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Silurian Brachiopods from the Cappanana Formation East of
Cooma, Southern New South Wales
DESMOND L. STRUSZ
Department of Earth and Marine Sciences, Research School of Earth Sciences, Australian National
University, Canberra ACT 0200 (correspondence to Dr D.L. Strusz, 97 Burnie Street, Lyons ACT
2606)(desmond-strusz@homemail.com.au)
Published on 21 February 2013 at http://escholarship.library.usyd.edu.au/journals/index.php/LIN
Strusz, D.L. (2012). Silurian brachiopods from the Cappanana Formation east of Cooma, southern New
South Wales. Proceedings of the Linnean Society of New South Wales 135: 1-17.
A collection of strongly tectonically distorted fossils from the Cappanana Formation east of Cooma,
NSW, is treated using digital imagery and computer software to restore the better-preserved specimens
to an approximation of their original shape and size, using previously published principles. The fauna is
dominated by brachiopods, all as dissociated valves, but also includes some encrinurid trilobites and a
few corals. Ten brachiopod species are recognised, of which seven - Mesopholidostrophia bendeninensis,
Morinorhynchus oepikensis, cf. Apopentamerus clarkei, Atrypa cf. duntroonensis, Atrypoidea australis,
Howellella aff. elegans and Spirinella caecistriata - are identical to, or closely comparable with, published
species of late Wenlock to Ludlow age. Three taxa (Coelospira sp., a leptaenine and an eospiriferine) are
very rare and could be new, but the material is insuffi cient, and restored images as obtained by the methods
used here should not be used to erect new taxa. The fauna is enclosed in mudstone, probably a current-
winnowed slump deposit, so the original environment cannot be determined.
Manuscript received 26 October 2012, accepted for publication 20 February 2013.
KEYWORDS: Apopentamerus, Atrypa, Atrypoidea, brachiopods, Cappanana Formation, distortion digitally
removed, Howellella, Ludlow, Mesopholidostrophia, Morinorhynchus , Spirinella, Silurian, Wenlock.
INTRODUCTION
In 1999 the residents of ‘Lara’ property, 15
km east of Cooma, advised ANU geologists that
abundant fossils had been turned up during regrading
of the road west of the homestead. Dr Tim Munson
(the then collections manager) and I visited the
spot, and made a collection of the better preserved
material, overwhelmingly brachiopods. Fossils in
the Early Palaeozoic of southeastern New South
Wales between Canberra and the Victorian border are
generally moderately to strongly distorted. As many
are in shales and thinly bedded muddy limestones,
part of this distortion is a result of compaction, but the
major component is tectonic, the result of at least one
major compressional phase with consequent strong
folding and faulting. In Canberra this distortion is
much less, insuffi cient to make recognition of taxa
diffi cult (see, e.g., Strusz 1984, 1985). This cannot be
said of the region around Cooma, some 100 km to the
south, an area further complicated by the formation
of the Cooma Gneiss (part of the Murrumbidgee
Batholith). Consequently, there have been very few
taxa described from that region, and the ages of the
various stratigraphic units are not well constrained.
Distortion of the collected specimens is strong, as is
weathering, but at the time of collecting the fauna
was recognised as being clearly Silurian, and several
taxa were considered comparable with those known
from Canberra. The collection was therefore set aside
until more was known of the better-preserved Silurian
faunas in the Canberra - Yass region.
MATERIALS
The topographic map available to us at the time of
collecting was the fi rst edition Cooma 1:100 000 sheet
of 1969, and this did not show ‘Lara’ homestead or all
of its access road, so our estimate of the position was
very approximate. Satellite imagery is now readily
available, as is a more detailed topographic map (the
Numeralla 1:25,000 sheet of 2001), and this allows
2Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
greater precision. The material came from the north
side of the road on a curve, and is at approximately
GA065.907. On the second edition of the Bega-
Mallacoota 1:250 000 geological sheet (Lewis and
Glen, 1995) this locality lies in a folded and faulted
area within the Cappanana Formation (see Fig. 1).
The fossils are preserved as moulds in soft, yellow-
to buff-weathering pale olive-green mudstone;
the extensive weathering has partly to completely
obliterated fi ne detail. Some of the moulds are lined
with clusters of small crystals of iron oxide - possibly
derived from the overlying Colinton Volcanics -
and limonitisation is heavy in some specimens. All
the brachiopods occur as dissociated valves, the
trilobites as detached parts (moults?). The valves
of strophomenide and orthotetide taxa are generally
parallel to the original bedding, but the more strongly
biconvex atrypide and spiriferide valves can lie at
moderate angles to that bedding. This means that
initial compaction has had only a minor effect on the
shape of the strophomenides and orthotetides, but
more on the others. The appearance of the material
- fl at-lying weakly convex valves, more randomly
oriented strongly convex valves, often in discrete
layers - suggests current winnowing of soft, slumped
sediment, and so the original depth inhabited by the
fauna is uncertain.
PROCEDURES
Important in making reliable identifi cations of
such strongly distorted fossils is a means to reconstruct
their original appearance. Because of the fragility of
the specimens from ‘Lara’ even after impregnation
with acetone-diluted adhesive (‘Tarzan’s Grip’ in this
case), no attempt has been made to obtain latex casts,
but the natural moulds, when carefully prepared,
were generally quite suffi cient. Specimens were
photographed using a digital camera with x3 optical
zoom, under a mix of natural and artifi cial light,
without whitening. For the purposes of this paper, as
fi ne detail is generally poorly preserved, colour has
been retained. Removal of the effects of distortion
followed methods outlined by Cooper (1990) and
Rushton and Smith (1993), taking advantage of the
digital tools available in Adobe Photoshop. In essence,
the procedure is to determine the extent of distortion
by determining the strain ellipse (i.e. a distorted
initial circle), then calculate the factors to be applied
to return that ellipse to circularity. These factors are
then applied to images of individual fossils to restore
their original shape.
