MEMOIRS

OF THE

UEENSLAND MUSEUM

BRISBANE VOLUME 32
29 JUNE, 1992 PART 1

ANOPSILANA BARNARDI, A NEW SPECIES OF ESTUARINE CIROLANID
CRUSTACEAN ISOPOD FROM TROPICAL EASTERN AUSTRALIA

NIEL L. BRUCE

Bruce, N.L. 1992 06:29: Anopsilana barnardi, anew species of estuanne cirolanid crustacean
isopod from tropical castem Australia. Memoirs of the Queensland Museum 32(1): 1-8.

Brisbane. ISSN 0079-8835.

Anopsilana barnardi sp.nov., the third species of Anopsilana recorded from Australia, is
described from northeastem Queensland. A key to the Australian species is given anda list of
all Anopsilana species is provided.[] Jsopoda, Cirolanidae, new species, northeastern

Queensland, Southwest Pacific, taxonomy.

Niel L. Bruce, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101,

Anstralia; 10 January, 1992.

Anopsilana Paulian & Deboutteville, 1956, con-
tains thirteen species. Its occurrence is restricted
to the tropical regions of the world; about half the
species are recorded from troglobitic habits (both
freshwater and euryhaline), the others are all
recorded from mangrove or estuarine habitats.

Anopsilana barnardi sp.nov. was taken from a
mid-stream grab sample in the Murray River, and
il is therefore not directly associated with
mangroves as are the other Australian species of
Anopsilana.

Anopsilana Paulian & Deboutteville

Anepsilana Paulian & Deboutteville, 1956: 87. Bruce,
1981; 955, fig. 5i-e; 1986: 196; Kensley & Schotte,
1989: 124; Brusca, Wetzer & France (in press).

Troglocirolana Rioja, 1956: 447 [Type species:
Cirolana cuberisis Hay, 1903, by monotypy].

Haitilana Notenboom, 1981: 314 [Type species:
Haitilana radicicola Notenboom, 1981, original
designation).

TYPE SPECIES

Anopsilana poissoni Paulian & Deboutteville,
1956, original designation. The location of the
types was not given in the original publication, but
was cited as at the Institute Scientific de
Madagascar, Tsimbazaza, Antananarivo,
Madagascar by Brusca et al. (in press).

DIAGNOSIS

Fronal lamina usually elongate (in two Pacific
species as long as wide), antenor margin project-
ing or sessile; pentagonal or with anterior margin
rounded or truncate. Antennule with peduncular
urticles 1 and 2 usually coalesced. Pereopods all
ambulatory. Pleopod 2 of male with appendix
masculina basally or sub-basally inserted, usually

robust; endopods of pleopods 3-5 without
plumose marginal setae. smaller than exopod;
lamellar or thickened (A. barndardi, A. oaxaca).
Mouthparts and other characters as for Ciralana.

REMARKS

Anopsilana, with little question, constitutes a
polyphyletic assemblage of species, differing
from Crrolana primarily in the lack of marginal
setae on the endopods of pleopods 3 and 4. These
endopods are also reduced in size in comparison
to those of Cirolana and in at least two species are
thickened or fleshy. Other minor differences, none
of which are shown by all of the species placed in
the genus, are the fusion of peduncular articles |
and 2 of the antennule; article 3 of the antennule
peduncle being proportionally longer than in
Cirolana and the appendix masculina varying
from being relatively robust to slender and basally
inserted.

Estuarine species tend to be well pigmented and
have robust well-spined perecopods that are
generally similar to those of the Cirolana ‘parva
group’ or ‘tuberculate group’ of species.
Troglobitic species have relatively more slender
pereopods with, by comparison, reduced spina-
tion. Additionally all the troglobitic species lack
eyes,

The distribution of the genus is restricted to
tropical localities. The cavernicolous specics are
from the peri-Caribbean region except for one
species from Madagascar and one from Palau.
The open-water species are all estuarine, tropical
and all have been recorded from mangroves or in
the vicinity of mangroves, with the exception of
the poorly known Anopsilana luciae. Anopsilana
pustulosa and A. willeyi have extensive Indo-
Pacific distributions. Brusca ct al. (in press) report
that A. brown? is widely distributed in the Caribb-

wm

can and also in the tropical East Pacific in suitable
habirats,

KEY TO AUSTRALIAN SPECIES OF
ANOPSILANA

|. Body with nodules on dorsal surfaces of pereon,
pleon and pleotelson; pleotelson apex narrow-
ly rounded; endopods of pleopods 3-3 lamel-
Tarssrercevonvessaneusexcaistioniasistattorrevesrr={=yort=+ peta
Body without dorsal ornamentation, pleotel-
son apex wide, subtruncate; endopods of
pleopods 3-5 fleshy............4. bdmmardl sp.nov,
2, Frontal lamina anterior margin rounded: pleote!-
son abruptly narrowed; uroped rami wilh
dense mass of marginal setae........ A. pustulosa

Frontal Jamina pentagonal; lateral margins of
pleotelson not abruptly narrowed; uropod
rami With single rank of marginal
SOEBE.yscyeasprpesvgueceecanseguysreparerrepsereepeeth, Weilleve

Anopsilana barnardi sp.nov,

MATERIAL EXAMINED

HoLotyre: ¢ (5.4mm), Murray R., north Queensland,
18°14°S, 146°O1"B, 24 May 1978, salinity af 23-34%o,
depth 1.5m, mid-esiuary, mud bottom, coll. P. Davie
(QM W17263).

PARATYPES: 1300 (2.9-5.7mm, m = 3.9m, 5.7mm
specimen dissected), 142 2(35.0-5.9mm non-ovig,
m=4.0mm; 6.4mm ovig), manca (2.7mm), same data as
holotype (QM W17264),

DESCRIPTION

Male: Body about 2.8 times as long as greatest
width; dorsal surfaces without nodules, unorna-
mented, Cephalon with median rostral point
visible in dorsal view; dorsal interocular furrow
present; eyes together about 0.3 width of
cephalon. Pereonite 1 about 1.6 times as long as
pereonite 2; pereonite 2>3>4<5 = 6>7 in length;
pereonite 7 shortest. Coxae of pereonites 2-7 each
with distinet and entire oblique carina; coxae 4-7
with posteroventral angle becoming increasingly
acute, Pleonite 1 concealed by pereonite 7;
pleonite 3 with lateral margins posteriorly
produced and overlapping lateral margins of
pleonite 4 which are produced posteriorly beyond
pleonite 5. Pleotelson 0.65 as long us anterior
width; lateral margins convex, posterior margin
subtruncate, provided with 8 (or 9, see Variation)
spines, interspersed with 2 short setae between
each spine pair,

MEMOIRS OP THE QUEENSLAND MUSEUM

Antenniile peduncle articles 1 and 2 fused, ar-
ticle 3 about 1.2 times as long as fused articles 1
and 2; flagellum extending to middle of pereonite
1, with 8 articles, the distal 2 being minute, about
0.7 as long as peduncle. Antennal peduncle ar-
ticle 4 about 2.5 times as long as article 3; pedun-
cular article 5 about 1-2 times as long as 4;
flagellum extending to pereonite 4, composed of
about 17 articles. Frontal lamina pentagonal, ses-
sile; lateral margins diverging anteriorly, Man-
dible lacking lacinia mobilis; spine row with 9
spines; molar process very thin, with 13 or 14
teeth; palp article 2 with 12 serrate setae on dis-
tolateral margin, article 3 with 14 setae distal 5 of
which are longer than remaining 11 and feebly
secrate, others obviously serrate. Maxillule with
12 spines on gnathal surface of lateral lobe;
medial lobe With 3 stout moderately plumose
spines, Maxilla with 5 long setae on lateral lobe,
9 long setac on central lobe, medial lobe with 4
stout plumose setae proximally, distally 6 simple
setae, Maxilliped with simple setae only; endite
with 2 coupling hooks and 4 plumose setae.

Pereopod | basis with | long and 1 short seta at
posterodistal angle; ischium with 2 setae al
anterodistal angle, 2 small setae at posterodistal
lateral margin and 1 spine at posterodistal medial
Margin; merus with 4 short stout tubercular spines
on Jateral posterior margin and 3th longer spine
at posterodistal angle, posterior medial margin
with 2 acute spines; carpus With single acute spine
and 3 setae at posterodistal angle; propodus palm
with 2 acute spines and third stout spine and 2
setae opposing dactylus. Pereopods 2 and 3
similar to 1 but less robust. Pereopod 2 ischium
with 2 acute spines at anterodistal angle and 2
stout spines at posterodistal angle; merus with 2
large and 2 small spines at anterodistal angle,
posterodistal margin with 6 blunt spimes in 2
clusters; carpus with 3 spines at posterodistal
angle; propodus with single spine on palm.
Pereopods 4-7 essentially similar, becoming
longer posteriorly. Pereopod 7 ischium with
group of 5 spines at anterodistal angle, posterior
margin with 3 groups of spines; merus with
clusters of 6 and 4 spines at distal angles, 3 spines
set posteromedially; carpus with abundant spines
on distal margin (about 13), 1 and 2 spines set on
posterior margin; propodus with 2 pairs of spines
on posterior margin; some spines on anterior mar-
gins strongly pectinate: spines on posterior mar-
gin all simple. Penes small, inconspicuous (only
easily visible on dissected male paratype) papil-
lae, set submedially on posterior of sternite 7.

Pleopod 1 rami subequa! length, endopod slight-

A NEW SPECIES OF ESTUARINE CIROLANID ISOPOD 3

FIG. 1. Anopsilana barnardi sp.noy. A-D, holotype.
E-G male paratype. A, dorsal view. B, lateral view.
C, frons. D, pleonites in lateral view. E, penes, ster-
nite 7. F, antennule. G, antenna peduncle. Scale
represents 2.0mm.

ly less than half as wide (0.45) as exopod;
peduncle about 1.8 as wide as long, medial mar-
gin with 4 coupling hooks. Pleopod 2 with rami
subequal in length; appendix masculina broad,
0.3 as wide as endopod, arising basally and ex-
tending just beyond end of endopod. Pleopods 3-5
exopod with complete but faint suture; endopods
fleshy and small, about 0.5 as wide and 0.7 as long
as exopods. Uropod rami not extending beyond
posterior of pleotelson, exopod 0.85 as long as
endopod; exopod lateral margin with 9 spines,
medial margin with 6; endopod lateral margin
with 3 spines, medial with 7; peduncle with 2
spines at lateroventral angle; apices of both rami
weakly bifid; both rami with plumose marginal
setae among spines except for proximal three
quarters of lateral margin of endopod.

Female: Slightly larger than male, differs in
secondary sexual characters, Brood pouch made
up of overlapping oostegites arising from ster-
nites 1-5; embryos within dorsally oriented infold-
ing of sternites.

Colour: Pale brown in alcohol, dorsal surfaces
with brown chromatophores; eyes black.

Size: Males average at 3.9mm, non-ovigerous
females at 4.0mm.

Variation: There is considerable variation in the
number of spines present on the pleotelson and
uropodal rami. The commonest spine counts are:
pleotelson with 8 spines (47.6%) or 9 spines
(33.3%), occasionally 7 or 10; exopod lateral mar-
gin with 7-10 spines, 9 spines (50.0%) or 8 spines
(35.7%) commonest; exopod medial margin with
5 (71.4%) or 6 (19.0%) spines; endopod lateral

4 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Anopsilana barnardi sp.nov., male paratype. A, left mandible. B, right mandible, incisor. C, maxillule. D,
maxilla, E, maxilliped, F, pereopod 1. G, pereopod 2.

A NEW SPECIES OF ESTUARINE CIROLANID ISOPOD 5

FIG. 3. Anopsilana barnardi sp.nov., male paratype. A-D, pleopods 1-3, 5 respectively. E, apendix masculina
apex. F, pereopod 7.

margin with 3 (60%) or 4 (40%) spines, medial other Australian species by the lack of nodules or
margin with 5 (15.0%), 6 (57.5%) or 7 (27.5%) other cuticular ornamentation. The pleopods of
spines (the holotype, 10 males and 10 females Anopsilana barnardi have the endopod of
were examined for spine counts). pleopods 3-5 far smaller than those of A. pus-

tulosa Hale (see Bruce 1981, fig. 5i-l) or A. willeyi
REMARKS Stebbing (see Bruce 1986, fig 138), and further-
This species is easily separated from the two more the endopod is thickened and semi-opaque,

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Anopsilana barnardi sp.nov., male paratype.
A, uropod. B, posterior margin of pleotelson. C,
posterior ventrolateral angle of uropod peduncle.

not lamellar. There are only two other Indo- TyPES: Zodlogisch Museum, Amsterdam.
Pacific species of Anopsilana: A. lingua Bowman

& Tliffe, 1987, which is a blind freshwater cave Anopsilana browni (Van Name, 1936)
dwelling species from Palau, and A. luciae (Bar-
nard, 1940) a poorly described estuarine species DISTRIBUTION: Brackish and freshwater; Carib-
from South Africa. Anopsilana luciae differs by bean; Cuba and Belize; East Pacific: Costa Rica
having a rounded projecting frontal lamina, a (Kensley & Schotte, 1989).

narrow pleotelson apex and two longitudinal sub-

median carinae on the pleotelson. It is possible Types: American Museum of Natural History,
that some estuarine species currently placed in New York.

Cirolana may prove to belong to Anopsilana when
the pleopods have been examined. Anopsilana crenata Bowman & Franz, 1982
ETYMOLOGY

ay DISTRIBUTION: Freshwater pool in a cave, Grand
The species is named to honour the late Dr J.L. Cayman Island.
Barnard who has made an immense contribution

to the knowledge of Australian peracarid Crus-

TYPES: Smithsonian Institution.
tacea.
Anopsilana cubensis (Hay, 1903)
SYNOPSIS OF ANOPSILANA SPECIES
DISTRIBUTION: Caves in several localities in
Anopsilana acanthura (Notenboom, 1981) Cuba.

DISTRIBUTION: Known only from a well in Haiti. TyPEs: Smithsonian Institution.

A NEW SPECIES OF ESTUARINE CIROLANID ISOPOD 7

Anopsilana jonesi Kensley, 1987
DISTRIBUTION: Amongst mangroves, Belize.
Types: Smithsonian Institution.

Anopsilana lingua Bowman & Lliffe, 1987

DISTRIBUTION: Natural well on Peleliu Island,
Palau,

TYPES: Smithsonian Institution.

Anopsilana luciae (Barnard, 1940)
DISTRIBUTION: Estuarine, South Affica.

Types: South African Museum, Cape Town.
Anopsilana oaxaca Carvacho & Haasmann, 1984
DISTRIBUTION: Mangroves roots, Pacific coast of
Mexico; also Clipperton Island (Brusca et al. in
press).

TYPES: Institute of Biology, National
Autonomous University of Mexico.

Anopsilana poissoni Paulian & Deboutteville, 1956

DISTRIBUTION: Mitoho Cave, southern Madagas-
car.

TYPEs: Not stated, but see type species entry after
genus synonymy given here.

Anopsilana pustulosa (Hale, 1925)
DISTRIBUTION: Estuarine and mangrove habitats
from East Africa to Australia; generally within
the tropics (Bruce, 1986). Type locality: Cook-
town, Queensland.

TyPes: Australian Museum, Sydney,
Anopsilana radicicola (Notenboom, 1981)
DISTRIBUTION: Natural spring, Haiti-

TyPEs: Zodlogisch Museum, Amsterdam.
Anopsilana willeyi (Stebbing, 1904)

DISTRIBUTION: Estuarine and mangrove habitats
from East Africa to Australia, within the tropics

(Bruce, 1986). Type locality: Sri Lanka (Steb-
bing, 1904, as Ceylon).

TYPES: Not located.
LITERATURE CITED

BARNARD, K.H. 1940. Contributions to the crus-
tacean fanna of South Africa 12. Further additions
to the Tanaidacea, Isopoda and Amphipoda,
together with keys for the identification of hither-
torecorded marine and freshwater species, Annals
of the South African Museum 32: 381-515,

BOWMAN, T.E. & FRANZ, R. 1982. Anopsilana
crenata, a new troglobitic cirolanid isopod from
Grand Cayman Island, Caribbean Sea. Proceed~
ings of the Biological Society of Washington 95:
522-529,

BOWMAN, T.E, & ILIFFE, T.M. 1987. Anopyilana
lingua, a new freshwater troglobitic isopod from
the Palau Islands (Flabellifera: Cirolanidae).
Proceedings of the Biological Society of
Washington 100; 347-352.

BRUCE, N.L. 1981. Cirolanidae (Crustacea: Ilsopoda)
of Australia: Diagnoses of Cirolana Leach,
Metacirolana Nierstrasz, Neocirelana Hale,
Anopsilana Panlian and Deboutteville, and three
new genera - Natatolana, Politolana and Car-
tetolana, Australian Journal of Marine and Fresh-
water Research 32: 945-966.

1986. Cirolanidae (Crustacea: Isopoda) of Australia.
Records of the Australian Museum, Supplement
6; 1-239.

BRUSCA, R.C., WETZER,, R. & FRANCE, S.C. (in
press). A | monograph on the cirolanid isopods
(Crustacea: Isopoda: Flabellifera: Cirolanidac) of
the tropical eastern Pacific,

CARVACHO, A. & HAASMANN, Y. 1984. Isopodos
litorales de. Oaxaca, Pacifico Mexico. Cahiers de
Biologie Marine 25; 15-32.

HALE, H.M. 1925. Review of Australian isopods of the
cymothoid group. Pt 1. Transactions of the Royal
Society of South Australia 49: 128-185.

HAY, W.P, 1903. On a small collection of crustaceans
from the island of Cuba. Proceedings of the
United States National Museum 26; 429-435,

KENSLEY, B. 1987. Further records of marine isopod
crustaceans from the Caribbean. Proceedings of
the Biological Society of Washington 100: 559-
577,

KENSLEY, B. & SCHOTTE, M. 1989. ‘Guide to the
Marine Isopod Crustaceans of the Caribbean,’
(Smithsonian Institution Press: Washington,
D.C.). 308pp.

NOTENBOOM, J. 1981. Some new hypogean cirolanid

8 MEMOIRS OF THE QUEENSLAND MUSEUM

isopod crustaceans from Haiti and Mayaguana
(Bahamas). Bijdragen tot de Dierkunde 51: 313-
331,

PAULIAN, R. & DEBOUTTEVILLE, C.D. 1956. Un
cirolanide cavernicole a Madagascar [Isopode].
Mémoires de I’Institute Scientifique de Madagas-
car. Série A, 11: 85-88.

RIOJA, E. 1956. Estudios carcinologicos 35. Datos
Sobre algunos isopodos cavernicolas de la isla de

Cuba. Anales del Instituto de Biologia, Univer-
sidad de México 27: 437-462.

STEBBING, T.R.R. 1904. Gregarious Crustacea of
Ceylon. Spolia Zeylanica, Bulletin of the Nation-
al Museum, Ceylon 2: 1- 26.

VAN NAME, W.G. 1936. The American land and
freshwater isopod Crustacea. Bulletin of the
American Museum of Natural History 71: i-vii,
1-535.

SYSTEMATICS OF THE INTERTIDAL TRAPDOOR SPIDER GENUS [DIOCTIS

(MYGALOMORPHAE: BAR YCHELIDAE) IN THE WESTERN PACIFIC WITH A NEW

GENUS FROM THE NORTHEAST
TRACEY B, CHURCHILL AND ROBERT J. RAVEN

Churchill, T. B, & Raven, R, J. 1992 06 29: Systematics of the mtertidal trapdoor spider genus
Idioctis (Mygalomorphae: Barychelidae) in the Western Pacific with a new genus from the
Northeast. Memoirs of the Queensland Museum 32(1): 9-30, Brisbane. ISSN 0079-8835.

The circum-tropical trapdoor spider genus Idioctis included 6 species from Madagascar, east
to Hawaii. A new genus, Nihoa, endemic to the Leeward Islands, north of Hawaii, is named
and now includes /diectis hawaiiensix Raven, 1988, endemic to Necker Island, and a further
new species, Nihoa mahina sp.noy., endemic to Nihoa Island. Four new species of Idioctis
from the Western Pacific are described: J. yerlata from north-eastern Australia; /. ferrophila
from New Caledonia; J, marove from the Solomon Islands; and /. talofa trom Western Samoa,
raising the number of species to nine, For the first time, the male of the type species J. helva
Koch, 1874, from Fiji, is described, Variation in the shape of spermathecae in /dioctis is
described. (]Chelicerata, Mygalomerphae, Barychelidae, systematics, phylogeny.

Tracey B, Churchill and Robert J, Raven, Queensland Museum, PO Box 3300, South Brisbane,
Queensland 4101, Australia; 3 February, 1992.

Spiders of a number of genera occur beside the
sea. However, only few are intertidal to that their
burrows are covered at high tide, Best known
among araneomorphs are Desis and Paratheuma
(Desidae), the latter having been revised recently
(Beatty and Berry, 1988). In contrast, spiders of
only one mygalomorph genus, Idiectis Koch,
1874, are known to live in the intertidal zone.

The genus /dioctis was first described by Koch
(1874) including only f. kelva from Fiji. No refer-
ence was made to the habitat. The first report of
the intertidal habits of a barychelid was with the
description of Idiactis littoralis Abraham, 1924,
found in the intertidal mudflats of Singapore.
Much later, 1. helva was recorded from the littoral
zone in Western Samoa (Marples, 1951, 1955).
Soon after, another species, Atrophonysia inter-
tidalis Benoit & Legendre, 1968, (transferred to
Idioctis by Raven, 1985) was collected and
described from the littoral zone of Madagascar
and the Seychelles.

By 1988, six species of Jdioctis had been
desenbed: the type species, /. helve from Fiji and
Western Samoa (Marples, 1951, 1955): £ fit-
teralis Abraham, 1924 from Singapore; J, inter-
lidalis (Benoit & Legendre, 1968) from
Madagascar and the Seychelles (Benoit, 1978); 1.
amas Raven, 1988 from Christmas Island; /.
entwetok Raven, 1988 from the Caroline and Mar
shall Islands; and J. hawaitensis Raven, 1988 from
the Leeward Islands, near Hawaii.

After a revision of the genus (Raven, 1988), we
discovered Jdioctis in the intertidal or littoral
zones in northem Australia, New Caledonia, and

the Solomon Islands. Three new species are
described here. Concurrently, the first male and
more females of the type species, J. helva, were
taken from the type-locality of Ovalau, a small
island of Fiji. Females of /. helva were also col-
lected from the coast of the large island of Vanua
Levu, Fiji. With this new material we conclude
that a new species occurs in Westem Samos, We
also document variation in spermathecae for the
first time in barychelids.

Raven (1988) noted that Jdiectis hawaiiensis
differed from all other congeners in the wider
stemmum, undivided spéermathecae, and relatively
shorter eye group. Since then, material from
another island in the Leeward Islands has proved
to bea new species for which a new genus 1s here
created. This new genus recognizes the plesiomor>
phic nature of both J. fawaiiensis and its new sister
species.

MATERIALS AND METHODS

All abbreviations are standard for the Araneae
and explained in Raven (1984a). The width of the
eye group or the median ocular quadrangle (ab-
breviated as MOQ)is the distance between the hwo
most separated points in a line orthogonal to the
long axis of the spider. All measurements except
those for eyes are given in millimetres. Eye meas-
urements are taken from camera lucida drawings
made at < 50 magnification; any error was taken
as 0.02 mm, or 1mm on the enlarged figure. Eye
interspaces, which are measured along a line join-
ing the centres of the respective eyes, are given 35

10 MEMOIRS OF THE QUEENSLAND MUSEUM

diameters of an AME. Presence of leg spines are
specified by the number recorded with their posi-
tion (rare niimber occurring in parentheses) al-
though ranges can include zero. Spine positions
are as follows: dorsal, if on or close to the midline;
pro- or retrolateral, if spine bases are visible pro-
or retrolaterally when viewed dorsally; ventral, if
bases are visible when viewed ventrally, Spines
are considered weak, and probably the equivalent
of attenuate macrosetae of Coyle (1971: 329), if
only marginally thicker than other setae on that
surface. Despite being Weaker, they are treated as
spines because their number and position suggest
they are the weaker form of thick spines on other
species. Descriptions of scopulae give the extent
(measured from the distal end of the leg segment),
the density of scopula hairs, and the number of
rows of setae that part the scopula hairs.

Acronyms for museums are: AMNH, American
Museum of Natural History, New York; BMNH,
British Museum (Natural History), London;
NHW, Naturhistorisches Museum, Vienna; QM,
Queensland Museum, Brisbane. Other museums
are given in full,

SYSTEMATICS
Idjoctis Koch

Idiectis Koch, 1874: 4&4 (type species by monotypy
Idioctis helva L, Koch), Simon, 1892; 125; 1903:
914, Rainbow, 1912: 115, Roewer, 1942; 214. Bon-
net, 1957; 2286, Bagnols, 1983; 129, Main, 1985: 12,
Raven, 1985: 113, 1988: 2. Platnick, 1989: 93.

Atrophonysia Benoit & Legendre, 1968; 330 (type
species by original designation Alrophonysia inter-
tidalis Benoit & Legendre), First synonymized by
Raven, (985; 113. Brignoli, 1983: 130. Raven, 1985;
113, 1988; 2; Plamick, 1989: 93

DIAGNOSIS

Idioctis can be distinguished from Jdiephthalma
by the narrow sternum and nasrow fovea and
ssetenre of a strong rastellum, and from other
barychelid genera by the relatively short bat
strongly trapezoidal eye group and a narrow ster-
num. [t is also readily distinguished from Nihoa
gen. nov, and many other barychelids in the pallid
pattern-free abdominal dorsum and the elongate
stemum.

DESCRIPTION

Carapace hirsute, without pattem. Fovea very
broad, procurved or straight. Abdomen pallid,
without pattern. Rastellar spines few in line over-

hanging fang, not on raised mounds. Maxillac
usually with c,18-20 cuspules on inner anterior
comer} anterior lobe and posterior heel produced.
Labium wide, without cuspules. Sternum long,
narrow (aboul 1.5 times longer than wide, rarely
wider), with small, marginal sigilla, if evident.
Scopulae (2 ?) entire on metatarsi and tarsi | and
I; for full length of prolateral face of metatarsi III
and only rarely present elsewhere on segment;
widely parted by setae on tarsi If; only narrow
lateral bands of hair, if present. on tarsi IV; absent
on metatarsi TV. Scopuliform bmsh present on the
provemtral face of tibiae I and II of 2 2. Spines.
Leg 1: fe d0-4; ti pO-1 v2-7; me rO-1. Leg 2: fe
d0-4; ti pO-1 yO-4; me v1-2. Leg 3: fe p0-1, dO-5;
@ p6-30; ti p(2), r(2), vO-7; me pO-3, 10-3, v0-13.
& 4: fe dO-7; pa p3-16; ti r(2), vO-7; me p0-7,
rO4, y2-9. Paired claws of 22 with teeth on
lateral face (Fig. 3); claws of d¢ biserially den-
tate. Tarsi with clavate and filiform trichobothna,
3: tibia I with prolateral spur and megaspine,
more distal prolateral megaspine and associated
cuticular ‘thumb’; palpal bulb pyriform, Inter-
tidal, rarely in forests.

CHECKLIST OF SPECIES

Idivetis helva Koch, 1874, type species Fiji.

fdiovtis eniwetok Raven, 1988 Caroline & Marshall
Islands.

ldioetis ferrophila sp.nov, New Caledonia,

Airephenysia inrertidalis Benoit & Legendre, 1968
Madagascar, Seychelles,

Idioctis littoralis Abraham, 1924 Singapore.

Idiectis marove sp.nov, Solomon Islands «

Idioctis talofa sp.nov. Western Samoa.

Idioetis xmas Raven, 1988 Christmas L., Australia,

Idioctix yerlata sp.nov. North-eastern Australia.

DISTRIBUTION

Spiders of the genus /dioctis are now known
near beaches and in littoral zones in Fiji, Sin-
gapore (Abraham, 1924), Western Samoa
(Marples, 1951, 1955), Madagascar (Benoit &
Legendre, 1968), Seychelle Islands (Benoit,
1978), Christmas Island (Indian Ocean, Raven,
1988), Marshall and Caroline Islands (Raven,
1988), north-eastern Australia, New Caledonia
and the Solomon Islands, Habitats, life history
observations and biogeography of the genus will]
be discussed elsewhere-

REMARKS

This paper contains the first description of a ¢
of the type species. The diagnosis of Jdioctis given
by Raven (1988) is correct save for the removal of

INTERTIDAL TRAPDOOR SPIDERS 1

characters present only in J. hawaiiensis, here
transferred to Nikoa, gen. nov.

Idioctis helva Koch
(Figs 2-6, Table 1)

Idioctis helva L. Koch, 1874: 484, tab. 37, fig. 3 a,b.
Hogg, 1901: 241-242, fig. 26a. Marples, 1955; 453-
454; Raven, 1988: 4, figs 1-5.

MATERIAL EXAMINED

LECTOTYPE: @, (here designated), Ovalau I,, 17°41°S,
178°50'E, Fiji, Mus. Godeffroy No. 8097, deposited in
Zoologische Institut and Zoologisches Museum, Ham-
burg.

PARALECTOTYPES: | 2, 1] juvenile, same data, deposited
in BMNH.

OTHER MATERIAL

TopotyPes: 1d, Nageledamu, Ovalau I., 17°41°S,
178°50°E, Fiji, in mangrove root, 14.x1.1988, T.B.
Churchill, QM $11229. 592 9, Nageledamu, Ovalau I.,
Fiji, 14.xi.1988, T.B. Churchill, QM $12521-5; 62 9,
Lota I., Vanua Levu, 16°46°S, 179°20°B, Fiji,
20.x.1988, T. B. Churchill, QM $12526-30.

DIAGNOSIS
Spermathecae two, each consisting of two short

FIG. 1. Idioctis yerlata sp.nov., 2.

TABLE 1. Leg measurements of Idioctis helva, 3.
I ll ti IV
3.60 3, 2.56 3.36
1,92 144 1.76
3.04 2.16 3.52
2.32 2.16 2.96
1.36 1.20 1,20
12,24 9.52 12.80

Femur
Patella

Tibia
Metatarsus
Tarsus
Total

dissimilar lobes, ental lobe stout, ectal lobe
slender. 2 2 with c.18 thorns on patella II. 4-5
teeth on paired claws of 2 2. Rastellum of ten
thick (four as long as wide) coniform spines in
short distal line. Preening combs absent, 6d.
Tibia I with small distal spur and upeurved
megaspine, prolateral face with distal tiangular
cuticular process and short lower angular
downcurved spine. Bulb pyriform; embolus
broad, not spiralled; cymbium with two divided
lobes. Two rows of teeth on all paired claws.

TOPOTYPE MALE

Carapace 4.36 long, 3.32 wide. Abdomen 4.16
long, 2.40 wide. Total length, 10.

