Vertebrates
from Australia

Postilla 180

Fig. 9
Cosmine scale from the Receptaculites Limestone.
Scale = 0.1 mm.

moid scale differs from previously described
specimens in two major respects. The first is
size. Jarvik (1948) lists osteolepid scales in
the 2 x 4mm to 4 x 6mm range, and
porolepid scales are of generally equivalent
size or even larger (Bystrow, 1960). The
Australian specimen is much smaller, about
0.9mm in length. Secondly, the scale has no
overlap area. Previous reports of cosmoid
scales with no overlap area are unknown

to me.

These two characters preclude assign-
ment of this isolated scale to previously es-
tablished genera. It is most similar to rhipidis-
tian scales, and possesses the primitive
rhipidistian characters of thickness, rhombic
shape, and complete cosmine layer. These
are characters shared by early members of
both osteolepid and porolepid rhipidistians.

Osteolepid remains are unknown beneath
the Middle Devonian. Porolepid scales are
rare in the Lower Devonian. Kulezycki (1960)
reports them from the Polish Lower Devonian,
while Jarvik (1950), Orvig (1957) and
Schultze (1968) all report them from the Grey
Hoek of Spitsbergen, which is near the
Lower/Middle Devonian boundary.

Paleoniscida

Only a few scales of known paleoniscoid ori-
gin were preserved whole in the Receptacu-
lites Limestone residue. They are of two gen-
eral types, one of which can be identified as
Ligulalepis toombsi (Fig. 10). This genus was
originally described by Schultze (1968) from
the Taemas Formation of the Murrumbidgee
area, with the majority of the material from the
Spirifer yassensis limestone. Schultze lists
one whole scale and several fragments from
the overlying Receptaculites Limestone. L.
toombsi is not known from other sites.

The scales of Ligulalepis toombsi are flat,
with a rhombic outline. Their length is greatly
exceeded by their height. The sculptured
crown consists of variously joined, obliquely
running, ganoin-covered riblets. A very
marked, spoon-shaped process is set off
from the anterior edge at a sharp angle. The
inner side of the scale possesses two parallel
keels, between which is a broad groove. The
peg and socket are both strongly expressed.

The second group of paleoniscoid scales
is very different. These scales (Fig. 11) are
typically rhombic, but with very elongate

lal Devonian Vertebrates
from Australia

Postilla 180

Fig. 10

Ligulalepis toombsi, a paleoniscoid scale from the

Receptaculites Limestone. Scale = 1.0mm.

Shape. The height of these scales is 3 to 6
times that of their rostral-caudal dimension
(=length). The scales have a peg and socket
imbrication system that seems to involve not
Only the base, as is usual in paleoniscoids,
but the crown as well. This results in an out-
line that resembles an elongate, skewed
parallelogram.

The crown of these scales is set off from
the base by a groove, but there is no discrete
neck. The enamel layer covering the crown is
discontinuous, consisting of elongate ribs
that run parallel to the long dimension of the
Scale. The edges of the ribs are themselves
Sculptured with a fluted ornamentation. The
grooves between the enamel ribs are pierced
by numerous openings, presumably of the
pore canal system.

The base of the scale is very distinctive,
possessing two equal keels, one running
along each of the two elongate scale edges.
The double keel is reminiscent of that found
on Ligulalepis scales (Schultze, 1968) but dif-
fers in encompassing the entire scale base
rather than only the medial portion of the

base. A deep, perforated groove runs be-
tween the two keels.

The morphology of these scales is distinct
and differs from that of paleoniscoid scales
previously described from the Lower Devo-
nian (Schultze, 1968) and Middle Devonian
(Schultze, 1968; Gross, 1953). Their early
occurrence makes them especially interest-
ing, but paucity of material prevents their
formal description.

Distribution and Paleogeography

As early as the Late Silurian it is possible to
distinguish ecological preferences among
various groups of vertebrates. Denison
(1956) has identified two Late Silurian as-
semblages. Osteostraci and Anaspida
(with eurypterids) comprise a brackish-to-
freshwater group, while Heterostraci
Thelodonti, and Acanthodii comprise a
marine group. The Receptaculites Limes-
tone, both in lithology and invertebrate fauna,
strongly indicates a marine origin. Its verte-
brate microfauna is both younger and more

12 Devonian Vertebrates
from Australia

Postilla 180

Fig. 11
Unnamed paleoniscoid scales from the Recep-
taculites Limestone. Scale = 1.0mm.

diverse than those discussed by Denison,
but resembles the marine assemblage in the
presence of Acanthodii and Thelodonti.