Strain ellipse
Several slabs containing a number of fossils with
unambiguous traces of their original hinge line and
longitudinal axis of symmetry (or the equivalents
in the case of trilobite cranidia) were photographed,
taking care to include a linear scale. The images were
then oriented with the trace of lineation horizontal.
Prints were then used to analyse strain following
Wellman’s method (see Cooper 1990, pp. 323-324),
with the ‘control line’ parallel to the lineation. The
Figure 1. Location of the collection from near
‘Lara’ homestead, east of Cooma. A: regional map.
B: geology in the vicinity of ‘Lara’ (derived from
Lewis and Glen, 1995). The approximate position
of the locality is marked by x. The undifferentiated
Yalmy Group is considered to be laterally equiva-
lent to the whole Bredbo Group (Cappanana For-
mation to Rothlyn Formation).
3
Proc. Linn. Soc. N.S.W., 135, 2013
D.L. STRUSZ
resulting points, to be used to estimate the strain
ellipse, were then plotted on a millimetre grid in
Photoshop. In a separate layer, an ellipse was drawn
through them using the ellipse tool, adjusting the
shape of the ellipse until it was seen to be a best
fi t for the plotted points (see Fig. 2). For each slab,
the correction factors needed to restore the ellipse
to circularity (on the assumption that the contained
area within the original circle remained constant)
were then calculated, again following the methods
described by Cooper (1990).
There are steps in this procedure by which
inaccuracies are inevitably introduced - determining
the original axis of symmetry (not easy if ornament
is uniform or internal structures are subdued or
diffuse), fi tting an ellipse to the plotted points, and the
assumption of unchanged area occupied by the fossil
during distortion. Added to these is the likelihood
for the ‘Lara’ specimens that not all individuals were
equally affected by compaction (see below) or lay
precisely on the original bedding plane. Consequently,
the calculated correction factors for the selected slabs
varied signifi cantly. There were insuffi cient suitable
slabs for a calculated mean to be useful. Many fossils
were on pieces with too few suitable individuals for
the above procedure to be appropriate. Therefore it
was decided to use correction factors rounded from
the median, and see how well they worked for the
majority of specimens - a potentially iterative process,
but fortunately the chosen factors (reduction in the
direction of lineation x0.6, extension at right angles
to that of x1.4) worked well in nearly all cases, so did
not need changing.
Application to individual fossils
Inevitably most of the fossils on the photographed
slabs were not well oriented relative to the direction
of lighting, and there were numerous fossils, not on
those slabs, which were needed for reliable taxonomy.
Therefore chosen fossils were photographed either
individually or in small groups of similar orientation,
again with the inclusion of a scale - in this case, a 6
mm diameter disc of graph paper (meaning that one
diameter would always be oriented correctly relative
Figure 2. Specimen Lara 24A (containing MMF45320-45334) with symmetry axes (see text) superim-
posed on individuals labelled a to f. Scale is in centimetres. Lower right: a control line 80 mm in length
is drawn parallel to the direction of maximum extension, its ends forming the control points referred
to by Cooper (1990). For each individual, lines drawn through these control points, parallel to the two
marked axes of symmetry, intersect at the labelled points (and their diagonally opposite counterparts).
The shaded area is the ellipse which appears to best fi t these intersections.
4Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
to the direction of photography, and could be used
to determine magnifi cation during processing of the
image). Care was taken to save a fresh fi le for each
image at each step in the subsequent process. Each
image was brought to an appropriate magnifi cation,
then rotated to bring the traces of lineation to either
horizontal or vertical orientation. The correction
factors were then applied, and their effectiveness
checked. Errors arose because the direction of lineation
was not always easily seen (meaning orientation of
the image could be not quite right), or in the case
of taxa such as globose atrypides or spiriferides
because orientation before compaction was not
always parallel to bedding before compaction, so
that the resulting distortion was uneven. This second
source of error also made orientating small specimens
for photography uncertain. Where the ‘restored’
images were not quite symmetric, the process was
repeated using the saved fi les, with slightly changed
orientation, until an optimum result was obtained.
The second source of error mentioned above meant
that full symmetry could not always be attained even
with several attempts. The fi nal ‘restored’ images
were then cropped to remove surplus background
(and signifi cantly reduce fi le size), and adjustments
made to colour balance (the artifi cial light used was
a ‘white’ LED, which resulted in very blue-biassed
images), to bring the images as close as possible
to the originals. Images of very dark brown fossils
were made more useful by changing brightness and
contrast. These images were then used for systematic
study of the fauna.
Limitations
As can be seen from the above discussion, this is
not a precise procedure. Provided strain was uniform
across the outcrop, a fair approximation to the original
proportions of symmetric fossils can be obtained.
However, in most cases the strain will vary from
bed to bed, depending on variations in lithology and
distance from the nearest fold axis. In the case of the
material from ‘Lara’, nearly all the specimens were
loose, the result of grading of strongly weathered rock,
and little bedding (let alone folding) could be seen in
the road-bed, so this limitation certainly applied!
Original size is not restored using this
methodology - again, because of the uncertainties in
orientation and strain analysis - but the images will be
a reasonable approximation of that size. Restoration
of proportions (such as length to width ratio) is more
accurate, but still not perfect.
