Silurian Brachiopods from the Historic Woolshed Creek Area,

Canberra, Australia
DESMOND L. STRUSZ

Department of Earth and Marine Sciences, Research School of Earth Sciences, Australian National
University, Canberra ACT 0200 (desmond-strusz@homemail.com.au)

Published on 21 November 2011 at http://escholarship.library.usyd.edu.au/journals/index.php/LIN

Strusz, D.L. (2011). Silurian brachiopods from the historic Woolshed Creek Area, Canberra, Australia.
Proceedings of the Linnean Society of New South Wales 133, 33-51.

The brachiopod fauna of seven species from the Canberra Formation at Woolshed Creek near Duntroon,
Canberra, is revised. Four species were collected by W.B. Clarke in 1844 and represent the first formal
recognition of Silurian rocks in Australia. Referred to European species and genera by de Koninck in
1876, these are now recognised as Mesoleptostrophia (Mesoleptostrophia) oepiki, Morinorhynchus oepiki,
Apopentamerus clarkei n.sp. and Atrypa (Atrypa) duntroonensis. The last dominates the fauna, and was first
described by Mitchell and Dun in 1920; a Canberra atrypid compared by Strusz in 1985 to the Victorian
Early Devonian species Spinatrypa perflabellata is now known to be A. (A.) duntroonensis. Salopina
mediocostata, Hedeina oepiki, and Spirinella caecistriata are also present, together with uncommon
trilobites (mostly Batocara mitchelli), corals, gastropods and bryozoans, some of which are illustrated. The
fossils occur in repeated crowded layers which probably represent storm deposits. The age is most likely to
be Homerian (latest Early Silurian).

Manuscript received 10 June 2011, accepted for publication 15 November 2011.

Key words: Apopentamerus, Atrypa, brachiopods, Canberra, corals, Hedeina, Homerian,

Mesoleptostrophia, Morinorhynchus, Salopina, Silurian, Spirinella, trilobites.

much later led to an area along Woolshed Creek,

INTRODUCTION including the original outcrop, being registered as a
Geological Heritage Site. Clarke’s fossil collections
Woolshed Creek runs south to join the Molonglo were subsequently examined by L.G. de Koninck
River a little to the east of Royal Military College, of Leuven University (Belgium), and published by
Duntroon (eastern Canberra), where it is crossed by him in 1876-7. The collections were destroyed in
Fairbairn Avenue. Duntroon was the family home of the Garden Palace fire of 1882 in Sydney. Mitchell
Robert Campbell, a wealthy merchant and grazier and Dun recollected Clarke’s site, and in 1920
who first took up land along the Molonglo River in described Atrypa duntroonensis based on three rather
1825. The property was managed by James Ainslie; poor specimens. Öpik (1958, figs 19, 20), published
Campbell only lived at Duntroon after his wife’s death photos of the outcrop north of Fairbairn Avenue, at
in 1833, and died there in 1846. In 1844, Rev. W.B. which time layers crowded with brachiopods could
Clarke, pastor and geologist, visited the area during be seen. Unfortunately since then there has been
his travels in southern New South Wales on behalf much deterioration, so that the site is no longer as
of the colonial government, and undoubtedly stayed spectacular.
with Campbell (whom he would have known from his
period as headmaster of the Kings School from 1839 New material
to 1841, a school established partly at Campbell’s At the beginning of the 21st Century traffic along
instigation). While at Duntroon, Clarke collected Fairbairn Avenue became sufficiently heavy that the
fossils from the vicinity, and from Yarralumla a few ACT government proposed duplicating the existing
kilometres to the west. He recognised that these bridge across Woolshed Creek. This was to be sited
fossils were of undoubted Silurian age, a conclusion just downstream of the existing bridge, in a part of the
he published in 1848. This was the first published heritage site lacking outcrops - hence not disturbing
identification of Silurian fossils in Australia, and the existing exposure. Excavations for the bridge
 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

abutments were carried out in 2008, and yielded large and partly due to tectonic forces. The degree of
quantities of fossiliferous material. Opportunity was distortion is variable, probably because of different
taken to collect specimens, and these form the basis amounts of compaction depending on the amount
of the present paper. and nature of the silt enclosing the fossils. Shells
The fauna is dominated by A. duntroonensis. The are randomly oriented on the bedding planes, so
fossils occur in repeated layers crowded with shells, that analysis of size and proportions is feasible but
with mostly dissociated but unbroken or only slightly can only be approximate (see remarks in Strusz
damaged convex upwards valves indicating that 1985a:112-114).
most layers are probably storm deposits from below
normal wave base (Fig. 1). Occasional layers contain Stratigraphy and age
syringoporoid corals which appear to be in growth The Woolshed Creek locality was included by
position, and are less crowded with shells, suggesting Öpik (1958) in his Riverside Formation, which is
these were subject to less disturbance before being now regarded as part of the lithologically variable
buried by silt. Seven species of brachiopod have been Canberra Formation. Unfortunately there is no reliable
identified in the collections. Of these, only one - a evidence from conodonts or graptolites for the age
very rare smooth pentameride - is new, and overall of this unit, which must therefore be argued on the
the fauna is typical of the Canberra Formation, as basis of stratigraphic correlation with the succession
described by Strusz (1985a). Interestingly, four of at Yass, north of Canberra; for a fuller discussion see
the brachiopods (including the rare pentameride) Strusz (2010b). The most likely age for the Canberra
were collected by Clarke and identified reasonably Formation has been deduced to be late Sheinwoodian
accurately as European Silurian species by de Koninck to early Homerian. From mapping by the Engineering
(1876-7), who consequently (but unfortunately) Geology section of the former Australian Bureau of
neither described nor illustrated them. Mineral Resources (now Geoscience Australia), it is
All the material is distorted, partly by compaction likely that the Woolshed Creek locality is high in the

Figure 1. Large excavated slab of siltstone from Woolshed Creek, showing successive layers crowded
with Atrypa valves and shells, mostly moulds and convex-upwards.