Colour in alcohol. Carapace and legs yellow
brown, chelicerae red brown; abdomen dorsally
grey brown with dark ‘V" dorsally showing loca-
tion of heart; venter entirely yellow brown,

CITBRETIN, 30

12 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Distribution of /dioctis * and Nihoa & in the Western Pacific and Indian Ocean.

2aGum

FIG. 3. Idioctis helva, scanning electron micrographs, paired claws, lateral view. a, b, leg 1, d (a), 2 (b). c, d, leg
IV, 2.

INTERTIDAL TRAPDOOR SPIDERS 13

FIG. 5. Idioctis helva, dorsal view of spermathecae of
? 2. a-d, g, Ovalau Island, Fiji. a, $12525. b, $12521.
c, $12522. d, $12523. g, §12524. e, f, h-j, Lotu Island
(Vanua Levu), Fiji. e, 12527. f, $12528. h, $12526,
i, $12529, j, 12530. Scale line = 0.25mm.

FIG. 4. Presence ™ and absence @ of Idiactis helva at
coastal localities sampled in Fiji.

~ Bei
ese ly

'

FIG. 6. Idioctis helva, topotype ¢. a, cephalothorax and chelicerae, dorsal view, b, spinnerets, ventral view. c,
eyes, dorsal view. d, sternum, maxillae, labium and chelicerae, ventral view. e, palpal tibia, cymbium, and bulb,
retroventral view. f, tibia I, ventral view, showing distal spur and megaspine. g, tibia 1 showing spur, prolateral
view. Scale line = 1mm (a), 0.5mm (b-g).

id MEMOIRS OF THE QUEENSLAND MUSEUM

Carapace, Uniform covering of short black
bristles and fine grey hair, not obscuring cuticle;
margins with longer setae and lower sclerotised
shelf of hirsute chitin below main margin. An
irregular row of 15-20 setae anteromedially; 2-3
pairs of fine setae; numerous thick setae between
PME and ALE. Striae shallow, narrow, glabrous,
Tecognised as areas intervening grey bush of hair.
Fovea wide, recurved.

Eyes, Tubercle distinct. Group occupies 0.45 of
head-width. Three rows; back row recurved, Front
line of lenses of AME just behind line of back of
ALE and ‘iris' well back. Eye group front width,
back width, length = 35:43:25. MOQ front width,
back width, length = 21;29:14.
AME: ALE-PME:PLE = 10:11:7:10. Eye inter-
spaces: AME-AME, 0.5; AME-ALE, 0.5; ALE-
ALE, 1.6; PME-PLE, 0.2; PME-PME, 1.8;
ALE-PLE. 0.6.

Chelicerae, Wide band of brown selae and grey
hair prodorsally, two narrower bands laterally.
Promargin with 8 evenly spaced teeth, basomesal-
ly with 2 small teeth and 3 pranules. Inter-
cheliceral tumescence absent. Rastelluin consists
of 5 long thick pointed spines and c.5 thick sctae.

Labium, 0,64 wide, 0,28 long. Cuspules absent.
Labiosternal suture is two sigilla narrowed at their
medial amet

Mavillae. 1.04 long in front, 1.36 long behind,
1.64 wide; with c.S-8 pointed cuspules. Heel an-
gular; anterior lobe distinct.

Sternum. 2.16 long, 1.48 wide. Only posterior
sigilla evident as rounded glabrous marginal
areas.

Legs. Formula 4123. All segments, especially
femora, with mat of dark grey prostrate hairs; and
erect black setae. All legs of similar diameter,
Tibia 1 with small distal spur and upcurved
megaspine, prolateral face with distal triangular
cuticular process and short lower angular down-
curved spine, Preening combs entirely absent.

Scapula. Never so thick as to obscure cuticle,
individual hair bases distinct. Tarsi I-III ventrally
pallid, Tarsi:] and I entirely ventral, thin, entire,
full length; TJ, thin, divided by 8-10 setac; 1'V,
with 10-15 scopula hairs in narrow triangle on
proximal prolateral face. Claw tufts smaller than
claws. Metatarsi: I, thin for full length, distally
cuticle is pallid; IL, similar but with single thick
seta on ventrodistal edge; IL, proventrally with
fight brush; [V, bare, Thorn spines: 10 on
prolateral patella IE,

Spines. Leg 1: fe d4, ti v3+2 megaspines; me v1
weak, distal. Leg 2: fe d3_ti v4; me v1 weak, distal,
Leg 3: fe d4, #1}, pa p10 thorns, ti v5, me rl, v4.

Leg 4: fe d6, rlw; tir2, v7w; me v3 4 4 thick setac.
Paip: fe dSw.

Claws, 2 full scooped rows of 8-10 teeth in each
row on paired claws of leg [; claws of leg TV with
2 rows, 6 in each ovter row and 2 in each inner
row.

Trichobothria, In two rows, each of c.8 for 3/4
of ubiae; c.12 in proximally retrolateral row on
metatarsi, distally row is irregular and trichae are
longer; .c.3 clavate and 15 filiform on tarsi in band.

Palp. Bulb pyriform; embolus broad, not
spiralled, cymbium with two divided lobes,

Spinnerets. PMS 0.20 long, 0,08 wide, 0.08
apart, c.0.18 of basal PLS in diameter. PLS basal,
middle, apical, and total article lengths = 0.64,
0.26, 0.14, 1.04, respectively.

DiSTRIBLUTION AND HABITAT

i. helya is known from mangrove roots and
trunks on Ovalau Island, east of Viti Levu, Fiji,
and from coral rock on Lotu Island. south of
Vanua Levu, Fiji.

REMARKS

Females of J. helva differ from those of other
species by the wider configuration of the eyes and
the smaller PMS. Spermathecae consist of two
short almost amorphous lobes joined basally to
form 4 common aperture. Males differ from those
of I. eniwetok Raven by the absence of preening
combs.

Idioctis yerlata sp,nov-
(Figs 1, 2, 7-9, Table 2)

MATERIAL EXAMINED

Houoryre: #, Cape. Tribulation, 16°05"S, 145°26'E,
in mangroves, North Queensland, Australia, 28-
29.yiii.1988, R.J. Raven, T.B. Churchill, J.A, Gallon,
QM 87213.

PaRATYPES: 799%, Cape Tribulation, 16°05’S,
145°26°E, in mangroyes, North Queensland, 28-
29.viii, 1988, R.J. Raven, T.B. Churchill, J.A, Gallon,
QM $7204-9, $7212, S11181. 1%, Noah Head, S. of
Cape Tribulation, 16°08"S, 145°28'E, 30. viii. 1988, RJ.
Raven, T.B. Churchill, J.A. Gallon, QM $7206.

DIAGNOSIS

Spermathecae with one large medial lobe and
variable basal lobe ectally; basal lobe small with
common atrium with lateral lobe or separate Jobe
arising above base of medial lobe (Fig. 9). Three
closely spaced teeth on cheliceral promargin near
fang base. Numerous (28-30) thorn spines on
prolateral patellae II]. 5-7 teeth on paired claws of

INTERTIDAL TRAPDOOR SPIDERS 15

Cooktown

Cape Tribulation
Nosh Head

Port Douglas

Ellis B ach
WeClifton|Beach
\@® Yorkeys Knob

FIG. 7, Presence of Idioctis yerlata sp.nov. at coastal localities sampled in Queensland, Australia; inset enlarged

at right, presence fl and absence @,

leg 1, Preening combs absent. Rastellum is 12-14
long curved spines.

HOLOTYPE FEMALE

Carapace 5.36 long, 4.40 wide. Abdomen 8.80
long, 5.28 wide, Total length, 16.

Colour in alcohol and life. Carapace and legs
yellow brown, chelicerae ted brown, abdomen
uniform yellow brown.

Carapace. Uniform cover of fine prostrate wavy
grey hairs; 10-15 long erect setae on medial caput
lateral of centre line; c.4 pairs of fine foveal
bristles; 27 long erect black setae anteromedially;
TABLE 2. Leg measurements of /dioctis yerlata,

holotype 2

I Il Ti IV Palp
Femur 3.28 288 2.24 344 2.64
Patella 2.48 2.32 1.84 2.64 1.84
Tibia 248 2.24 1.84 344 = 1.64
Metatarsus 1.84 1.76 1,12 2.80
Tarsus 136 1.20 088 1.20 1.52
Total 1144 1040 7.92 13.52 7.64

7 long black anteriorly directed bristles and
numerous fine black setae between PME; 2 long
recurved and 10-15 shorter black bristles between
ALE; caput raised; striae distinct. Fovea broad,
straight with recurved ends.

Eyes. Tubercle low. Group occupies 0.33 of
head width. Three rows; back row more or less
straight. Eye group front width, back width, length
= 44:58:34. MOQ front width, back width, length
= 23:42:17. AME:ALE:PME:PLE = 7:13:7:15.
Eye interspaces: AME-AME, 1.3; ALE-ALE, 3.4;
AME-ALE, 1.6; PME-PLE, 0.1; PME-PME, 4.0;
ALE-PLE, 1.6.

Chelicerae. With moderately dense bush of
black hairs dorsally and in lateral band, rastellum
consists of 12-14 long strong spines; promargin of
furrow with two very large and five smaller teeth,
and six small teeth basally.

Labium. 0.92 wide, 0.48 long, without cuspules,
separated from sternum by narrow groove.

Maxillae. 1.60 long in front, 2.04 long behind,
0.96 wide; with 6-9 spindle-shaped cuspules in

it MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. /dioctis yerlara sp.nov. holotype 2. a, cephalothorax and chelicerac, dorsal view. b, abdomen, dorsal view.
c, eyes, dorsal view. d, sternum, maxillae, labium and chelicerae, ventral view, e, spinnerets, ventral view, f.
abdomen, ventral view. Scale line = 2 mm (a, b), Imm (a), 1).Smin (c, ¢).

close group on inner angle; anterior lobe indis-
lunct, posterior “heel” produced.

Sternmune. 3-12 Jong, 7.00 wide; sigilla evident as
shallow depressions in margin.

Legs. Formula 4123. Legs I and Il much thicker
than TI] and [V; tarsi as fat as distal metatarsi.
Dorsal tarsi and meiatarsi with pile of short
straight grey hair. Tibia through tarsus IV asetose,
almost glabrous, With numerous short coniform
spines on prolateral patellae Tl. TV. Preening
combs consist of group of 8 spines on metatarsi
III, outer pair and middle pair long and slender,
others short, thick; eight also on metatarsi I'V, two
long slender setae separated by shorter spines on
ventral edge of metatarsi TV.

Scapula. Full and entire on metatarsi and tarsi I,
Tl; divided by setae on metatarsi and tarsi III: one

narrow prolatera) band on tarsi TV; elsewhere ab-
sent. Palpal tarsal scopulac entire.

Spines, No spines on tarsi. Leg 1: fe 0, (d11 (hick
setae); ti pl weak, v7 weak; me v1 distally with
thick bristle. Leg 2: me v2 distally lv with thick
bristle. Leg 3: pa p28-30; tibia yl + Gw, me dis-
toventral 2 long medial with 4 on cach side and
one between 2 long. Leg 4: fe 0, pa p11-16, ti v5,
me ©. Palp, ti v1 or v0.

Claws, Paired claws with 5-7 teeth on ectally
displaced keel on leg I; 3 teeth on leg I; 2 on leg
IV; palpal claw without tecth. All paired claws
extending beyond tufts, except on palp.

Trichebothria. c-8 for 3/4 of tibiae: ¢,12 on
metatarsi; 5 clavate and 12 filiforni on tarsi,

Spermathecae, Two, each with one large medial
and one ectal lobe,

INTERTIDAL TRAPDOOR SPIDERS 17

FIG, 9. Idioctis yerlata sp.nov., dorsal view of sper-
mathecae of 99. a, 87204. b, $7207. c, 87212. d,
$7213, e, 11181. f, $7206, g, 87208. Scale line =
0.25mm,

Spinnerets. PMS 0.48 long, 0.22 wide, bases
0.10 apart, c.0.24 of basal PLS in diameter. PLS
basal, median, and apical total article lengths =
1.14, 0.60, 0.20, 1.94, respectively.

DISTRIBUTION AND HABITAT

Known only from mangroves and loose coral
rubble in the intertidal zone near Cape Tribulation,
north Queensland, Australia. A trapdoor burrow
like that of /dioctis was noted in the intertidal zone
on a mangrove tree just south of Port Douglas,
about 30km north of Cairns, north Queensland.

ETYMOLOGY

The specific epithet is an aboriginal work mean-
ing oyster, which is often the impression conveyed
by the door of /. yerlara burrows.

REMARKS

Females differ from all species of /dioctis in the
larger relative size (3.4 umes an AME diameter)
of the ALE-ALE interspace,

Idioctis ferrophila sp.nov,
(Figs 2, 10-12, Table 3)

MATERIAL EXAMINED

Houotyre: 2, Port Boisé, 22°20'S, 166°59'E, New
Caledonia, ironstone boulder in water on beach,
25.x.1988, R.J. Raven, T.B. Churchill, QM §13537,
PARATYPES: 79 2, Port Boisé, 22°20'S, 166°59'E,
New Caledonia, ironstone boulder in walter on beach,
25.x.1988, R.J. Rayen, T.B, Churchill, QM 812511-3,
512515-7, $13538; 32%, same data, 6,ix,1990, P.
Goloboff, N. Platnick, R.J. Raven, AMNH.

DIAGNOSIS

Spermathecae with large medial lobe and short
basal lobe with axes perpendicular, Thorn spines:
c.30 on prolateral patellae IJ, c.11 small on

proximal prodorsal patella ['V. 3-5 teeth on paired
claws of g 2. Rastellum a line of 10 short conical
spines, not on mound, on edge above fang. dd
unknown.

HOLOTYPE FEMALE

Carapace 5,28 long, 4.12 wide. Abdomen 8.48
long, 4.88 wide. Tota] length, 17.

Colour in alcohol and life. Carapace yellow
brown with light brown mottling, chelicerae dark
reddish brown-burgundy; legs, sternum, maxillae,
and labium yellow brown; abdomen light yellow
brown with slightly darker medial band over heart.

Carapace, With fine gray hairs forming uniform
covering; hairs become larger and paler on mar-
gins and darker and thicker near anteromedial line.
Striae glabrous, narrow, barely distinct. Narrow
sclerotised and hirsute ledge below edge of
carapace. Fovea very slightly procurved. About
10 thick and several finer anteromedial setae: two
long setae between PME; 5-8 long recurved and
several shorter setae and fine hairs between ALE.
Clypeus absent.

Eyes. Tubercle low. Group occupies 0.32 of
head-width. Three rows; back row procurved. Eye
group front width, back width, length = 44:55:32.
MOQ front width, back width, length = 25:33:19.
AME:ALE:PME:PLE = 10:14:8:14. AME-
AME, 0.4, AME-ALE, 0.8, ALE-ALE, 1.7, PME-
PLE, 0.3, PME-PME, 2.2, ALE-PLE 1.0.

Chelicerae. Porrect with prodorsal and lateral
bands of long fine hairs. Slight ridge parallels
upper cheliceral face. Rastellum a line of 10 short
conical spines, not on mound, on edge above fang.

; i
Bi ae
e*

ind inte ial

FIG. 10. Presence and absence @ of Jdioctis fer-
rophila sp.nov. at coastal localities sampled in New
Caledonia.

18 MEMOIRS OF THE QUEENSLAND MUSEUM

=~

Sas

Se

PIG. 11, Idioctis ferrophila sp.nov., holotype 2 . a, cephalothorax and chelicerae, dorsal view. b, eyes, dorsal view.
c, spinnerets, ventral view. d, sternum, maxillae, and labium, ventral view. Scale line = Imm (a, d), 0.5mm (b,

c).

Furrow promargin with 7 thick teeth, basomesally
wath 1 minute and three small teeth,

Labiwn. 0.88 wide, 0.44 long. Labiosternal su-
lure a narrow groove.

Maxillae, 1.52 long in front, 2.16 long behind,
1.04 wide; with c.4-6 cuspules on inneredge. Heel
distinct; anterior lobe, indistinct.

Sternum, 2.80 long, 1.88 wide, With slightly
darker and thicker hairs on margin; otherwise fine
grey hairs medially. Sigilla shallow, indistinct.

Legs. Formula 4123. I, If thicker than II, IV.

TABLE 3. Leg measurements of Idioctis ferrophila,
holotype ?

[ i
2.88 2.68
240 2.16
2.08 2,04
1.60 1.52
1.12 1,12

10,08 9.52

Palp
2.24
1.68
1,36

2.08
1.52
1.36
1.36
0.80
7.12

Femur
Patella

Tibia
Metatarsus
Tarsus
Total

1.44
6.72

Covered with fine grey hairs mixed with black
setae; fine ‘furry’ grey hairs form narrow band
near trichobothria on tibiae and metatarsi.
Metatarsi I, I with distinct darker pair of setae
distoventrally. Thorn spines: c.30 on prolateral
patellae [iI, c.11 small on proximal prodorsal
patella IV.

Scepula. Metatarsi and tarsi I, U1, full, entire but
not obscuring cuticle and separate hairs distinct.
Metatarsi: IIT, one wide band prolaterally, ventral-
ly and retrolaterally with setae and spines; IV,
absent. Tarsi: III, full, parted by band 6-8 setal
rows wide; IV, only small cluster in proximal
proventral corner.

Spines. True spines only ventrally on palpal
tibiae and distal metatarsi TIT, TV; 6-8 thicker setae
on dorsal femora. Leg 1, 2: none except tibia v2
(w) and v4, respectively. Leg 3: thom spines plus,
ti V6 (w), me v8 on distal edge plus 1 thick and 4
thinner medially. Leg 4: thorn spines plus, tibia v7

INTERTIDAL TRAPDOOR SPIDERS he

FIG. 12. /dioctis ferrophila sp.nov., dorsal view of
spermathecae of 2 2, a, $7211. b, §12513. c, 812517.
S125 1. e,$12512. f,$13538. g,$13537. Scale line
=0.25mm.

(w); me, v8 distal (4 close enough to be almost a
comb) and 3 thinner proximad. Palp, ti v5.

Claws. 5 short (leg I) to 3 (leg TV) teeth on paired
claws, palpal claw bare. Tufts dense, notenclosing
claws, divided.

Trichobothria. In 2 rows, each of c.6 for 3/4 of
length of tibiae; c.6-7 in short diagonal row on
metatarsi; 5 clavate and c.14 filiform on tarsi in
two triangular bands, Trichobothria associated
with fine grey hairs.

Spermathecae. Two, each with two lobes,
medial lobe broad, lateral lobe small digitiform.

Spinnerets. PMS 0.22 long, 0.08 wide, 0.08
apart, c.0.12 of basal PLS in diameter. PLS length
of basal, middle, apical, and total articles = 0.76,
0.22, 0.18, 1.92, respectively.

DISTRIBUTION AND HABITAT

Idioctis ferrophila is known from ironstone
boulders on the beach at Port Boisé at the south
eastern corner of New Caledonia. A trapdoor bur-
row with a spider like that of Idioctis was noted in
intertidal rocks just south of Goro, 5-7 km east of
Port Boisé. The specimen was not collected. At
both locations, the boulders and rocks had water
around their bases at low tide.

ETYMOLOGY

The specific epithet is latin derived for iron
(ferros) and greek love (philos) and relates to the
original collection of this species from only
ironstone boulders in southern New Caledonia.

Idioctis talofa sp.nov.
(Figs 2, 13-15, Table 4)

MATERIAL EXAMINED

HOLOTYPE: 2, Upolu, 13°55"S, 172°45'W, close to

high tide mark, Western Samoa, B.J. Marples, BMNH
1974.129.1.

PARATYPES: 29 2, Upolu, 13°55'S, 172°45°W, close
to high tide mark, Western Samoa, B. J. Marples,
BMNH 1974.129.2. 1492 and eggs from Nu'ulopa
Islet, Western Samoa, B. J. Marples, BMNH 1974.135.
12, ‘Samoa’, 19,ii,1882, Kauf, Godeffroy collection in
NHW.

DIAGNOSIS

Spermathecae two, each consisting of two
similarly sized lobes. one lobe pointed. 18-20
thorn spines on prolateral proximal corner of
patella I; c.9 on patella IV; prolateral distal
corner of femur [V with 2-3 short spines. 4 teeth
on all paired claws. Preening combs absent. Ras-
tellum is 11 thick curved spines in single line with
bases touching.

HOLOTYPE FEMALE

Carapace 5.00 long, 4.08 wide. Abdomen 7.17
long, 4.00 wide. Total length, 15.

Colour in alcohol. Carapace orange brown,
darker on caput, legs orange brown. Abdomen
dorsally pallid with purplish tinge and ventraily
pallid.

Carapace. With light uniform covering of sil-
very brown hairs, none in striae, light bush on
margins, numerous brown bristles beside 10 thick
and several smaller anteromedial bristles, 6-8
thick bristles between PME, c.10 between AME.
Fovea distinctly procurved; clypeus absent.

Eyes. Tubercle low, distinct. AME pigmented.
Group occupies 0.35 of head-width. Three rows;
front row widely separated from middle row, back
row slightly procurved, Eye group front width,
back width, length = 40:52:33, MOQ front width,
back width, length = 24:35:19, AME:ALE:PME:

|; | |

STERN SAMOA

172°W.

FIG, 13, Localities of Idioctis talofa sp.nov, in Wester
Samoa ®-

20 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 14. /dioctis talefa sp.noy., holotype 2. a, cephalothorax and chelicerae, dorsal view. b, spinnerets, ventral
view. c, sternum, maxillae, Jabium and chelicerae, ventral view. Scale line = 1mm (a, c), 0.5mm (b).

PLE = 10;12:8:14. Interspaces: AME-AME, 0.5,
AME-ALE, 1.0, ALE-ALE, 1.9, PME-PLE, 0.2,
PME-PME, 2.4, ALE-PLE, 1.3.

Chelicerae. Band of moderately long brown
bristles prodorsally and narrower band laterally.
Rastellum is c. 11 thick curved spines in single line
hanging over long cheliceral edge, with bases
touching. Furrow promargin with 8 teeth, all bases
separate but most anterior teeth bases very close,
basomesally with 6 small teeth.

Labium. 0.92 wide, 0.84 long. Separated from
sternum by narrow groove,

Maxillae. 1.44 long in front, 1,96 long behind,
1.20 wide; with 5-8 pointed cuspules on inner
angle. Heel rounded; anterior lobe just distinct.

Sternum. 3.24 long, 1.96 wide. Widest between
coxae J, Sigilla small, marginal, indistinct.

Legs. With uniform covering of moderately long
brown hairs, Scopula. Metatarsi and tarsi 1, [1
entire, full; metatarsi HI, only on proventral face;

metatarsi TV absent; tarsi If with narrow band of
4-6 rows parting scopula; tarsi IV thin, short hairs,
only on proventral face. Thorn spines: 18-20 on
prolateral proximal corer of patella IM; c.9 on
patella IV; prolateral distal corner of femur IV
with 2-3 short spines. Preening combs absent.
Claws, All paired claws with 4 distinct teeth;
palpal claw bare. Claw tufts not enclosing claws.
Spines. No true spines dorsally or laterally ex-
cept for femora and patellae I, [V. Spines on
ventral legs generally weak, strong only on distal
metatarsi If] and IV but no so close as to form
comb, Leg 1: ti p3, me-0. Leg 2: ti v3, me v1, Leg
3: pa pl&, ti vow, me v9. Leg 4, fe p3, pa p6, ti
yvéw, me v5 short plus 4-6 thinner. Palp: ti pl, v4.
Trichobothria. In two rows, each of c.11 for 3/4
of tibiae; c.9 in curving row on metatarsi; 3 clavate
and c.15 filiform on tarsi.
Spermathecae, Two, each consisting of two
similarly sized lobes, one lobe pointed.

INTERTIDAL TRAPDOOR SPIDERS 21

FIG. 15. Idioctis talofa sp.nov., dorsal view of sper-
mathecae of @ @, a-h. Scale line = 0.25mm.

Spinnerets. PMS 0.36 long, 0.12 wide, bases
0.06 apart, c.0.19 of basal PLS in diameter. PLS
basal, middle, apical, and total article lengths =
0.74, 0.26, 0.18, 1.18, respectively.

DISTRIBUTION AND HABITAT

Known from laval rock on beaches on the island
of Upolu, and the islet of Nu’ulopa, Western
Samoa (Marples, 1955).

ETYMOLOGY

The specific epithet is a Samoan word meaning
‘welcome’, in appreciation of the warm reception
given to TBC.

REMARKS

Differs from /. helva in the spermathecal lobes
being equal in size, anterior most teeth in
cheliceral margin not fused basally, and has 1-2
thorns on prolateral distal femur IV. Of the 17
adults examined, one from ‘Upolu’ has only 4-6
spines on prolateral patella III of both legs. In all
others, the number varies from 10-20 on a single
specimen.

Idioctis marovo sp.nov.
(Figs 2, 16-18, Table 5)

MATERIAL EXAMINED

HOoLotyPE: ?, Uepi Island, Marovo Lagoon, Western
Province, Solomon Islands, Western Pacific, 8°27’S,
157°56’E, intertidal limestone beach rock, 7.vii.1990,
T.B. Churchill, QM $17121.

PARATYPE: 5@ ¢, same data as holotype, 7.vii.1990,
T.B. Churchill, QM S17118-20, QM S17122, QM
$17123.

DIAGNOSIS

Spermathecae two, each consisting of large ter-
minally enlarged lobe with smaller one attached
to the outer base of the former. Thorn spines: 25

TABLE 4. Leg measurements of Idioctis talofa,
holotype 2
I Il Il IV
3.36 2.80 2.36 3.42
2.28 2.16 1.72 2.58
2.12 196 1.64 3.17
164 140 1.28 2.75
0.88 1.08 080 1.00 1.32
10.28 9.40 7.80 12.92 6.84

on prolateral proximal corner of patella III; 10-16
on patella IV; prolateral distal corner of femur IV
with 5 short spines. 2-4 teeth on paired claws.
Rastellum of 10 strong spines in single line with
bases touching. Preening combs a whorl of spines
on metatarsi II, 1V. ¢ ¢ unknown.

Palp
2.52
1.64
1.36

Femur
Patella

Tibia
Metatarsus
Tarsus
Total

HOLOTYPE FEMALE

Carapace 3.92 long, 2.96 wide. Abdomen 5.28
long, 2.96 wide. Total length, 12.

Colour in alcohol. Carapace yellow brown,
orange brown on caput; legs yellow brown;
chelicerae red brown. Abdomen pallid but dorsal-
ly with two purple brown markings at anterior and
posterior ends.

Carapace. With light uniform covering of sil-
very brown hairs, none in striae; light bush on
margins; numerous brown bristles beside 10 thick
and several smaller anteromedial bristles, 6-8
thick bristles between PME, c.10 between AME.
Fovea very slightly procurved.

Eyes. Tubercle low, distinct. AME pigmented.
Group occupies 0.48 of head-width. Three rows;
front row widely separated from middle row, back
row straight. Eye group front width, back width,
length = 32:40:27 MOQ front width, back width,
length = 21:27:13. AME:ALE:PME:PLE =
8:12:6:13. AME-AME, 0.8, AME-ALE, 0.9,
ALE-ALE, 1.3, PME-PLE, 0.1, PME-PME, 1.9,
ALE-PLE, 0.9

Chelicerae. Band of stiff silver bristles prodor-
sally and two narrow bands retrolaterally. Rastel-

Western Province

eee
tT

Guadalcanal Way
12°8

168"E. I

[SOLOMON ISLANDS== |

156°E. 160°E. 164°E.

FIG. 16. Location of Jdioctis marovo sp.nov. at Uepi
Island in the Western Province (inset), Solomon Is.

ny MEMOIRS OF THE QUEENSLAND MUSEUM

ay"
HAT

Ar i or WZ
fa MH,
AEN iA i
A NN A

Me iy
fi 44/ f aie
\ Un Ath 1
H | it ATARI
Wy Wilh! i ¢ \
4 1
LOREEN oR
Te

f

PIG. 17, Idioctis marovo sp.nov. holotype ¥. a, cephalothorax and chelicerae, dorsal view. b, abdomen, dorsal
view. c, abdomen, ventral view, d, stemum, maxillae, labium and chelicerae ventral view. e, eyes, dorsal view,
f, spinnerets, ventral view. Scale line = 1mm (a-d), 0.5mm (e, £).

INTERTIDAL TRAPDOOR SPIDERS 23

FIG, 18. Jdioctis marove sp.nov., dorsal view of sper-
mathecae of ? 9..a, 817119. b, $17123. c, 817122. d,
317121. e, $17120. & $17118. Scale line = 0.25mm.

lum of 10 strong spines in single line with bases
touching overhang cheliceral edge. Furrow
promargin with 7 teeth, all bases separate but most
anterior teeth bases very close, basomesally with
4 small teeth,

Labium. 0.64 wide, 0.28 long. Separated from
sternum by distinct narrow groove.

Maxillae. 1,00 long in front, 1.44 long behind,
0.76 wide; with 5 pointed cuspules on inner angle.
Heel rounded; anterior lobe just distinct.

Sternum, 2.24 long, 1.52 wide. Sipilla small,
oval, and marginal, increasing in size posteriorly.

Legs. With uniform covering of moderately long
brown hairs. Scopula: metatarsi and tarsi I, U,
entire, full; metatarsi I, only on proventral face:
tarsi III, with band of 4-6 rows of setae parting
scopula; metatarsi and tarsi TV absent. Thom
spines: 25 on prolateral proximal corner of patella
TH; 10-16 on patella IV; prolateral distal corner of
femur IV with 5 short spines. Preening combs a
whorl of spines on metatarsi IL, TV.

Spines. No true spines dorsally or laterally but
3-5 long fine black setae on femora I. Spines on
ventral legs generally weak, strong only on distal
metatarsi Il and IV but no so close as to form
comb. Leg 1: fe 0, pa 0, i v2w, me 0. Leg 2: fe 0,
pa O, ti v2, me vlw. Leg 3: fe 0, pa p24 thorn

TABLE 5. Leg measurements of /djectis marovo,
holotype ?

I Palp
4.64 3,60 4.16
3,12 2.48 1.84
3.20 2.64 2.84
2.32 2.40
1.60 1.68

14.88 12.80

4.88
3.28
3.28
2.56
2.32
16.32

Femur
Patella
Tibia
Metatarsus
Tarsus
Total

2.08
10.56

spines, ti v2w, me v10, Leg 4: fe p5 thorn spines,
pa p10-16, ti 0, me v7 in distal whorl. Palp, ti rl,
v4 + 3w.
Claws, Paired claws with 2-4 distinct teeth; pal-
pal claw bare. Claw tufts just enclosing claws.
Trichobothria, In two rows, each of c.11 for 3/4
of tibiae; c.15 in curving row that splits into two
rows for the distal half on metatarsi; 6 clavate and
c.16 filiform that split into two rows each two
wide, two centrally and distally, on tarsi.
Spermathecae. Two, each consisting of large
terminally enlarged lobe with smaller one at-
tached to the outer base of the former.
Spinnerets. PMS 0,26 long, 0.14 wide, bases
0.06 apart, c.0.30 of basal PLS in diameter. Basal,
middle, apical, and total article lengths of PLS =
0.74, 0.12, 0,14, 1,00, respectively.