As shown by Hughes and Jell (1992), and Rushton
and Smith (1993), this is a very useful tool in assessing
the relationships of taxa which have been described
from strongly distorted specimens, particularly when
there are many individuals available. As in the current
case, it is also useful when dealing with a distorted
fauna in which many of the taxa have been previously
described from good material from other localities.
Known taxa can often be identifi ed with confi dence.
However, it is not reasonable to base new taxa just on
images from which distortion has been removed. This
is particularly so when trying to discriminate between
species of the same genus, where size and proportions
are often critical and not infrequently overlap.
RESULTS
Ten brachiopod species have been recognised,
of which four can be confi dently identifi ed with
known taxa; another three are closely comparable
with known taxa but because of insuffi cient material
cannot be identifi ed with certainty. One species can
be placed in a known genus but not species, and the
fi nal two (represented by only a few specimens) can
be assigned to subfamilies. The fauna also contains
encrinurid trilobite fragments, mostly librigenae, and
a few moulds of rugose corals, including (Fig. 3) a
small syringaxonid with a massive columella.
The fauna of the Cappanana Formation farther
north, near Bredbo, has been listed previously, mostly
at the generic level, but the only species described
from the formation is the rugose coral Rhizophyllum
interpunctatum de Koninck, 1876, from Rock Flat
Creek southeast of Cooma. Richardson and Pickett
(p. 61 in Pickett 1982 ) commented that ‘precise
correlation is not possible until identifi cations have
been carried to specifi c level for as many forms as
possible, but the overall aspect of the fauna resembles
most closely that of the Canberra Group, and it is most
likely to be of Late Wenlockian or Ludlovian age.’
Of the brachiopods described here, the known
species elsewhere range in age from mid-Wenlock to
Figure 3. MMF45247, calical mould of an unde-
termined syringaxonid rugose coral.
5
Proc. Linn. Soc. N.S.W., 135, 2013
D.L. STRUSZ
early Ludfordian, with several known from Wenlock
rocks in Canberra but not above, and only one not
previously known below the Ludlow (it enters at Yass
at about the Wenlock-Ludlow boundary). This is not
enough for a defi nitive determination of the age of
the Cappanana Formation, but a Wenlock age is more
likely than not.
SYSTEMATIC PALAEONTOLOGY
Classifi cation follows that in the six volumes of
the revised Treatise on Invertebrate Paleontology,
part H (Williams et al. 1997-2007). References to
suprageneric taxa can be found in the relevant parts
of that work.
The specimens used in this study have been lodged
with the Geological Survey of New South Wales, their
catalogue numbers being prefi xed with MMF. Other
depositories cited are the Research School of Earth
Sciences, Australian National University (ANU), the
Commonwealth Palaeontological Collection held
by Geoscience Australia (CPC), and the Australian
Museum (AMF). The dimensions quoted are for
specimens as digitally restored, and are at best only a
reasonable estimate of the dimensions of the original
undistorted valves.
Abbreviations used are:
Ls - length of shell (normally the ventral valve)
Ld - length of dorsal valve
Lwmax - length to position of maximum width
Ws - shell width (except for alate species)
Wc - width of corpus (i.e. of the main body of alate
taxa)
Wh - width of hinge (including alae where relevant)
in strophic shells
CM - width of cardinal margin in nonstrophic shells
In lists of dimensions, italicised numbers indicate
estimated values (e.g. doubled from measured half-
width).
Class STROPHOMENATA Williams et al., 1996
Order STROPHOMENIDA Öpik, 1934
Superfamily STROPHOMENOIDEA King, 1846
Family RAFINESQUINIDAE Schuchert, 1893
Subfamily LEPTAENINAE Hall and Clarke, 1894
Leptaeninae, gen. et sp. indet.
Fig. 4
Material.
MMF45315 - ventral internal and dorsal external
moulds.
Description.
Outline transverse, subquadrate with rounded
margins, concave anteromedially; maximum width at
Figure 4. Leptaeninae, gen. et sp. indet., MMF
45315; a, b, dorsal external mould as collected,
and restored; c, d, ventral internal mould as col-
lected, and restored; e, posteromedian area of 4d,
in grey-scale with increased contrast, enlarged to
show the unusually shaped ventral muscle fi eld.
6Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
about 2/3 length. Ventral valve convex, dorsal valve
gently resupinate, with abrupt dorsal geniculation;
trail short. Dorsal surface radially ribbed, relatively
irregularly rugate; ribs rounded, 4-5 per mm
marginally. Ventral beak broad, fairly low; interarea
narrow, concave; delthyrium triangular, open,
pseudodeltidium obscure; foramen not seen.
Teeth narrow, diverge from median line at about
50°. Muscle fi eld subquadrate, with strong anterior
and lateral reentrant, extends to about 1/3 valve length;
in cross-section trough-shaped, gently impressed
medially and moderately raised laterally. Adductor
scars small, elongate elliptical, smooth. Floor of fi eld
not strongly ridged. Low, narrow myophragm starts
at mouth of anterior reentrant in muscle fi eld, extends
to a little beyond mid-length. No other details visible.
Dorsal interior unknown.
Dimensions Ls 5.0 mm, Ws 9.5 mm, Wh c. 8.6 mm,
Ls/Ws 0.53, Wh/Ws c. 0.9.
Remarks
This single individual cannot be easily assigned
to a leptaenine genus, particularly in the absence
of details of the dorsal interior. It does not closely
resemble any described species of Leptaena. A
weakly concave anteromedian margin is unusual but
not unknown in Leptaena - see Hoel (2005, Figs 2K,
P). The Siluro-Devonian leptaenine Glossoleptaena
Havlíček, 1967, has a concave anterior margin, but
this is associated with a distinct sulcus; also it has
much weaker shell ornament. The outline of the
ventral muscle fi eld in the ‘Lara’ specimen is quite
distinctive (see Fig. 4e), perhaps approached only by
that of Glossoleptaena pixis (Kelly, 1967).