34 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

formation, which would indicate an early Homerian Pholidostrophiidae? gen. et sp. indet. Strusz,
age (see Strusz and Henderson 1971, Henderson and 1985a:111, Fig. 6.
Matveev 1980). M. (Mesoleptostrophia) oepiki (Strusz); Strusz
2010b, Figs 2, 3K-M.

SYSTEMATIC PALAEONTOLOGY Type material

Holotype CPC24751, paratypes CPC24744-
Specimens registered with the Australian 24750, 24752-24783, 24877, 24880-24883,
Museum, Sydney, have numbers prefixed AM F, those 24903-24905, 24917-24922. Canberra Formation,
with the Department of Earth and Marine Sciences, Fyshwick, ACT. Wenlock.
Research School of Earth Sciences, Australian
National University are prefixed ANU, and those Woolshed Creek material. AM F.110137-110141,
in the Commonwealth Palaeontological Collection, 110193-110196, 110212-110214, 110225-110226,
Geoscience Australia, Canberra, are prefixed CPC. 110253-110254, 110259, 110263, 110268, 110276-
Geoscience Australia and the Research School of 110280, 110316-110318, 110350, 110368, 110413-
Earth Sciences, Australian National University, also 110418, 110423-110424.
hold unregistered bulk material.
Classification follows that in the Treatise Diagnosis (new)
on Invertebrate Paleontology, part H (Revised). Moderately concavo-convex Mesoleptostrophia
References to supra-generic taxa can be found of medium to large size, uniformly costellate, with
in the relevant parts of the Treatise. Only Atrypa long narrow alae, denticulation to 1/3 width of corpus;
duntroonensis and the new pentameride are fully low ridges posterolaterally bounding subtriangular,
described. posteriorly strongly impressed ventral muscle field,
lateral to which valve floor is coarsely tuberculate;
fine ventral myophragm; prominent notothyrial
Class STROPHOMENATA Williams et al. 1996 platform continuous with dorsal myophragm and pair
Order STROPHOMENIDA Öpik, 1934 of often prominent curved muscle-bounding ridges.
Superfamily STROPHOMENOIDEA King, 1846
Family LEPTOSTROPHIIDAE Caster, 1939 Discussion
Genus MESOLEPTOSTROPHIA Harper and The description given by Strusz (1985a)
Boucot, 1978 is generally adequate, except that the outline is
Subgenus MESOLEPTOSTROPHIA Harper and erroneously described there as elongate (instead of
Boucot, 1978 transversely) semi-elliptical. Ls/Ws (excluding alae)
varies in the Fyshwick material between 2/3 and
Type species 3/4. The material from Woolshed Creek agrees well
Mesoleptostrophia kartalensis Harper and with that from Fyshwick, although in some bands the
Boucot, 1978, p. 68 [= Strophodonta (Leptostrophia) distortion is greater.
explanata Paeckelmann and Sieverts, 1932, non Two species of M. (Mesoleptostrophia) are now
Sowerby, 1842]. Early Devonian, Turkey. known from the Silurian of the Yass-Canberra region
(see Strusz, 2010b). M. (M.) oepiki differs from the
Mesoleptostrophia (Mesoleptostrophia) oepiki somewhat younger M. (M.) quadrata (Mitchell, 1923)
(Strusz, 1985) in its long, slender alae separated from the less quadrate
Fig. 2 corpus by weaker reentrants, coarser ribs, no anterior
sulcus in gerontic shells, a fine ventral myophragm,
Synonymy and curved dorsal muscle-bounding ridges. M. (M.)
Leptaena compressa, J. de C. Sowerby; de oepiki is readily distinguished from the similarly
Koninck 1876:27; non Orthis compressa alate Mesopholidostrophia bendeninensis (Mitchell,
J. de C. Sowerby, 1839, which is 1923) by its stronger ornament, weaker convexity,
Mesoleptostrophia (Mesoleptostrophia) stronger ventral and dorsal muscle-bounding ridges,
compressa - see Cocks 2008:70; de Koninck and stronger notothyrial platform.
(transl. Dun in David, David and Dun) The large dorsal valve (CPC24884) considered
1898:22. by Strusz (1985a) to be a probable pholidostrophiid is
Leptostrophia (Leptostrophiella) oepiki Strusz, distinctive in that its strong muscle-bounding ridges
1985a:110-111, Figs 4-5. and myophragm are supplemented by two distinct

Proc. Linn. Soc. N.S.W., 133, 2011 35

 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

Figure 2. Mesoleptostrophia (Mesoleptostrophia) oepiki (Strusz, 1985); a, AM F.110193, ventral internal

mould; b, AM F.110194, ventral internal mould; c, AM F.110141, incomplete ventral internal mould
showing long narrow ala and concave lateral margin; d, AM F.110413, incomplete dorsal internal mould
clearly showing bifid cardinal process, denticulate cardinal margin; e, AM F.110415, incomplete dorsal
internal mould showing relatively robust myophragm and muscle-bounding ridges; f, AM F.110317,
crushed gerontic dorsal internal mould showing more complex muscle field, including curved bounding
ridges and short anderidia - compare CPC24884 (Strusz 1985a, fig. 6).

36 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

ridges within the adductor muscle field. One large

dorsal internal mould from Woolshed Creek (AM
F.110317, Fig. 2f) has a similar complex of structures,
and I now think it is most likely that the Fyshwick
specimen is a gerontic M.(M.) oepiki.
The specimens collected by Clarke from
“Duntroon” and reported (but not figured) by de
Koninck (1876) can confidently be referred to M.
(M.) oepiki.

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, p. 291. Ludlow, Bohemia.

Morinorhynchus oepiki Strusz, 1982

Fig. 3

Synonymy
Strophomenes pecten, Linnaeus; de Koninck
1876:28; non Anomia pecten Linnaeus,
1767, which is Coolinia pecten - see Cocks,
2008:99; de Koninck (transl. Dun in David,
David and Dun) 1898:22.
Morinorhynchus oepiki Strusz, 1982:119-122,
Figs. 14-15; Strusz, 1985a:111-112, Fig. 7;
Strusz, 1985b:681-682, Figs 3.1-9; Strusz
2003:31-32, Fig. 20; Strusz, 2010b, Figs 2,
3 K’-M’.
Morinorhynchus oepiki?; Strusz, 1985b:681-682,
Fig. 3.10.