DISTRIBUTION AND HABITAT

Known from limestone beach rock and the root
bases of coconut trees in the littoral zone on the
island of Uepi, in the Marovo Lagoon, Western
Province, Solomon Islands.

ETYMOLOGY

The specific epithet is taken from Marovo
Lagoon, Solomon Islands (where the spiders were
collected), which has been nominated for World
Heritage listing. In the face of increasing pressure
from mining and logging activities, its pristine
state is a credit to the wisdom of the local people.

REMARKS

Females of /. maroye differ from those of all
other species except /. eniwerok in that patella III
is clearly shorter than tibia III, and from 1.
eniwetok in the presence of numerous thorn spines
on patella I'v.

Nihoa gen.nov. Raven & Churchill

DIAGNOSIS

Nihoa can be distinguished from Idioctis by the
wider sternum, the medially located teeth on the
paired claws of 2 and the absence of a second
row of teeth on the claws of d d. The two genera
are readily distinguished by the presence of dis-
tinct pattern dorsally on the abdomen of Nihou,
Nihoa also differs from both Idioctis and Idioph-
thalma in the absence of a rastellum and from
Rhianodes Raven in the absence of a groove on
the anterior face of the maxillae,

DESCRIPTION
Carapace hirsute, without pattern. Fovea very

24 MEMOIRS OF THE QUEENSLAND MUSEUM

broad, straight to slightly procurved. Abdomen
with distinct pattem or mottling. Rastellum ab-
sent. Maxillae usually with c.10-20 cuspules on
inner anterior corner; anterior Jobe indistinct,
posterior heel produced. Labium wide, without
cuspules. Stemum long, narrow (about 1.4 times
longer than wide, rarely narrower), with small,
marginal sigilla, if evident. Scopulae (? 2) entre
on metatarsi and tarsi I-DT, in distal half mixed
with setae on metatarsi III, and one small tiangle
proventrally on metatarsi IV, if present; dense and
divided by narrow band of setae.

Spines (rare value in parentheses). Leg |:
femur, pO d0-4 10, patella, 0, tibia, p0-1 v2-7. me
pO r0-1. Leg 2: femur, p0 d0-4 r). patella pO, tibia
p0-1 y0-4, metatarsus 0 y1-2. Leg 3: femur pO0-1
JO-5 10, patella p6-30, tibia pO(2) rO(2) vO-7,
metatarsus pO0-3 r0-3 vO-13. Leg 4: femur pO d0-7
r0, patella p3-16, tibia p0.d0.r0(2) v0-7, metatarsus
p0-7 r0-4 v2-9,

Paired claws of 2 ? with teeth on medial keel:
claws of dd with or without one row of teeth.
Tarsi with clavate and filiform trichobothria. d 3:
tibia I with prolateral spur and megaspine, more
distal prolateral megaspine and associated
cuticular ‘thumb’; palpal bulb pyriform; embolic
tip flanged.

TYPE SPECIES
Nikhoa mahina sp.nov.

REMARKS

Raven's (1988) inclusion of Idioctis hawaiien-
sis in Idioctis was presumably based upon the
notion that it was the only genus that could be on
such remote islands. He did identify that the
species was the sister group of all other /dioctis
species but lacked data on the ¢ of the type
species to confirm the generic diagnosis.

DISTRIBUTION AND HABITAT
Known from Nihoa Island in the northern group
of the Leeward Islands from terrestnal habitats,

INCLUDED SPECIES

Nihoa mahina sp.noy,Nihoa 1, Leeward Islands, NW
Pacific

idioctis hawaitensis (Raven, 1988)Necker t., Leeward
Islands, NW Pacific

Nihoa mahina sp.noy, Churchill & Rayen
(Figs 19-22, Table 6)

MATERIAL EXAMINED
HoLotyPe: o, Nihoa I., 22°10'N, 163°10°W. Leeward

Islands, northwest Pacific Ocean, 40ft.(12.2m) above
sea-level, on rocks near camp in moonlight, 15.11.1981,
§. Conant, Bernice P. Bishop Museum,

ParRaTyres: % (allotype), Nihoa I., 22°10'N,
163"10'W, Leeward Islands, Hawaii, in sleeping bag,
vi.1981, S. Conant; 2, Nihoa L., 2.11.1981, drowned in
seep pool, S, Conant; ¢, Miller Valley, Nihoa I,
vi. 1982; all in Bernice P, Bishop Museum.

DIAGNOSIS

Paired claws of d ¢ with teeth. Upper cuticular
process on tibia I of dd at base of spur. Sper-
mathecae two coniform mounds.

HOLOTYPE MALE

Carapace 10.00 long, 9.67 wide. Abdomen
10.00 long, 6.83 wide. Total length, 24.

Colour inalcohol, Carapace, legs and chelicerae
red brown. Abdomen dorsally yellow brown with
medial longitudinal discontinuous band of brown
flanked laterally by somewhat paired, brown spots
anteriorly and recurved brown bands posteriorly.
Abdomen ventrally yellow brown with four small
brown marks almost equidistant, centrally. Ster-
num, maxillae, and labium red brown. Spinnerets
yellow brown.

Carapace. Fine silvery hairs form uniform
covering intermixed with numerous setae; line of
6 thick-and several finer anteromedial bristles, and
several thick bristles between ALE; striac
glabrous. Fovea slightly procurved.

Eyes, Tubercle low. Group occupies 0.24 of
head width. Eyes in three rows. Eye group: front
width, back width, length = 71:82:60. MOQ front
width, back width, length = 44:60:31.
AME:ALE:PME:PLE = 16:22:16:24. Eye inter-
spaces: AME-AME, 1.1, AME-ALE, 1.0, ALE-

FIG. 19. Localities of Nihoa in the Leeward Islands. N,
mahina sp.nov. on Nihoa l. and N. hawaiiensis
(Raven) on Necker I,

INTERTIDAL TRAPDOOR SPIDERS 25

FIG. 20. Nihoa mahina sp.nov. holotype d . a, cephalothorax and chelicerae, dorsal view. b, eyes, dorsal view. c,
sternum, maxilla, labium and chelicerae, ventral view. d, spinnercts, yentral view. e, abdomen, dorsal view. f,
abdomen, ventral view. Scale line = 2mm (a, c, ¢, f), Lmm (b, d).

ALE, 2.3, PME-PLE, 0,1, PME-PME, 2.4, ALE-
PLE, 1.1.

Chelicerae. With prodorsal and lateral bands of
setae and fine silver hairs. Rasiellum only a line
of 20-25 long pointed brisiles on distal edge. Fur-
row promargin with 8 thick teeth and 3 smaller
teeth; 12 small teeth basomesally. Intercheliceral

tumescence absent.

Labium. 1.76 wide, 0.72 long. Labiosternal su-
lure a distinct groove.

Maxillae. 2.96 long in front, 3.60 long behind,

1.60 wide; with c.12 cuspules on inner edge. Heel
distinct; anterior lobe indistinct,
Sternum. 5.60 long, 4.08 wide. Sigilla marginal,

round, and posterior pair largest.

Legs. Formula 4123. Tibia I with distal,
prolateral spur and short megaspine; base of spur
with broad cuticular process below which is one
(rarely two) spine. Scopulate surface of tarsi pal-
lid.

Scopula. Leg 1, full, thick, entire and undivided
on metatarsi and tarsi I, If. Leg IL scopula extends

26 MEMOIRS OF THE QUEENSLAND MUSEUM

a a
=== s.

FIG, 21. Nihoa mahina sp.nov. holotype ¢ . a, tibia I, prolateral view showing distal spur and megaspine, hair and
scopula omitted. b, tibia I showing distribution of hair and scopula. c, palpal bulb. d, palpal tibia, cymbium and
bulb, ventral view. e, palpal bulb. Scale line = 1mm (a, b, d), 0.5mm (c, e).

to retrolateral side; metatarsi II], thin on distal half
extending to prolateral side with setae intermixed;
tarsi II, entire, thick with setae intermixed;
metatarsi IV, with distal prolateral 1/4 with thin
scopulae; tarsus ITV with setal band 2-4 rows
across intermixed with thin scopulae.

Spines. Leg 1: fe p2, pa 0, ti p2+2, v3+2, me 0.
Leg 2: fe p2, pa 0, ti pl, v2, me 0. Leg 3: fe p2, 13,
pa 0, p2+1w, ti p2, r2, v5, me p4, r5, v10. Leg 4:
fe p2, d2, 12, pa 0, tir5, v10, me p6, r5, v17. Palp:
pa 0, ti vl.

Palp. Bulb with distally flanged embolus. Cym-
bium with two deeply divided lobes, retrolateral
face slightly convcave.

Claws. 2-4 (leg I) to 0 (leg IV) teeth in medial
keel on paired claws. Tufts dense, enclosing
claws.

Trichobothria. c.8 for half length of tibia, c.12
on metatarsi in single curving row; two bands each
of c.4 small clavate and c.10 filiform on tarsi.

Spinnerets. PMS 0.84 long, 0.36 wide, bases

0.32 apart, c.0.33 of basal PLS in diameter. PLS
basal, middle, apical, and total article lengths =
1.12, 0.64, 0.52, 2.28, respectively.

ALLOTYPE FEMALE

Carapace 10.00 long, 8.33 wide. Abdomen
10.33 long, 7.33 wide. Total length, 25.

Colour in alcohol. Carapace and legs orange
brown, chelicerae red brown. Abdomen dorsally
yellow brown with medial longitudinal discon-
tinuous band of brown, flanked laterally by some-
what paired, brown spots anteriorly and recurved
brown bands posteriorly. Abdomen ventrally yel-
low brown with four small brown marks almost
equidistant, centrally. Sternum, maxillae, and
labium orange brown. Spinnerets yellow brown.

Carapace. Fine silvery hairs form uniform
covering intermixed with numerous setae well
developed between head region and fovea; striae
glabrous; fovea straight; c.30 setae between AME,
c.14 between PME, c.40 long setae along medial

INTERTIDAL TRAPDOOR SPIDERS 27

A i
Aye

ny eit! NUG A

My i el

PANTY ii at

Vai

Ny il }

FIG, 22, Nikoa mahina sp.nov. paratype %, a, cephalothorax and chelicerae, dorsal view. b, eyes, dorsal view, c,
spermathecaé. d, sternum, maxillae, labium and chelicerae, ventral view. e, spinnerets, ventral view. f, abdometi,
dorsal view. g, abdomen, ventral view. Scale line = 2mm (a, d, f, g), Imm (b, €), 0.5mm (c).

caput; lateral edge of carapace with several
anterior and posterior directed lines of setae with
dense fine silvery hairs. Clypeus absent.

Eyes. Tubercle low. Group occupies 0.30 of
head width. Eyes in three rows. Eye group front
width, back width, length = 72:81:55. MOQ front
width, back width, length = 45:55:35.

AME:ALE:PME:PLE = 20:23:12:25. Inter-
spaces: AME-AME, 0.4, AME-ALE, 0.8, ALE-
ALE, 2.6, PME-PLE, 0.1, PME-PME, 1.8,
ALE-PLE 0.8.

Chelicerae, With prodorsal and lateral bands of
setae with fine silver hairs. Rastellum absent, c.35
strong bristles in line on distal edge. Furrow

2 MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 6, Leg measurements of Nikea mahina,
holotype 3
I II
833 «68.16
533 4.83
6.17 3.33
583 5.66
Tarsus 350 383 2.83 3,16
Total 29.16 27.8) 24.12 32.32

promargin with 8 thick teeth and 3 smaller teeth;
6 basomesal granules.

Labium. 1.76 wide, 1.12 long. Labiosternal su-
ture broad,

Mazxillae. 2.72 long in front, 3.68 long behind,
1.60 wide; with c.10-12 cuspules on inner edge.
Heel angular; anterior lobe, indistinct.

Sternum. 5.44 long, 3,76 wide, Sigilla: anterior
and medial pair not evident; posterior pair small,
round, marginal.

Legs. Thorn setae and preening combs absent.
‘Tarsi broad, tarsi I c.60% of length wide; tarsi IV
distally incrassate, distal width c,1.4 times basal
width.

Scopula. Very dense on legs [, Ll: metatarsi and
tarsi I, If, full, thick, entire, undivided; metatarsi
U1, thin on distal half with thick setae intermixed;
larsi TI, entire, thick with 3-5 setae intermixed;
metatarsi IV, thin in distal prolateral 1/4, divided;
tarsi [V, thick, full, divided by narrow setal band
2-4 rows across and few setae intermixed, setal
brush dense distally.

Spines. Leg 1,.0. Leg 2: 0, save ti pl. Leg 3: fe
p2w, r2w, pa p2 thorns, Hi p2, rl. v3, me p3, r3, v7.
Leg 4: fe 0, pa 0, tirl, v4, me p2, rl, v9. Palp, fe
plw, pa 0, ti p3w, vow.

Claws. 4 (leg I) to 0 (leg IV) teeth on paired
claws on medial keel. Tufts dense, just enclosing
claws, Palpal claw very small, curved, bare. _

Trichobothria, In wo rows each of ¢.8 for half
length of tibia; c.11 on metatarsi in single curving
row; c.3-5 clavate and 12-14 filiform in each of
two bands on tarsi.

Spermathecae. Two coniform mounds.

Spinnerets. PMS 0.96 long, 0.44 wide, bases
0.16 apart, c.0.53 of basal PLS in diameter, PLS
basal, middle, apical, and total article lengths =
L.12, 0.44, 0.48, 2.04, respectively.

Tl IV
6.16 9.50
4.83 4.83
5.00 6.83
5.33 68.00

Palp
3.50
3.16
3,33

Femur
Patella
Tibia
Metatarsus

2.350
14,49

DISTRIBUTION AND HABITAT
Known only from the island of Nihoa in the
Leeward Islands, northwest of Hawaii.

ETYMOLOGY
‘The generic name refers to the type locality. The
species name is a Hawaiian word for moon, given

TABLE 7. Leg measurements of Nihoa mahina,
allotype 2
I Il
6.66 6.50
466 416
3.83 4,00
3.50 3,16
2.00 2.33
20.65 20.15

Ti IW
5.66 7.33
3.50 4.66
3.33 5.50
3.50 5.33
1.83 2.66

17.82 25.48

Palp
5.00
3.50
2.66

Femur
Patella

Tibia
Metatarsus
Tarsus
Total

3.16
14,32

that the type specimen was collected in the moon-
light.

REMARKS

Males of NV. mahina differ from those of WN,
hawaiiensis in the presence of teeth on the claws
of Jeg I and in the basal position of the cuticlar
process on tibia I, Females differ in the coniform
spermathecae, in having 4, rather than 2, teeth on
the claws of leg I, and in the darker abdomen
dorsally.

Nihoa hawaiiensis (Raven), new comb,
(Fig. 19)

Idivctis hawatiensis Raven, 1988:6; Platnick, 1989-93,

MATERIAL EXAMINED

Hovotyre: d, Necker I., Leeward Islands, north-west
Pacific, 23°35'N, 164°42’ W, B.H. Bryan Jr, 29.vi.1923,
AMNH.

PARATYPE; 9, same data as holotype, AMNH.

OTHER MATERIAL

The types and also from Necker I., Leeward Islands: 3,
23.vi.1982, S. Conant, 42 9, Annex Hill, alt. 82m.,
24.vi.1984, S. Conant; all in Bernice P. Bishop
Museum, Hawaii.

DIAGNOSIS

Paired claws of dd without teeth, Upper
cuticular process on tibia of ¢ ¢ prolateral, well
above spur. Spermathecae two long domed
mounds.

DISTRIBUTION, BURROW AND HABITAT

The spiders were found under rocks in burrows
on very sparsely vegetated hillsides on Necker
Island, one of the Leeward Islands, northwest of
Hawaii.

SPERMATHECAL VARIATION IN /DIOCTIS

This paper includes the first documentation of
spermathecal variation in 3 genus of the family
Barychelidae. Shape of spermathecae has been

INTERTIDAL TRAPDOOR SPIDERS 28

used widely in mygalomorphs to distinguish be-
tween species (e.g., Schiapelli & Gerschman de
Pikelin, 1962; Forster & Wilton, 1968: Raven,
19$4a). As such, spermathecal shape has been
considered a relatively stable and useful diagnos-
tic character. Rarely are exceptions well sup-
ported. In his excellent and detailed revision of
fuagrus, Coyle (1988) showed extensive vana-
tion in the shapes of spermathecae of one species,
£, mexicanus. On the other hand, Rayen (1984b,
1990) found in the Aname macilata group
(Nemesiidae) and Tritiame (Barychelidae).

respectively, that spermathecae are almost con-
stant within a species group. Moreover, Raven &
Churchill (1991) noted that one shape is
widespread in barychelid genera and may be the
family synapomorphy.

Among mygalomorphs, barychelids have
received little attention from taxonemists or
ecologists, presumably because they build cryptic
burrows. They are also under-represented in
museum collections. Hence, from the limited
material available variation in shape of sper-
mathecae has been unclear but assumed to be
minimal. In /dioctis, we find that this variation is
usually too great to be useful as a specific charac-
ler on is own, even though it was stable in species
of Encyocrypta .

Recognition of amumber of species in which the
shape of the spermathecae cannot be diagnostic
muy seem unwarranted, In a major revision of
Pacific barychelids (Raven & Churchill, in prep.),
a plesiomorphic spermathecal shape has been
identified and found to be widespread. However,
as with /dioctis, barychelid males do show
mutually exclusive differences where none were
evident among their (unequivocally) conspecific
females, Hence, interspecific differences can be
sufficiently clarified with non-sexual somatic
characters.

ACKNOWLEDGEMENTS

We sincerely dedicate this paper to Ms Julie
Gallon, who assisted with the collection of /diactis
yerlata, Julie has been missing since 2 August,
1990, and we will continue to miss her dearly, This
research, visits to European museums, and collec-
trons made in New Caledonia, Fiji, the Solomon
Islands, and Singapore were funded by an
Australian Research Council grant to RIR, The
authors are indebted to the generous hospitality of
Dr Jean Chazeau of O.R_S.T.0.M,, Nouméa, New
Caledonia. Ms Clare Bremner and Mrs Bronwyn
Mitchell made the excellent figures 1, 8, 11, 14,

17, 20-22; RJR did the rest. We kindly thank Mr
Joseph Koh, and Associate Professor Dennis Mur-
phy of the University of Singapore, for their
hospitality and assistance to RJR in acquiring
comparative material of J. Jittoralis. Mr John
Ravenscroft kindly assisted in the search for /diac-
tis helva, in Fiji, We are grateful to Dr G, Rack,
Zoologisches Institut and Zoologisches Museum,
Hamburg, for the loan of types of Idioctis heiva,
to DrJ. Gruber, Naturhistorisches Museum, Vien-
na, and Mr P.D. Hillyard, British Museum
(Natural History), London for the loan of Idiactis,
material from Samoa, and to Ms Sabina Swift,
Bernice P. Bishop Museum, Honolulu, for send-
ing the new material of Nihoa. The invitation of
Dr Harry Pamaby to TBC to accompany the
Australian Museum expedition to the Solomon
Islands is gratefully appreciated. TBC warmly
thanks the local people of the Marovo Lagoon area
for their co-operation and hospitality. We are
grateful to Dr S. Conant for data on habitats of
Nihoa mahina and Nihoa hawaitensix in Hawaii.

LITERATURE CITED

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BENOIT, P.L.G. 1966. Les Barychelidae-
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BENOIT, P.L.G. & LEGENDRE, R. 1968, Un
barychélide nouveau du Madagascar:
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BRIGNOLI, P.M. 1983, ‘A catalogue of the Arancac
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COYLE, F,A. 1971, Systematics and natural history of
the mygalomorph spider genus Antrodiaetus and
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of the Museum of Comparative Zoology 141,
269-402,

30 MEMOIRS OF THE QUEENSLAND MUSEUM

1988. A revision of the American Funnel-web
mygalomorph spider genus Evagrus (Araneae,
Dipluridae). Bulletin of the American Museum of
Natural History 187:203-292.

FORSTER, R.R. & WILTON, C.L. 1968. The spiders
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HOGG, H.R. 1901. On Australian and New Zealand
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KOCH, L. 1874. ‘Die Arachniden Australiens, nach der
Natur beschrieben und abgebildet”, Vol.1, pp.473-
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MAIN, B.Y. 1985. Mygalomorphae. In Walton, D.W.
(ed.), “Zoological Catalogue of Australia, 3.
Arachnida; Mygalomorphae, Araneomorphae in
part, Pseudoscorpionida, Amblypygi, and
Palpigradi’, (Australian Government Publishing
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MARPLES, B.J. 1951. Mygalomorph spider in Samoa.
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1955. Spiders from Western Samoa. Journal of the
Linnean Society of London, Zoology 42:453-504.

PLATNICK, N.I. 1989. ‘Advances in Spider
Taxonomy 1981-1987: A supplement to
Brignoli’s A catalogue of the Araneae described
between 1940 and 1981’. (Manchester University
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POCOCK, R.]. 1903. On the geographical distribution
of spiders of the order Mygalomorphae. Proceed-
ings of the Royal Society of London 24(1):340-
368,

RAINBOW, W.J, 1911, A census of Australian
Araneidac. Records of the Australian Museum
9(2):107-319,

RAVEN, R.J. 1984a,. The Australian curtain-web

spiders (Ischnothelinae: Dipluridae: Chelicerata),
Australian Joumal of Zoology, Supplementary
Series 93:1-102.

1984b. A revision of the Aname maculata species
group (Dipluridae, Araneae) with notes on
biogeography, Journal of Arachnology 12:177-
193, tables 1-5,

1985. The spider infraorder Mygalomorphae
(Araneae): cladistics and systematics. Bulletin of
the American Museum of Natural History 182.
21-180, tables 1-9.

1988. A revision of the mygalomorph spider genus
Idioctis (Araneae, Barychelidae). American
Museum Novitates 2929: 1-14,

1990. A revision of the Australian spider genus Trit-
tame Koch (Mygalomorphae: Barychelidae) and
a new related genus. Invertebrate Taxonomy 4:
21-51.

RAVEN, R. J. & CHURCHILL, T.B. 1991. A revision
of the mygalomorph spider genus Encyocrypta
Simon in New Caledonia (Araneae:
Barychelidae). In Chazeau, J. and Tillier, S. (eds),
‘Zoologica Neocaledonica’. 2. Mémoires du
Muséum national d’ Histoire naturelle, A, 149:31-
86.

ROEWER, C.F. 1942. Katalog der Araneae. (Paul Budy
Bd.: Bremen) 1:i-viii + 1-1040.

SCHIAPELLI, R. D. & GERSCHMAN DE PIKELIN,
B.S. 1962. Importancia de las espermatecas en la
sistematica de las arafas del] suborden
Mygalomorpha. Physis (Buenos Aires) Secc. C,
23 (64):69-75,

SIMON, E, 1892, ‘Histoire naturelle des araignées’.
1(1): 1-256. (Roret: Paris).

1903, “Histoire naturelle des araignées’. 2(4)
Supplément Général :875-1080. (Roret: Paris).

SHIPWRECK AND HISTORICAL SURVEY: CHILCOTT ISLET. CORAL SEA
R.A. COLEMAN

Coleman, R.A. 1992.06 29: Shipwreck and historical survey: Chilcott Islet, Coral Sea. Memoirs
of the Queensland Museum 32(1); 31-53, Brisbane, ISSN 0079-8835.

Many Coral Sea recfs appearing on modern charts were first discovered by unsuspecting
sailing ships that struck the reefs and were wrecked in the | Sth and 19th centunes. [n 1845 the
teak, India-built barque, Coringa Packet, was totally wrecked on a reef surrounding a small
sand islet subsequently named Chilcott Islet after the ship’s Captain.

In the Jate 1870's, Chilcott Islet was briefly mined for guano (phosphate) by F.B. Beaver &
Co, of Melboume. During 11-12 December. 1991. the author located wreckage believed to be
the remains of Coringa Packet and conducted a brief preliminary s , The wreck is
considered to be archacologically significant because of its potential to illustrate technical
detail of a poorly-documented vessel type (‘Country vessel’/Opium Clipper) and of Indian
shipbuilding practice during the first half of the 19th century. [_| shipwreck, Indian shipbuild-
ing, guano mining, archaeology, history, Coral Sea.

R.A. Coleman, Queensland Museum, PO Box 3300, South Brisbane, Queensland 410),

Ausiralia; 14 January, 1992,

The Coringa Islets, comprised of Chilcott Islet
and South West Islet, lie respectively in 16°57°S
by 150°00°E and 16°59°S by 149°53'E in the
Coral Sea.

The group is named after the sailing barque
Coringa Packet, which was wrecked there in
1845. Chilcott Islet, where the wreck actually
occurred, is named after the ship's Captain, F.
Biggar Chilcott.

The Coringa Islets are within the Cormga-
Herald National Nature Reserve which was
proclaimed under the National Parks and Wildlife
Conservation Act 1975 on 16 August, 1982. The
Reserve is managed by the Australian National
Parks and Wildlife Service {(ANPWS).

The Queensland Museum administers the Com-
monwealth Historic Shipwrecks Act 1976 in the
Commonwealth Coral Sea Territory on behalf of
the Department of Arts, Sport, Environment, and
Territories (DASET). An essential administrative
duty of the Museum, under the Act, is to locate,
identify and assess the Australian maritime
heritage resource and make recommendations to
the Minister for its better appreciation, protection
and preservation,

On the basis of archival research, it was deter-
mined that the Cerixga Packer wreck had the
potential to meet criteria for additional protective
measures and warranted preliminary field inves-
tigation.

An opportunity was presented for the author and
a volunteer assistant to join a multidisciplinary
scientfic charter organised by Dr James Charley
of the Botany Department, University of New

England, Armidale, Dr Charley obtained the
necessary permit to conduct scientific research
within the Reserve. The charter vessel, TSMV
Kanimbia was to visit a number of reef and islet
systems in the area including Lihou, Tregrosse,
Diamond, Magdelaine, Coringa and Herald for a
total voyage time of fourteen days,

Although it was intended to Jocate and examine
two nineteenth century wrecksites at the eastern
end of Lihow Reef, whilst there bad weather
caused the abandonment of that part of the pro-
gram. However, the position of a recent, uniden-
tified wreck was noted but persistent rain squalls
prohibited our approaching the site with safety.
Only its two masts and central stack are visible
above water and it appears to be a cargo vessel ar
bulk carrier of approximately 120-150m in length.
The condition of the mast tops suggest that the
wreck occurred within the past 10-20 years.

Whilst ashore at Turtle Islet, Lihou Reef,
material was discovered on the beach near the
waterline at the northern end, which may warrant
an investigation of the surrounding reef when the
opportunity presents itself. The material consisted
of a stone (suggestive of ship’s ballast), a lip
fragment of anineteenth century bottle, an uniden-
tified small circular copper attachment plate and
seyetal small pieces of iron (which were sugges-
tive of hoop iron from wooden casks).

Our main objective was to locate and examine
the wreck of the Coringa Packet at Chilcott Islet.
Due to the constraints of a very tight schedule,
only one and one half days could be spent there,
onc third of which time was only partially produc-

32 MEMOIRS OF THE QUEENSLAND MUSEUM

“,
\CORAL

| CORINGA-HERALD NATIONAL
\ NATURE RESERVE 43

ee

LIHOU REEF NATIONAL
NATURE RESERVE

Ss
-\_ ISLANDS
bee:

‘

a
~~ TERRITORY

\
Dione

fe =e LS SS

AUSTRALIA

FIG. 1. Location of Coringa-Herald National Nature Reserve.

33

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY

“QATOSAY SU} UIYIIM S}a[s] BSULIOD Jo VOLO] ‘7 ‘OLA

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4 MEMOIRS OF THE QUEENSLAND MUSEUM

live due to hazardous ses conditions on the reef
flat.

HISTORY OF CORINGA PACKET

Today, little is known of the early history of the
Coringa Packet. However, shortly after the
Britsh Expeditionary Force was sent to resolve
the opium issue in China in 1840 (an action now
generally referred to as the ‘Opium Wars’), she
was for a period engaged in the Opium Trade. At
that time she was described as being rigged as a
bri

2:

The vessel, under the command of F, Biggar
Chilcott, arrived in Sydney on 11 March, 1845 and
anchored off Campbell’s Wharf having departed
Calcutta the 11th December, Madras the Sth
January, Trincomalee the 14th January, and
Hobart Town the 2nd March. She carned twelve
passengers and a general cargo.”

Advertisements in the Sydney Morning Herald
during March, 1845 announcing her availability
for cargo for Ceylon and Madras described the
vessel as ‘Teak Built’.

‘These items of information suggest that she was
built and registered in India, probably Calcutta, as

YOR CEYLON AND MADRAS.
7 - FENHE TEAR WARQLE
CORINGA PACKET,
Captaln Chileett: wil. sail for
the abort porte about the let
April For freight or passage spply to the
Ceptais, on board; ur fn ‘
mls LYALL, SCOTT,

YN CO.

FIG. 3. Advertisement in the Sydney Morning Herald
during March 1845

a “Country Vessel’ (India based, under European,
American, or Parsee ownership: a concession
begrudgingly given by the East India Company,
which had earlier held the monopoly on East In-
dian wrade.). Correspondence subsequent to her
wreck, and the quantity of return cargo belonging
to Chileon, suggests thai she was owned by Chil-
colt and possibly under charter to the Honourable
East India Company as an ‘Extra Ship’, This
would explain why she was not listed in Lloyd's
Register. The H.E.I, Co. maintained its own in-
surance, Phipps * fists most ships built in, and
operating from, India up to 1839. He makes no
mention of Coringa Packet, which may have been
built in 1840-1,

For reasons explained Jater in this report, it is
important to discover where the ship was built and
archival research will need to continue into the
problem. '

When Coringa Packet departed Sydney bound
for Ceylon and Madras with a mixed cargo and a
consignment of mail on 26 April, 1845, she was
listed as a 230 ton barque. Four passengers were
officially recorded as being on board: Lieutenant
Blackhall, Ensign Bloomfield, Mr E. Lord, and Mr
Henry Wan Deerlin. A large part of the craw were
Indian seamen or ‘Lascars’ making a total com-
plement of forty. Lyall, Scott, and Company acted
as her agents in Sydney.

Chilcott, upon his arrival in Sydney, had been
the bearer of correspondence addressed to the
prominent pastoralists and horse breeders James
and William Macarthur of Camden from their
agent in Calcutta, G.A. Plaistowe and the auction
house, Cockerell and Company. The Macarthurs
had found a ready market for good horses among
the officers of the Bengal Army and other affluent
Europeans in India and had recently sent a ship-
ment to Calcutta on the East Indiaman Blundell,
Plaistowe reported, *...there appears a greal
demand for these horses, and they have been ex-
pected and talked of, three months before our
arrival, and 1 hope they may go off well. Auction
sales appear the best and most fair way of each
horse attaining its value, and it is arranged that the
16th and 2131 December, be the days of sale - half
each day. I could sell several privately, well, but!
find that would prejudice the remaining ones aj
auction, wherein purchasers Would say the best
had been taken, and the culls, only, sold publicly.
lam advised, also, from Cockerell’s House, to
such a course.’