Very few leptaenids have been described from
the Australian Silurian, none being similar to the
present species. An indeterminate Leptaena from the
Walker Volcanics in Canberra was fi gured by Strusz
(1982, p. 119). The single ventral internal mould
has a semi-oval outline and a small subtriangular
ventral muscle fi eld. Bracteoleptaena pannucea
Rickards and Wright, 1997 has a weakly developed
trail, and a fairly large cordate ventral muscle fi eld
with evenly curved muscle-bounding ridges to either
side of but not anterior to the fi eld. The valve fl oors
are noticeably papillate. Leptaena compitalis Strusz,
2003 has a fairly strong trail, and the ventral muscle
fi eld is small, rhomboid in outline with a concave
front contained within the well developed muscle-
bounding ridges.
Family EOPHOLIDOSTROPHIIDAE Rong and
Cocks, 1994
Genus MESOPHOLIDOSTROPHIA Williams, 1950
Type species
Pholidostrophia (Mesopholidostrophia) nitens
Williams, 1950. Wenlock, Gotland.
Mesopholidostrophia bendeninensis (Mitchell, 1923)
Fig. 5
Figure 5. Mesopholidostrophia bendeninensis
(Mitchell, 1923); a, b, MMF45238, ventral inter-
nal mould with well defi ned muscle fi eld, as col-
lected, and restored; c, d, MMF45329a, ventral
internal mould with poorly defi ned muscle fi eld,
distinct ventral process, and strong reentrants in
front of narrow alae, as collected, and restored; e,
f, MMF45250, incomplete dorsal internal mould
as collected, and restored.
7
Proc. Linn. Soc. N.S.W., 135, 2013
D.L. STRUSZ
Synonymy
Mesopholidostrophia bendinensis (Mitchell,
1923); Strusz, 2003, pp. 10-17, fi gs 7-10, cum syn.;
Strusz, 2010b, pp. 149-150, fi gs 3Q-S..
Type material
Lectotype AM F28789. Bowning Creek, near
Bowning, NSW. “Lower Trilobite Bed” = Black Bog
Shale; Ludfordian (Upper Silurian).
New material
MMF45233, 45236-45238, 45243, 45244,
45250, 45260-45262, 45273, 45303, 45310-45314,
45329-45331.
Description
Outline weakly to moderately transverse, semi-
oval, alate; alae narrow, reentrant at junction with
corpus of shell shallow or absent. Largest shell 16.3
mm long. Greatest width at 0.35-0.4 Ls. Profi le
concavo-convex, ventral valve moderately to strongly
convex, dorsal valve weakly to moderately concave.
Ornament absent, or weak with moderately irregular
rounded ribs, and sometimes sparse, irregular, weak
concentric wrinkles. Ventral interarea low, fl at,
apsacline; dorsal interarea not seen. Delthyrium open;
notothyrial structures not seen. Cardinal margin on
both valves denticulate to 0.4-0.5Wc.
Ventral muscle fi eld variably well expressed,
moderately to strongly impressed into valve fl oor
posteriorly, extends to about 0.4Ls; outline anteriorly
poorly defi ned, subtriangular to bilobed. Adductor
scars obscure, or weak, small, elongate, subparallel.
Muscle-bounding ridges divergent, straight, faintly
to moderately tuberculate, little if at all raised above
posterolateral valve fl oor. Ventral process small,
divided by shallow triangular furrow. Myophragm
fi ne to obscure. Valve fl oor outside muscle fi eld
variably tuberculate.
Cardinal process lobes elongate, narrowly
triangular, nearly parallel, separated by narrow slot
and supported by small, low notothyrial platform;
other details obscure. Notothyrial platform
continuous with socket ridges, muscle-bounding
ridges and myophragm. Inner socket ridges low,
slightly curved, tuberculate, weakly divergent from
cardinal margin. Muscle-bounding ridges curved,
initially fairly strongly divergent then anteriorly
only moderately so (at about 25-30°), tuberculate.
Myophragm long, narrow, may be slightly swollen
anteriorly. Notothyrial platform, muscle-bounding
ridges and myophragm combine to form anchor-
shaped structure posteriorly enclosing moderately
divergent, oval, smooth adductor scars. Remainder of
valve fl oor moderately tuberculate.
Dimensions
specimen Ls Wc Wh Ls/Wc Ls/Wh Wh/Wc
MMF45311 16.3 17.3 ≥21 0.94 ≤0.78 ≥1.2
MMF45329 10.5 14.5 16.3 0.72 0.64 1.12
MMF45238 15.0 16.3 ≥16.7 0.92 ≤0.9 ≥1.02
Remarks
There are three fairly similar species of
‘stropheodontid’ brachiopods in the Silurian of
southeastern Australia which had to be considered
when identifying this form: Mesoleptostrophia
(Mesoleptostrophia) quadrata (Mitchell, 1923),
Mesoleptostrophia (M.) oepiki Strusz, 1985, and
Mesopholidostrophia bendeninensis (Mitchell,
1923). The fi rst of these (from the Ludlow of Yass) is
fairly small, and can be recognised externally by its
subquadrate outline and the usually strong reentrants
where the relatively short alae join the sides of the
shell corpus; its ribs are fi ne and subdued. Internally
it has a large, triangular, posteriorly impressed
ventral muscle fi eld bounded by straight tuberculate
muscle-bounding ridges. The notothyrial platform,
inner socket ridges, dorsal muscle-bounding
ridges, and myophragm are all well developed.