The early vertebrate record in Australia is
sparse. There is no pre-Devonian record.
Hills (1958) listed a limited fauna from Aus-
tralia’s Lower Devonian (originally consi-
dered to be Middle Devonian), much of
which was collected at localities near the
material described here. It included several
arthrodires, petalichthyids, and the dipnoan
Dipnorhynchus. This genus is also known
from the Lower Devonian of Germany
(Lehmann and Westoll, 1952).

Since Hills’ work, several new discoveries
have added to the fauna known from the
Lower Devonian of Australia. In 1968
Schultze reported a microvertebrate fauna
containing elements very similar to those
reported here. It included Onychodus,
Ohiolepis, and Ohioaspis in addition to the
new paleoniscoid Ligulalepis, which was de-
scribed in detail. Gross (1971) described the
thelodont Turinia australiensis and undeter-

mined acanthodians from the Lower Devo-
nian of western Australia. Turner (1973)
noted an as yet unpublished report of Turinia
sp. from the Lower Devonian of the Toko
Syncline of Australia.

The Receptaculites Limestone fauna is
notable among known Emsian/Eifelian as-
semblages for its remarkable taxonomic di-
versity. It includes components of faunal as-
semblages found at other, widely distributed
Lower Devonian localities, all within a single
assemblage. Skamolepis fragilis is previ-
ously known only from the Grey Hoek of
Spitsbergen and Latvia (Karatajute-Talimaa,
1978). Cheiracanthoides has been found at
its original localities in North America (Wells,
1944) and a variety of localities near the
Ostsee in Germany (Gross, 1973). The
rhipidistian scale is unique.

The subassemblage of Onychodus,
Ohioaspis, and Ohiolepis, which is often as-
sociated with ptyctodontid tooth plates, has a
wider distribution. The original association
was described by Wells (1944) from Ohio,

12} Devonian Vertebrates
from Australia

Postilla 180

Fig. 12

Known distribution of Lower Devonian vertebrate
assemblages with composition similar to that of the
Receptaculites Limestone. Base map of southern
hemisphere in Lower Devonian from Smith, Briden,
and Drewry, 1973.

Devonian Vertebrates
from Australia

Postilla 180

Indiana, and Kentucky. It was subsequently
reported by Orvig (1969b, p. 317) from the
Onondaga of New York State and by Schultze
(1968) from the Murrumbidgee area of Aus-
tralia. Schmidt (1961) and Orvig (1969b) re-
ported Ohiolepis scales from the Heisdorf
beds of Germany.

Conclusions

The diverse fauna from the Receptaculites
Limestone includes species found in a vari-
ety of other, widely separated locations. This
occurrence suggests two conclusions. First,
it confirms that Australia was not faunally iso-
lated in the Lower Devonian and that the
fauna itself had wide geographic distribution.
A plot of known occurrences of the fauna on
a paleogeographic map of the Lower Devo-
nian (Fig. 12) suggests that the localities oc-
curred in shallow epicontinental marine seas

of the southern hemisphere and that latitude
was not a limiting factor of distribution. Sec-
ondly, the occurrence of diverse members of
the known Lower Devonian vertebrate fauna
in a single locality suggests that the fauna
forms a uniform whole that cannot be sepa-
rated into subgroups with different en-
vironmental, stratigraphic, or geographic
implications.

Acknowledgments

| would like to thank the following people who
have helped in various stages of this project:
Keith S. Thomson, Brian D. E. Chatterton,
Susan Turner, and Valentina Karatajute-
Talimaa.

The photographs were prepared at Har-
vard University’s SEM Laboratory, NSF
Set-up Grant BMS-7412 494.

Literature Cited

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Murrumbidgee River, south of Yass, N.S.W. J. Proc. R. Soc. New South Wales 92:115-128.
Bystrow, A. P. 1960. The microstructure of skeleton elements in some vertebrates from Lower Devonian

deposits of U.S.S.R. Acta Zool. 40:59-83.

Chatterton, B. D. E. 1971. Taxonomy and ontogeny of Siluro-Devonian trilobites from near Yass, New

South Wales. Palaeontographica 137(A): 1-108.

Denison, R. H. 1951. Late Devonian fresh-water fishes from the western United States. Fieldiana, Geol.

11(5):221-261.

1956. A review of the habitat of the earliest vertebrates. Fieldiana, Geol. 11(8):359-457.

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lew 2,

1953. Devonische Palaeonisciden-Reste in Mittel-und Osteuropa. Palaeontol. Z. 27:85-112.

—— 1969. Lophosteus superbus Pander, ein Teleostome aus dem Silur Oesels. Lethaia 2:15-47.

Z. 45(3/4):97-106.

1971. Unterdevonische Thelodontier-und Acanthodier-Schuppen aus Westaustralien. Palaeontol.

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The Author

Emily B. Giffin. Department of Geology
Wellesley College, Wellesley, MA 02181.