‘Good horses will sell well here, but itis great
folly in shippers sending unbroken horses. The
first question | am asked here, speaking of horses
- What kind of cattle? Are they broken fo saddle,
side-saddle, hamess, double or single? I have hor-
ses that are not broken upon the Catalogue; then,
they say, What a pity - you should always bring
horses here, thoroughly broken, and they will meet
ready sale and realise good prices.’

Chilcott was not slow to realise the potential for
profit and when Coringa Packet finally cleared
Sydney Heads there were three horses on board as
part of the cargo consigned under the Captain’s
name, His speculation may have been influenced
by one of his passengers, Lieutenant Blackhall,,
who had been sent to Australia to inyestigate the
potential supply of horses and other livestock to
the British military in India.

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 35

What then took place is best described by Chil-
cott himself;

‘On the Sth May, passed Wreck Reef: the wind
hanging far to the eastward prevented us from
weathenng Alert Reef and shoals in the vicinity;
bore away to the westward to clear these dangers.”

‘May 7. Passed through the group of islands
knowo by the name of Tregoss’s Islands
{Diamond Islets in the Tregrosse reef system];
shaped a course N.W. by W. with the intention of
sighting Lizard Island, and at 9:45 P.M. while
sailing at the rate of eight Knots an hour, the ship
struck with a tremendous crash, on a reef, and
slewed broadside on; sounded the well, found
seven feet water -cul away the mast and let 20 the
anchor, for fear the ship might beat out into deep
water and sink, as we could nel make oulanything
about us buta foaming sea: the vessel] continued
striking yrolently, and the sea making a complete
breach over all, and, as you may suppose, daylight
was most anxiously looked for; and to our great
joy, a small island or sand bank was observed to
the S.W., distance about half-a-mile; (fortunately
the mast had gone over the side without injury to
the boats). We then rigged a pair of shears, and
with great difficulty got the long boat out; all
hands next employed in procuring water and
provisions, and Landing it on the island”

“At night, all hands left the wreck, and slept on
shore; at daylight, went again to the wreck, which
we found had suffered much during the night, and
got the water and provisions out; remained on the
island until the 21 stof May, when we then decided
to man the three boats and proceed for Torres
Straits, in the hope of falling in with Her Majesty's
ship Fly, or secking some means for the rescue of
the remainder of the crew left on the island, com-
posed of twenty-four souls. We left on the tsland
800 gallons of water, and provisions sufficient for
four months; and T trust, ere that time expires,
stime measures will be taken to remove them-

*..We had a boisterous passage in the boats,
from the wreck to the Barrier Reef, and lam sorry
tosay, we Lost the gig, which upset on the night of
the 23rd, and Mr. E. Lord, passenger, was
drowned; the remainder of the crew were picked
up by myself and the cutter, under the charge of
the chief officer,”

Chilcott and his group of fourteen sailed the
remaining two boats through the Barrier Reef to
Booby Island, on the western side of the Torres
Swait, arriving on the 31st. There, they not only
found sustenance in the cave Jater known as the
‘Post Office caye', but anoiher group of
shipwrecked people. These were the survivors of

the Hydrabad which had been wrecked a few days
earlier near Murray Island and who had arrived! at
Booby the day before.®

Tt was immediately obvious to Chilcott and Cap-
lain Robertson of Hydrabed that although a ship
might be expected to call into the island al any
time, the huge number of people who must suryiye
on the limited supplies in the cave could nol
remain there for long. Also, Chilcott knew that to
notify the authorities of his temaining crew on the
islet, he must quickly convey the news to a port
from which ships sailed to Sydney. Chilcott and
his group were exhausted after eleven days in the
boats. With a volunteer crew and several of his
passengers, Captain Robertson immediately set
out for Port Essington in their long boat.

In the meantime, the twenty-four Lascars lefl on
Chilcott Islet had reached 2 decision. The con-
struction of araft from materials salvaged fron the
wreck had begun before Chilcott and the others
had departed in the boats. Having little faith in the
boats’ chances of success, twenty-one of Theos
decided to attempt to teach the mainland. They set
out twe days behind Chilcott, on the 23rd of May.
leaving three less venturous men on the Isler.

Two days after Robertson's long boat left
Booby Island, on 2nd June, the schooner Shani-
rock, Captain G. Browning, called in and took on
board Chilcott and the fifty remaining survivors
of the two wrecks, Their first port of call was Port
Essington where, on the 7th June, they found
Robertson had safely arriyed, Chilcott had left
letters at Booby Island for Captain Blackwood of
HMS Fly advising of the wrecks, the survivors’
progress to Port Essington, and of the situation of
the Lascars left at Chilcott Islet.

The Fly with her small tenders Prince George
and Midge had been conducting surveys off the
south coast cf New Guinea and had become
separated from the Midge and the ship’s second
gig. Captain Blackwood dispatched the Prince
George lo Booby Island for intelligence, as that
was their emergency rendezvous, and on 2 June
the Fly weighed anchor and followed.

J. Beete Jukes. Naturalist on Fiy recorded, ‘On
nearing Booby Island [on the Sth of June), we saw
the appointed signa) for good news flying, and
found that our boats, having missed us on the coast
of New Guinea, after a vain attempt to reach
Caedha, or Bramble Key, had run for this little
islet We had left provisions here in the early part
of the year forany ship- wrecked people that might
come in their boats [the N.S,W. Government
maintained supplies in the “Post Office cave").
Availing themselves of these, and procuring a

MEMOIRS OF THE QUEENSLAND MUSEUM

36

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CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 3

fresh supply of water near Port Lihou, in En-
deavour Strait, fa bay on the south side of Prince
of Wales Island, twelve miles to the cast. The
Coringa Packet and Hydrabad survivors had evi-
dently used all the cask water in the cave. The only
wher water on the island is in a smal) rock catch-
ment which is difficult to locate and few knew
about its existence al thal time], our peuple had
Icft the second gig on Booby Island, and gone on
in the Midge to Port Essington. We also found
intelligence that two large vessels had been
wrecked - one the Hyderabad, in the Cumberland
Passage, near Murray Island, another the Coringa
Packet.’

‘The crews and passengers of these, consisting
altogether of seventy or eighty people, had also
visited Booby Island, and gone on to Port Es-
sington, partly in their boats and partly in a small
schooner [Shanirock] that happened to pass by.
Among the passengers by the Hyderabad were
some ladies from Van Diemen’s Land, on their
way to India, whose acquaintance we had had the
pleasure of making when in Hobarton’

The Fly immediately sailed to Port Essington
amiving on the 12th of Jane. They found that
Shamrock had already sailed on the 10th for
Ceylon with Chilcott, his mates, and two or three
hands. The remaining shipwreck survivors,
‘amounting to nearly 70 persons were now
residing [there], and as it was impossible for the
garnson to support this number much longer, and
there was no probability of any other vessel touch-
ing for some months, the “Fly” was under the
necessity of taking them to Sincapore....'

Port Essington was out of the way and of no
commercial interest to merchantmen coming
through Torres Strait. Most sailed directly
northwest from Booby Island for Coupang in
Timor or through one of the Indonesian straits to
Singapore.
‘On the 11th of June’, Captain J.B. Pain of the
brig Spy reports, *...1 took eleven of her crew fall
that remained of Coringa Packet’s Lascars who
had sailed on the raft] off the nethermost [north-
ernmost] of Sir Everard Home's Islands [at the tip
of Cape Grenville], where they had been for fif-
tcen days, subsisting on shell-fish and what little
water they could find in the cavities of the rocks;
three: died the morning | fell in with them, and the
others were in a dreadful state of exhaustion.
These are the only survivors of twenty-one men
who left the wreck on a raft on the 231d of May,”

Tt would appear that a mistake in dates was made
here and that the Lascars had arrived at the Home
Islands on the 30th of May. According to Captain

John Hews of the Maid of Athens, Pain picked up
the surviving Lasears on the 15th of June. And
Pain varies his figures in subsequent reports to say
that he rescued ten, not eleven men. that eleven,
not ten had died, and that they had been im the
Home islands sixteen, not fifteen days.'”

Captain Pain had been busy, On the 3ist of
May, he had picked up the survivors of the Mald
of Athens, which had been wrecked, just after
midnight on the Mth, southeast of Sir Charles
Hardy Islands, which are a couple of miles east of
Home's Islands.'' His delay of several days in
picking up the Lascars a few miles away must
have been occasioned by his standing by the wreck
of Maid af Athens to salvage what he could.

Arriving at Booby Island on the 17th of June,
Pain found that Chilcott’s groupand the Aydrabad
survivors hac already departed for Port Essington
and that Fly had proceeded after them. Pain Ieft a
letier at the Booby Island “Post Office’ advising
of his rescue of cleven of the Lascars and the
wreck of Maid of Athens. A copy of his letter was
entrusted to the Captain of the brig Joxepit
Wheeler, then anchored at Booby enroute to
Manila. Spy then proceeded with her passengers
directly to Coupang arriving there on the 4th of
July. "Asa consequence, those at Port Essington
were unaware that only three men remained at the
Coringa Packet wreck.

Prince George, with some of the survivors on
board, parted company with Fly at Port Essington
on the 18th of June and proceeded to Sydney by
the western route. The cutter arrived at Sydney on
the 13th of August bnnging news of the ship-
wrecks and the mistaken advice that twenty-four
Coringa Packet Lascars were awaiting rescuc. ~

Governor Gipps immediately called | for tenders
for a vessel to proceed to the reef, *...and a
schooner called the “Frolic” was chartered for the

rpose. The Government however required that
she should be commanded by a competent
navigator and Mr. Yule [Commander of the sur-
yey vessel HMS Bramble which was at Sydney
undergoing a refit] was requested to examine the
candidates: the former master of her, [William
Pilfold], broke down at once, and out of 6 or 7
others examined only one understood how to work
outa common set of sights for time!: the owners
were ulso very anxious for the master of the
“Fralic” who had been a long time-in their employ
to continue in her, but this Mr. Yule would not
consent to unless a competent “navigator” went
with him, and the only one who had passed a
salisfaciory examination cbjected 10 go in a sub-
ordinate capacity: all this occasioned considerabic

38 MEMOIRS OF THE QUEENSLAND MUSEUM

Port
Essington

Gulf
if

Ql °
Carpentaria

¥ Booby Is.

Chilcott Islet

FIG. 5. Chart showing relative positions of Chilcott Islet, Home Islands, Booby Island, and Port Essington.

delay but at length a person [Mr G. Levian] was
found who proved fit & willing to go as “mate &
navigator” with the former Skipper, and on the
19th of August the “Frolic” sailed.’"4

‘In the mean time however the “Heroine”
whose master, McKenzie [Martin MacKenzie], a
very clever enterprising little fellow had also ap-
plied, tho’ in vain, for the contract, sailed for Java
and in her passage touched at Chilcott’ § Bank
[Islet] a fortnight before the “Frolic”... "15 Mac-
Kenzie took the three remaining Lascars on board,
dug up a chest of silver plate and other valuables
which Chilcott had buried, recovered the mails,
and, as was his habit, planted several coconuts so
that in future, unsuspecting mariners could see the
low-lying islet, before sailing on to Port Essington
where (it is presumed) he lodged a claim for
salvage.'® In fact, MacKenzie was an excellent
navigator and must have derived a great deal of
satisfaction from having shown the bumbling
bureaucrats in Sydney, and particularly
Lieutenant Yule, who had insulted his sen-
sitivities, a thing or two. He now had a double- axe
to grind. Enroute to Port Essington with the men
and goods he had rescued, Heroine struck a rock
in the middle of Endeavour Strait where Yule had
surveyed previously and declared free of dangers.
Now who was the incompetent navigator?

Of course, the Frolic found no one or thing to

rescue when they finally arrived at Chilcott Islet.
Captain Pilfold’s log book explains what took
place:

‘Saturday, 6th September, 1845. - At9:30 A.M.,
sighted the wreck of the Coringa Packet, on Chil-
cott Island, a sand key, bearing S.W.., distant three
leagues. At 10:15 landed on the island (north end),
and saw the place where a tent had been erected,
but no trace of the shipwrecked crew. I then sear-
ched for the mails and property said to have been
buried about ten paces from the tent (no parts of
which were left), but I found a large hole, about
that distance, in which I supposed them to have
been buried; in this hole many pieces of tarpauline
were strewed about, which bore the appearance of
having been lately dug up, as did the hole itself. I,
Mr. G. Levien [the mate], and three of my crew,
then searched the island throughout, to see if we
could find any letter, mark, or notice, by or
through what means these men left the island; and
after a fruitless search of near three hours, gave up
all hope of gaining any clue of their destination,
and returned on board. The Coringa Packet is now
lying with her head to the eastward, and by all the
appearances, in which I am borne out by the
opinion of my crew, the copper has been stripped
off on the starboard side, and the whole of the
rigging, masts, and everything moveable, have
been taken or carried away. There is a derrick

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 39

rigged out of the main hatchway, and the hatches
all off. On the island everything valuable has been
taken away; no part of the raft, spars, or casks, of
any description are remaining - saving a piece of
one of the lower masts [probably the one which
was cut away shortly after the wreck] and an old
scuttle butt, a few rags of canvas, and clothing of
the crew.’

Checking at the Raine Island beacon for the
missing survivors, Pilfold found a letter which had
been left by MacKenzie with the confusing news
that Heroine had rescued the mails and goods but
only three Lascars. Frolic then proceeded ta
Booby Island where he found Captain Pain’s ac-
count of the rescue of all that remained of the
twenty-one on the raft and letters explaining the
others’ departure for Port Essington. He then
sailed on to Port Essington where he found two
Lascars (one had joined the crew of Herome),
Chilcott’s valuables, and the mail which he sub-
sequently took on to Singapore in accordance with
his charter with the New South Wales Govern-
ment.’

Goyemor Gipps advised the Secretary of State,
W_E. Gladstone, in a dispatch dated June 16, 1846,
that he chartered the Frolic for £294/19/11. He had
every reason to believe, he wrote, that the East
India Company directors would gratefully take the
charge upon themselves, and he sent Gladstone the
charter documents and the details of the Frolic’s
services: a letter from the Resident Councillor at
Singapore reporting the Frolic's arrival; a certifi-
cate that the men and the Frolic’s owners had done
their work satisfactorily; and a declaration by the
treasurer and auditor of New South Wales that the
charter cost the N.S.W. Government £249/19/11.

The East India Company responded that it held
itself accountable for £240 for the hire of the
Frolic and considered that the balance, £54/19/11,
was the value of provisions shipped on board the
Frolic and intended for the Lascars at the wreck.
But since the provisions were not applied for this
purpose, but were landed at Port Essington and
retained there, it was clear that the East India
Company was not accountable for them. Earl
Gray, who had become Secretary of State, thought
that the outstanding £54/19/11 imi ght be con-
sidered chargeable to Port Essington. ”

SUMMARY DETAILS OF
CORINGA PACKET

Barque
Teak built [probably India, possibly Calcutta,
c.1840-41]

Tonnage - 230
Formerly engaged in Opium Trade
Copper sheathed, apparently iron-fastened,
Captain - F. Biggar Chilcott
Sydney Agents - Lyall, Scott & Company
Insurers - Apparently, H.E.I, Co,
Departed Sydney - 26 April 1845 [after a depar-
ture on 14 April aborted due to adverse winds]
Destination - Ceylon and Madras via Tortes Strait
Wrecked - 7 May 1845, Chilcott Islet, Coringa
Islets, Coral Sea
Cargo consigned by J.[?] B, Chilcott [was this a
muss-print, or did Captain Chilcott have a rela-
tive as a partner?)
S00 bags flour
30 logs cedar
20 kegs beef
1 case bird skins
3 horses
9 rams
30 sheep
25 tons coal
Cargo consigned by John Lord
20) bags flour
Cargo consigned by J.C. Lyall
12 cases cigars
Mails consisting of 249 letters for London; 400
Sydney. and 118 foreign letters, and 415
newspapers for Calcutta; 124 letters, and 152
papers for Madras; 66 letters and 102 papers lor
Bombay; 26 letters and 35 papers for Ceylon; |
letter for Mauritius: and 1 letter for the Cape o!
Good Hope
Passengers
Lieutenant Blackhall
Ensign Bloomfield
Mr E. Lord [drowned when the gig over-
turned]
Me Heary Van Deerlin [some confusion
here, Francis Stewart, Chief Officer of
Aydrabad, reported that Chilcott, two pas-
sengers, and a Miss Beech arrived al
Booby Island along with eleyen of the
ship's hands”’].

GUANO MINING ON CHILCOTT ISLET

I did not have the opportunity to examine the
entire islet during our December 1991 visit. How-
ever, Dr Charley reported having seen two
prepared blocks of guano on top of the islet near
the middle of the southeastern side. The ANPWS
Coringa-Herald National Nature Reserve: Plan
of Management (p.8) briefly states, ‘Relics of
guano mining during the 1860°s remain on Chil-

40) MEMOIRS OF THE QUEENSLAND MUSEUM

cot Tslet’, but does hot say what the “relics” might
be. My research indicates that, officially, guano
inining did not commence on Chilcott Island uotil
1877 and that it probably did not continue for yery
long,

A Mr Gilbert Carver Roberts claimed in a letter
to the Earl of Derby (then British Secretary of
State for Foreign Affairs), dated 9 March, 1878,
that he had been the first discoverer of guano on
Chilcott Island during a voyage sometime sub-
sequent to 1871.7!

The only official license to mine guano at Chil-
cott was issued to F.E. Beaver of Melbourne by a
letter dated 5 February 1877. The license was for
a period of seven years and cost Beaver £100. He
had initially applied to mine the Willis Islets to the
north, This was obviously a speculative applica-
tion as he shortly afterwards wrote that he had
been misinformed as to the potential of the Willis
Islets and he wished to change his application to
include Chilcott and the Herald Cays.** Obvious-
ly, he had visited the Willis Islets after his applica-
tion, and prior to its approval, and had
subsequently discovered a potentially richer har-
vest at Chilcott and the Heralds.

A story, which appeared in the Cooktown
Courier of 28 February, 1877 entitled ‘The
Douglas Tragedy” and tn the Cooktown Herald of
the same date entitled ‘Most Homble Murders at
Chilcot Island’, gives us a graphic description of
an incredible occurrence at Chilcott Isict. (Later,
the Courier story appeared in the Bundaberg Star
and Burnett River Times of 23 March, 1877 from
which the following is quoted).

It is interesting to note here that Cooktown was
then the centre of government administration in
north Queensland due to its proximity to the Pal-
mer goldficlds. Cairns had no hospital nor
newspaper and its most senior policeman was
Sergeant Armstrong who submitted a report to his
supenor, Inspector Clohesy, al Cooktown. The
Cooktown Courier journalist writes:

‘The Blackbird [steamer], which arrived here on
Saturday from Caims, landed three wounded men,
who were sent to the [Cook District] Hospital, and
who appeared to. be hacked and cal in a most
terrible manner. The story they tell is one which
can hardly be equaled by the most imaginative
writer of fiction,”

“Tt appears that the Douglas, schooner, belong-
ing to Messrs, F.E. Beaver and Co., of Melbourne,
landed a general cargo in Cairns, and then
proceeded on the 18th January to Chilot [sic]
fsland, lying about 200 miles due east, and worked
for guano by that firm. (Beaver had jumped the

gun. He was not informed by Lord Carnaryon’s
Private Secretary of his successful application for
a license to mine guano on Chilcott and Herald
Cays until he received a letter dated 5 February
1877°°).

‘In order to obtain a favourable wind the
Douglas went south as far as Flinders Passage,
anchoring on the way under Drink [Dunk] Island,
near Cardwell. While there a canoe with three
blacks came off and were invited on board. They
were supplied with tobacco, pipes, &c, and given
to understand that they were wanted to get some
companions, and to go with the vessel on her
voyage. They apparently agreed, went back and
brought another man. The new comer had such a
disagreeable appearance that the captain refused
to take him, and left with the three first. On arriv-
ing at Chilot [7 February], asmall detached island,
about a mile and a-half long by one broad, they
found the Alexandra, brig. belonging to the same
firm, half toaded, Mr, Beaver, one of the firm, who
was on board the Douglas, not having enough
colored labor to screen the guano fast enough,
determined to go back by the Alexandra, half
loaded as she was, and to fill up with cedar at the
Daintree. The brig accordingly lefton the moming
of the 10th, and - the fact is worth noticing - she
took nearly all the firearms of the Douglas, and
the blacks were present when the transfer was
effected,’

‘That evening [actually the l4th] night fell with
an over-cloudy sky, botsterous wind, and every
appearance of rain. Two white men, Cochrane and
McIntosh, were on shore in a hut which had been
erected, and which was stored with water and
rations; two of the blacks were also with then. In
the early part of the night, while the white men
were sitting in ther hut, the blacks seemed to have
killed them by simultaneous and treacherous
strokes, the attitude of the bodies indicating that
they could hardly have been aware of the death-
stroke. Then, armed with the half axes which had
served for the murder, the two blacks swam
through the tempestuous sea off to the vessel,”

‘No one on the schooner suspected evil, and all
the whites seem to have been asleep; probably the
black was waiting for his comrades, Softly they
stole about their murderous work. Two white men
were usleep on the deck, and both were struck so
that they made no sound. One (Shaw) says that he
knew nothing till he came to his senses two hours
afterwards, waking in a sort of dream finding
himself one mass of clotted blood, and ch
all over the head and arm. What saved him was he
had wrapped a rug and thick flour bag over his

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 4i

shoulders as he lay down, and the bag was dinted
with the blows of the blunt axes. Thanks Lo the
darkness of the night, he managed to crawl into
the forecastle, although a black spying him as he
went, aimed a blow at him, which missed, Think-
ing he was too far wounded to be worth troubling
about, the murderers left him, and he managed to
craw] aft through the hold and get into the cabin”
‘But I must go back. After the blacks had left the
two men, Troy and Shaw, for dead on the deck,
they went down into the hold where another sailor,
Purcell, was sleeping, and attacked him. He was
fearfully chopped on the face. head, and arm, one
finger was cut off, and a huge gaping gash made
in his back. Him they left for dead, but he sub-
sequently crawled through the hold aft into the
cabin, Meanwhile Deasy, the acting second mate,
who was asleep in the forecastle, heard Purcell cry
outand rushed out on deck. Never suspecting what
had happened, he thought that the mooring chain
had parted and shouted to that effect. Ina moment
he saw a blackfellow by his side with an uplifted
axe over his head. He dodged the black and sung
out “Captain, the blacks are murdering us.” Then
all rushed on him. How he escaped is a miracle.
He has numerous slight flesh wounds and a severe.
chop on the arm, but only the most wonderful
agility and pluck saved him, Once the murderers
had him down on his back on the deck, and two
paused to let the third get a chop at him. Even this
he managed to dodge by shifting his leg, escaping
with a flesh wound on the inside of the thigh.
While this was going on, the mate awakened by
the noise, rushed past and got into the forerigging
where another man had escaped. Deasy struggled
out of the grasp of the murderers ind ran for the
forecastle, one black following him. Getting his
knife owt, which up to that moment he had not been
able to draw, he struck at his assailant but missed
the stroke, and stnking on the axe, lost his knife.
Then, picking up a small grindstone lying there,
he struck the black and staggered him, thus
managing to get into the forecastle. A hurried
search showed him there was no weapon to be
found, and he came out again to make a rush for
the rigging. In his haste and in the darkness, he
rushed for the post side, where one of the blacks
was part of the way up and another on the bulwark
preparing to ascend, with the intention of atlack-
ing the mate and Lawrence on the foreyard. Deasy
sprung past the black on the bulwark, and grappled
the one on the ngging; but before he could wrest
the axe out of his hand, the second black wounded
Deasy on the heel, Finding that the next moment
he wauld be killed, he scrambled up and reached

the foreyard, when he cut blocks with the mate’s
knife, and the men used them as weapons to keep
back the blacks, who, after a while, made nu
attempt to ascend,”

“During this struggle, which occupied less time
than I take in writing about it, the Captain,
awakened by the noise, came up, and as he laid his
hand on the top of the companion, it was chopped
by a blow from an axe. He retreated into the cabin,
where he remained with his son, and was sub-
sequently found by the two sorely-wounded men,
Shaw and Purcell. They vainly endeavored to load
a pistol, striking matches, but not daring to light a
lamp. But the flowing blood clogged the pista] and
damped the powder [poetic license by the Bun-
daberg Star editor, it was a revolver which fired
cartridges), and they could do nothing. The
steward had shut himself up in the galley: three
men were on the foreyard, Deasy almost fainting,
and lashed to prevent falling; and poor Troy lay
on the deck near the galley.’

‘There was a sort of lull, The men on the
foreyard thought that all hands, except themsel-
ves, the captain, and his boy, were dead; and the
blacks compelled to pause in their active attack,
began to look for the bodies of their victims. Shaw,
by this time, had crawled away - and, on searching
the hold, they found Purcell also gone; there
remained only Troy lying motionless near the
galley, How long he had recovered his senses no
one could tell, bat he was not dead, The murderers
came to where he lay, and with one blow of an axe
chopped off his foot. The steward, trembling in his
galley, heard the poor fellow groan, ‘Oh God, I'm
punished now.’ Then they chopped his hody and
clove his head, till all Jife - all semblance even of
humanity - was battered ovt of him.'

‘So the terrible hours of the night wore on, and
day began to dawn; the three blood-stained
demons holding the deck - the steward hidden in
his galley - the three men on the yard - the captain
and his boy in the cabin, with the wo poor
wounded men weltering in their blood beside him.
The grey light of morning made objects visible,
and the blacks thought to finish their work. Pick-
ing up stones and picces of coal from the hold, they
began to pelt the men on the yard, who dodged the
missiles as best they could. These two blacks
ascended the rigging with their axes, while the
third remained on deck pelting the whites. These,
compelled to disregard the stones, confined them-
selves to keeping the axes at bay with their sling-
blocks, Then the blacks found the steward was in
the galley: one went to guard the companion while
the other burst open the galley door. The steward

42 MEMOIRS OF THE QUEENSLAND MUSEUM

jumped through the other door, rushed to the com-
panion, dodged the blow aimed at him by the
guard, and tumbled below.’

‘Now there was hope for the whites, Daylight
was brightening and an un-wounded man had
reached the cabin where there was 4 revolver and
ammunition, But deliverance was not for some
time. For nearly an four the men on the foreyard
had to keep at bay two of the blacks who were
assailing them, while the third kept guard over the
companion, cunningly shielding himself from the
loaded revolver of the steward. At last an in-
cautious movement of the guard exposed his head,
and the next second a bullet went into his brain.
The two blacks exchanged a hurried sentence in
their own language, and one went to pick up his
fallen comrade. The sailors in the foreyard dropp-
ed down the ngging, The mate first on deck took
up a handspike and staggered the third man with
a blow on the head, and the others closed around
him. The one who had gone to the dead guard left
him, saw the game was up, and jumped overboard.
Two of the blacks were now dead, and the steward
emptied his revolver at the third one while he
swam, but did not succeed in hitting him. They
saw him land on a rock, and put off in a boat after
him, but he jumped again into the sea, and al-
though they waited for him he was never seen
again.

eThen the survivors wentto the Island and found
the bodies of their comrades in the hut, and made
sail for Cairns with the wounded, Arrived there
[18 February], an inquiry was held, and the three
worst wounded were sent to our hospital, where,
under the skilful treatment of Dr Ahearne, a sur-
geon of no mean ability, they are progressing as
well as could be expected.’

With minor vanations, the three newspaper
reports agree in the principal details.

How long after this Beaver continued his opera-
tions at Chilcott Islet is not known. The editor of
the Cooktown Herald, with libellous gusto, first
attacked the editor of the Courier saying that the

‘niggers from Dunk Island’ were his “friends' and
then accused Captain Harris of kidnapping the
three natives and cailed fora Government inquiry.
The whole sequence of unpleasant events. may
have prompted Beaver to forget guano mining
altogether.

Prom what we can leam from the Douglas.
episode, at least one hut was constructed on the
islet. It was furnished with a water tank and sup-
plies, “...and arrangements made for the regular
working of the deposits." Three men, Patrick Troy,
Humphrey Coghalin, and Alexander McIntosh,

were buried there. Therefore, some evidence of
the event may remain on the islet.

CHILCOTT’S LOCATION OF THE
CORINGA PACKET WRECK

Captain Chilcott was quite specific as to the
location of the islet and the position of the wreck.
He reported, ‘The position of the danger on which
the Coringa Packet was wrecked is as follows: -
Lat. 16°52'30" S., long. by the means of 20 lunar
observations 149°55'45" E., the chronometer
showing a longitude of 149°55° E.., variation of the
compass 9 degrees E.; rise of tide at springs 5 feet;
the island about 10 feet above the level of the sea,
surrounded by a reef dry in some places at low
water, the wreck of the vessel bearing N.E. by half
a mile; and the whole of this danger does not
exceed 4 1/2 miles in circumference, Another
small island we observed to the S_W_ about four
miles [South West Islet].° He also stated that the
current Was ‘running constantly to the north-
westward.**4

MODERN DESCRIPTION OF CHILCOTT
[SLET

Australian Chart AUS 6/2 (corrected to 1991)
gives a position of 16°56'25"S, 149°59°5S"E
based upon an “Observation Spot’ at the Eastern
end of the islet Magnetic Variation 8°15'E (1984)
increasing about 2° annually.

The islet is described as ‘An elongated pear
shaped cay with long axis onented NE-SW. 800m
long and 255m wide, Beachrock, piled up at a
moderate to steep slo » fringes | the S. shore line.
Maximum elevation 5m ASL.">

FACTORS AFFECTING SHIPWRECKS IN
THE CORAL SEA

In general, isolated islands, reefs and cays
abstract the flow of water resulting in an ac-
celerated current around the reefs and cays. Some-
times an eddy forms on the lee side. (This refers
to deflection of the main, deep northwesterly cur-
rent prevalent throughout the Coral Sea. The
northwesterly trend of surface current, which is
less affected, is still evident in the reef-top shal-
lows and, combined with the predominant winds
and seas from the southeast, generally dictate the
distribution of wreck maternal deposited there).
Over the Coral Sea Plateau, the tidal range indi-
cated is 2m_

On average. two or three cyclones per annum

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 43

affect the Western Coral Sea, usually between THE SITES
December and April. Although infrequent, tropi- ; -
cal cyclones are the greatest hazard in the region. Two separate sites were located and it is con-

Gales occur infrequently throughout the year. sidered highly probable they are both associated
. ; with the same wreck.