Mesoleptostrophia (M.) oepiki (Wenlock, Canberra)
can be distinguished from it by its longer, thinner alae
separated from the valve corpus by weaker reentrants,
coarser ornament, a fi ne ventral myophragm, and
curved dorsal muscle-bounding ridges. It also tends
to be larger. Mesopholidostrophia bendeninensis
is closer to the latter in size, and in having long
slender alae and weak reentrants. It differs from
both species of Mesoleptostrophia in its very weak
external ornament, a ventral muscle fi eld which is
less obviously triangular and is often bilobed, weak
ventral myophragm and muscle-bounding ridges,
weak inner socket ridges, notothyrial platform and
muscle-bounding ridges, and a longer myophragm
which may expand slightly anteriorly.
The material from ‘Lara’, where many
features are not well preserved, is so similar
to both Mesoleptostrophia (M.) quadrata and
Mesopholidostrophia bendeninensis that it is very
unlikely to be a new species of either genus. The
very weak external ornament and often signifi cant
convexity alone suggest the latter, and internal
features, particularly the form of the ventral muscle
fi eld and the relatively weak dorsal structures, confi rm
the specifi c identity.
8Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
Since the revision of this species was published,
a locality low in the Bowspring Limestone Member
at Yass (locality GOU57 - see Strusz 2005) yielded a
few specimens. This locality is below the Ancoradella
ploeckensis zone, and could be either earliest Ludlow
or latest Wenlock.
Order ORTHOTETIDA Waagen, 1884
Suborder ORTHOTETIDINA Waagen, 1884
Superfamily CHILIDIOPSOIDEA Boucot, 1959
Family CHILIDIOPSIDAE Boucot, 1959
Subfamily CHILIDIOPSINAE Boucot, 1959
Genus MORINORHYNCHUS Havlíček, 1965
Type species
Morinorhynchus dalmanelliformis Havlíček,
1965. Ludlow, Bohemia.
Morinorhynchus oepiki Strusz, 1982
Fig. 6
Synonymy
Morinorhynchus oepiki Strusz, 1982; Strusz
2003, pp. 31-33, fi gs 20-21, cum syn.; Strusz, 2010b,
pp. 140-150, fi gs 3K’-M’.
Type material
Holotype CPC20987, paratypes CPC20415-
20419, 20988-20994. Molonglo Valley, Canberra,
ACT. Walker Volcanics; probably Homerian,
Wenlock.
New material
MMF45234, 45240, 45246, 45249, 45252-
45255, 45264-45267, 45271, 45274, -45277, 45298-
45302, 45304, 45320-45327, 45335-45340, 45348-
45353, 45355.
Figure 6. Morinorhynchus oepiki Strusz, 1982; a-d, MMF45298, a, b, ventral external mould as collected,
and restored; c, d, ventral internal mould as collected, and restored; e, f, MMF45255, incomplete dorsal
external mould showing large chilidium extending posterior to hinge line, as collected and restored; g, h,
MMF45264b, dorsal internal mould as collected, and restored.
9
Proc. Linn. Soc. N.S.W., 135, 2013
D.L. STRUSZ
Description
Shell gently to moderately biconvex, ventral
valve moderately more convex than dorsal valve;
outline semi-elliptical, Wh ca 0.95Ws, Ls ca 0.75Ws
but very variable, greatest width at about 0.4Ls.
Anterior margin rectimarginate to moderately sulcate,
dorsal sulcus most prominent in large shells. Ventral
umbo low, interarea low, fl at, strongly apsacline;
pseudodeltidium large, moderately convex; foramen
small, apical. Dorsal umbo low, interarea very
low, chilidium broad, convex, projects posteriorly.
Ornament unequally parvicostellate, with strong,
rounded ribs separated by wider fl at-bottomed
interspaces crossed by fi ne growth lines; at 5 mm
radius, at least 20 ribs in 5 mm.
Teeth small, dental plates well developed, upright,
gently curved outward, divergent at about 90°. Muscle
fi eld obscure. Socket plates well developed, curved
outwards, strongly divergent, fused with bilobed
cardinal process; myophragm weak, broad. Further
details obscure.
Dimensions
specimen Ls Ws Wh Ls/Ws Wh/Ws
MMF45266 11.5 18.0 17.5 0.64 0.97
MMF45298 17.5 23.5 22.8 0.74 0.97
MMF45326 22.0 25.0 23.0 0.88 0.92
MMF45353 15.8 20.5 18.8 0.77 0.92
Remarks
The specimens from east of Cooma are
morphologically (allowing for the poorer preservation)
the same as Wenlock specimens from the Canberra
Formation (Strusz, 1985) and those from the Ludlow
of Yass (Strusz, 2003). As noted by Strusz (2003), M.
oepiki is now known to reach a larger size than the
type specimens from the Walker Volcanics indicated.
The largest new specimen is MMF45326, with Ls 22
mm, Ws 25 mm, comparable with MMF610 (Ls 17
mm, Ws 26.1 mm) from Bowning. Rib spacing is on
the low side, but within the previously known range.
Unlike elsewhere, where Morinorhynchus oepiki
is a minor part of the fauna, the collection from ‘Lara’
is dominated by this species.