Type material
Holotype CPC20987, paratypes CPC20415-
20419, 20988-20994. Walker Volcanics, Canberra.
Late Wenlock.

Woolshed Creek material Figure 3. Morinorhynchus oepiki Strusz, 1982; a,

AM F.110165, 110297, 110335, 110346, AM F.110297, incomplete and rather strongly dis-
110369-110370 torted ventral internal mould; b, AM F.110346,
small ventral internal mould; c, AM F.110335,
Diagnosis dorsal external mould; d, AM F.110370, dorsal in-
Thin, nearly planoconvex to slightly resupinate ternal mould.
Morinorhynchus with fine unequally parvicostellate
Remarks
ornament (Strusz 2003).
The few small specimens from the Woolshed
Creek excavations are very similar to those described

Proc. Linn. Soc. N.S.W., 133, 2011 37

 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

by Strusz (1985a) from the Canberra Formation in

Fyshwick, and the remarks made therein still apply. It
is an uncommon species at all known localities.
This is almost certainly the species from
“Duntroon” referred by de Koninck (1876) to Anomia
pecten Linnaeus, 1767, a species which has since
been referred to the related and quite similar genus
Coolinia Bancroft, 1949.

Class RHYNCHONELLATA Williams et al., 1996

Order ORTHIDA Schuchert and Cooper, 1932
Suborder ORTHIDINA Schuchert and Cooper, 1932
Superfamily ENTELETOIDEA Waagen, 1884
Family DRABOVIIDAE Havlíček, 1950
Subfamily DRABOVIINAE Havlíček, 1950
Genus SALOPINA Boucot in Boucot et al., 1960

Type species
Orthis lunata J. de C. Sowerby, 1839. Ludlow,
Shropshire.

Salopina mediocostata Strusz, 1982

Fig. 4

Synonymy
Salopina mediocostata Strusz,1982:111-114, Figs
5-7; Strusz 1984:125-126, Figs 2-3; Strusz,
1985a:108, Fig. 3; Strusz 2002:67-70, Figs
11-13; Strusz, 2010b: Figs 4, 5 M-P. Figure 4. Salopina mediocostata Strusz, 1982; a, AM
F.110130, small ventral internal mould - compare
Type material Strusz, 1985a, fig. 3C; b, AM F.110372, small ven-
Holotype CPC20337, paratypes CPC20253- tral internal mould; c, AM F.110227, incomplete
20336, 20338-20344, 20932-20942. Walker dorsal internal mould - compare Strusz, 1985a, fig.
Volcanics, Wenlock, Canberra. 3G; d, AM F.110374, dorsal internal mould.

Woolshed Creek material

AM F.110127-110132, 110227, 110255, the presence of the enlarged median costella on the
110371-110374, 110396-110397. dorsal valve, characteristic of the Walker Volcanics
specimens, cannot be confirmed.
Remarks
This species is not common at Woolshed Creek,
and nearly all of the specimens are quite small - the Order PENTAMERIDA Schuchert and Cooper, 1931
largest measured is AM F.110227 with a width of 7.2 Suborder PENTAMERIDINA Schuchert and
mm, but most are less than 4 mm across. However, Cooper, 1931
they are closely comparable with previously published Superfamily PENTAMEROIDEA M’Coy, 1844
specimens, especially the similarly distorted Family PENTAMERIDAE M’Coy, 1844
Fyshwick material (Strusz 1985a) e.g. compare the Genus APOPENTAMERUS Boucot and Johnson,
ventral internal mould of AM F.110130 (Fig. 4a) with 1979
CPC24731 (1985a, fig. 3C), and the dorsal internal
mould AM F.110227 (Fig. 4c) with CPC24913 Type species
(1985a, fig. 3G). Unfortunately all available external Apopentamerus racinensis Boucot and Johnson,
moulds are incomplete and not well preserved, so 1979. Wenlock, Wisconsin, USA.

38 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

Diagnosis (new) trilobate), ‘..suggests that Sulcipentamerus is more

Smooth non-lobate pentamerine, moderately closely related to Pentamerus than to Harpidium.’
to strongly biconvex with ventral valve usually (Rong et al. 2007:249). It seems to me, from the above,
deeper than dorsal valve; outline transversely oval that Apopentamerus is best treated as a valid genus
to pyriform; ventral median septum long, supporting distinct from both Pentamerus and Harpidium.
narrow spondylium; inner hinge plates subparallel,
long, their junction with outer hinge plates smooth, Apopentamerus clarkei n.sp.
without flanges. Fig. 5