Tropical depressions occur usually two or three

times per year. These create conditions similarto SITE ONE (Chil.01)

cyclones but are less violent.2’ Based upon Chilcott’s description of his course,

/
Back réef slope
with numerous

patch reefs gt Front reef slope

i
|

]
/ Deep reef flat
/ Shoal
/ /
/ I
! \ Reef /
AA 7 crest |
\ \ / /
\ \ / /
“Dee reef flat Reais (
ad >~ Chilcott Islet
/
\\ \ “Beach rock”
+ en aS 2 ms
The ) /
Patch reefs som
0 1 2 Kilometres

Se Bathymetric contour

FIG. 6. Chilcott Islet. Approximate positions of sites are numbered,

+4 MEMOIRS OF THE QUEENSLAND MUSEUM

(NW by W after passing through the Diamond
Islets), and the position he gives for the wreck, ("/:
mile NE of the Islet), we had a good indication of
where on the reef to begin our search. Our ex-
perience with other wrecks in similar situations
led us to confidently expect wreck material on the
reef flat scattered in a northwesterly direction
from the point of initial impact.

After first exploring the beachrock at the base
of the beach on the northeast point of the Islet -
where miscellaneous iron remnants, u small piece
of coal, and broken bottle glass were found - a
“bottle tree’ was constructed on the centre of the
beach facing NE as a visual reference point.

Proceeding in the zodiac onto the deep reef flat,
a reciprocal back-sight was taken on the “tree”
using 4 monocular compass. At 225° magnetic,
and approximately 30m inside the breakers on the
reef crest, a reference buoy wis dropped in 2m of
water. Immediately, the wreck's anchor chain was
seen by a diver some 10m southwest of the refer-
ence buoy.

A second flag buoy was attached to the centre
of the chain and, bearing in mind the predominant
winds and seas from the SE and the aurthwesterly
direction of the current, the first buoy was trans-
ferred to the northwestern extreme of the chain to
give the surface swimmers a Visual reference line
of potential material distribution.

The zodiac was anchored over the centre of the
chain and Kanimbia contacted by hand-held VHF
radio. The vessel recorded their own GPS position
and by our holding our boat's metal fuel tank
overhead as a radar beacon, they were able to
record a radar range and beanng on us and the NE
end of the Islet as well as its NW and SE sides.
From the zodiac, magnetic compass bearings of
218° and 236° were recorded to the centre of the
beach slope on the SW and NE sides of the Islct,
respectively. With this information, a reasonably
accurate GPS latitude and Jongitude of the site was
calculated as 16° 55,650" S. by 150° 00,712' E

The SE nun of the chain was traced as far as the
breakers would allow the divers to proceed. A pile
of chain was found near that end (which then
trended SSE for 7.5m before ending). The anchor
deployed by Chilcott was not sighted. It should be
found just off the outer edge of the reef crest
amidst the breaker build-up.

The line of the chain was recorded with tapes
and underwater compass. The greater length of
chain lies on an alignment of 300" magnetic or 15°
west of due northwest.

It being joo dangerous to work on the site at high
tide due to the swell and wave chop in the shallow

water on the reef flat, and the timing of the lows,
being at approximately 0700 and 0800 for the two
mornings we had at the Islet (Kanimbla was to sail
at 1000 the second morming), it was decided not
to altempt a regimented area survey due to lack of
time and trained personnel. Therefore, our swim
search must be considered to be only a cursory
examination of the area.

The objectives were to gain an oversight of
material quantity, type, and distributional trend_
Also if was intended to recover a small number of
items which might assist in confirming the site as
that of Coringa Packer and to assist in the inter-
pretation of the material distribution sequence.

ftems located included two rudder pintles (one
won, One bronze from the top of the midder), a
boom nng and goose-neck (possibly from one of
the ship's gaff booms), a sandstock brick (a bed of
brick was often used beneath ship's galley stoves
as Insulation for the wooden deck below. Galley
stoves were niust often placed in the forecastle
area of the ship), two large rectangular dressed
stones (not recovered, one measured
250% 300 * 550mm having one end cut at an
angle), two ballast stones, an iron ngging block
strap with hook, a few iron fastening pins (two
recovered), broken bottle glass (fram base of
green case bottle), an abraded piece of timber, a
bob-stay plate (mounted on the stem for the attach-
ment of the lower bowsprit brace or bob-stay” *) a
large iron thimble (used te form an eye in a bight
of rope), and several unidentified iron objects (not
recovered), Six short sections of studded anchor
chain averaging 300mm, being loose on the reef,
were also recovered for study,

The survey of Site One was curtailed by the
necessity to examine and briefly record a second
site discovered on the Islet itself,

SITE TWO (Chil.02)

The southeast side of the Islet (normally the
weather-side) is almost entirely built of broken
slabs of beachrock thrown up by cyclones. The
sock extends from water level to the crest of the
Isletat a height of approximately 6-8m above low
water level.

Mid-way along this side, and at the top of the
beachrock, storms have deposited several large
ship's timbers.

The timbers are very weathered and the type of
timber is not obvious from its surface appearance
(although one piece, a bowsprit, has the ap-
pearance of conifer), Reference to scantling tables
of the period should give us an idea of the size of
ship from which they came. A sample was sub-

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 45

\

Prayaeling
Wind & curtent

% |ron Strap
W/Thru -
(60am tong)

2 Large fron Obxoct
x Stone Bloch (Measured)
* Brick (R), iron fasteners, etc,

% Bottle Glass (A)

% Bobstay Plate (A)

Booy 2.

mm

Ballast stories (R)
A Boom ring &
% goasaneck (Fy

Timber (FR)
Storie block

“SRG. to N. Beach
e 236" Mag.

Chilcott Islet

Site (Chil. 01)

11-12 December 1997
R, Coleman,

% Pinties (R)
x

(Dist. to let approx, 10m)

/ ARG, 10S. Beso

ra 2)a" Mag

FIG. 7. Plan of Site One.

sequently submitted to the Queensland Forestry
Department for positive identification of the tim-
ber species to determine where the vessel was
built.

All of the material examined indicated that these
were the remains of the upper bow section of a
reasonable-sized vessel,

There were a number of cant futtocks and top
timbers, a bowsprit, fragments of unidentified
worked timbers, and a few planks, all of which had
been iron-fastened.

Two iron knees (one broken) were located half-
way down the beachrock and these also appear to
be from the bow section as the upper arms are at
an angle, or ‘cant’ to the flat of the lower arms.
The knees were also iron-fastened.

At the water's edge, below the knees, were
found two large pieces of tumbled cast iron, pos-
sibly parts of hawse pipes (through which the
anchor chain passes through the ship’s hull at the
bow), or chain pipes (through which the anchor
chain passes through the deck into the chain locker
below). Other unidentified pieces of iron were
observed lodged in the lower rocks.

Time did not permit precise recording. How-

ever, general measurements were taken of a few
timbers and everything was photographed in situ
using a photographic scale.

One top timber was recovered so that it could be
accurately recorded for comparison with scantling
tables and as a sample for timber identification.

DISCUSSION

It seems reasonably safe to assume that Site One
is part of the Coringa Packet, since it was located
where Chilcott said it should be and no other site
was located in the general area.

Initial examination of the material from the site
indicates that it is consistent with what we should
expect from a vessel of Coringa Packet’s vintage
and size. The recovered piece of timber proved to
be Tectona grandis (teak) which is further confir-
mation that the_site is that of the ‘teak built’
Coringa Packet.”

We have no record of another wreck in the
Coringa Islets nor in the Tregrosse Reef/Diamond
Islet complex to the southeast from which material
might have floated. This lack of recorded wrecks
is not to be taken as conclusive. Several ships of

46 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 9. Site Two. Bowsprit. Length 9.4m (30feet 10inches), base dimension 38cm (1 foot 3inches) square.

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 47

22 - aaa

G. 10. Site Two. fron knees.

FI

FIG. 11, Site Two. Iron objects found at water level.

MEMOIRS OF THE QUEENSLAND MUSEUM

48

"OY, POILOIPUT oe s}[oq BuTUajse} Wo enpisal UOT “(soyour ¢°Z]) UNUSTE
yutod-prur je paprnout ‘(sayour [ 1) WUIQgZ papls ‘(sayoutp JoafQ]) WIC] *¢ Yue] ‘Jaquiy do} paraAodal JO UOTIDES PUL SMOTA INO “OMY, MWS “ZI “OW

WW)

V-V NOLL
O1LOaS

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 49

the period were listed as ‘missing’ in the Coral Sei
with no location given. However, although anum-
ber of India-built vessels were engaged in trade
with Australia, it would be an extremely unusual
coincidence to find two teak-built vessels wrecked
in the same location within a few hundred metres
of each other.

On first examination of the location of the
material at Site Two, itis difficult, bul not impos-
sible, to reconcile as being part of the bow section
of the Site One wreck.

Tt would be an unusual storm to carry material
from Site One to the southwest and south of Lhe
islet before it had been carried tu the north beyond
the Islet by winds, seas, and currents that would
have generally prevailed for some months after the
wreck in May. However, itshould be recalled iron,
elass, and coal were found on the NE tip and
slightly around the north and south sides of the tip
of the Islet. in a santhwesterly direction from the
reefiop site.

A very shallow, smooth, beachrock shelf abuts
the tip of the islet extending for a radius to the east
and south of some 350- 100m-. In normal weather
conditions the seas would be rough there. ltis very
unlikely that material salvaged by Chilcott or
MaeKenzie or building materials of the guano
miners would have been brought ashore in ex-
posed, rough shallows at that point. Rather, they
would have landed just around the tip on the
sheltered northern beach and apposite to the nor-
mally protected anchorage. Therefore, iL1s conjec-
tured that the material found on the tip of the Istet
is indeed flotsam and, at least the iron items, being
of little value to any of the groups on the Islet, have
heen coincidentally transported there while at-
tached to pieces of floating wreckage during a
storm from the northeast.

The two rudder pintles found in the water some
200m southwest of the anchor chain are nearly
identical in size having an overall length of 53cm
(21 inches) and an intemal breadth of c. 2icm (8+
inches). The iron pintle was the higher of the two
on the rudder, iron being used above the waterline.
‘The bronze pintle, being of similar size, was the
next below and just at or below the waterline. The
pin of the iron example was c. 5em longer than that
of the bronze to facilitate alignment when install-
ing the rudder. While the overall width of the two
pintles appears slightly small for a 230 ton vessel,
they tell us something of the shape and style of
rudder used and consequently provides some infor-
mation concerning the shape and construction of
the stem of the vessel. The most informative pintle
dimension is that of the pin chameter (roughly

measured wt ¢. 50min) which was the more critical
in relation to the size of vessel. Steel” gives a
recommended pin diameter of I’ inches
(47.6mm) for vessels between 201 and 330 tons
which would include Ceringa Packer at 230 tons.

By reference to the location of the pintles on the
Site One plan (Fig.7), one will see that at least the
top part of the wreck’ s rudder, with the two upper-
most pintles still affixed, became detached from
the ship and floated some 200m from the site, to
the southwest toward the Islet, before breaking up
and a portion sinking from the weight of the metal.
Generally, this would indicate that the ship was
suffering a major structural break-up, The sub-
stantial timher of the rudder tiller was socketed
into the upper end of the nidder post, ILthen passed
through the stern of the ship between the large
timbers of the transom framing and swung in an
are directly below the main deck beams. For the
pintle pins to become disengaged from their
mating gudgeons, the horizontal tiller either had
to break the transom timbers, virtually destroying
the stern of the ship, or snap off itself allowing the
rudder to move vertically a minimum distance of
25¢m (the length of the pin of the iron, or upper-
most pintle), This, combined with the presence of
ballast stones and the dressed stone blocks in the
immediate vicinity of the stretched anchor chain
indicates that the ship Was coming apart Where it
was held by the anchor and chain. Therefore, the
odds in favour of the upper bow section, which
due to the absence of heavy objects was reasonab-
ly buoyant, floating to the southwest and eventual-
ly being thrown up on the rocks before it could
float to the northwest beyond the islet, are greatly
improved.

Again referring to the Site One plan. one will
observe that the lie of the majority of chain is
SE-NW as is the axis of the majority of loose
eer. Chilcott states that Ww, ‘hen the ship struck,

.slewed broadside on....’*’ There can be little
soubt that her head had come about from her
sailing course of northwest by west to the north-
east and she was then port side to the reef, the edge
of which runs to the northeast at that port, Chil-
cott then cut away the mainmast to lighten ship
and to reduce windage and the pendulum effect
and dropped an anchor. He says the sea was
‘making a breach over all.” Swinging to the
anchor at the dictates of the southeast wind und
sea and the current from the same direction, and
grounding in the shallows of the reef, the ship, for
a time rested (according to Pilfold’s observations
four months after the wreck) with her head to the
east. Pilfold's reference to the copper being

0 MEMOIRS OF THE QUEENSLAND MUSEUM

removed from her starboard side indicates that she
was then heeled onto her port bilges making it
difficult to remove copper from that side.**

A possible explanation of subsequent events is
that, in time. spring tides and associated swells
may have lifted her sufficiently for the ship to
swing her head to the southeast and her stern to
the northwest thus aligning the anchor chain in
that direction and allowing time for a concentra-
tion of heavy matenal to be deposited to the
northwest of the chain. The chain may have parted
(which would explain the jumbled pile of chain at
the SE end) and the resultant tremendous force
upon the forward-mounted windlass may have
contributed to the upper section of bow, or at least
pieces of it, being wrenched from the weakened
hull. As the remainder of the hull was eventually
carried to the northwest, the last of the chain
dribbled from the below decks chain locker.

The top timber recovered from Site Tyo has
heen identified as Tectona grandis (teak)*

As it seems that the timbers at Site Two do date
from 1845, it is remarkable that the guano miners
in the 1870°s. who were renown wreck
scavengers, had not used them for firewood of to
construct their hut and facilities. Perhaps they had.
aun abundant supply of smaller firewood and
lighter plank that required less effort to gather. Or,
did Beaver & Co, abandon guano mining on the
island after the Douglas massacre thus occupying
the Islet only too briefly to cause much distur-
bance?

No copper alloy fastenings were observed on
cither site, which Is an unusual coincidence in the
case of nineteenth century wrecks; nor for that
fatter, were any metal sheathing or tacks, which
are generally encountered.

The country vessel, Valetta {1825}, excavated
by the author in 1983, was iron fastened but copper
sheathed aver timber sheathing. Faith in the resis-
tance of teak to the ravages of Teredo navalis, plus
the faot that almost all metalwork used in Indian
shipbuilding was imported from England or
Europe** (and thus expensive), would seem to
have justified the economies of less expensive iron
fasteners and the cheaper expediency of using teak
sheathing to insulate the iron fasteners from the
electrolytic effect of the copper sheet. Phipps
made a number of references to India-built vessels
being sheathed with copper shect over teak
sheathing boards and implied that copper fasten-
ing of country vessels was an exceptional and
expensive luxury.

Also, the suggestion ts, that ‘country’ skippers
and owners were quite happy with teak sheathing

alone. But, in order to procure cargoes from mer-
chants accustomed to the security of their cargo
being carried in the generally dry, copper sheathed
European-built vessels, they had to make the con-
cession. Captain Pilfold of the Frolic poinredly
stated that the copper sheathing from the
side of the Coringa Packet had been removed by
the time he arrived, (Obviously by MacKenzie
who scems to have salvaged everything pos-
sible**), Pilfold, by citing witnesses, Was ensuring
that he would not be accused of breaking his
contract by keeping salvaged material for his own
fit,
O The use of conifer or other straight, reasonably
clear softwood far masts and spars, including
bowsprits, was normal practice in all shipbuilding.
Indeed, pine spars from New Zealand and
Australia were a frequent export cargo of north:
bound ships of the period. However, Phipps”’
states that at least the lower masts and large spars,
including the bowsprit. of country vessels were
frequently made of teak and sometimes *poon'
from Malacca, the upper masts and lighter spars
being made from imported European, American,
or Chinese pine,

The identification of the timber of the ‘tap
timber’ (the uppermost section of a frame or rib),
being an intemal structural member. is certainly
more conclusive as to the ship's origin, Country
yessels” frames were -almost always made of either
Sissoo (Dalbergia sissoo Roxb.) or Saul (Skarea
robusta Roxb.) both of which were obtained from
the forests north of the Ganges.“’ However,
Phipps also mentions that, as those timbers be-
came less available in the late 1830's, more ves-
sels were built entirely of teak (Tectona grandis).

From the small amount of dimensional data
collected on the Site Two timbers and the size and
form of the bowspnil (including the unusual man-
ner in which it was stepped), it would seem that
the ship was designed to carry an enormous
amount of sail (the relatively large crew caried by
Coringa Packet muy lend support to this theory).
The rather fine stern indicated by the slightly
narrow breadth of the rudder pintles (which
reflects the width of the sternpost) also suggests
that the ship was built for speed. These were not
common features of the ordinary country vessel
where speed was sacrificed for cargo volume.
Rather, they were essential features of “Opium
Clippers’ whose cargo took up less space. Their
speed allowed them to make two or more annual
round trips to China to the ordinary country
vessel's one, Speed was also necessary to avoid
Chinese government war-junks wishing to stop

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 51

FIG. 13. The Calcutta-built Opium Clipper Waterwitch with Fi exceptionally long bowsprit, jib-boom, and tall,
raked masts allowing a tremendous amount of sail to be set.

the illegal trade or pirates attracted either by the
valuable opium on the outward voyage or the large
amount of specie carried on the return voyage. For
this reason, opium clippers were heavily armed.

A high percentage of opium clippers built in
India were constructed at Calcutta. Phipps”® lists
seven being built there between 1830 and 1839.
Lubbock claims that three were built at Bombay
and a half-dozen at Moulmein. Essentially they
were built to either of two designs. The first was
a copy of a very famous American privateer. The
second was specially designed for the trade by Sir
Robert Seppings, the Surveyor of the Royal Navy.
These were generally barque rigged. Of those built
at Moulmein, only one was to the Seppings design.
The others were copies of fast Mediterranean
‘fruit’ schooners.” If Phipps’ or Lubbock’s
figures are anywhere near accurate, it can be seen
that purpose- built opium clippers were a rare
breed indeed.

It is obvious that Site One does not represent the
primary site of a wreck. Rather, it is a loose trail
of scattered, isolated material. One would expect
a concentration of copper sheathing, ballast, coal,
and other heavy material such as spare anchors,
rudder gudgeons, etc. to be evident. Although it

seems that MacKenzie salvaged everything he
could from the wreck, it is predicted that much
more material will be found to the northwest in the
deeper water, (say, 10m), off the back of the deep
reef flat where it was carried by floating portions
of the ship’s hull. Some material, associated with
the bow section, may be found to the southwest
along the southern shore of the islet.

There are three possible scenarios:

1. Both Sites One and Two are part of the same
wreck.

2. Site Two is material from a wreck which
floated to Chilcott from elsewhere.

3. Another wreck has occurred on Chilcott reef
further to the south and opposite the Islet in an area
which was not investigated.

However, it would appear that the presence of
Tectona grandis at both sites is too consequential
to ignore and that the first scenario must be the
favoured option.

In the short term, further study of the material
and data already collected may resolve a number
of questions, but much more remains to be learned
from the sites. In the interim, it is important that
these preliminary findings be brought to notice
and I would strongly recommend that future

52 MEMOIRS GF THE QUEENSLAND MUSEUM

pnvule, lourtst and scientific visitors and the
Royal Australian Navy (frequent yisitors) be ad-
vised against indiscriminate disturbance of any
material remains in the Reserve unless they are
conducting authorised archaeological research.

From an archaeological standpoint, there is no
question that the Coringa Packet wreck and the
guano mining aspect of the islet demand further
investigation.

Whilst it is difficult to envisage the practical
long-term preservation of the timbers and other
material of Site Two in situ, it is critical to the
study of ‘country vessels’ that they be precisely
recorded and sampled. With this data, it may be
possible to reconstruct a portion of the bow section
using computer graphics, which will provide us
with hitherto unknown details of aneven rarer ship
style, the ‘Country-built Opium Clipper’, There-
fore, it is recommended that, at the first oppor-
tunity, the material at Site Two be and
then, as a matter of expediency and economics,
removed to a safe place where they can be studied
at leisure, Itis estimated thatan archaeologist with
4 team of five volunteers assisting could ac-
complish this in two days if working during the
northwest monsoonal period (say, late November-
December), Considerable care would have to be
exercised in the removal of the timbers because
many of them are extremely fragile and vital in-
formation may be lost if they are damaged in
transport.

In future, a controlled survey of the reef flat, and
the discovery and recording of the pnmary under-
water site, may contribute greatly to the meagre
information currently known.

Itis likely that a recommendation will be made
to the Federal Minister, DASET, that, after con-
sultation with ANPWS, a Protected Zone similar
ta the one at Porpoise Cay (Wreck Reef} be
declared under the Commonwealth Historic
Shipwrecks Act 1976. This would allow protective
and management measures, specific to the nature
of the resources, to be put into place.

Possible justifications for the declaration of a
Protected Zone are as follows:

1. Little is known about the design or construc-
tion techniques employed in ‘country vessels.’
This type of ship played an important role in early
Ausiralian maritime history. Moreover, if Corin-
ga Packet proves to have been built to one of the
rare ‘Opium Clipper’ designs (of which only afew
were built immediately prior to the introduction of
steam, iron hulls and thus prior to the consequent
ilecline of Indian shipbuilding) - and the chances

are excellent that she was - then the recording and
study of her remains are even more important,

2. Historical aspects of both the Coringa Packet
story and that of the guano miners are of sufficient
significance to warrant closer study. The material
remains of both episodes can contribute to the
interpretation and appreciation of these aspects.
Their contribution would be in a physical sense as
a part of the Reserve's resources, and in an
academic sense as part of Australian and East
Indian maritime history.

3. The remains are cultural heritage resources
located within an administered Reserve which
already has some management mechanisms in
place. These unique resources require additional
protective and management regimens specific to
their nature.

4. Site Two, which has considerable potential to
provide new information, is extremely vulnerable
to natural attrition and unwitting, casual destruc-
tion or disturbance by visitors to the Reserve.

ACKNOWLEDGEMENTS

I wish to express my gratitude to Mr Max Allen
of Max Allen Chasters, without whose generous
assistance the field work would not have been
possible. Funding for field work was provided
from the Commonwealth Historic Shipwrecks
Programme, DASET. I would also like to thank
Basry Muldowney for his assistance in the field.
Figures |, 2 and 6 courtesy ANPWS,

REFERENCES

(1) Lubbock, B. “The Opium Clippers’, Brown. Son &
Ferguson, Glasgow, 1933:193.

(2) Shipping Gazette & Sydney General Trade List, 15
March 1845, Hereafter=SG&SGTL.

(3) Phipps, J. ‘A Collection of Papers, Relative to Ship-
building jn India,,,", (Typescript), Scott and Co.,
Calcutta, 1840,

(4) SG&SGTL, 15 March 1845.

(5) SG&SGTL, 16 Angust 1845,

(6) The so-called ‘Post Office’ for which Booby [sland
became so famous was in fact nothing more than
4 box placed upon the top of the island near a
flagstaff. The Post Office was not moved inta the
“Post Office cave’ until] 1857, Coleman, R.A.,
"History of Boaby Island’ (in prep).

(7) Jukes, J.B. “Narrative of the Surveying Voyage of
H.M.S, Fly’, T.&W. Boone, London, 1847, vol, 1;
301-2,

{8) Allen, J. & Corris, P,, teds), ‘The Journal of John
Sweatman: a nincieenth century surveying voyage

CHILCOTT ISLET SHIPWRECK AND HISTORICAL SURVEY 53

in North Australia and Torres Strait’, Univ. of Qld
Press, St Lucia, 1977:60; SG&SGTL, 16 August
1845.

(9) Nautical Magazine, 1845, 14:720.

(10) SG&SGTL, 17 December 1845.

(11) Sydney Morning Herald, 11 & 17 December 1845,
Hereafter = SMH.

(12) Nautical Magazine, 1845, 14:719-20.

(13) Allen & Corris, op. cit.:56.

(14) Ibid.:57.

(15) Ibid.

(16) SMH, 1 April 1846.

(17) SG&SGTL, 17 December 1845.

(18) Ibid.; SMH, 1 April 1846.

(19) For this correspondence, see Historical Records of
Australia, series 1, 24:513, 757; 25:99, 329-31,
438; Legislative Council Votes and Proceedings,
2 September 1854; New South Wales Archives
51/11617, 45/8262.

(20) SG&SGTL, 16 August 1845.

(21) New South Wales Archives, AO 4/1620;548-54,

(22) Ibid.:456.

(23) Ibid.

(24) Nautical Magazine, 1845, 14:556-7; SG&SGTL,
16 August 1845.

(25) Australian National Parks and Wildlife Service,

“‘Coringa-Herald National Nature Reserve: Plan of
Management’, Government Printer, Canberra,
1988:43.

(26) Ibid.:11 citing Orme,1977.

(27) Ibid.:15. For material conservation considerations,
see Ibid.:14, ‘Salinity and Turbidity’.

(28) Underhill, H.A. ‘Masting & Rigging the Clipper
Ship & Ocean Carrier’, Brown, Son & Ferguson,
Glasgow, 1946:76.

(29) Queensland Forest Service report ref.820-67, 15
January 1992.

(30) Steel, D. ‘The Elements and Practice of Naval
Architecture’, (1977 reprint), Sim Comfort As-
sociates, London, 1805:Folio LIII.

(31) SG&SGTL, 16 August 1845.

(32) SG&SGTL, 17 December 1845.

(33) Queensland Forest Service report ref.820-67, 15
January 1992.

(34) Phipps, op. cit. 1:7-8, citing a letter from James Kyd
to Admiral Drury, 1808, and elsewhere.

(35) SG&SGTL, 17 December 1845.

(36) Phipps, op. cit. 2:14-15 and elsewhere.

(37) Ibid.:45-7.

(38) Ibid.,1:113-5.

(38) Lubbock, op. cit.:17.

(40) Ibid:facing 96.

Sa MEMOIRS OF THE QUEENSLAND MUSEUM

ANOMALOPUS PLUTO INGRAM, A POORLY
KNOWN SKINK FROM CAPE YORK PENINSULA.
Memoirs of the Quéensland Museum 32(]): 54:- The taxon
Anomalopus pluto Ingram, 1977, a limbless, fossorial skink
was based on a specimen collected at McDonnell crossing,
Cockatoo Ck, 115i 5 of Bamaga, Cape York Peninsula
(1133'S, 142°26°E) on 14 July, 1975. Since that time, the
species has been encountered infrequently.

In 1980, an Australian Museum (AM) party collected 6
specimens on 16-17 July (AM R94360, McDonnell Ck Cross-
ing, 1 km E of main N/S Peninsula road, 11°33'S, 142"27"E;
AMR94361-2, R94484-5, R1 12004, 15km E of Heathlands
settlement on Captain Billy Ck road, 11°45'S, 142°41"E).
Hleven years later, 4 specimens of A. pluto were collected by
officers from the Queensland National Parks and Wildlife
Service (QNPWS).One specimen was collected on 6 Seplem-
her on the road edge during road reconstruction, c.127km E of
Heathlands ranger base (11"40°06"S, 142°45°30"E). Two days
later, 8 to 10 specimens were uncovered ina 30m? area at Eliot
Ck (11'24'40"S, 142"24'43"B) during earth moving activities.
Of these, only the three Jeast damaged specimens were col-
lected. In December 1991, QNPWS (Far Northern Region)
epproached the Queensland Museum (QM) to assess the Eliot
Cx A. plute sile, an area of proposed redevelopment. | visited
Eliot Ck on 18 January 1992 but found no A. pluto. However,
four specimens were recorded from 3 other localities: SO(im
W of Heathlands ranger base (11°45'S, 142° 35'E),
QMJ54050 (17 Jan,), 54082 (19 Jan.), Lem B of Heathlands
ranger base (11° 45'S, 142°36'E), QMJ54083, 54213 (19
Jan.); Captain Billy Cx road, 22.25 km [rom Heuthlands
ranger base (11°39"S, 142°44°E) (24 Jan,), one seun bul not
callected.

Habitats

Ingram (1977) recorded QMIZ6261 (the holotype) under
loaf litter in a small patch of monsoon forest. AMR94360,
from | kim af the type tocaliry, way found under a piece of
wood on the verge of the rond bordering @ ‘vine and serub
thicket’ (Greer & Cogger, 1985), Rainforest communities al
the type locality are small, isolated patches on erey loamy
sand, surrounded by tall open forest dontinated by Ewcalyprus
tetrodonta (David Jones, pers. comm.). AMR94361-2,
R94484-5 and R1L12004 were found in low dense heath on
white sand; one beneath a lignotuber and the others from a
pile of debris besidea track (Greer & Cogger, 1985). The three
QNPWS specimens from Eliot Ck came from tal! open forest
of mainly bloodwood, stringy barks, Xanthostemon
xerophylus, acacias, scattered livistona palms and pandanus,
iranwood with 3 grassy understory and. a moderate scattering
of shrubs (Mike Provic, pers. comm). These were within a
few metres of a large termite mound in powdery grey Sand.
The fourth QNPWS specimen came from thick heath on
red/yellow powdery sand in Melaleuca viridiflara; Banksia
denata, jacksonias; acacias and xanthormhoeas scrub (Mike
Provic, pers. comm.) with abundant Jarge red termite mounds.
QMJ54040 and J54082 were collected in a stand of epacrid
scrub (Leucopogon sp.), 100m from a smull sirearn, rom grey
foamy-sand beneath a small termite ridden log which sup-
ported an active termite mound. QMJ54083, 154213 were in
a similar situation but the logs, while termite-ridden, did not
Support active termite mounds. The specimen fram Captain

Billy road was in norophyll vine forest (Duyid Jones, pers.
comm.) an grey loamy-sand, in asyociation with a termile-nid-
den log supporting an active termite mound.

Overall, there appears to be no corelation of vegetation
lypes at collection sites. However, A. plute has always been
found in sandy substrates.

Notes

A, pluto is known from 15 preserved specimens in the
collections of the QM (5), AM (6) and QNPWS (4). These are
from 7 localities, all within the Heathlands area of Cape York
Peninsula. Following Thomas & McDonald’s (1989) clas-
sification, McDonald etal. (1991) assigned A, pluto a ranking
of 3K becanse of its limited distribution and the lack of
information. However, A. p/ute localities are within a range
of less than 1(X) km; thus, according to their criteria, a rank of
2K is correct. While il is impossible to assess abundance of 4.
pluto, the many specimens encountered at Eliot Ck suggests
that it may be common in areas of fayourable habitat.

Lite information is available on diet, Stomach contents of
QMI54083 contained only termite fragments, not surprising
from the close association between A. pluto and active termite
nests at several Jogalities. Absence of sand particles in the
stomach of QMJ54083 may suggest this specimen had been
feeding on termites within their burrows (E. Dahms, pers,
comm.). Whether A. pluto is a specialist termite feeder or a
more general feeder is unknown.