Class RHYNCHONELLATA Williams et al., 1996
Order PENTAMERIDA Schuchert and Cooper, 1931
Suborder PENTAMERIDINA Schuchert and
Cooper, 1931
Superfamily PENTAMEROIDEA M’Coy, 1844
Family PENTAMERIDAE M’Coy, 1844
Genus APOPENTAMERUS Boucot and Johnson,
1979
Type species
Apopentamerus racinensis Boucot and Johnson,
1979. Wenlock, Wisconsin.
cf. Apopentamerus clarkei Strusz, 2011
Fig. 7
Synonymy
cf. Apopentamerus clarkei Strusz, 2011, pp. 36-
39, fi g. 5.
Material
MMF45256, 45305, 45306, 45328.
Discussion
These four small shells are clearly a smooth
pentameroid, but only the exterior (completely lacking
ribs, but one has a few concentric wrinkles) and the
ventral interior are known. The ventral median septum
is very long, and supports a narrow and upwardly
fl aring spondylium. There is considerable similarity
with smaller specimens of Apopentamerus clarkei,
recently described from the Canberra Formation,
although the specimens from ‘Lara’ are even smaller
Figure 7. cf. Apopentamerus clarkei Strusz, 2011;
a, b, MMF45256, ventral internal mould as col-
lected, and restored; c, d, MMF45305, ventral in-
ternal mould as collected, and restored.
10 Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
(compare especially MMF45305 with Strusz 2011,
fi gs 5e, h, and MMF45256 with fi g. 5c). I am
reasonably confi dent that this is the same species, but
the material from ‘Lara’ is insuffi cient for certainty.
Order ATRYPIDA Rzhonsnitskaya, 1960
Suborder ATRYPIDINA Moore, 1952
Superfamily ATRYPOIDEA Gill, 1871
Family ATRYPIDAE Gill, 1871
Subfamily ATRYPINAE Gill, 1871
Genus ATRYPA Dalman, 1828
Subgenus ATRYPA Dalman, 1828
Type species
Atrypa reticularis Linnaeus, 1758. Ludlow,
Gotland.
Atrypa (Atrypa) sp. cf. duntroonensis Mitchell and
Dun, 1920
Fig. 8
Synonymy
cf. A. (Atrypa) duntroonensis Mitchell and Dun,
1920; Strusz, 2011, pp. 39-44, fi gs 6-9, cum syn.
Material
MMF45269, 45281, 45302, 45318, 45319.
Discussion
Available material is very limited, with no well
preserved ventral valves. Until the recent redescription
of A. (Atrypa) duntroonensis by Strusz (2011), based
on abundant new material from the Wenlock of
Canberra, only one species of Atrypa was reasonably
well known from the Silurian of southeastern
Australia: A. (A.) sp. cf. dzwinogrodensis Kozłowski,
1929 of Strusz (1984, 2007) from the Late Wenlock?
to Ludlow of Yass. The two highly variable species
are very similar externally, A. (A.) duntroonensis
tending to be somewhat more elongate, with the
dorsal valve more often anteromedially extended as
a broad ventrally directed tongue. Rib shape, density
and mode of increase are the same in the two species.
In the dorsal valve, A. (A.) duntroonensis has a deeper
cardinal pit, the sockets have distinct median ridges,
and diverge at about 110-120°, and the myophragm is
less prominent than in A. (A.) sp. cf. dzwinogrodensis.
MMF45318 is the only available dorsal internal
mould, and has a broad but rather low myophragm.
The socket plates are short but robust, and the sockets
(whose fi ner details are obscure) diverge at more than
140°. On this basis, the few specimens from ‘Lara’
are most comparable with A. (A.) duntroonensis, but
specifi c identity must remain uncertain.
Suborder LISSATRYPIDINA Copper, 1996
Superfamily LISSATRYPOIDEA Twenhofel, 1914
Family LISSATARYPIDAE Twenhofel, 1914
Genus ATRYPOIDEA Mitchell and Dun, 1920
Subgenus ATRYPOIDEA Mitchell and Dun, 1920
Type species
Meristina (?) australis Dun, 1904. Ludlow,
Molong, NSW.
Atrypoidea (Atrypoidea) australis (Dun, 1904)
Fig. 9
Synonymy
Atrypoidea (Atrypoidea) australis (Dun, 1904);
Strusz 2007, pp. 322-331, fi gs 16-21, cum syn.
Material
MMF45258, 45263, 45270, 45272, 45280,
45307-45309, 45232, 45233.
Figure 8. Atrypa (Atrypa) sp. cf. duntroonensis
Mitchell and Dun, 1920; a, b, MMF45302b, dor-
sal external mould as collected, and restored; c-
f, MMF45318, c, d incomplete dorsal external
mould as collected, and restored, e, f, dorsal inter-
nal mould as collected, and restored.
11
Proc. Linn. Soc. N.S.W., 135, 2013
D.L. STRUSZ
Discussion
The limited available material is mostly not well
preserved, but fi ts well within the morphological
limits of this highly variable species, and there can
be little doubt of its identity. The one good ventral
internal mould, MMF45263, is very like AMF17432
(Strusz 2007, Fig. 18C) in outline. With Ls = 18.5
mm, Ws = 17.0 mm, CM = 10.5 mm and Lwmax ≈
7mm, it fi ts well within the published plots of Ls:
Ws and CM:Ws, and differs only in the relatively
posterior position of maximum width (it is just
within the limits of the published plot of Lwmax:
Ls). Its internal structures are also quite comparable
with the few known interiors (e.g. ANU9736, Strusz
2007, fi g. 17L). The same can be said of the dorsal
valves - MMF45308 is close to the dorsal valves of
AMF29186 and 29195 (Strusz 2007, fi gs 17G,19C),
and MMF45309 to AMF29183, 29184 (Strusz 2007,
fi gs 17E, F) and 29188 (fi g. 18B).