Discussion Synonymy
The diagnosis is based on the original diagnosis Pentamerus oblongus, J. Sowerby; de Koninck
of Boucot and Johnson, plus the discussion of 1876:31; Jenkins 1879:26, 30, pl. 6, fig. 3;
pentamerine relationships by Rong, Jin and Zhan Mitchell 1887:1198; de Koninck (transl.
(2007) analysed below. Dun in David, David and Dun) 1898:24-25.
Boucot and Johnson erected Apopentamerus Pentamerus australis M’Coy; Etheridge
for a Wisconsin pentameride species differing from 1892:51-52, non M’Coy 1877 (which is
Isovella Breivel and Breivel in Antsigin et al., 1970, Meristella - Gill 1951:32).
in the presence of a spondylium, and from Harpidium
Kirk, 1925, in its significantly longer ventral median Etymology
septum. The dorsal interior is as in Pentamerus In honour of the Rev. W.B. Clarke, who first
J. Sowerby, 1813. Sapel’nikov (1985) considered collected this species from Woolshed Creek.
Isovella and Apopentamerus to be synonyms of
Harpidium. Type Material
Boucot, Rong and Blodgett (in Kaesler, 2002), Holotype AM F.110260, Paratypes AM
treated Isovella as a subgenus of Harpidium, F.110168, 110298-110299, 110314-110315, 110343-
characterised by a moderately long ventral median 110345, 110358, 110398-110399.
septum and substantial thickening in the ventral umbo.
They considered Apopentamerus to be a synonym of Other Material
H. (Isovella), and Sulcipentamerus Zeng, 1987, to be ANU46537, from Yass locality KC48 (see
a third subgenus of Harpidium characterised by its Strusz 2002), is tentatively identified as A. clarkei.
elongate and markedly ventribiconvex shell, and a
short ventral median septum. Distribution
Sulcipentamerus has been extensively revised on Canberra Formation, Canberra; basal Bowspring
the basis of large collections by Rong, Jin and Zhan Limestone and probably Yarwood Siltstone Member,
(2007) and Jin, Harper and Rasmussen (2009). In Yass. Early Homerian to early Ludfordian.
their discussion of generic relationships, they noted
that Isovella has laterally projecting crural bases Diagnosis
(the flanges of Boucot et al. in Kaesler 2002) at the Large strongly pyriform ventribiconvex
junction of inner and outer hinge plates - flanges not Apopentamerus with prominent ventral beak, long
seen in Harpidium, Apopentamerus, and Pentamerus ventral median septum, narrow spondylium, very
itself. As this is a feature previously recognised as long subparallel inner hinge plates.
taxonomically useful, Rong et al. (2007: 248) noted
‘This implies that Apopentamerus is either a valid Description
genus or a subgenus of Harpidium and that Isovella is Moderately sized, smooth, ventribiconvex shell
most likely to be a genus independent of Harpidium.’. of pyriform to subtriangular outline with prominent
These authors also pointed out that Boucot et al. postero-dorsally directed ventral beak, low dorsal
(in Kaesler 2002) regarded Sulcipentamerus as a beak. Ventral area small, triangular, gently concave,
subgenus of Harpidium on the assumption that it sharp-edged; delthyrium partly closed by concave
lacked trilobation. However, the strong variability crescentic pseudodeltidium. Anterior commissure
shown by the large Chinese collections (including (where preserved) appears to be broadly and very
species with a weak tendency in large shells to gently sulcate, and in one dorsal valve there is a
trilobation), and also by collections of Pentamerus possible faint sulcus. The largest, rather strongly
oblongus J. de C. Sowerby, 1839, from Estonia distorted, specimen is 25.4 mm long and about 44
and Norway (ranging from non-lobate to markedly mm wide.

Proc. Linn. Soc. N.S.W., 133, 2011 39

 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

Figure 5. a-g, Apopentamerus clarkei n. sp., Canberra Formation, Woolshed Creek, probably early
Homerian; a, b, holotype AM F.110260, ventral internal mould in ventral and postero-ventral views, the
latter clearly showing the narrow spondylium; c, d, paratype AM F.110345, damaged ventral internal
mould in ventral and posterior views, the latter revealing the small pseudodeltidium; e, f, paratype AM
F.110344, small ventral internal mould in ventral and posterior views; g, paratype AM F.110168, dorsal
internal mould showing the very long subparallel inner hinge plates; the anterior median furrow sug-
gests the possibility of a similar external furrow. h, ?Apopentamerus clarkei, ANU46537, Yarwood Silt-
stone Member, Black Bog Shale, Yass (locality KC48), early Ludfordian; incomplete ventral mould in
ventral view, showing the long septum supporting a narrow spondylium.

Ventral median septum high, extending to between hinge plates anterior to mid-length in some
between 1/3 and 2/3 but generally about half valve specimens.
length; spondylium narrow, upwardly flaring, of
uncertain length but apparently not reaching mid- Discussion
length. There may be a little thickening umbonally in From the previous discussion of generic
large specimens. Teeth unknown. relationships, these few specimens are most likely
Inner hinge plates subparallel and fairly close, to be either Sulcipentamerus or Apopentamerus. The
extending along valve floor to near anterior valve former is highly variable externally, with some species
margin; posteriorly they are gently medially convex, being noticeably trilobate while others are non-lobate;
and merge smoothly with more strongly convex the shell is markedly ventribiconvex, the dorsal
outer hinge plates. Crural bases obscure; no flanges. valve sometimes flat or concave, and more generally
Dental sockets narrow, diverge at about 90°. Outer bears a gentle sulcus. Apopentamerus is much less
socket ridges flat-topped. Low, narrow median ridge strongly ventribiconvex, and characteristically has a

40 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

faint median furrow on each valve, giving a slightly A. (Atrypa) duntroonensis Mitchell and Dun, 1920
emarginate outline. Moreover, the general shell shape Figs 6-9
in Sulcipentamerus is elongate, even subcylindroidal,
whereas Apopentamerus is generally transverse, Synonymy
often subpyriform. Internally, the median septum in Atrypa reticularis, Linnaeus; de Koninck
Sulcipentamerus is shorter than in Apopentamerus, 1876:35; de Koninck (transl. Dun in David,
while the spondylium is wider and relatively long. David and Dun) 1898:26.
On balance, therefore, I consider the Woolshed Creek ?Atrypa? hemisphaerica, J. de C. Sowerby; de
specimens belong in Apopentamerus. Koninck 1876:35; de Koninck(transl. Dun
This species is undoubtedly that identified by in David, David and Dun) 1898:26-27.
de Koninck (1876) as Pentamerus oblongus, an
equally smooth northern hemisphere species from
which it clearly differs in its wide pyriform shape.
The incomplete dorsal internal mould from near
Hattons Corner, Yass, figured by Jenkins (1879, pl.
6, fig. 3) as Pentamerus oblongus is almost identical
to specimens AM F.110314. 110315 and 110344.
Etheridge (1892:1198) thought Jenkins’ specimen
could be identified with Pentamerus australis M’Coy,
1878, but that species, from the Early Devonian of
Lilydale, Victoria, was transferred to Meristella by
Gill (1951:320) and is clearly unrelated. ANU46537,
a single incomplete ventral internal mould from Yass
locality KC48 and previously thought unidentifiable,
is now recognised to be probably A. clarkei, and is
figured here (Fig. 5h).
Apopentamerus clarkei differs from the type
species A. racinensis Boucot and Johnson, 1979, in a
rather longer ventral median septum, and an upwards-
flaring spondylium. From A. szechuanensis Rong et
al., 2007, it differs in its much greater convexity,
gently sulcate anterior commissure, and lack of
median furrows on ventral and most dorsal valves. It
differs from A. muchuanensis Rong et al., 2007, also
in its longer ventral median septum and inner hinge
plates.