A. pluto has an elongute right lung and no right oviduct.
Other lizards that have lost the right oviduct produce a brood
size af one and A, plido should also follow this pattern (Greer,
1984),

Tissues from QMJ54213 are deposited in the Evolutionary
Biology unit, South Australian Museum,

The work was done with a grant from the Royal Gengraphi-
cal Society of Queensland, Kate Couper, Wayne Longmore,
Paul Feeney, Martin Cohen, Stove Williams and Trevor Mc-
Leod assisted in the field. Ross Sadlier supplied details of
Australian Muscum specimens. David Jones, Allen Greerand
Mike Provic gave helpful advice,

Literature Cited

Ingram, GJ. 1977. A new legless skink Anomalopus pluta
from Cape York Peninsula, Queensland, Victorian
Naturalist 94: 52-53,

Greer, A.B. & Cogger, 1G. 1985, Systematics of the reduce-
limbed and Jimbless skinks curteatly designed ta the
genusAaomalopus (Lacertilia: Scincidae). Records ofthe
Australian Museum 37(1); 11-54,

Thomas, M.B. & MeDonald, W.3.F. 1989. ‘Rare unit threst-
ened plants of Queensland’. 2nd ed. (DPI: Brisbane)
McDenald, K.R., Covacevich, J-A., Ingram, GJ, & Couper,
P.J. 1991. The status of frogs and reptiles. Pp. 338-345.
In Ingram, GJ. & Raven, RJ. (eds), ‘An was of
Queensiand's frogs, reptiles, birds and mammaals."

(Board of Trustees, Queensland Museum: Brisbane),

Greer, 4.E. $989. ‘The biology and evolution of Australian
lizards’. (Surrey Beutly & Sons; Chipping Norton)
Hipp.

PJ. Couper, Queensland Muse, PO Box 3300,
South Brisbane, Queensland 4/01, Australia; 20
February, 1992.

A NEW SPECIES OF SKINK OF LERISTA FROM QUEENSLAND AND A
RE-APPRAISAL OF L. ALLANAE (LONGMAN).

PATRICK J, COUPER AND GLEN J. INGRAM

Couper, P.J. & Ingram, G.J. 1992 06 29: A new species of skink of Lerista from Queensland
and a re-appraisal of L. a/lanae (Longman). Memoirs of the Queensland Museum 32(1):
55-59, Brisbane, ISSN 0079-8835.

L. colliveri sp.nov. is found in NE and Central Queensland. It has been confused with L.
allanae (Longman) but, besides the strickingly different colour-pattern, it can be readily
distinguished by the presence of a nubbin for a foreleg whereas L. allanae has no forelegs,
With L. colliveri separated from L. allanae, the range of the latter species is oven more
restricted than thought. This rare species has only been found between Clermont and Capella,
Central Queensland, within the area 22°50-23°06’S and 147°38-147°54’E. It apparently has
not been met with since 1960.01Scincidae, Lerista allanae, Lerista colliveri, new species,
Queensland.

Patrick J. Couper and Glen J, Ingram, Queensland Museum, PO Box 3300, South Brisbane,

Queensland 4101, Australia; 14 February, 1992.

The status of Lerista was clarified by Greer
(1967, 1986). Since 1972, the species of the genus
have received much attention in the central and
western States because of the work of Glen Storr
(see Wilson & Knowles, 1988. for references). In
that time, however, the species of Queensland
have not been as intensely studied.

For Queensland, Storr (1972) clarified the
status of L. muelleri (Fischer) and Cogger (1975)
noted 9 species of Lerista from the State. Greer
(1979) described a new species L. ameles from
NE Queensland and Cogger et al. (1983) listed
the synonymies for all species, Greer et al. (1983)
added three new species - L. cinerea, L. storriand
L. vitrata - from NW, NE and Central Queensland
and Storr (1986) erected L. aericeps aericeps
nov., from SW Queensland and the Northern
Territory.

In 1988, Wilson & Knowles reviewed all
Australian terrestrial reptiles and noted 15
species of Lerista from Queensland. However, in
1990, Greer said L, aericeps aericeps Storr was a
junior synonym of L. xanthura Storr. Even so,
Storr (1991a) was unconvinced and preferred to
retain L. aericeps as a valid species, Storr(1991b)
also disagreed with Greer’s (1990) and Wilson &
Knowles’s (1988) concept of L. orientalis (de
Vis); he divided the taxon into three allopatric
species, two of which were new: L. zenulata and
L. ingrani,

Recently, Coyacevich & Couper (1991b) listed
17 species from Queensland and also provided
maps (1991a) of 1S of the species that were
represented in the Queensland Museum collec-
tion. In this paper, we add a further new species
from NE and Central Queensland that has been

confused with L. allanae (Longman), We alse
ré-appraise the status of the rare skink, L. allanae.

Measurements are in millimetres, Specimens
prefixed by J are housed in the Queensland
Museum, by R are in the Australian Museum,
Queensland regions follow Ingram & Raven
(1991),

Lerista colliveri sp.nov,
(Figs 1-3)

Leristaallanae Wilson and Knowles, 1988: 291, photo
no.523; Cogger, 1983, 1986: pl.672; Covaceyich &
Couper, 199 1a: fig. 305

MATERIAL EXAMINED

Houoryee: J16181 Hughenden (20°51'S, 144°12°E),
CQ, J. Devecchi.

PARATYPES: J33128 Battery Stn (19°26'S, 145°51"E),
NEQ; J33123 Harvey Ra, Mingela (19°41'S,
146°20'E), NEQ, A. Taplin, Mar 1979; 533124 Fan-
ning R Stn, c.1,5 km from hsd (19°44'S, 146°26"B),
NEQ, K.R. McDonald, F. Parker, 1 Mar 1980;
333127 Fanning R Sta, 1.5km from hsd (19°44'S,
146°26'E), NEQ, K.R, McDonald, F. Parker, 1 Mar
1980; J46266-7 Mingéla Ra, Fanning R Stn
(19°44°S, 146°27'E), NEQ, K.R. McDonald, F.
Parker, 1 Mar 1980; R62452 Red Falls at the Basalt
Wall, 60km due WNW Charters Towers (19°56'S,
145°44E”), NEQ, E, Cameron, H, Cogger, A, Greer
and P. Webber, 16 Jul 1977; R113515-39 area inand
near Basalt Wall in vicinity of Reed Falls, Lolwarth
Ck, (19°56'S, 145°446') CQ, A.E. Greer, B.N. Ar-
nold, D. Kent, R, Sadlier, G. Shea, 10-12 Sep 1984;
J33125-6 Toamba Stn (19°58’S, 145°34"E), NEQ,
B.C. Lawrie, D, Locke, 1 Jun 1981; 145648 Dun-

36 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Lerista colliveri (holotype J16181). Above: FIG. 2. Distribution of Lerista colliveri (*) and L.
dorsal view of head. Below: lateral view of head. allanae (+).

FIG. 3. Lerista colliveri Red Falls, NEQ (Glenn Shea),

A NEW LERISTA SKINK 57

raven St, c.80km NW Hughenden at Stewart Ck
(20°30'S, 143°55"E), CQ, A. Rozefelds and J.
Covacevich, 7 Aug 1985; J16182-3 Hughenden
(20°51'S, 144°12°E), CQ, J. Devecchi.

DIAGNOSIS

A large (maximum snout-vent length 84) Leris-
fa with a movable eyelid; foreleg reduced and
represented by a nubbin; and hindleg with one
clawed digit. Most similar to ZL. allanae but dis-
tinguished by colour-pattern (continuous, dark,
longitudinal lines dorsally and laterally; ventral
scales with or without dark flecks vs longitudinal
lines of dark spots dorsally and laterally, all
ventral scales dark edged).

DESCRIPTION

Snout-vent length (mm): 41-84 (N 12, mean
67.4). Length of appendages etc. (% snout-vent
length): hindleg 7-9 (N 12, mean 7.6), tail 111-
127 (N 4, mean 118.5), head length 9-13 (N 12,
mean 10.0), head width 6-8 (N 12, mean 6.7).

Nasals broadly in contact. Prefrontals widely
separated. Frontoparietals narrowly (N 7) to
moderately (4) separated but sometimes touching
(1). Usually 2 nuchals contacting the panetals,,
rarely 3 (N 12, mean 2.1). Supraoculars 3, first
two in contact with the frontal. Supraciliaries 5
(N 28), Loreals 2, second smaller. Presuboculars
2, second the smaller. Supralabials 6, fourth
below eye. Infralabials 6, Midbody scale rows 18
(N 28). Paravertebral scales §1-94 (N 13, mean
87.7). Foreleg almost absent, represented by a
nubbin in a dimple, Hindleg with one clawed
digit, Lamellae under longest toe 5-7 (N 12, mean
5,9).

Dorsa)] and lateral ground colour tan with a
pattern of more or less continuous, dark, lon-
gitudinal lines; the lines are formed by individual
scales having a dark central streak; on lower
flanks the lines are broken into dark-edged scales.
Ventral ground colour cream; chest and belly
immaculate to well-marked with dark flecks; un-
derside of tail boldly to obscurely patterned with
longitudinal lines of dark flecks. Head shields
dark-edged and mottled; lips barred; Joreals and
temporals marked by a dark streak breaking up
postenorly.

DISTRIBUTION
From just east of Townsville, NEQ, southwest
to the Hughenden area, CQ.

ETYMOLOGY
Named for Mr Stanley Colliver OBE (1908-

1991) for his generosity and outstanding con-
tributions to the Queensland Museum.

REMARKS

L. colliveri was included in L. allanae by Cog-
ger (1983, 1985), Wilson & Knowles (1988) and
Covacevich & Couper (1991a). The two species
are similar. However, L. colliveri has distinct
forelegs represented by nubbins while L. allanae
has no protruding forelegs; they also differ in
colour-pattem, The two species are separated by
about 400km.

According to the criteria of status in Ingram &
Raven (1991: 337), £. colliveri should be listed
as ‘3K’.

Lerista allanae (Longman)
(Figs 3-5)

Rhodona allanae Longman, 1937:167. Holotype
J6180 from Retro Station, Capella, CQ.

MATERIAL EXAMINED
J12232 Clermont (22°50'S, 147°38'E), CQ, C. Vallis,
Jan 1960; J6040, 6179-80, 6238, 6308, 6429-30 Retro

FIG, 4. Lerista allanae (holotype 36180). Above: dar-
sal view of head. Below: lateral view of head.

38 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 5. Lerista allanae (holotype J6180) from Retro Station, Capella, CQ (Garry Cranitch).

Stn, Capella, via Clermont (22°52S’, 147°54’E), CQ,
P.C. Allan; R13819 20 miles W of Capella (23°06’S,
147°43'E), CQ, J.R. Slevin, Aug 1929.

DIAGNOSIS

A large (maximum snout-vent length 88) Leris-
ta with a movable eyelid; foreleg absent (usually
represented by a shallow depression); and
hindleg with one, clawed digit. See diagnosis of
L. colliveri for further differences from that
species.

DESCRIPTION

Snout-vent length (mm): 61-88 (N 7, mean
76.3). Length of appendages etc. (% snout-vent
length); hindleg 6-10 (N 7, mean 7.7), tail 100-
104 (N 2, mean 102), head length 9-10 (N 7, mean
9.0), head width 6-8 (N 7, mean 6.4).

Nasals broadly in contact. Prefrontals widely
separated. Frontoparietals narrowly (N 4) to
moderately (2) separated but rarely touching (1).
2-4 nuchals contacting the parietals (N 7, mean
2.4). Supraoculars 3, first two in contact with the
frontal. Supraciliaries 4 or 5 (N 8, mean 4.5).
Loreals 2, second smaller. Presuboculars 2,
second the smaller, Supralabials 6, fourth below
eye. Infralabials 6. Midbody scale rows 18, rarely
20 (N 8, mean 18.3). Paravertebral scales 78-89
(N 7, mean 85.1). Foreleg absent, represented
usually by a shallow depression. Hindleg with

one, clawed digit. Lamellae under longest toe 8-9
(N 8, mean 8,6).

Dorsal and lateral ground colour tan to gray-
brown; back and flanks marked by longitudinal
lines of dark spots. Ventral ground colour pale
with chest and belly strongly marked by dark
edged scales; dark edging particularly prominent
under tail. Head shields dark-edged, obscure mot-
tled; lips barred; loreals and temporals marked by
dark streak breaking up posteriorly.

DISTRIBUTION
Between Clermont and Capella, CQ, within the
area 22°50-23°06’S and 147°38-147°54'E.

REMARKS

Longman (1937) described Rhodona allanae
from three specimens. He designated J6180 as
holotype but left unwritten the registration num-
bers of the two paratypes. Covacevich (1971) said
the paratypes were J6040 and J6238 and that
J6308 was a questionable alternative to J6238,
However, while there is no doubt about J6040,
both the dates of registration of J6238 and J6308
(28 June 1937 and 30 November, 1937 respec-
tively) are after the date of publication of
Longman’s paper (24 June 1937) and thus unlike-
ly to have been examined for the paper, Actually,
the second paratype must be J6179, which was
registered with the holotype and is noted as
‘paratype’ in the register.

A NEW LERISTA SKINK 459

According to the criteria of status in Ingram &
Raven (1991; 337), L. allanae should be listed as
“2E’. McDonald et al. (1991) rated it as ‘3R’-
However. they included specimens of L, colliveri
in their concept of LZ. allanae. With L. colliveri
removed, the species is only known from over a
restricted range of about 60km and has been
collected only in the years 1929 by JR. Slevin
(see Slevin, 1955), 1936-1938 by Mrs P.C. Allan
(inferred from the same dates of registration in
the Queensland Museum register) and 1960 by C.
Vallis. In the last 25 years, despite intensive sear-
ching between Clermont and Capella by many
collectors, L. allanae has not been found again.

L. allanae is apparently sympatric with two
other Lerista : L, punctatovittata (J46739-41) has
been taken from c.8-10km E of Capella at
Gregory Mine turnoff (23°05’S, 148°06"E) and L.
fragilis has been collected nearby (J6181-3, 6231
Retro Stn, Capella, via Clermont; J46742-3,
46746 c.8-10km E of Capella at Gregory Mine
tumoff),

ACKNOWLEDGEMENTS

We thank Steve Wilson, Gregory Czechura and
David Nebauer (Queensland Museum) and Allen
Greer (Australian Museum) for their advice and
help: Ross Sadler (Australian Museum) for the
loan of specimens in his care; and Glenn Shea
(University of Sydney) and Garry Cranitch
(Queensland Museum) for their photography,

LITERATURE CITED

COGGER, H,G, 1975. ‘Reptiles and amphibians of

Australia.” Ist ed. (Reed: Sydney). 584pp.
1983. Tbid. 3rd ed, 660pp.
1986, Ibid, 41h ed, 688pp.

COGGER, H.G, CAMERON, E.E. & COGGER, H.M.
1983. Amphibia and Reptilia. Zoological
Catalogue of Australia 1: 1-313.

COVACEVICH, J. 1971, Amphibian and reptile type-
specimens in the Queensland Museum. Memoirs
of the Queensland Museum 16: 49- 67.

COVACEYVICH, J.A. & COUPER, PJ. 1991a. The
reptile records. Pp. 45-140. In Ingram & Raven
(1991).

1991b. Checklist of reptiles. Pp, 356-359, Ibid,
GREER, A,E. 1967. A new genenc arrangement for

some Australian scincid lizards, Breviora 267;
1-19.

1979, A new species of Lerista (Lacertilia: Scin-
cidae) from northern Queensland, with remarks
on the origin of the genus. Records of the
Australian Museum 32: 383-388.

1986, Diagnosis of the Lerista bipes species-group
({Lacertilia: Scincidae), with a description of a
new species and an updated diagnosis of the
genus, Records of the Western Australian
Museum 13: 121-127.

1990, The taxonomic status of Lerista aericeps Storr
1986 with a diagnosis of the Lerista orientalis
species group, Ibid. 14; 443-448,

GREER, A.E., MCDONALD, K.R. & LAWRIE, B.C.
1983. Three new species of Lerista (Scincidac)
from northerm Queensland with a diagnosis of the
wilkinsi? species group. Journal of Herpetology
17: 247-255,

INGRAM, G.J. & RAVEN, R.J. (eds) 1991. ‘An atlas
of Queensland's frogs, reptiles, birds and mam-
tals,’ (Board of Trustees, Queensland Museum:
Brisbane). 391 pp.

LONGMAN, H.A. 1937. Herpetological notes.
Memoirs of the Queensland Museum 11; 165-
168, pl.14,

MCDONALD, K.R,, COVACEVICH, J,A,, IN-
GRAM, G.J. & COUPER, P.J. 1991, The status
of frogs and reptiles. Pp. 338-345. In Ingram &
Raven (1991).

SLEVIN, IR. 1955, Notes on Australian amphibians.
Proceedings of the California Academy of Scien-
ces 28: 355-392,

STORR, G.M. 1972. The genus Lerista (Lacertilia;
Scincidae) in Western Australia, Journal of the
Royal Society of Western Australia 54: 59-75,

1986. A new species of Lerista (Lacertilia’ Scin-
cidae) with two new subspecies from central
Australia. Records of the Western Australian
Museum 13: 145-149

1991a. Revision of Lerista aericeps (Lacertilia:
Scincidae) of central Australia, Ibid, 15: 365-370.

1991b. Revision of Lerista orientalis (Lacertilia:
Scincidae) of northern Australia. Ibid. 15: 413-
417,

WILSON, S.K, & KNOWLES, D.G. 1988,
“Australia's reptiles: a photographic reference to
the terrestrial reptiles of Australia,” (William Col-
lins! Sydney). 447 pp.

60 MEMDIRS OF THE QUEENSLAND MUSEUM

A NESTING RECORD FOR COERANOSCINCUS
RETICULATUS (GUNTHER). Memoirs of the Queensland
Museum 32(1); 60, 1992:- On 5 March, 1992, a
Coeranoseincus reticulatus nest containing § eggs was found
in moist soil beneath a rotten rainforest log in the Mistake
Mountains, SEQ (27°5S8’S, 152°22°E). The log measured
107% 20cm (length X width) and was in broad contact with
the ground surface and situated in full sunlight (12.25pm), The
eggs, just visible below the ground surface, were buried in
loose soil in a 2.5em deep depression. The substrate consisted
of a soft, loamy, basaltic soil with a pH of 6.37, Surface air
temperature was 27.6"C and the soil temperature within the
nest was a cooler 20.0°C.

The eggs varied in length from 23.71-28.94mm (mean
27.22mm) and width from 14.0-18.55mm (mean 16.59mm),
The size of the eggs seemed surprisingly large for a species
with a maximum snout-vent length (SVL) of 195mm. Only
one egg was accurately weighed. [t measured
26,05 x 16.88mm and weighed 3.85gm. Six of the eggs were
regular, oval shaped; one was slighty immegular oval; and one
was distinctly bean shaped, Seven of the eggs appeared to be

predominantly cream with black crossbands; the back and tail
had an obscure, chequered effect of cream, black and brown:
the pale venter scales were strongly marked with black edges
producing a series of dark longitudinal lines running along the
flanks, belly and under-surface of the tail,

Greer & Cogger (1985) examined shell thickness of ovidu-
cal eggs in C. reticulatus and suggested the species was
oviparous, Our record confirms their hypothesis and the C,
reticulatus eggs from Mistake Mountains represent the first
nesting record for the species.

However, it is not possible to determine whether or not the
8 eggs found were a single clutch. From preserved museum
specimens of C. reticulatus, McDonald (1977) recorded 3-6
oviducal eggs(N=4) in the months October-December. Thus,
8 eggs might be a single clutch but it is also possible that the
nesting site was shared by more than one female. Communal
nesting is well known in scincid lizards and is summarised by
Greer (1989).

The fieldwork on which this paper was based is part of a
study on the rainforests of SE Queensland funded by the
National Rainforest Conservation Programe,

FIG. 1. Coeranoscincus reticulatus egg (QM J54649) and hatchling (QM J54647).

healthy; the eighth egg had a small patch of fungus on the
shell.

Four eggs were carefully removed from the nest and
returned to the laboratory for incubation in vermiculite. Two
of these developed fungus and were preserved in 10% for-
malin (registered as Queensland Museum (QM) 154649 (Fig.
1) and 54650). When opened QM J54650 contained a well-
developed, cream coloured embryo (SVL 39.2mm, total
fength 71.52mm) with conspicuous dark crossbands on the
anterior half of the body.

The two eggs left incubating hatched on 2 Apmnil, 1992 (QM
J54647) and 5 April, 1992 (QM J54648). Hatchling OM
J54647 (Fig. 1) (SVL 59.86mm, total length 113,.34mm)
emerged from an egg measuring 28,07 * 18.55mm. Hatchling
QMJ34648 (SVL 58.24mm, total length !11.98mm) emerged
from an egg measuring 26.05 *16.88mm. Both hatchlings
were similarly patterned: the head and neck regions were

Literature Cited

Greer, A.E. 1989. ‘The biology and evolution of Australian
lizards’ (Surrey Beatty&Sons:Chipping Norton). 264pp.

Greer, A.E. & Cogger, H.G. 1985. Systematics of the reduce-
limbed and limbless skinks currently assigned to the
genus Anomalopus (Lacentilia: Scincidae). Records of the
Australian Museum 37(1): 11-54.

McDonald, K-R. 1977. Observations on the skink
Anomalopus reticulatus (Glinther) (Lacertilia; Scin-
cidae). Victorian Naturalist 94(3): 99-103,

Couper, P.J., Queensland Museum, PO Box 3300,
South Brisbane, Queensland 4101, Australia; Whittier,
J., Department of Anatomy, University of Queensland,
Queensland 4072, Australia; Mason, R.T., Department
of Zoology, Oregon State University, Corvallis,
Oregon, USA; Ingram, G.J., Queensland Museum; 13
April, 1992.

JALIBANG 2 AND THE DISTRIBUTION OF PECKED FACES IN AUSTRALIA

BRUNO DAVID, DAVID CHANT AND JOSEPHINE FLOOD

David, B., Chant, D. & Flood, J. 1992 06 29; Jalijbang 2 and the distribution of pecked
faces in Australia. Memoirs of the Queensland Museum 32(1): 61-77. Brisbane. ISSN
0079-8835,

Archaeological excavations at the Jalijbang 2 rockshelter, near Katherine, Northern Ter-
ritory, show that in this area pecked faces and Panaramitce type peckings are unlikely to
pre-date the mid-Holocene. These results, however, are unlikely to shed light on the
antiquity of pecked faces elsewhere in arid and semi-arid Australia, for the Cleland Hills,
Durba Hills, Dampier, Sturt Creek, and Jalijbang pecked faces appear to be regionally
distinctive. Itdoes, however, show that the so-called Panaramifee artistic style consists of
a broad, highly generalised set of conventions which cannot be purcly understood as of
having great antiquity] reck art, Wardaman, Northern Territory, Australian prehistory.

Bruno David, Department of Anthropology and Sociology, University of Queensland,
Queensland 4072, Australia; David Chant, Department of Social Science, University of

Queensland, Queensland 4072, Australia; Josephine Flood, Australian Heritage Commis-
sian, PO Box 1567, Canberra, Australian Capital Territory 260], Australia, 9 November,

199]

In a secluded sandstone gorge, 150km south-
west of Katherine, N.T. (Fig. 1), are found a
series of highly unusual engraved figures. When
they were first observed by archaeologists
during the mid 1980s, the figures were deseribed
asa minor but striking component of a broader
regional body of rock art dominated by non-
figurative and track peckings and figurative
paintings (Lewis & McCausland, 1987). The
figures themselves consist of four large
anthropomorphs pecked into a near-yertical
sandstone wall, three of which have their faces
depicted (Figs 2-4), A further three pecked faces
are located on nearby boulders (Figs 5 & 6), The
site in which these occur is known as Jalijbang
(spelled ‘Jalikpany’ by Lewis & McCausland,
1987) by the local Wardaman Aboriginal people.
The pecked faces on the wall and boulders are
all depicted in frontal view, and are reminiscent
of the pecked faces from Cleland Hills (1000km
to the south), which are commonly believed to
be of great antiquity (Edwards, 1968). Lewis &
McCausland (1987: 78), entertaining the idea of
prehistoric inter-regional contacts between
people from the Jalijbang region, Cleland Hills,
and from other regions containing similar
engravings, concluded that:

“whether or not there is any relationship between
the Jalikpany figures and the complex
anthropomorphic figures and/or face engravings
elsewhere cannot, at this time, be determined.’

This paper is an attempt to address this issue
und to investigate the antiquity of the Jalijbang

figures, Furthermore, if the Jalijbang engravings
can be shown to be closely related stylistically to
the Cleland Hills engravings, then the antiquity
of the former may shed light on the antiquity of
the latter.

INVESTIGATING THE ANTIQUITY OF
THE JALITBANG PECKINGS

Jalijbang 2 is a small rockshelter with a high
ceiling near the downstream end of a small
gorge, The site itself has a curious setting in that
itis bounded to the east by a vertical sandstone
rock face, to the south by massive sandstone
boulders (evidence of past roof-falls), and to the
west and north by a seasonal creek, The creek
bed itself consists of high quality ochre pebbles
and quartzite and sandstone blocks. A small
pocket of soft sediments has been preserved in
the area bounded by the creek, the rock wall and
the boulders, and this was made possible only
because the boulders themselves protected it
from the creeks northward waterflow during the
wet season. Water rushes past the soft deposits
at Jalijbang 2 during the wet season, detouring
around the massive boulders, and flooding the
pocket of soft sediments with quiet waters which
deposit new silts during times of flood. It is in
this protected pocket that archaeological sedi-
ments are found, preserved by the periodic addi-
tion of silts (Fig, 7),

The large boulders near the soft sediment con-
tain numerous engravings. Most of these consist
of abraded greoves and pecked bird tracks, But

62 MEMOIRS OF THE QUEENSLAND MUSEUM

te
JALUIBANE Y|MGALARR|

° STURT GREEK

o CLELAND HILLS

QURBA HILLS

FIG. 1, Australia showing locations mentioned in text.

near the soft deposits, where the boulders meet
the creek bed, lies a medium-sized boulder with
a pecked face (Fig. 5). Two other pecked faces,
as well as peckings of dingo-forms, occur on
boulders in a rockshelter 20m upstream (Fig. 6).
These boulders appear to be part of the massive
sandstone boulder scree which begins at the up-
stream rockshelter, and end at the downstream
end of Jalijbang 2, They are likely to have been
deposited as a result of either instantaneous or
prolonged roof-fall events.

The peckings on these boulders all occur on
the boulder’s upper or side surfaces (Fig. 8).
None have been found on underlying boulders.
Consequently, all of the boulder engravings are
likely to post-date the roof-fall. Further, if the
roof-fall itself was a prolonged event, then the
engravings themselves were likely to have been
undertaken after the /atest episode of roof-fall,
given that engravings only occur on the uppermost
boulders. In excavating at Jalijbang 2, we there-
fore aimed at dating the roof- fall in order to
obtain a maximum antiquity for the boulder
engravings, which include three pecked faces

(two of which occur in the upper rockshelter)
reminiscent of the Cleland Hills pecked faces.

In July and August 1989, one of us (B.D.)
undertook a small excavation at Jalijbang 2,
immediately beneath the four pecked
anthropomorphs which adorn the sandstone
wall (Fig. 7). Sixteen juxtaposed 50cm % 50cm
squares were excavated until compact sedi-
ments were reached, after which excayation of
only four squares continued’. All materials
were excavated in bucket spits following the
natural stratigraphy (Johnson, 1979). Stone ar-
tefacts and other cultural materials greater than
2cm maximum dimension were recorded in
three dimensions and bagged separately, as
were all hearth stones,

Although the Jalijbang 2 sediments were well
stratified, most of the individual stratigraphic
laminations resulting from the alluvial deposi-
tion of silt could not be identified in situ, as
variations in texture and colour were minimal.
"The surface sediments from the other 12 squares were

excavated so that loose surface sediments would not
fall into the main pitand contaminate older materials.

PECKED FACES IN AUSTRALIA 63

FIG. 2. Pecked anthropomorph located on a wall at
Jalijbang 2.

Five stratigraphic units were revealed from the
excavation (Fig. 9):

Stratum 1: loose surface sediments. Fine tan-brown
silts, extremely loose, containing small amounts of
organic materials. Sediments are uniformly very fine.

Stratum 2: immediately underlies Stratum (SU) 1. SU
2 consists of a number of ashy and/or charcoal-rich
lenses within a broader silt layer. These lenses are
given separate identification (SU 2b, 2c and 2f), and
most are spatially discontinuous across the excava-
tion. SU 2 proper is a fine, silty sediment, finely
laminated in some places, but the lamination was too
indistinct to record except for sub-stratum 2a. The silts
contain very small amounts of organic matter (includ-
ing charcoal). Sediments are light tan-brown in colour.
The separate sub-strata are:

SU 2a, being a 2mm thick lens of slightly darker silt;
SU 2b, a fairly well defined layer of grey ashy silt
(poor in charcoal);

SU 2c, an extremely well defined charcoal concentra-
tion, containing burnt stones. It is very well defined in
square E6 especially. In this layer were found the first
evidence of pebbles (all burnt), many of which are fire
cracked. SU 2c is black in colour, and is almost pure
charcoal;

FIG, 3, Pecked anthropomorph located on a wall at
Jalijbang 2.

SU 2d, a fine gravel lens, with silty matrix around it.
It is present in square E6, but it is ill-defined else-
where;

SU 2e, a greyish orange silt;

SU 2f, an extremely well defined charcoal lens. It is
very ashy and black in colour.

Stratum 3: extremely compact gravel and pebble layer,
orange-brown in colour. Pebbles range in size from
small (<Icm) to approximately 25cm maximum size.
The pebbles and gravel making up SU 3 appear similar
to the materials which currently make up the adjacent
creek bed, The boundary between SU 2 and SU 3 is
well defined.

Stratum 4; decomposing sandstone, mottled reddish
yellow. Intersection with overlying SU 3 is ill-defined,
spanning approximately 3cm.

Stratum 5: solid sandstone. This is the basal
stratigraphic unit consisting of large boulders with
relatively flat surfaces. Excavation could not proceed
into or below Stratum 5 because of the size of the
boulders.

RADIOCARBON DATES
Two radiocarbon dates were obtained:

ad MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Pecked anthropomorph Jocated on a wall at
Jalijbang 2

a. 260+ 60BP (New Age; Conventional Age is
250+-60BP) (Wk-1551). This date is from a good
charcoal sample from Square E6, XU 6, and
dates the uppermost hearth excavated (SU 2c)
("C permil = -26.6). This hearth is a well-
defined charcoal and hearth stone lens which is
assumed to have been created as an archaeologi-
cally instantaneous event,

b, 1040+60BP (New Age; Conventional Age
is 1010+60BP) (Wk-1550). This dates the lower-
most hearth (SU 2f), and comes from a good
quality charcoal sample from Square F6, XU 10
(°C permil =~-26.0).

OccuPaATION AT JALUBANG 2

Human occupation at Jalijbang 2 began some-
time before 1040+60BP. Low density deposition
rates of stone artefacts are found throughout the
deposits, although peak rates occur in associa-
tion with the two hearths (around 260BP and
1040BP) (Table 1). Only two small fragments of
bone were found (both in post-260BP times),
whilst hearth stones and heat-shattered burt

rocks are associated with the uppermost hearth.
Ochre occurs, but it is difficult to associate its
presence with artistic activities, as the creek bed
itself contains vast numbers of ochre pebbles,
and periodic floods can be expected to deposit
very small fragments of ochre with the deposited
silts. Ochre pebbles were also used as hearth
stones during use of the uppermost hearth. No in
situ ochre fragments exhibit any signs of use
wear (Appendix 1).