Order ATHYRIDIDA Boucot, Johnson and Staton,
1964
Suborder Incertae Sedis
Superfamily ANOPLOTHECOIDEA Schuchert,
1894
Family ANOPLETHECIDAE Schuchert, 1894
Subfamily COELOSPIRINAE Hall and Clarke,
1895
Genus COELOSPIRA Hall, 1863
Type species
Leptocoelia concava Hall, 1857. Lochkovian,
New York.
Coelospira sp. indet.
Fig. 10
Material
MMF45317, 45334.
Discussion
The two diminutive specimens are damaged and
incomplete ventral valves of undoubted anoplothecid
morphology. Both show strong curved ribs fl anking a
more fi nely ribbed fold. The one internal mould is not
well enough preserved to show details. Nevertheless
the basic morphology indicates this form could only
belong to Coelospira.
Figure 9. Atrypoidea (Atrypoidea) australis (Dun, 1904); a, b, MMF45263, ventral internal mould as col-
lected, and restored; c, d, MMF45308a, dorsal internal mould as collected, and restored; e, f, MMF45309,
dorsal internal mould as collected, and restored.
12 Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
Coelospira has been recorded from the late
Wenlock to early Ludlow of the Yass-Canberra region
(C. cavata Strusz, 1982), and Cadia in central NSW
(Coelospira sp. Percival in Rickards et al., 2001). The
one incomplete ventral valve from Cadia differs from
those of C. cavata only in having fewer, stronger
ribs on the fold. In this respect the specimens from
‘Lara’ more closely resemble the Cadia form, but lack
of material displaying the range of morphological
variability means the specifi c relationships of the
Cadia and ‘Lara’ forms remain unknown.
Order SPIRIFERIDA Waagen, 1883
Suborder SPIRIFERIDINA Waagen, 1883
Superfamily CYRTIOIDEA Frederiks, 1924
Family CYRTIIDAE Frederiks, 1924
Subfamily EOSPIRIFERINAE Schuchert, 1929
Eospiriferinae gen. et sp. indet.
Fig. 11
Material
MMF45282-45286.
Description
Only ventral valves are known, represented by
relatively small external and internal moulds. Outline
transverse spiriferoid, length 3/4 - 7/8 width, with
Figure 10. Coelospira sp. indet.; a-d, MMF45334,
a, b, incomplete ventral internal mould as collect-
ed, and restored, c, d, incomplete ventral external
mould as collected, and restored.
Figure 11. Eospiriferinae gen. et sp. indet.; a-d,
MMF45285, a, b, ventral external mould showing
well developed capillate ornament, as collected,
and restored, c, d, ventral internal mould with
very long parallel dental plates, restored, and as
collected, e, f, MMF45284a, damaged ventral in-
ternal mould with shorter parallel dental plates,
as collected, and restored.
13
Proc. Linn. Soc. N.S.W., 135, 2013
D.L. STRUSZ
prominent umbo, incurved beak, rounded lateral and
anterior margins; greatest width somewhat anterior
to mid-length. Valve strongly convex, without sulcus
or ribs; anterior margin rectimarginate. Micro-
ornament of rounded capillae separated by somewhat
wider interspaces, Eospirifer-like, increasing by
intercalation. Concentric micro-ornament absent or
too fi ne to be preserved.
Dental plates parallel, long, extending to or
beyond mid-length. Muscle fi eld obscure, but weak
myophragm between dental plates in some specimens,
starting anterior to apex. Teeth small, triangular. One
internal mould shows a small delthyrial plate. No
visible pallial markings.
Dimensions
specimen Ls Ws Ls/Ws
MMF45284 6.0 6.8 0.88
MMF45285 5.3 7.2 0.74
MMF45286 6.5 8.4 0.77
Discussion
In the absence of dorsal valves the generic
identity of this form cannot be determined, but it is
clearly a smooth eospiriferine. It is close to Eospirifer
Schuchert, 1913, especially in the form of the micro-
ornament, but species of that genus generally have
robust divergent extrasinal dental plates, and usually
have at least a shallow sulcus. In the absence of a
sulcus, it cannot be said whether the dental plates in
the ‘Lara’ form are intrasinal or extrasinal. Moreover,
Eospirifer lacks a delthyrial plate or deltidium. Close
to Eospirifer is Endospirifer Tachibana, 1981, which
is known from the Silurian of Canberra and Yass,
differing in having a well developed deltidium. It
has fairly long and only moderately divergent dental
plates which, however, lie outside a narrow but
generally well developed sulcus. Nurataella Larin,
1973 (= Baterospirifer Rong, Su and Li, 1984) lacks
a sulcus and has a rudimentary delthyrial plate, but
the dorsal valve is of low convexity, and there are
short extrasinal dental plates. Moreover, its capillae
are more like those of Myriospirifer Havlíček, 1978,
which are broadly rounded to fl attened, and increase
exclusively by splitting. The only eospiriferine known
to me with parallel dental plates is the diminutive
Nanattegia Strusz, 2010a, from the Silurian of Yass,
but that has concave fl anks on the very convex ventral
valve, a deep, narrow sulcus, and no apical delthyrial
structures.
Suborder DELTHYRIDINA Ivanova, 1972
Superfamily DELTHYRIDOIDEA Phillips, 1841
Family DELTHYRIDIDAE Phillips, 1841
Subfamily HOWELLELLINAE Johnson and Hou,
1994
Genus HOWELLELLA Kozłowski, 1946
Type species
Terebratula crispa von Hisinger, 1826. Wenlock,
Gotland.