Order ATRYPIDA Rzhonsnitskaya, 1960

Suborder ATRYPIDINA Moore, 1952
Superfamily ATRYPOIDEA Gill, 1871
Family ATRYPIDAE Gill, 1871
Subfamily ATRYPINAE Gill, 1871
Genus ATRYPA Dalman, 1828 Figure 6. Atrypa (Atrypa) duntroonensis Mitchell
Subgenus ATRYPA Dalman, 1828 and Dun, 1920; a, b, paralectotype AM F29202
in dorsal and ventral views - original of Mitchell
Type species and Dun, pl. XVI, figs 9, 12; c, paralectotype AM
Anomia reticularis Linnaeus, 1758; Ludlow, F29201 in ventral view - original of pl. XVI, fig.
Gotland. 8; d-f, lectotype AM F29203 in dorsal, ventral and
posterior views - original of pl. XVI, fig. 10. Note
the significant distortion of the ventral valve vis-
ible in the posterior view of the lectotype.

Proc. Linn. Soc. N.S.W., 133, 2011 41

 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

Type material
Mitchell and Dun did not designate a holotype,
and figured four specimens, which are thus syntypes.
The originals of Plate XVI, figs 8-10 are registered
with the Australian Museum as AM F29201-29203;
the original of fig. 11 is missing. I here designate
F29203, the original of Pl. XVI, fig. 10, as lectotype.
The remaining syntypes thus become paralectotypes.

Type locality
Mitchell and Dun described their locality as
‘near Duntroon homestead....’ The only fossiliferous
outcrops in that area are in Woolshed Creek, which
flows south into the Molonglo River just east of
the Royal Military College, Duntroon. De Koninck
reported that the specimens collected by Rev. W.B.
Clarke in 1844 and examined by him came from
Duntroon, and Mitchell and Dun were quite certain
that their species was the same as that collected by
Clarke. The outcrop where preservation matches that
of the type specimens is just north of Fairbairn Avenue
where it crosses Woolshed Creek; this locality is now
a designated Geological Heritage Area.

New material
Large numbers of specimens have been collected
from the excavations (see Introduction) about 20
m along strike from the type locality: only those
providing useful information for the redescription
of this species have been registered. These are AM
F.110119-110126, 110145-110150, 110154-110155,
110166-110167, 110173-110192, 110207-110211,
110217, 110228-110231, 110235-110251, 110257-
110258, 110264-110265, 110269-110272, 110282-
110294, 110300-110310, 110325-110331, 110336-
110342, 110347-110349, 110359-110366, 110375-
110386, 110400-110409, 110419.

Horizon, age
Figure 7. Atrypa (A.) duntroonensis; plots of shell Canberra Formation, Wenlock. In the absence of
length Ls, hinge width Wh, and the ratio of shell conodonts and graptolites, the precise age cannot be
length to width against shell width Ws - the cross determined, but as discussed above the most likely age
in the first two is the lectotype. The significant for the Woolshed Creek locality is early Homerian.
change in slope in the plot of Wh:Ws suggests that
maturity, indicated by an increasingly wide cardi- Diagnosis (new)
nal margin, is reached at a width of about 12-13 Small to medium sized biconvex to dorsibiconvex
mm. Atrypa with short marginal frills, impressed ventral
Atrypa duntroonensis; Mitchell and muscle field with raised anterior rim, relatively
Dun,1920:270-271, pl. XVI, figs 8-12. delicate teeth and crural plates, and dental cavities
Atrypa sp. cf. perflabellata (Talent, 1963); Strusz retained in adult shells.
1985a:112-114, figs 8A-S.
Spinatrypa sp. cf. perflabellata; Strusz 2010b, Description - exterior
figs 6, 7U-W. Adult shells biconvex to dorsibiconvex,
moderately to strongly swollen, with outline varying

42 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

Figure 8. Atrypa (A.) duntroonensis; a, b, AM F.110210, ventral internal mould and latex replica showing
strong vascular impressions; c, AM F.110207, ventral internal mould; d, AM F.110251, incomplete large
ventral internal mould with weakly expressed muscle field (see also Fig. 9b); e, AM F.110328, partly
exposed steinkern in posterior view with very convex dorsal valve, 3-ridged cardinal process; f, AM
F.110419, latex replica of two adjacent ventral valves with deeply impressed anteriorly scalloped and
somewhat raised muscle fields (see also Fig. 9c); g, AM F.110269, incomplete laterally compressed ven-
tral internal mould showing impressions of narrow triangular teeth; h, AM F.110383, incomplete strong-
ly laterally compressed juvenile ventral internal mould, showing relatively narrow cardinal margin; i,
AM F.110342, moderately convex steinkern in lateral view; j, AM F.110147, longitudinally compressed
juvenile dorsal internal mould; k, AM F.110408, laterally compressed juvenile dorsal internal mould; l,
AM F.110329, dorsal internal mould - cardinalia enlarged below to show corrugated sockets and outer
hinge plates; m, AM F.110124, large dorsal internal mould with robust crural plates; n, AM F.110407,
large distorted dorsal internal mould. The infill of dental cavities is clearly visible in Figs 8c, d, g, h.