Although stone artefacts occur below the bot-
tom hearth (1040+60BP), their numbers are very
low. These low rates are associated with SU 3
and 4, the gravel and sandstone layers below the
alluyial silts and hearths, The gravels themselves
indicate the presence of creek flow in this section
of the site sometime in the past (in pre-1040BP
times}, although the large boulders were present
to the immediate south during that time (as indi-
cated by the underlying SU 5 boulders in the
excavations). An extension of the depth-age
curve dates the uppermost surface of the in situ
sandstone boulders (SU 5) to 2880BP. although
the presence of a different stratigraphic unit be-
tween it and the dated silts (SU 2) means that this
date is likely to be a very general indication only
of the age of the roof-fall which resulted in the
boulder deposit. Consequently, this extrapolated
date should be used with caution, and is
presented here as a very general indication only
of the order of magnitude which we can expect
the age of the roof-fall underlying the cultural
deposits to be. It is not known if cultural
materials occur below the boulders, as excava-
tion could not proceed below them.

Otner DATES

Two other independent sources indicate a mid-
to late-Holocene antiquity for the end of the
roof-fall expressed stratigraphically as SU 5:

1. On boulders located in a rockshelter 20m
upstream from Jalijbang 2 (at Mennye-ya), a
large, exfoliated boulder contains pecked dingo-
forms on its upper surface. These boulders. were
likely to have been deposited during the same
roof-fall event(s) which resulted in the creation
of the scree-slope at Jalijbang 2, and dated to
approximately pre-2880BP by reference to the
depth- age curve. As dingoes first arrived in
Australia around 3500 years ago (Solomon &
David, 1991), the peckings must have been un-
dertaken sometime after that time. This does not
inean that the boulders were necessarily
deposited after this time, but does indicate that
atleast some of the art on them were done in mid-

PECKED FACES IN AUSTRALIA 65

FIG. 5, Weathered pecked face from creek-bed boulder. Jalijbang 2 (see Fig, 7 for location of boulder).

TABLE 1. Deposition of materials, Squares E6 and F6, All rates are per mi per 100 years.

to late-Holocene times. The other peckings on
this and nearby boulders show a similar degree
of patination (moderate to heavy) as the dingo
peckings, indicating probable comtemporaneity,

2. A cortex sample was extracted from a near-
by boulder by Alan Watchman (pers. comm.,
1988). The cortex was examined by Watchman

| squane [oer ff ef 3 fa fis fs fs —
ae ee por | | 0.22 |1989AD-2608P |
es [oe ess emo no | [00 | as [a2 fpoope —
pms fen feat fue [3a |_| _}__1_ 05260-10405 —_

[is | <0i | iss | 36 |

a [sa | 100 | a0 |_|

res Pan [ana [an i
Fio-ii| iss | 12000 [oesoo | | | 996 [1467.6 [io4opP
Fe Tartan os] a PT t05 -onn

1=CHARCOAL (KG). 2=LITHICS (#). 3=LITHICS (G). 4=BONE (G), 5=HEARTH STONES (#).
=OTHER BURNT ROCK (KG), 7=RESIDUE (KG), 8

=a

=YEARS IN SPITS

(1990), who found a thin, microscopic layer of
organic-rich materials. at its base. The organic
materials were extracted and dated through
AMS, where a date of 40104300 was obtained
(Watchman, 1990). As the sample came from
near the base of the cortex, if dates the begin-
nings of cortex development, and hence when

66 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 6, Pecked faces, A, -N=Dampier. B-C=Jalijbang boulders. D-G=Cleland Hills. H=Durba Hills. After Dix
(1977) and Lewis & McCausland (1987).

PECKED FACES IN AUSTRALIA 67

FIG. 7. Plan of Jalijbang 2. Fine-dashed area is large
boulder scree.

the boulder probably became exposed to the

elements. 4010+300BP is therefore likely to rep-

resent the antiquity of the boulder’s deposition

to its current position,

Discussion OF ANTIQUITY OF BOULDERS AT
JALUBANG 2

Three independent investigations al] point toa
mid- to late- Holocene antiquity for the end of
the roof-fali which resulted in the massive
boulder scree at Jalijbang 2. The in situ
sandstone boulders located in the excavation
were themselves tentatively dated to around
2880BP by extension of the depth-age curve.
A major problem with this dating ts that it
assumes a steady sedimentation rate below the
lowest radiocarbon date, and hence, by itself,
the resultant chronology can only be taken as
a general approximation at best. When com-
pared to the cortex date (4010+300BP) and the
presence of pecked dingo-forms on boulders

from theadjacentscreedeposits(post-3500BP),
however, the depth-age calculation appears to be
of the correct order of magnitude. The roof-fall
at Jalijbang 2 appears to have ended around 3000
to 4000 years ago.

ANTIQUITY OF PECKED PACES

Given a likely mid- to late-Holocene antiquity
for the end of the roof-fall at Jalijbang 2, itis also
likely that the pecked faces and other engravings
on the boulders are of a similar antiquity. The
pecked face in the creek bed at the base of the
boulder scree (Fig. 5) is also unlikely to pre-date
the mid-Holocene as the annual waterflow in the
creek can be expected to have obliterated the
pecking were if to have greater antiquity. It is
notable that during pre-3000BP times precipita-
tion was significantly higher than at present,
making it even move unlikely that this pecking
would have survived (Chappell & Grindrod, 1983).

As all of the boulder engravings at Jalijbang
2 are located on the upper and side surfaces of
the uppermost boulders only, it is also Jikely that
all engravings past-date the latest episode of
roof-fall (3000 to 4000BP). These engrayings
include pecked faces and bird tracks and
abraded grooves.

The peckings on the rock wall, however, are
much mote difficult to date. Itis possible that the
roof-fal] which terminated around 3000 to 4000
years ago resulted in the creation of the rock wall
itself (and hence the peckings postdate this time),
but it is also possible that the current rock wall
existed before the roof-fall. If the former was the
case, the peckings on the wall would be roughly
contemporaneous with the boulder engravings.
It is possible, however, that the wall engravings
are older, although they possess similar patina-
tion to the adjacent boulder peckings (supporting
general contemporaneity, although degree of
patination can vary widely even within localised
areas). In any case as will be shown below the
boulder pecked faces are stylistically different
from the wall ones, and therefore the antiquity of
the former cannot be used to date the latter on
stylistic grounds. In short, the boulder engray-
ings probably date to the mid- to late-Holocene,
whilst the antiquity of the wall engravings is
unknown.

We investigate below the possiblity of deter-
mining the antiquity of other pecked faces from
arid and semi-arid Australia by investigating
stylistic distributions of pecked faces, We
specifically ask the question, are the dated
(boulder) pecked faces at Jalijbang 2 of a purely

68 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. All of the abraded and pecked figures located on the boulders at Jalijbang 2. All scales are in centimetres.

Continued on next page

Y
uv

local style, or are such faces found elsewhere
also; if the latter, can the antiquity of the Jalij-
bang 2 faces reveal information on the antiquity
of pecked faces elsewhere?

To address these questions, we undertake
preliminary analysis of all published pecked
faces from Australia which have a basic
resemblance to those of Jalijbang.

THE REGIONAL OCCURRENCE OF
PECKED FACES

WaARDAMAN COUNTRY
(JALUBANG AND YINGALARRI)

The pecked faces from Jalijbang can be
divided into two groups, those on boulders and
those on the wall. The three faces found on the

PECKED FACES IN AUSTRALIA 69

FIG. 8. Continued from previous page. All of the abraded and pecked figures located on the boulders at Jalijbang

2, All scales are in centimetres.

/ ——
i@] x
— No WY ce ial
L ee te
oO 10
—_———J
SF
\
&
= °

rock wall are very similar to each other: they all
have concentric circles for the eyes, ears, noses,
and broadly cylindrical outlined faces. They also
have full or partial bodies. The faces pecked on
boulders, however, exhibit neither bodies nor
ears, and consist only of non-concentric eyes.
Only one of the boulder pecked faces has a nose.
The boulder faces are generally smaller than
those located on the wall, and were created by a
small number of separate, pecked lines (6, 7 and
7 for the boulder faces, compared to the 14, 20
and 26 lines that make up the pecked faces on the
wall). In short, the Jalijbang pecked faces can
be divided into two distinct stylistic groups, the

? These numbers refer to the separate lines that make
up the faces of each figure only.

mS ee

structurally complex wall faces, and the more
generalised boulder ones.

A pecked face was also recently rediscovered
by the authors at Yingalarri, 20km east of Jalij-
bang (and still within Wardaman country). This
pecking (see Flood et al., in press), like the
Jalijbang boulder ones, lacks the structural com-
plexity of the Jalijbang wall peckings (see Table
2 for traits present).

CLELAND HILLs

Edwards (1968) reported on the occurrance of
fifteen pecked faces from the Cleland Hills,
1000km south of Jalijbang. Of these, five figures
are illustrated in his and subsequent reports (Ed-
wards, 1968; Dix, 1977) (many of the others are
badly weathered and difficult to distinguish).
The Cleland Hills faces have been linked stylis-

me MEMOIRS OF THE QUEENSLAND MUSEUM

@ CHARCEAL
© SANOSTONE

FIG. 9. Section drawing, Jalijbang 2,

tically to those from Jalijbang, Dampier, Sturt
Creek and elsewhere (e.g. Dix, 1972, 1977;
Lewis & McCausland, 1987), and have been
considered to have considerable antiquity by
virtue of their advanced state of patination. Yet
the Cleland Hills faces contain a suite of charac-
teristics not found in pecked faces elsewhere.
These faces are generally pear- shaped or heart-
shaped with noses depicted whilst their mouths
aré shown as straight or curved lines (at Jalijbang
mouths on the wail faces are represented as
single dots). Eyes consist of concentric circles,
but unlike the Jalijbang wall faces, they were
executed by a simall number of separate pecked
jines only (7, 8, 9, 10 and 10 for the five faces
Ulustrated in Edwards, 1968 & Dix, 1977). We
will return to these points below.

DAMPIER
The pecked faces from the Dampier region of

10

20

70

Western Australia are relatively varied in style.
One of their dominant features is the repre-
sentation of hair on many figures, although this
is not always so. Other characteristics are con-
centric circle eyes, and the enclosure of the head
in an outline. The nose is sometimes represented,
but ears are not and heads are never pear-shaped.
The Dampier faces include figures created by
numerous separate lines (up to 32) as well as
some created by a few lines only (as few as
seven). The former are more common.

Dursa HILts

The Durba Hills (W.A.) face reported by Dix
(1977) exhibits hair and concentric circle eyes,
but nose, ears, body, and head enclosure are
missing. It is reminiscent of the Dampier faces,
and this is especially so of the hair and of the
parallel lines below the eyes.

PECKED FACES IN AUSTRALIA 71

FIG, 10, MDS map based on eight qualitative yari-
ables.

Sturt CReek

The Sturt Creek pecking has, like the Cleland
Hills faces, a limear mouth. The head is enclosed
and attached to a full body (Crawford, 1968;
Walsh. 1989). Itis in this way reminiscent of the
Jalijbang 2 wall peckings, although unlike the
latter, it does not have concentric circle eyes,
ears oF nose.

REGIONAL VARIABILITY

The Jalijbang 2 boulder and wall pecked faces
are stylistically different from each other, espe-
cially with respect to the presence or absence of
concentric circle eyes, nose, ears, and associated
bodies. Similarly, whilst the wall faces are com-
posed of many separate pecked lines, the boulder
faces contain considerably fewer lines (Table 3).
The pecked face Jocated on an exposed boulder
it Yingalarri is stylistically similar to the boulder
faces from Jalijbang 2.

To the south and southwest of Jalijbang, how-
ever, are found a number of pecked faces which,
ulthough superficially resembling the Jalijbang
faces, are regional variations of the same theme
(pecked faces). Such peckings are characteristic,
although minor, components of some arid and
semi-arid zone pecked assemblages. Since they
are not found in more humid zones, their shared
presence in arid Australia has often entertained
the possibility of inter-regional interactions link-
ing the places where they are found in a vast
desert social network. But before this idea can be
properly entertained, the presence of shared

stylistic traits should be demonstrated. The
presence of pecked faces in different areas, by
itself, cannot be taken to signify a single artistic
convention - pecked faces are not necessarily
structurally complex endeavours, and only a
limited number of traits (e.g. presence of eyes
and head outline) are needed to make a picture
resemble a face, In other words, the presence of
non-essential lines (¢.g. ears are not needed to
make. face-form), and/or of particular combina-
tions of features, are needed to be shared before
two geographically separate bodies of pecked
faces can be deemed to be represented by asingle
stylistic convention.

In examining the similarities and differences
between the known pecked faces of Jalijbang,
Yingalarri, Cleland Hills, Durba Hills, Dampier
and Sturt Creek. we began by identifying
whether certain traits were present or not, These
traits are:

1. hair or head-dress;

2. nose;

3, ears;

4, linear mouth (as opposed to a dot or an
enclosed area);

5. concentric circle eyes (a circle around a dot
is not considered a concentric circle);

6. pear or heart shaped head:

7. head enclosed by an outline:

8. associated body.

The traits themselves were chosen to highlight
stylistic variation. It was noticed that some traits
were present in all pecked faces (e.g. eyes),
whilst others were present in some only. The
traits chosen specifically represent the major
traits present in a number of peckings but not all
of them, A Multi- Dimensional Scaling (MDS)
statistic was subsequently undertaken to reveal
how the separate faces cluster stylistically with
each other, on the basis of the combination of the
eight identified traits. Our main aim was to deter-
mine whether or not the faces from each region
clustered together and, if so, did these clusters
separate out conspicuously from the other
regions.

The variables used in the MDS were the eight
traits enumerated in Table 2. MDS is a set of
mathematical techniques which enables the
mapping of units (pecked faces) as points in
geometrical space, reducing the number of
dimensions (eight in our case) to two. It enables
the user to locate the units in a spatial configura-
tion - a ‘map’ (Fig. 10) (Alvey etal., 1982: 122).
Ulsaner (1978: 5) writes: ‘Once we have located
the ... points in (multi-dimensional) space, we

72

TABLE 2. Traits present in the pecked faces analysed.

LINEAR | CONC. | PEAR-SHAPED
MOUTH | EYES HEAD

satwauL | |X
praLwate | |X
3. JAL WALL

4. JAL BOULDER

5. JAL BOULDER

6. JAL BOULDER

11.CLELAND HILLS
12. CLELAND HILLS

seek to determine the hidden structure, or
theoretical meaning of this spatial
configuration’.

The importance of MDS is that it provides a
way of reducing the data to two dimensions.
These two dimensions represent the ‘hidden
structure’ of the data. Identification of key dif-
ferences between the pecked faces was deter-
mined by identifying those that stand out or
cluster together on the MDS map. To do this,
statistical proximity values between units (peck-
ed faces) are computed. “The larger the dis-
similarity (or the smaller the similarity) between
two objects, as shown by their proximity values,
the further apart they should be in the spatial
map’ (Kruskal & Wish, 1978: 7).

The results of the MDS shows a high level of
regional clustering. As expected, the Jalijbang 2
wall and boulder peckings separate out. The
Yingalarri figure, also located within Wardaman
country, cluster with the Jalijbang 2 boulder
faces. The Cleland Hills faces separate out well,
as do the Dampier and Durba Hills faces. The

MEMOIRS OF THE QUEENSLAND MUSEUM

HEAD
ENCLOSED

BODY

ae
al

Sturt Creek figure is similar to one of the Dam-
pier figures, but all other pecked faces-are
regionally discrete (Fig. 10). In short, the arid
and semi-arid zone pecked faces consist of a
series of regionally specific figures, differen-
tiated by the presence or absence of particular
traits. It is the combination of such traits which
gives each region its stylistic particularity. Un-
fortunately, the published information and il-
lustrations of these pecked figures do not reveal
enough information to determine whether or not
there are also regionally specific metrical dif-
ferences. We have nevertheless briefly at-
tempted to gather information on this question
by quantifying the following:

a. the ratio of the distance between the eyes
(taken from centre of eyes) to width of face at
that plane;

b. the maximum width to height ratio (the
former being at a parrallel to the eye plane, and
the latter perpendicular to width);

c. the number of separate pecked lines making

PECKED FACES IN AUSTRALIA 73

‘TABLE 3. Quantitative data.

i
LINES

5. JAL BOULDER 0.54

IT.YINGALARRI | 0.36. |_os7| 8 |
&, CLELAND HILLS
9.CLELANDHILLS | 0.50 | 1.00 | 10 |
H0.CLELANDHILLS | _o.ss_| 1.00 | Ww _|
1.CLELANDHILLS | 0.56 | 090 | 9 |

12, CLELAND 1.20
0.71 l

| 8
|_i6_|
| 8

_

DBE=DISTANCE BETWEEN EYES. FW= FACIAL
WIDTH . W=WIDTH. H=HEIGHT

up the faces (if a body is present, itis not included
in the quantification),

The MDS undertaken on this data set does not
reveal the distinct regional character revealed by
the previous analysis (Table 3, Fig. 11). Never-
theless, the Cleland Hills faces separate out well
from the other peckings, accentuating their
regional specificity, Detailed quantitative
analysis will have to await the availability of
further quantitative data from each of the pecked
figures.

DISCUSSION

The rock art located in Wardaman country,
Northern Territory, is widespread and stylistical-
ly diverse. A very small component of the War-
daman rock art assemblage consists of pecked
faces, some of which also have associated torsos.
Within Wardaman country, the Jalijbang and
Yingalarri pecked faces are superficially
reminiscent of the Cleland Hills faces. These
have often been infered to have considerable
antiquity because of their stated advanced state
of patination. Published photographs, however,

show that some of these peckings are moderately
patinated only (e.g. Edwards, 1968). In any case
patination, especially of peckings in open sites,
is not necessarily an indication of great age, as is
testified by the existence of patinated post-con-
tact graffiti. Similarly, the Jalijbang pecked faces
have been claimed to be of considerable antiq-
uity, but there is no strong reason to believe this
is necessarily the case, On the contrary, the peck-
ed faces found on boulders in Jalijbang 2 and in
a nearby rockshelter (Mennye-ya) are likely to
date to the last 3000-4000 years. Because these
figures are stylistically distinct from pecked
faces from other parts of arid and semi-arid
Australia, however, their antiquity cannot in this
way be used to infer the ages of pecked faces
elsewhere.

Furthermore, the engravings. found on the
Jahjbang 2 boulders also include pecked bird
tracks and linear designs and abraded grooves.
Such peckings have often been characterised as
typical components of the so-called Panaramitee
engraving tradition, Such engravings are often
believed to have considerable antiquity (e.g.
Maynard, 1977), but at Jalijbang 2 they appear
to have a maximum age of 3000-4000 years BP.
Abraded grooves and pecked bird tracks, how-
ever, have also been found at the Yingalarri 1
excavations (Mulvaney, 1975), where they were
dated to at least SOOOBP. Abraded grooves are
also known to have been created in the Jalijbang
and Yingalarri areas during recent times (Flood

FIG. 11, MDS map based on three quantitative vari-
ables.

74 MEMOIRS OF THE QUEENSLAND MUSEUM

etal.,in press). Itis therefore likely that peckings
and abraded grooves were part of a long artistic
tradition in Wardaman country, spanning the
period from pre-SOO0BP times to the ethnohis-
toric present. This pattern a to be simular
to that found by Nobbs & Dorn (1990) in the
Olary region to the south, where cation-ratio
dates revealed a broad range of non-figurative
and track peckings spanning approximately 30
000 years of prehistory (but see Watchman, in
prep.), In the Wardaman and Olary regions at
least, the ‘Panaramitee’ (as defined stylistically
by Maynard, 1977) has had a long history,
broken only in recent times by the White in-
truders who constrained the local people access
to the lands in which they engraved and painted.

In short, although some of the engravings
found in Wardaman country have been dated to
pre-5000BP times by Mulvaney (1975). there is
no evidence that the pecked faces from Jalijbang
2 have any great age. Although their precise
antiquity remains a matter for speculation, ex-
cayations at the site point towards a likely mid-
to late-Holocene antiquity, We say this through
circumstantial evidence only:

|. the occurrence of extensive rcof-fall pre-
dating approximately 3000BP may indicate that
the main, pecked wall at Jalijhang 2 did not exist
before that time;

2, boulder peckings occur only on the upper
and side surfaces of boulders believed to have
been deposited in their current positions during
the mid- to late-Holocene. Given the absence of
peckings fromthe undemeath surfaces, the peck-
ings are likely to have been undertaken only after
the boulders were deposited in their current posi-
ons:

3. the pecked face on the boulder in the creek
bed at Jalijbang 2 has not been totally obliterated
by water action, implying that it may not be of
great antiquity:

4. other nearby peckings, also associated with
the roof-fall boulders relating to points | and 2
above, contain peckings of dingoes, which be-
cause of their known antiquity in Australia must
post-date approximately 3500 to 4000 years BP;

5. analysis of cortex on a boulder near Jalij-
bang 2 indicates that cortex development began
approximately 4000 years ago. Because of its
proximity to Jalijbang 2, it is likely that this
boulder resulted from the same roof-fall activity
as the boulders currently forming the scree-slope
into the site itself (points 1 and 2 above), in
support of a similar antiquity for the roof-fall

itself (creation of the boulder surface where cor-
tex development ensued).

Although the Jalijbang 2 boulder peckings
(and wall peckings to a lesser extent) are more
likely to be of mid- to Jate-Holocene age, it has
not been possible to gain information on the
antiquity of other, similar engravings from the
Northern Territory and Western Australia, The
latter are stylistically distinct from the pecked
faces found in Wardaman country, with each
area possessing regionally-distinctive pecked
faces. These findings question the previously
widely-stated belief that pecked faces from
many paris of the Australian arid and semi-arid
zones were similar and presenting material
evidence fora broad, interactive social network
which linked central Australia into a more or less
unified “desert culture’. On the contrary, this is
nowhere reflected in the distribution of pecked
faces which emphasise the regional specificity
of artistic traditions in Australia’s arid core.

ACKNOWLEDGEMENTS

We would like to thank the Wardaman people
for allowing us to excayate at Jalijbang,
Earthwatch for financial sepport and fieldwork
assistance, and A.LAT.S.LS. for funding the
carbon dates. Bryce Barker, [an McNiven, Philip
Jones and an anonymous ‘referee made useful
comments on an earlier draft.

LITERATURE CITED

ALVEY,N., GALWEY, N. & LANE, P, 1982. ‘An
Intreduction to GENSTAT”. (Academic Press:
London).

CHAPPELL, J. & GRINDROD, A. (eds) 1983.
‘CLIMANZ: Proceedings of First CLIMANZ,
February 1981’. (R.S.Pac.S., A.N.U.; Canber-
ra).

CRAWFORD, I.M. 1968. ‘The Art of the Wandjina’.
(Oxford University Press; Melbourne).

DIX, W. 1972. Rock art along the stock route, Hemi-
sphere 16(7): 18-25.

1977. Facial representations in Pilbara rock engray-
ings. Pp. 277- 285, In Ucko, P.J.(ed.), ‘Form in
Indigenous An’. (A.LA.S.: Canberra).

EDWARDS, R. 1968. Prehistoric rock engravings at
Thomas Reservoir, Cleland Hills western central
Australia. Record of the South Australian
Museum 15(4): 647-670.

FLOOD, J., DAVID, B. & FROST, R. in press.
Dreaming into Art, In Morwoad, M. (ed.), “Eth-

PECKED FACES IN AUSTRALIA 15

nography and Art’ (preliminary title), (A.A.S.:
Canberra).

JOHNSON, I. 1979. The getting of data, (Unpublished
PhD thesis, Australian National University,
Canberra).

KRUSKAL, J.B. & WISH, M. (eds) 1978, ‘Multi-
dimensional Scaling’. (Sage Publications:
Beverley Hills).

LEWIS, D. & McCAUSLAND, B. 1987. Engraved
human figures and faces from Wardaman
county, eastern Victoria River district, Northern
Territory. Australian Aboriginal Studies
1987(1): 67-79.

MAYNARD, L. 1977. The archaeology of Australian
Aboriginal art. Pp. 83-100. In Mead, MLS. (ed,)
‘Exploring the Visual Arts of Oceania’. (Univer-
sity of Hawaii Press: Honolulu).

MULVANEY, D.J. 1975. ‘The Prehistory of
Australia’. (Penguin: Ringwood).

NOBBS, M. & DORN, R. 1990. Age determinations
for rock varnish formation within petroglyphs.
Rock Art Research 5(2): 108-46.

SOLOMON, 8S. & DAVID, B. 1991, Middle Range
Theory and actualistic studies: bones and din-
goes in Australian archaeology. Pp. 234-256, In
Solomon, S., Davidson I. and Watson D. (eds),
‘Problem Solving in Taphonomy’ Tempus 2:
234-56 (Anthropology Museum, University of
Queensland: Brisbane).

ULSANER, E.M. 1978. Editor's Introduction. Pp.
5-6, In Knuskal & Wish (1978).

WALSH, G. 1989. ‘Australia's Greatest Rock Art’.
(E.J Brill/R. Brown & Associates: Bathurst).

WATCHMAN, A. 1990. What are silica skins and
how are they important in rock art conservation?
Australian Aboriginal Studies 1990 (1): 21-29_

im prep. An assessment of cation-ratio dating. (Un-
published report. A-.A-T.S.I.S.; Canberra).

APPENDIX |
List of materials excavated from Jalijbang 2.

|
tw
te
—]
Nn
ma
—]
bh

37
ret athe te) i al oar

ee wo | ost | | pas 3.8 |
| 10 | | sa] 2] | 66 | os] | | os] 28 |
TO TED CT) MT
Pa | sit s | 98[ s6[ [os | [| | | oad 2s
| is | iat 5 | 29] 20] | soo | or] [| | 025] 2.5
pw | o7] 3 | os] a} | ao | orf | 6 |
}—— oon | — =a |. |
| fisoo | |
Ey
etal riswo | [|] 01 a0] 3]

I

5
“
eo]
c

cs 8
| 20 | <oi{ 2 | 02] 10] [sso | | | | or a7| 3 |
jar | of or | oat | fs [Ta

MEMOIRS OF THE QUEENSLAND MUSEUM

76

APPENDIX | cont.

w|
my) a ai] oh mpm] eon} om] st a
Los]

SIa1s a
S

SP mympy ol7|2 t
ie oat Mal ost oe mn} Alo =)
_ ae|uinles
+

PECKED FACES IN AUSTRALIA 77

APPENDIX 1 cont.

ss
>
=
a

[ou
| o1_|
| 3.6 | 242A
| 3.9 | 2428
| s.1_|
| 3.9

4

4

3

Nn
_

9

_

2
2E
2E
2E
2E
2E
2F
2F
2E

_

| 36 | 344

iy ra

11
3.6
3.9
3.
4.0
1.2
2.5
3.2
3.7
3.8
2.6
3
&
6
3.6
2.2

1=CHARCOAL (G). 2=LITHICS (#). 3=LITHICS (G). 4=OTHER ORGANICS (G). 5=BONE (G). 6=OCHRE
(G) (NOTE DATA PRESENTED FOR E6 ONLY). 7=CaCO3 ROOTS (G). 8=HEARTH STONES (#).
9=OTHER BURNT (HEAT-SHATTERED) ROCKS (G). 10=AREA EXCAVATED (M°) (NOTE DATA
PRESENTED FOR E6 ONLY). 11=MEAN THICKNESS OF XU (CM) (NOTE DATA NOT PRESENTED
FOR PERIPHERY SQUARES). 12=LAND SNAILS (#). 13=LAND SNAILS (G).

MEMOIRS

OF THE

(QUEENSLAND MUSEUM

BRISBANE

© Queensland Museum
PO Box 3300, South Brisbane 4101, Australia
Phone 06 7 3840 7555
Fax 06 7 3846 1226
Email qmlib@qm.qld.gov.au
Website www.qm.qld.gov.au

National Library of Australia card number
ISSN 0079-8835

NOTE
Papers published in this volume and in all previous volumes of the Afemoirs of the
Queensland Museum maybe teproduced for scientific research, individual study or other
educational purposes. Properly acknowledged quotations may be made but queries regarding
the republication of any papers should be addressed to the Editor in Chief. Copies of the
journal can be purchased from the Queensland Museum Shop.

A Guide to Authors is displayed at the Queensland Museum web site

A Queensland Government Project
Typeset at the Queensland Museum

REVISION OF SARMATIUM DANA (CRUSTACEA:BRACHYURA: SESARMINAE}
WITH DESCRIPTIONS OF THREE NEW SPECIES

P.1.F, DAVIE

Davie, P.J.F. 1992 06 29: Revision of Sarmatium Dana (Crustacea:Brachyura: Sesarminae)
with descriptions of three new species, Mempirs of the Queensland Museum 32(1): 79-97.

Brisbane. ISSN 0079-8835.

Sarmatium Dana, 1851, is revised and re-diagnosed, Crabs of this genus are primarily
characterised by; carapace deeply vaulted; ocular peduncle swollen basally, cornea constricted
and reduced; upper surface of palm of male cheliped bearing a series of transverse grooves
separating transverse swollen ridges. Sarmatium is now considered to contain five species: S.
crassum Dana, 1851; 8. germaini (A, Milne Edwards, 1869); 5. striaticarpus sp.nov.; S, hegerli
sp.nov,;and§, wridentatus sp.nov, They are easily separated by the different patterns of ridges
and grooves on the upper surfaces of the palm and carpus of the male cheliped. []Crustacea,
Brachyura, Grapsidae, Sesarminae, mangroves, Indo-West Pacific, new species.

Peter JF. Davie, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101,

Australia; 3 February, 1992.

Sarmatium Dana, 1851, has a very confused
history. Established for asingle species 8. crassum
Dana, 1851, it was enlarged considerably by
Tesch (1917) who included six other Indo-West
Pacific species and two Atlantic species, Seréne
& Soh (1970: 398) pointed out that the two Atlan-
lic species, S. curvatus H, Milne Edwards, 1837,
and S. pectinatus H. Milne Edwards, 1853, ure
rightfully placed into the genus Metagrapsus H.
Milne Edwards, 1853. Tesch (1917) had pre-
yiously considered Metagrapsus as a junior
synonym of Sarmatium. Features to separate the
two genera were given by Seréne & Soh (1970).
The six Indo-West Pacific species placed in Sar-
matium by Tesch: S. integrum A. Milne Edwards,
1873; S. inermis de Man, 1887; S. indicum A-
Milne Edwards, 1868; .S. punctatum A. Milne
Edwards, 1873; S. biroi Nobili, 1905; and S.
fryatti Tesch, 1917 (=rotundifrons A. Milne Ed-
wards, 1869), are now all included in Neosar-
maltium Serene & Soh, 1970. Tesch (1917),
following de Man (1891), considered Sarmarium
vermaini (A. Milne Edwards, 1869) to be a junior
synonym of §. crassum but S. germaini was later
shown to be a yalid species by Sertne & Soh
(1971).