Howellella sp. aff. elegans (Muir-Wood, 1925)
Fig. 12
Synonymy
Howellella sp. aff. elegans (Muir-Wood, 1925);
Strusz, 1982, pp. 132-134, fi gs 27A, C-E; Strusz,
1985, p. 117, fi gs 12A, 13; Strusz 2010b, fi gs 6, 8A-
C
Material
MMF45239, 45241, 45242, 45259, 45279,
45287-45297, 45341-45344, 45345-45347.
Description
Ventral valve fairly strongly convex, outline
spiriferoid, umbo wide and fairly prominent, lateral
and anterior margins rounded; Ls/Ws and Wh/Ws
both about 0.8. Ventral sulcus well developed, defi ned
by two strong, sharply rounded ribs; At most, one pair
of subdued lateral ribs arises anterior to beak. Dorsal
valve moderately convex; beak low, fold prominent,
rounded to slightly fl attened medially, fl anked by
strong furrows. Inner pair of lateral ribs lower but well
developed, rounded, slightly curved laterally. Outer
pair of lateral ribs low, developed only anterolaterally;
there can be a very faint third pair of ribs developed
only marginally. Ventral interarea concave, apsacline;
delthyrium open - structures otherwise obscure.
Dorsal interarea very low, weakly anacline. Micro-
ornament of fi mbriate growth lines.
Teeth small, dental plates robust, upright,
divergent forward, extra-sulcal, extend forward to
0.3-0.4Ls. Myophragm low, commences at about
level of cardinal margin, extends forward to about
mid-length. Muscle fi eld obscure. Sockets small,
widely divergent; inner socket ridges well developed,
outer hinge plates narrow, steep, crural bases short,
supported by triangular crural plates convergent on
low notothyrial platform. Cardinal process small, its
ventral face bilobed (MMF45290, 45341, 45346) or
trilobed (MMF45342). Myophragm weak or absent;
muscle scars obscure.
14 Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
Discussion
While preservation of detail in these small
specimens is rather poor, in all that can be seen they
are the same as the material from the Walker Volcanics
and Canberra Formation of Canberra .
Superfamily RETICULARIOIDEA Waagen, 1883
Family RETICULARIIDAE Waagen, 1883
Subfamily RHENOTHYRIDINAE Gourvennec,
1994
Genus SPIRINELLA Johnston, 1941
Type species
Spirinella caecistriata Johnston, 1941. Late
Wenlock, Yass, NSW.
Spirinella caecistriata Johnston, 1941
Fig. 13
Synonymy
Spirinella caecistriata Johnston, 1941; Strusz
2011, pp. 45-46, fi g. 12, cum syn.
Material
MMF45245, 45248, 45268, 45278, 45316,
45354, 45356, 35357.
Figure 12. Howellella sp. aff. elegans (Muir-Wood, 1925) Strusz, 1982; a, b, MMF45287a, ventral internal
mould as collected, and restored; c, d, MMF45288a, ventral internal mould as collected, and restored;
e-h, MMF45342a, dorsal internal mould, e, f, as collected, and restored, g, tilted posteriorly (restored)
to show cardinalia, enlarged in h; i-m, MMF45344, i, j, dorsal external mould as collected and restored,
and k, enlargement showing fi mbriate ornament, l, m, dorsal internal mould as collected, and restored;
5 mm scale bar applies to a-g, i, j, l, m, 2 mm scale bar applies to h, k.
15
Proc. Linn. Soc. N.S.W., 135, 2013
D.L. STRUSZ
Remarks
These few small specimens are not well
preserved, most showing evidence of post-burial
compaction prior to tectonic distortion. Nevertheless
they show all the characteristics of the species, and
the proportions fi t well within the limits of variability
reported by Strusz (2010a). Ventral valves have a
weak ventral sulcus, fairly prominent beak, fi mbriate
micro-ornament, strong, divergent dental plates and a
weak myophragm; two internal moulds show a well-
Figure 13. Spirinella caecistriata Johnston, 1941; a-e, MMF45278, ventral valve external and internal
moulds, a, b, as collected, c, d restored, e portion of c reoriented and enlarged to show fi mbriate orna-
ment; f, h, MMF45354a, incomplete ventral internal mould as collected, and restored; g, j, MMF45357,
incomplete ventral mould with large apical callist, viewed postero-ventrally, as collected and restored; j,
k, MMF45316a, dorsal internal mould as collected, and restored; 5 mm scale bar applies for all except
e, to which 2 mm scale bar applies.
16 Proc. Linn. Soc. N.S.W., 135, 2013
SILURIAN BRACHIOPODS FROM SOUTHERN NEW SOUTH WALES
developed apical callist - e.g. compare MMF35357
with Strusz (2010a, fi g. 18G). The one dorsal internal
mould is damaged by compaction, so no useful
comparison can be made.
Dimensions
specimen Ls Ws Wh Ls/Ws Wh/Ws
MMF45278 9.0 12.0 11.5 0.75 0.96
MMF45354 4.8 5.5 4.5 0.87 0.82
ACKNOWLEDGEMENTS
This paper was prepared in the Research School of
Earth Sciences (ANU) in my capacity as a School Visitor,
and also as a Research Associate of the Australian Museum,
Sydney, and is a contribution to IGCP591. I would like
to thank Dr T. Munson (now with the Northern Territory
Geological Survey) for his assistance in collecting the
material, Dr I. Percival and Dr R.A. Glen (New South
Wales Geological Survey) for assistance in sourcing an
appropriate geological map of the area, and Dr J. Laurie
(Geoscience Australia) for his careful review of the fi rst
draft of the paper.
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