Proc. Linn. Soc. N.S.W., 133, 2011 43

 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

but variability is high. The two largest specimens are

ventral valves with Ls 18.0 mm, Ws 24.0 mm, and
Ls 23.0 mm, Ws 21.3 mm. Four steinkerns and two
whole shells including the lectotype are between 15.0
and 22.0 mm wide, and their Ts/Ws varies between
0.46 and about 0.7. The least swollen specimens tend
to be nearly equibiconvex, whereas in large swollen
shells the dorsal valve can be twice the thickness
of the ventral valve. In transverse profile ventral
valve is medially rounded, may be weakly carinate
posteriorly, and has almost planar flanks, while dorsal
valve is evenly rounded. Anterior commissure gently
to fairly strongly uniplicate. In longitudinal profile
ventral valve is most convex posteriorly, flattening
anteriorly, frequently forming very shallow sulcus
which can extend dorsally as a tongue. Marginally,
where growth lamellae are very crowded, valve
surface often turned strongly dorsally. Dorsal valve
evenly convex, frequently with flattened to upturned
margins. Dorsal umbonal region with median
depression which dies out anteriorly. Ventral beak
low, wide, suberect to gently incurved, generally
closely adpressed over extremely low dorsal beak.
Large parabolic delthyrium occupies most of small
apsacline ventral area, and mostly truncates beak.
Ribs low, 5-12 but mostly 6-8 in an arc of 5 mm at
5 mm radius, slightly coarser medially than laterally.
Ventral valve has two raised ribs postero-medially,
from length of about 6 mm separated by very fine
single mid-rib. Increase is usually by splitting, always
at growth lamellae, and most often episodic at radii
of about 6-7 and 12-13 mm; there may be some
intercalated ribs towards adult shell margins. Growth
lamellae faint umbonally, then low, about 1 mm apart,
becoming very crowded in narrow marginal zone,
where they may form short oblique frills up to 1 mm
Figure 9. Atrypa (A.) duntroonensis; detailed views
long. The lamellae curve forward in inter-rib troughs,
of external and internal structures; a, AM F.110348,
but do not form spinose outgrowths. A narrow skirt
incomplete external mould showing short growth
(about 2-3 mm) extending dorsally from ventral valve
lamellae, and short marginal frill; b, AM F.110251,
margin is occasionally preserved.
latex replica of large ventral valve in antero-dor-
sal view, showing dental cavities (that on the right
Interior
was occupied by a bubble, so clearly shows the
Teeth transversely elongate, of moderate
significant size); c, AM F.110419, latex replica of
size, separated by narrow transverse grooves from
large ventral valve in antero-dorsal view, showing
cardinal margin; well-preserved moulds show weak
slit-like dental cavities, raised anterior margin to
corrugations. Dental cavities slit-like cones, widely
muscle field.
divergent, reduced but always present even in large
shells. Muscle field medium to large, flabellate to
from rounded to shield-shaped or subquadrate.
subtriangular, variably but often strongly impressed,
Cardinal margin highly variable in width, and weakly
longitudinally furrowed with anteriorly raised rim
to fairly strongly curved but not incurved. Most adult
which may be finely grooved. Adductor scars small,
shells are 15 to 20 mm wide, with mean Ls/Ws 1.05
slightly raised either side of median depression
(overall mean is 1.066); a plot of Ls/Ws against Ws
posteriorly placed within muscle field. Pedicle callist
suggests a trend to less elongate shells with growth,
weak, separated anteriorly from muscle field by fine

44 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

concentric ridge; a few specimens show what appears Gratsianova and Yolkin (2001) in reverting to the
to be a small pedicle collar at apex of delthyrium. original assignment, pending the results of the
Mantle canal system saccate, variably but sometimes present study. The relationship between the Silurian
strongly impressed on valve floor. and Pragian taxa is not clear, although on the basis of
Cardinal pit deep, triangular in juveniles but slot- internal structures one could speculate that the latter
like in adults, with fine ridges at apex forming cardinal could have been derived from the former.
process, and floor raised a little above general valve A. (A.) duntroonensis is clearly the species
floor. Crural plates robust in large shells, fine in small identified by de Koninck (1876-7) as A. reticularis.
shells, forming deep triangular impressions in internal While the destruction of Clarke’s collections precludes
moulds. Sockets narrow, widely divergent, somewhat absolute certainty, I am confident in the light of the
arcuate in all but smallest valves, corrugated; inner variability shown by the current collections from
socket ridges flat-topped, often also corrugated; outer Woolshed Creek that the specimens identified by de
socket ridges fine, overhang sockets. Fairly prominent Koninck as A. hemisphaerica were juvenile A. (A.)
forward-expanding myophragm in juveniles, usually duntroonensis.
with fine median ridge, frequently becomes subdued As noted by many authors up to and including
or obscure in large shells. Mantle canal system Copper (2004:35), distinguishing between species
obscure. Two medium-sized worn steinkerns show and even genera of Silurian atrypids can be difficult,
dorso-medially directed spiralia of 9-10 whorls; no and there has been considerable disagreement in
further details are visible. deciding the boundaries between, and content of, the
various genera and subgenera that have been erected.
Ontogeny In general, I follow Copper (in Kaesler 2002; 2004).
Juvenile shells lenticular, rounded to elongate, Doing so, however, leads me to recognise in the
with sharp nearly straight ventral beak, narrow curved present species characters used to distinguish several
cardinal margin, biconvex to somewhat dorsibiconvex Silurian genera: Atrypa itself, Protatrypa Boucot,
profile. Ventral umbonal region generally moderately Johnson and Staton, 1964, Gotatrypa Struve, 1966,
to fairly strongly carinate posteriorly, dorsal umbo Rugosatrypa Rzhonsnitskaya, 1975, and Oglupes
with narrow sulcus. Margins without crowded frills Havlíček, 1987.
or skirt. External sculpture strongly imprinted on The dorsibiconvex shape of most larger A. (A.)
valve floors in thin-shelled juveniles, generally lost in duntroonensis, in which the anterior part of the
thicker-shelled adults. Dental cavities well developed, ventral valve is flattened and even gently sulcate, is
diverging at 90-120°, ventral muscle field generally very typical of A. (Atrypa) but can also be seen in
not noticeably impressed. Sockets prominently some Oglupes - e.g. O. visbyensis Copper, 2004 (see
corrugated. The size at which shell morphology Copper’s Pl. 12, figs a-e). Generally, though, Oglupes
changes from juvenile to adult appearance is highly is a very globose form with a more even longitudinal
variable, with some relatively large shells retaining convexity to the ventral valve, and more prominent
an almost juvenile appearance apart from greater ribs.
convexity and more deeply impressed ventral muscle The Llandovery Protatrypa is small and
field. Particularly for ventral valves, however, the lenticular, with a carinate ventral valve, very subdued
change from juvenile to adult morphology is usually growth lamellae, and dental cavities in adult shells
very marked. - the last not generally known in Atrypa. Juvenile
A. duntroonensis thus resemble adult Protatrypa,
Discussion but adults can be readily distinguished by greater
Morphological variation is considerable and, convexity, more prominent growth lamellae, and the
while overlain by post-burial distortion, it is apparent presence of short marginal frills.
that in common with many species of Atrypa much Gotatrypa is fairly small, biconvex to
of this is inherent variability. With the material now dorsibiconvex (Copper 2004, Pl. 10, fig. Bb), with
available it is quite clear that A. (Atrypa) duntroonensis short frills as in A. duntroonensis, but dental nuclei
is conspecific with the material from the Canberra rather than dental cavities, and deltidial plates in
Formation of Fyshwick compared by Strusz (1985a) juveniles. Moreover the ventral valve does not
with Spinatrypa perflabellata Talent, 1963, from become flat or gently sulcate anteriorly, and the
the Pragian of Victoria. In my discussion in 1985 I growth lamellae are projected as short spines in the
considered Talent’s species to be Atrypa rather than inter-rib troughs.
Spinatrypa, while in my recent biostratigraphic Rugosatrypa is not well illustrated and the
summary (Strusz 2010b) I chose to follow Talent, original description is brief, but it would appear to

Proc. Linn. Soc. N.S.W., 133, 2011 45

 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

differ from Atrypa in smaller size, a lenticular profile, Type species

regular rather widely spaced concentric growth rugae Anomia crispa Linnaeus, 1758. Ludlow,
but low growth lamellae not forming frills or a skirt, Gotland.
the presence of deltidial plates in adults, a weak
pedicle callist, and prominent dental cavities. A. Hedeina oepiki Strusz, 2010
(A.) duntroonensis differs from it in a more swollen (Figs 10-11)
profile, no concentric rugae or deltidial plates, and
reduced dental cavities in large shells. Synonymy
On balance, therefore, I think the present species Howellella sp. aff. elegans (Muir-Wood, 1925);
is best retained in Atrypa (Atrypa), differing from Strusz 1982 partim (Fig. 27B only).
other species by consistently retaining distinct dental Cyrtiidae gen.? et sp. nov.; Strusz 1985a:115-
cavities in adult shells. 117, Figs 11-12.
Hedeina oepiki Strusz, 2010a:103-104, fig. 13;
Strusz, 2010b: Figs 6, 8 I-K.
Order SPIRIFERIDA Waagen, 1883
Superfamily CYRTIOIDEA Frederiks, 1924 Type material
Family CYRTIIDAE Frederiks, 1924 Holotype CPC24671, paratypes CPC24831-
Subfamily EOSPIRIFERINAE Schuchert, 1929 24870. Canberra Formation, Fyshwick ACT;
Genus HEDEINA Boucot, 1957 Wenlock.

Figure 10. Hedeina oepiki Strusz, 2010; a, b, AM F.110252, ventral external mould with shape of dental
lamellae shown by infill of the delthyrial cavity, and enlargement showing capillae preserved in the
sulcus; c, AM F.110390, longitudinally compressed ventral internal mould with weak ribs on flanks; d,
AM F.110355, ventral internal mould; e, AM F.110160, ventral internal mould; f, AM F.110135, distorted
dorsal internal fold; g, h, AM F.110262, dorsal internal mould and enlargement showing ctenophorid-
ium; i, AM F.110133, incomplete dorsal internal mould with well developed myophragm. 2 mm scale bar
applies to Figs 10b, h.

46 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

Strongly distorted small specimens with weak

lateral ribs can be difficult to distinguish from
Endospirifer anxius Strusz, 1982, in which one or
rarely two faint lateral ribs are sometimes present.
That species, however, has a suboval rather than
subtriangular outline, less divergent dental plates
which are barely extrasinal, and no notothyrial
platform.

Superfamily RETICULARIOIDEA Waagen, 1883

Family RETICULARIIDAE Waagen, 1883
Subfamily RHENOTHYRIDINAE Gourvennec,
1994
Genus SPIRINELLA Johnston, 1941
Figure 11. Hedeina oepiki Strusz, 2010; plot of shell
length Ls against width Ws for specimens from the Type species
Canberra Formation at Woolshed Creek (●) and Spirinella caecistriata Johnston, 1941. Late
Fyshwick (x). Wenlock, Yass, NSW.
Woolshed Creek material
AM F.110133-110135, 110142-110144, 110151- Spirinella caecistriata Johnston, 1941
110152, 110156-110162, 110169-110171, 110197- Fig. 12
110206, 110219-110224, 110252, 110262, 110267,
110273-110274, 110295-110296, 110319-110324, Synonymy
110354-110357, 110387-110395, 110420-110421. Meristina (?) australis Shearsby 1912:112-113,
non Dun 1904.
Distribution, age Spirinella caecistriata Johnston, 1941:161-
Canberra Formation and Walker Volcanics, 167, Pl. VII, figs 1-11; Johnson, Boucot
Canberra; Wenlock (late Sheinwoodian? to and Murphy 1976: Pl. 28, figs 17-27;
Homerian).

Remarks
The specimens from the Woolshed Creek
locality show even greater variation in the degree
of distortion than those described from Fyshwick.
Allowing for that, and the effects it will have on any
statistical comparison, the present material agrees in
every important respect with Hedeina oepiki from
Fyshwick (respective means for specimens from
Fyshwick and Woolshed Creek: Ls/Ws 0.64 / 0.74,
Wh/Ws 0.71 / 0.80, Wf/Ws 0.31 / 0.27). The ribs vary
in form and number, but those flanking the ventral
sulcus are always noticeably more prominent than the
next lateral pair, and the dorsal fold is strong, its crest
somewhat flattened. Internally, the ventral myophragm
varies from low to quite strong; the dental plates are
robust, clearly extrasinal, and extend to about one-
third valve length. In the dorsal valve the crural plates
are strongly recessive, and there is a low notothyrial
platform supporting an apical ctenophoridium. There
can be no doubt that populations of just one species Figure 12. Spirinella caecistriata Johnston, 1941;
occur at Woolshed Creek and Fyshwick. A comparison a, AM F.110234, small incomplete dorsal inter-
with the younger species H. bruntoni can be found in nal mould; b, AM F.110266, large dorsal internal
Strusz (2010a). mould - compare Strusz, 1985a, fig. 14C.

Proc. Linn. Soc. N.S.W., 133, 2011 47

 SILURIAN BRACHIOPODS FROM WOOLSHED CREEK

Strusz 1984:144-147, Figs 18-19; Strusz ACKNOWLEDGEMENTS

1985a:117-118, Fig. 14; Strusz 2005:29-30,
Fig. 1; Strusz, 2010a:112-117, Figs 18-20; I would like to thank Luke Williams (Guideline ACT
Strusz, 2010b, Figs 6, 8V-X. site engineer for the Woolshed Creek bridge construction)
Reticulariopsis silurica Strusz, 1982:134-136, for his willing and interested cooperation during sample
collection and subsequent transport of bulk material to
Fig. 28.
ANU at Black Mountain, Canberra. Cooperation by the
Office of the Environment and Heritage, ACT Department
Type material of Territory and Municipal Services, was also excellent -
Holotype AM F39376. Paratypes AM F39378, my thanks for interesting discussions with Euroka Gilbert in
39379 and figured topotype AM F39377 have since particular. Material was collected by John Laurie and Peter
been lost - see Strusz (1984, 2005). Yass Formation Butler of Geoscience Australia, and myself. This paper
(Cliftonwood Limestone Member), Yass, NSW; has benefited greatly from discussion with Paul Copper on
Homerian. Silurian atrypid discrimination. I am also grateful to Ian
Percival for reviewing the initial draft of the paper.The work
was done in the Department of Earth and Marine Science
Woolshed Creek material
(ANU) in my capacity as a School Visitor, and also as a
AM F.110163-110164, 110172, 110215-110216, Research Associate of the Australian Museum, Sydney.
110232-110234, 110266, 110311-110313, 110367,
110410.
REFERENCES
Diagnosis (Strusz 2010a)
Suboval, moderately ventribiconvex Spirinella
Antsigin, N.Y., Breivel, I.A., Breivel, M.G., Varganov, V.G.,
with prominent ventral umbo, erect to slightly
Nasedkina, V.A. and Yanet, F.E. (1970). Pogranichnye
incurved beak; interarea concave, weakly apsacline, otlozheniya mezhdu ordovikom i silurom na srednem
not well delineated laterally. Teeth small, triangular; i prilegayushchei chasti severnogo Urala i ikh
dental plates long, moderately divergent, continued paleontologicheskaya kharakteristika. Materialy po
anteriorly by grooves of vascula media; ventral paleontologii Urala, Instituta Geologii i Geokhimii,
muscle field generally somewhat impressed, elongate, Uralski Filial, Akademiya Nauk SSSR, Sverdlovsk, 3-
longer than dental plates; delthyrial plate or apical 12, 52-56, Pls I-III.
thickening small, crescentic; crural plates narrow, Bancroft, B.B. (1949). Welsh Valentian brachiopods. In
triangular, more or less convergent downwards, rest Welsh Valentian brachiopods and the Strophomena
posteriorly on small notothyrial platform; lanceolate antiquata group of fossil brachiopods. (Ed. A. Lamont)
dorsal adductor field and myophragm. pp. 2-10, Pls I-II. Preprint from The Quarry Managers’
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Yorkshire.
Remarks
Boucot, A.J. (1957). Revision of some Silurian and
Spirinella caecistriata is a minor component of Early Devonian spiriferid genera and erection of
the Woolshed Creek fauna. The few specimens are Kozlowskiellinae, new subfamily. Senckenbergiana
not well preserved, but are clearly conspecific with Lethaea 38, 311-334, Pls 1-3.
the similarly distorted specimens from Fyshwick. In Boucot, A.J. and Johnson, J.G. (1979). Pentamerinae
particular, compare the dorsal internal mould AM (Silurian Brachiopoda). Palaeontographica, Abteilung
F.110266 (Fig. 12b) with CPC24876, figured Strusz A 163, 87-129.
1985a, Fig. 14C. The largest measurable specimens Boucot, A.J., Johnson, J.G. and Staton, R.D. (1964). On
are over 16 mm wide, with Wh/Ws varying some atrypoid, retzioid, and athyridoid Brachiopoda.
considerably around 0.73 - this compares well with Journal of Paleontology 38, 805-822.
the data from undistorted material in Strusz (2010a, Boucot, A.J., Martinsson, A., Thorsteinsson, R.,
tables 8 and 9). Walliser, O.H., Whittington, H.B. and Yochelson, E.
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trilobite Batocara mitchelli, rugose and tabulate corals from the Yass Basin. Palaeontographica, Abteilung A
of which a tryplasmatid, a columnar halysitid and a 167, 77-119, Pls 1-16.
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of the corals are shown in Fig. 13.

48 Proc. Linn. Soc. N.S.W., 133, 2011

 D.L. STRUSZ

Figure 13. Moulds of corals and trilobites associated with the brachiopod fauna at Woolshed Creek; a-
b, Tryplasma sp., a, AM F.110218, b, AM F.110256; c-d, Halysites sp., AM F.110351, columnar corallum
and enlargement of one smooth-sided palisade showing macrocorallites and intervening microcorallites;
e-f, Entelophyllum? sp., AM F.110261; g-h, undetermined diminutive cyathaxoniid?, g, AM F.110153, h,
AM F.110281; i, cheirurine pygidium cf. Cheirurus sp. Chatterton and Campbell, 1980, AM F.110422; j,
Uriarra kausi Chatterton and Campbell, 1980, AM F.110411, pygidium; k-l, Batocara mitchelli (Foerste,
1888), k, AM F.110275, pygidium, l, AM F.110136, small incomplete cranidium; m, n, indet. calymenid
AM F.110332, small pygidium in dorsal and posterior views. All scale bars 5 mm long.
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