Seréne & Soh (1971) considered Sarmatium to
consist of two species, S. crassumand S. germaini.
It is apparent however, after examining the larger
amount of material available for the present study,
that what Serene & Soh (1971) thought to be S.
crassum from Singapore is in fact a new species.
In addition, field work carned out m northern
Australia has revealed two hitherto unknowns
species. With the five species here recognised, the

genus concept for Sarmaiiam can now be much
better understood,

Abbreviations used in the text are; AM,
Australian Museum, Sydney; MNHN, Muséum
national d°Histoire naturelle, Paris: ZRC,
Zoological Reference Collection, National
Museum of Singapore; NNM, National Natural
History Museum, Leiden; QM, Queensland
Museum, Brisbane; SMP, Senckenberg Museum,
Frankfurt; USNM, United States National
Museum, Washington; ZMG, Zoological
Museum of Goetingen (collection housed at
SMF); ZMH, Zoological Museum of Hamburg;
ZMK, Zoological Museum Copenhagen. The
descriptions for this paper were prepared using the
DELTA computer system for generating
taxonomic descriptions (Dallwitz & Paine, 1986).

Measurements given in the text are of the
carapace breadth (measured at the widest point
including lateral spines) followed by length, Leg
segments were measured in a straight line to give
maximum dorsal length and so are not always the
maximum possible length, and this should be
borne in mind when using the ratios, The exact
limits of the width of the hind margin are also
sometimes difficult to determine and in this work
they were defined by the point at which the lateral
carapace suture mects the rear margin. Gonopod
terminology follows that of Sakai & Yatsuzuka
(1979),

Sarmatium Dana, 1851

Sarmentinet Dana, 18514:288; 185 )b; 25); 1852; 357-8;

RO MEMOIRS OF THE QUEENSLAND MUSEUM

Kingsley, 1880: 212; Seréne & Soh, 1970: 397;
1971: 237; Saka, 1976: 664.

Sarmatium (in part): De Man, 1887: 659; Alcock, 1900:
426; Tesch, 1917: 213, 258; 1918: 115; Crosnier,
1965; 74,

Sesarma (Sarmatium) (in part): De Man, 1929: 111;
Tweedie, 1936: 67.

TYPE SPECIES
Sarmatium crassum Dana, 1841.

DIAGNOSIS

Carapace sub-quadrate, slightly wider vhan
long; greatest width behind exorbital angles:
deeply vaulted, depth c,0,8- 0.9 X carapace width
in males; regions moderately defined:
anterolateral margins regularly convex; with 1-3
teeth behind the exorbital angle; front deflexed,
c.35- 0.4% carapace width, 0.5 x fronto-orbital
width, pre-orhital teeth obsolete. Ocular peduncle
swollen basally, cornea constricted and reduced.
Basal segment of antennal peduncle with a well
developed outer rounded tongue; antennal flagel-
lum small and entering orbit. Basal antennular
segment swollen. Inter-antennular septum nar-
row. 0.2- 0.3 * width of front, Median post-frontal
lobes distinct, very broad, lateral lobes nat clearly
differentiated. Third maxilliped merus 0-7-
0.8 X length of ischium; merus longer than wide;
outer margin convex; antero-external angle not
produced; exopod narrow, barely visible in frontal
view. Chelipeds subequal: upper surface of palm
with a series of transverse grooves separating
swollen ridges; outer surface of palm without
median longitudinal row; dorsal surface of dactyl
in males with at Jeast one large, broad, chitinous
tubercle, and some small pointed chitinous
tubercles. Legs: merus anterior margin with an
acute sub-distal spine, unarmed terminally; carpus
with accessory Carinae. on upper surface. G1 long,
reaching just past suture between sternites. 2/3;
moderately stout; completely calcified; palp ob-
vious but not separated from stem, rounded, cal-
cified: apical process corneous, strongly
produced, straight; gonopore terminal or slightly
displaced towards the dorsal surface. G2 short,
relatively narrow, tapeting, twisted, tip blunt.
Male abdomen relatively narrow, third segment
slightly the wider or sub-equal to first; first seg-
ment does not cover entire width of sternum
between 4th pereiopods. Base of penis slightly
calcified, just visible beneath setae adjacent to
margin of third segment,

REMARKS

Sarmatium is separated from the two closest
genera Neosarmatium and Metagrapsus by two
major characters; 1, ocular peduncle swollen
basally, comea constricted and reduced; 2, upper
surface of palm of male cheliped with a series of
transverse grooves separating swollen ndges.

KEY TO THE SPECIES OF SARMATIUM

1, Guter yentral border of cheliped palm with a
marked longitudinal ridge extending onta
fixed-finger (even on juveniles and females);
male cheliped with superior outer surface of
palm bearing 7-8 relatively evenly spaced
transverse grooves; dactylus with three large,
strang chitinous jubercles on proximal hall,
followed by 17-18 small acute chitinous
tubercles; carpus with a finely granular area
distally, without trace of oblique fine stria-
tions. Maximum size c.28mm..,
seulivnbep'gosentsde germaini A. Milne Edwards, 1869

Outer ventral border of cheliped palm and
fixed-finger without a longitudinal ridge; male
cheliped with superior outer surface of palm
bearing less than 7 strong transverse grooves;
daclylus vaniously armed, but not as above;
upper face of carpus sometimes with oblique
fine striations, Maximum size 28mm ....,......2

2. Palm of male cheliped with 4 major transverse
grooves on upper outer surface. Dactylus with
four strong chitinows tubercles on pam
half. Maximum $12¢¢.25MM....-.0: er eersaed

Palm of male cheliped with transverse grooves
not as conspicuous. but with a large, smooth
or granular swelling occupying most of upper
ouler surface. Dactylus with only one major
chitinous tubercle on proximal half, Maximum
Sie 1 S-Di sasssepeckassstianesboneid teas Tsouers dent 4

3. Upper surface of carpus of cheliped with large
patch of Uny, flattened, squamous granules
behind articulation distally; these may be
aligned into short fine rows near outeredge but
never appear as transverse striations. Ridges
and grooves on upper surface of palm all sub-
parallel. First proximal tooth on dactyl of
— ‘ances sii ere from articula-
tion.. revere ..crassum Dana, 1851

Upper surface of carpus of cheliped with broad
patch of c.25 long, fine, transverse striations

REVISION OF THE GENUS SARMATIUM 81

between articulation and spine of inner angle.
Upper surface of palm with proximal most
corrugated ridge separated from next broad
groove by a triangular space. First proximal
tooth on dactyl of cheliped placed almost on
edge of proximal end ....,..striaticarpus sp.nov.

4. Male cheliped has dorsal margin of dactyl, with
l large forwardly directed, chitinous tubercle
On outer edge, proximally. Superior outer face
of palm with 2-3 narrow, smooth grooyes dis-
tally, followed by a deep concavity, and a large
swelling back to the posterior margin, the dis~
tal slape of which is long and covered in well
separated, medium sized, low round granules,
Upper surface of carpus with 9 prominent
transverse ridges distolaterally, Maximom
size c.20mm............hegerli sp.nov.

Male cheliped has dorsal margin of dactyl bear-
inga single, large, elongate, slightly oblique,
medial tooth, proximally, Superior onter face
of palm wilh very broad, deep, smooth, obli-
que depression, a very large rounded medial
transverse swelling, and a narrow proximal
band of seyen very fine, long transverse
tidges. Lipper surface of carpus without
transverse ridges. Maximum size c.l13mm
cssssqeeeneeseerses AM GenTATS SP.NOV.

Sarmatium crassum Dana, 1851
(Figs 1A, 2, 3A-C)

Sarmatium.crassum Dana, 185 lb: 251; 1852: 358-9, pl.
23, fig. 1; H. Milne Edwards, 1853: 189; De Man,
1887: 651; Bamard, 1955: 28, fig.9; Crosnier, 1965:
74-6, figs 121-124, pl. 5, fig. 1; McNeill, 1968: 78.
- Serene & Soh, 1970: 397, 405 (name in list);
Fishelson, 1971; 128, 130; Serene, 1973: 125-6;
Hartnoll, 1975; 308, 318, 322-23; Holthuis, 1977:
174-5.

nat Sarmatium crassunt Seréne & Soh, 1970, pl. 4C,
D; 1971: 237-40, fig. 2, pl. 2 [= 5. striaticarpus sp.
nov.)

? Sarmatium crassum: Nobili, 1899: 505; Alcock, 1900:
426-27, Tesch, 1917: 215,

TYPE SPECIMEN
Lost.

TYPE LOCALITY
Upolu, Samoa.

MATERIAL EXAMINED _
NNM 24830, 1d (18.8%16.5mm), Melita Bay, En-

tedebiz, Dahlak Arch., L, Rode lee, Eritrea, 14.4.1962.
J.S.R.8. Expedition. MP-B10475, Id
(13.2% 12.0mm), Nosy Be, Madagascar, mangroves.
A. Crosnier. MP-B10474, 1d (12.8 11.5mm), Route
de Ducas, 4km de Noumea, P. Fourmanoi, 7.8.1971-
QM Wi6819. 1d (8.0X7.1mm) 1 imm, 2
(8.7%7.4mm), Harmer Ck, far N Queensland ,
11°50'S, 142°57°E, mouth of river in mudbank at low
water, P. Davie & J. Short, 31.10.1990. QM W16829_
1? (18.5% 15.7mm}, same data as OM WLES19, OM
W6791, 20 d (14.8 13.2; 21.1% 19.3mm), Curtis 1.
ar Tide 1, MEQ, 23°46'S, IS1°15'E, 17.Dec, 1975, P,
Saenger, mud in Riuzephora. QM W6804, 1d
(20.8% 180mm), 2m from mouth of Auckland Ck, nr
Gladstone, MEQ, 23°51"S, 151°14’E, 17.12.1975, P.
Saenger, mud in Avicennic. QM W15228, 82d
(21.5192; 21.0 19.7; 19.5% 17.3; 17.0% 14.7;
17.) % 15.2; 17,2 X15,2; 13.9% 12.5; 11.8 10.4mm),
4992 (24,220.5; 21.4%18,2: 18.0% 15.2;
11,.9%10.0mm). Cailiope R. and Auckland Ck,
Gladstone, MEQ, 23°51°S, 151°16@°E, Queensland
Electricity Commision Sarvey, 1974-1983, P. Saenger.
QM W5387. 12 (15,6134mm), Mary and Susan
River Heads, Hervey Bay, SEQ, P. Davie, 25.7.1975.
QM W5357,1¢0 (12.6 10.6mm), Moon Ck, Fraser 1,
SEQ. 25°1L'S, 153°04'E, 21.7.1975, P. Davie, R. Tim-
mins. QM W5382, 1d (19.1% 17.3mm), Eli Ck, Her-
vey Bay, SEQ, 25°17°S, 152°49°E, 26,7.1975, P.
Davie. QM W5351, 12 (17.6% 15.1mm), Pulgul Ck,
S of Urangan, Hervey Bay. SEQ, 25°19'S, 152°54°E,
19,7.1975, P, Davie- QM W5287. 12
(24,3 %20.3mm), small branch of Serpentine Ck, SEQ,
27°24'S, 153°07°E, Sept 1972, B, Campbell et al.
NNM Unreg., 1¢ (13.9 12.lmm), Snellius Exped..
16,9,1930, Bongao, Tanitawi, Sulu Arch. ZMK K4410,
12 (23.6% 19.4mm), Tahiti.

DESCRIPTION

Carapace: c.1.1 X broader than long in males,
1.2 in females, Fronto-orbital width c.0.75-
0.85 X carapace length. Carapace deeply vaulted:
convex in both directions but only slightly from
side to side. Depth c.0.8 * carapace width (c.0.7
in females). Regions moderately defined;
mesogastric well defined: cardiac distinct; intes-
tinal distinct. Lateral margins subparallel; slight-
ly concave. Anterolateral margins regularly
convex; with two blunt teeth behind the exorbital
angle: exorbital angle blunt, only slightly project-
ing. First anterolateral tooth similar in size to
exorbital angle but with longer margin; second
anterolateral tooth blunt; minute, almost obsolete.
Front ¢.0.36-0.39% carapace width; c..5X
fronto-orbital width: moderately deflexed; with
shallow median emargination and bilobed; lateral

82 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. A, Sarmatium crassum Dana, upper face of palm ofd chela (QM W15228). B, S. striaticarpus sp.nov.,
upper face of palm of 3 chela (holotype, ZRC 1970,1,23,14). C,D, S. hegerli sp.nov., upper and outer faces of d
chela (holotype, QM W9698). E, 5. unideniatus sp.nov., outer face of male chela (holotype, AM P31822),

angles obtuse, blunt; pre-orbital teeth obsolete;
lateral margins slightly diverging posteriorly.
Post-frontal lobes distinct; median lobes distinct-
ly broader than laterals; lateral lobes not clearly
differentiated. Branchial ridges prominent form-
ing a series of short broken granular striations;
last continuous, parallel with posterolateral mar-
gin, and finishing over coxa of fourth walking leg.
Posterior margin c.0.45-0.5 X carapace width.
Carapace surface smooth, shining, punctate; setae
arranged sparsely on branchial lines. Upper orbi-
tal border minutely granular; straight; inner angle
rounded. Lower orbital border straight; evenly
granular. Inner orbital tooth present; well
developed; equilateral triangular, ocular peduncle
swollen basally, comea constricted and reduced.

Basal segment of antennal peduncle with a well
developed outer lobe in form of rounded tongue.
Inter-antennular septum narrow; 0.22-
0.24 width of front,

Third maxilliped: Merus 0.7-0.8 X length of is-
chium. Suture between merus and ischium slight-
ly sloping inward. Ischium inner margin
microscopically granular. Palp articulates medial-
ly on distal margin of merus. Exopod narrow,
barely visible in frontal view; flagellum normal.

Chelipeds: Subequal; large and robust. Merus
posterior border with minutely granular stria-
tions; distinct subdistal spine; Jower border
granulate; anterior border convex, coarsely
granulate; carpus with a small spine at inner angle;
inner margin unarmed, striated, rounded, ventral-

REVISION OF THE GENUS SARMATIUM 83

FIG. 2. Sarmatium crassum, 6 (QM W15228). A, dorsal view. B, chela. C, upper surface of palm of chela. Scale

line in mm.
ly with row of small granules and short proximal
oblique crest bearing row of long setae; granules
present on inner face of carpus just below inner
angle; outer margin striated, with narrow patch of
sparse, very short setae; upper surface micros-
copically striated proximally, smooth and
wrinkled medially, behind articulation distally a
large patch of tiny, flattened, squamous granules
which become aligned into fine rows near outer
edge. Palm upper surface with a series of
transverse grooves separating swollen ridges, dis-
tal margin of upper surface of palm bearing row
of 10-13 pectinated teeth along edge of articula-

tion, proximal to this is a series of 6 broad, swol-
len, rounded ridges, relatively evenly separated,
ridges 3/4 and 5/6 joined by low rim top and
bottom; fourth ridge broader than others; fifth
ridge low and with row of 15-20 granules along
its lower distal edge; superior margin with a series
of moderately large granules, but without con-
spicuous setation. Outer surface of palm smooth
and punctate; without median longitudinal row,
without a ventral ridge. Outer surface of palm
naked, small patch of setae at base of fingers,
fringe of setae at insertion of dactyl. Inner surface
of palm with sparse large flat smooth granules,

84 MEMOIRS OF THE QUEENSLAND MUSEUM

lmm

0.5 mm

FIG. 3. Male first gonopods (setae removed). Sarmatium crassum Dana. A, abdominal view. B, C, lateral views
of apex. S. striaticarpus sp.nov. D, abdominal view. E, sternal view. 5. hegerli sp.nov. F, abdominal view. G,

H, lateral views of apex.
largest and most conspicuous near gape, but not
forming a strong vertical crest. Immovable finger
slightly flattened on outer surface; moderately
long. Length cutting edge c.0.45 length
propodus in males (0.5 in females). Ventral bor-
der of chela straight, or slightly convex. Dorsal
surface of dactyl of males bearing 4 large, broad,
chitinous tubercles in proximal three-fifths, first
set away from joint about same distance as to
following tooth, a small patch of setae in the
space; second tooth larger than first; third and
fourth teeth largest, similar sized; teeth 1-4 set
increasingly obliquely, highest towards outer
face; followed in distal quarter by a row of 12-14
small closely set chitinous tubercles of even size.

Fingers pointed; curved slightly inwards; a
moderate gape between cutting margins in males.

Walking legs: Medium length; compressed and
flattened; second pair slightly the longest, c.1.5-
1.7% maximum carapace width. Third leg: merus
c.2,.4-2.6 % as long as wide; carpus c.2.6-2.8 X as
long as wide; propodus ¢.2.3- 2.6%as long as
wide. Dactyli about equal to length of propodi;
slender and almost straight; terminating in acute
chitinous tips. Propodus with an accessory carina
on inferior proximal portion of upper surface.
Meni of legs 1-3 with scattering of small distally
directed prickles; dorsal and ventral borders of
meri granulate. Setae short, predominantly on
upper distal face of carpi, and on upper half of
propodi, and continuing onto dactyli in thin rows.

REVISION OF THE GENUS SARMATIUM 85

Male abdomen: Relatively narrow; third seg~-
ment slightly the widest. Slightly calcified base
of penis adjacent to margin of third segment.
Segments three-five slightly tapering. Width seg-
ment three 4, 1-4.7 X length (relatively narrower tn
smaller males). Segment six not elongated; 1.45-
1.6 wider than long. Telson nearly subequal in
Jength to segment 6; 1.25-1.5 ¥ longer than wide
(relatively longer in large males); evenly
rounded.

Gonopeds: G) moderately stout; slightly
curved. Inner-dorsal margin distally curved in-
ward, Dorsal surface of stem flattened; complete-
ly calcified. Palp poorly developed, not separated
from stem, large, broad, rounded, calcified. Outer
dorsal margin of stem convex. Distal part of the
stem broad but narrowing. Apical process
present; corneous; strongly produced; straight.
Gonopore slightly displaced towards the dorsal
surface. Setae long, simple, dense, obscuring
structural detail. G2 short, relatively narrow,
lapering. twisted, tip blunt,

COLOURATION

Fawn-brown, chelipeds and legs biscuit-
coloured with orange tinge (Bamard, 1955);
carapace purple in life with yellow patches,
thoracic legs yellow with purple patches (Cros-
niet, 1965).

DISTRIBUTION

With certainty from only the following
localities: Samoa (Dana, 1851b, 1852); Durban
Bay, South Africa (Barnard, 1955), Madagascar
{Crosnier, 1965); Dar es Salaum, Tanzania
(Martnoll, 1975); Eritrea, Red Sea (Fishelson,
1971; Holthuis, 1977): Sulu Archipelago, In-
donesia (present record); eastern Queensland,
from Brisbane north to 11 50'S on Cape York
(McNeill, 1968; and present records); New
Caledonia (Seréne, 1973); Tahiti (present record).

Questionably from; Sumatra (Nobili, 1906 - no
diagnostic characters were given, so may also be
atributable to 8S. striaticarpus sp.nov.); Nicobars
(Alcock, 1900. single? specimen); ‘Pacific®
(Tesch, 1917, single? specimen).

HABITAT

In the mangroves (Crosnier, 1965; McNeill,
1968). “An uncommon species in creek
mangrove, where specimens were collected from
the Ceriops and mixed zones” (Harinoll, 1975).
Mangroves; mouth of river in mudbank at low
water, mud in RAlzophora, mud in Avicennia
(present records),

REMARKS

Seréne & Soh (1970) presented data and figures
scparating what they believed to be Sarmatium
cressum from §. germaini. Unfortunately their S.
crastum specimens represent a new species very
closely allied to S. crassum, but differing in a
number of distinctive characters (see later). The
accurate identity of S. crasswm is slightly
problematic as. being a Dana species, the type
was destroyed in the Chicago fire of 1871. This
means that itis difficult to be absolutely certain
what is the true §. crassum. After considering
Dana's description, | consider the Asian
specimens previously attributed to §. erasswn to
be the new species, and it is here named §,
striaticarpus. Sarmatium crassum differs from §.
striaticarpus on the following grounds:

1. The shape and disposition of the ridges and.
grooves on the upper surface of the palm are
different. In §. crasmwn the ridges are all sub-
parallel] and this is evident in Dana’s Fig. 1d;
whereas in 5, striaticarpus the proximal mest
corrugated ridge is separated from the next broad
groove by a triangular space, Also there is a slight
groove separating the distal row of pectinatious
teeth in Dana’s figure which is also true of my S$.
crassum material but not present in S. striaticar-
pus,

2. The first proximal tooth en the dacty! of the
cheliped is placed slightly distally from the ar-
ticulation in §. cresswm and this also seems true
of Dana’s figure although it is not clear; in S.
sirjaticarpus it is placed almost on the very edge
of the proximal end.

3. So far, 8. strianearpus is only known for
certain from vicinities north of the equator. To
remove all doubt over the identity of §..crassum
the erection of a neotype would be necessary. |
am reluctant to do this because I have not sees
specimens from the type-locality (Samoa), Al-
though the Asian specimens certamly represent a
new species (5S. striaticarpus), slight reservations
must remain about the identity of the specimens
here referred to S. crassum. The shape of the male
abdomen in Dana’s figure differs a little from that
of the present material, and it is difficult to ascer-
tain if this difference is due to prowth changes
(Dana‘s spectmen was quite small), inaccuracies
in the figure, or if it does in fact represent a real
difference at the species Jevel. In this last case a
new name would be required for the material here
identified as S, crassum.

Sannalium striaticarpus sp.nov. can further be
separated from §. crassum on the following
grounds: the small chitinous tubercles on the dis-

86 MEMOIRS OF THE QUEENSLAND MUSEUM

tal half of the dactyl of the cheliped are greater in
nurnber and more closely spaced in §. crassum; S.
Striaticarpus has a broad patch of c.25 long, fine,
(ransverse striations, mostly continuous but some
subdivided, on the upper surface of the cheliped
carpus between the articulation and the spine of
the inner angle, in 5. crassum the upper surtace is
microscopically stated proximally, smooth and
wrinkled medially, and with a large patch of tiny,
flattened, squamous granules behind the distal
joint, which become aligned into fine rows near
outer edge.

Sarmatium striaticarpus sp.nov.
(Figs 1B, 3D,E, 4)

Sesarma (Sarmatium) crassum: Tweedie, 1996: 67-8.

Sarmatium crassum: Seréne & Soh, 1970, pl. 4C, Dy
1971; 237-40, fig. 2, pl. 2; Sakai, 1976: 664-5.

? Sarmatium sp. (atf. S, crassum Dana): Sakai, 1936:
173, text-fig. 7.

MATERIAL EXAMINED

Hovortyre: ZRC 1970.1.23.14, 1d (24.3% 23.0mum),
Johore Straits, C.L. Soh, 21.1,1970.

Pararypes: USNM 73213, 12 (16.2% 14,0mm),
lloilo, Panay I., Philippines, H.C. Kellers, U.S, Navy,
April 1929, Ho Ilo Eclipse Exped.

DESCRIPTION

Carapace: c.J,1* broader than long. Fronto-
orbital width c.0.75 x carapace length. Carapace
deeply vaulted; evenly convex bongitudinally, flat
from side to side, Depih c,0.8-0.9X carapace
width. Regions well defined; mesogastric well
defined: cardiac distinct; intestinal indistinct,
defined laterally by branchio-intestinal grooves.
Posterolateral margins divergent posteriorly;
moderately concave, Anterolateral margins
regularly convex; cristate; with three blunt, for-
wardly directed teeth behind the exorbital angle
{third reduced to trace only). Front
¢.0.36X carapace width; 0.5% fronto-orbital
width; moderately deflexed; witha broad median
emargination; lateral angles obtuse; pre-orbital
teeth obsolete; lateral margins slightly diverging
posteriorly. Medial post-frontal lobes very broad,
well defined, laterals almost obsolete; without
clumps of setae. No short ridge on first
epibranchial tooth. Branchial ridges not
prominent; relatively short. Posterior margin
¢.0.5 X carapace width, Carapace surface smooih,
shining, punctate; setae arranged sparsely on
branchial lines. Upper orbital border evenly
granular; straight and slightly oblique; inner angle

rounded. Lower orbital border straight; smooth,
Inner orbital tooth present; well developed; equi-
lateral triangular. Antennal flagellum very small,
entering orbit. Orbital hiatus open, Basal segment
of antennal peciincle with an elongate, rounded
outer lobe. Basal antennular segment moderately
swollen, Inter- antennular septum c.0.23 x width
of front.

Third maxilliped: Menus ¢.0.7 X length of is-
chium. Suture between metus and ischium slight
ly sloping inward, Ischium sub-triangular: inner
margin smooth. Palp articulates near outer distal
margin of merus. Exoped narrow, barely visible
in frontal view; c.0.3 X width of ischium; flagel-
lum normal. :

Chelipeds: Subequal; large and robust. Merus of
male without stridulatory ridge; posterior border
minutely granulate; distinct subdistal spine;
lower border granulate; anterior border coarsely
granulate medially. Carpus with a spine at inner
angle; inner margin with narmow granular bead;
granules present on inner face of carpus just
below inner angle; outer margin with a series of
smooth granular striations: between articulation
and spine of inner angle is a broad patch of ¢.25
long, fine, transverse striations, mostly con-
tinuous but some subdivided. Palm upper surface
with a series of transverse grooyes scparating
swollen ridges: a distally swollen margin bearing
row of 10-12 pectinated teeth along edge of ar-
liculation, largest near superior margin. minute
distally; behind this a wide groove; then a ridge,
itself subdivided by a narrow groove into a very
narrow distal ridge and a wide proximal ndge;
behind this a sedes of two very wide swollen
ridges separated by a broad concavity, but both
ridges connected by a narrow margin top and
bottom: behind this a triangular depression with
its apex dorsally, and then a pair of short, close-
ly-set comb-like ridges. Outer surface of palm
smooth and punctate; with slight depression at
base of fixed finger; without ventral ridge. Outer
surface of palm naked; with small patch of setae
at base of fingers just finging articulation. Inner
surface of palm with a low granular vertical crest
of 5-6 granules largest ventrally: and an oblique
crest of about 6 large granules running from
below vertical crest to upper part of fixed finger,
Immovable finger slightly flattened on outer sur-
face: moderately long; length cutting edge
c,0.45 X Jength propodus, Ventral border of chela
slightly convex. Dorsal surface of male dactyl
with 15-17 chitinous tubercles; 4 large, widely
spaced tubercles, set obliquely, on the proximal
half, becoming longer but no higher distally; in

REVISION OF THE GENUS SARMATIUM 87

FIG. 4. Sarmatium striaticarpus sp.nov., 3 (holotype, ZRC 1970.1.23.14). A, dorsal view. B, chela. C, upper

surface of palm of chela. Scale line in mm.

distal half a row of 12 small closely set sharp
tubercles, forwardly pointed, becoming minute
distally; proximal portion of inner edge without
distinct granules. Fingers pointed; curved in-
wards; a moderate gape between cutting margins.

Walking legs: Medium length; compressed;
slender; first three pairs all of similar length but
third pair slightly longest (c.1.7 X maximum
carapace width). Third leg: merus c.2.8 Xas long
as wide; carpus c.2.6 X as long as wide; propodus
c.2.6 X as long as wide; dactylus c.0.8 X length of
propodus. Dactyli slender and slightly recurved;
terminating in acute chitinous tips. Propodus with
an accessory carina on inferior proximal portion

of upper surface. Leg segments smooth, anterior
margins minutely granular; fringed with short,
soft, setae, also arranged in thin tufts in rows on
anterior and posterior faces.

Male abdomen: Relatively narrow; third seg-
ment widest. First segment broad, only slightly
narrower than third segment. Segments 3-5
slightly tapering. Width segment three
c.4.3 Xlength. Telson longer than preceding seg-
ments, only slightly longer than sixth segment;
c.1.4Xlonger than wide; evenly rounded.

Genopods: G1 moderately stout; slightly
curved. Inner-dorsal margin evenly curved onto
palp. Dorsal surface of stem flattened; completely

88 MEMOIRS OF THE QUEENSLAND MUSEUM

calcified. Palp well developed, separated from
stem, large, narrow. rounded, calcified. Outer
dorsal margin of stem convex. Distal part of stem
broad but narrowing. Apical process present; cor-
neous; strongly produced; straight. Gonopore dis-
placed towards the dorsal surface. Setae long,
simple, lying around cormeous tip and apical part
of stem obscuring structural detail. G2 short,
evenly tapering, moderately twisted, apically
pointed.

COLOURATION
Palm of the cheliped brownish red (Seréne &
Soh, 1971}.

DISTRIBUTION

Singapore (Tweedie, 1936; Seréne & Soh, 1971;
present records); Southern Okinawa, Japan
(Sakai, 1936, 1976); Philippines (present record).
Relatively common in the mangroves of Sin-
gapore and Malaysia (Seréne, 1973, referring to
this species as 3. crassurn).

HABITAT
Mangrove swamp (Sertne, 1973; Sakai, 1976;
present records).

ETYMOLOGY

Named in reference to the charactenstic stria-
tions on the upper surface of the carpus of the
cheliped,

See ‘Remarks’ for &. crasstnz.

Sarmatium germaini (A, Milne Edwards, 1869)
(Figs 5, 6A-C)

Sesarma germant A, Milne Edwards, 1869: 28; De
Man, 1887: 651; 1891: 51,

Sarmatinum germaini: Seréne & Soh, 1970: 397, 405;
1971: 238, fig. 1, pL. t (1-4); Soh, L978: 11, pl. da.

MATERIAL EXAMINED

LECTOTYPE. MP-B3668 (Dry), 12 (15.0 16.6mm),
Poulo Condore, South China Sea, coll. M. Germain.
PaRALECTOTYPES, MP-B10472 (Dry), id
(13.3% 12.4mm), Poulo Condore, South China Sea,
coll, M. Germain, (On the posterior portion of the
carapace is written ‘428° and below this ‘GS° in faded
black ink - specimen is dry, carapace intact but Jeft
cheliped, various legs and abdomen are detached.)

MP-B10473, 1d, same collection data as BIO472.
badly broken.
Orner Materia. ZRC 1970.2,20.3. I2

(27.8 25.6mm}, Johore Straits, Singapore, C.L. Soh,
Feb. 1970. MP-B10471. 1d (26.0% 244mm), Port
Swettenham, Malaysia, coll. Saseknmar, 22,10,1968.
MP-B21576, 1 (19,9 17.4mm), Serangoon R., Sin-
fapore, mangroves, 28.8.1965, MP-B21574, 1d
(14,.2%13.1mm)12 (16.9% 15.3mm), Johore Baharu,
1°23'N, 103°44°E, mangroves, 10.7.1965, QM
W14858, 1d (27.9 26.8mm), Mandai Swamps, Sin-
papore, [°25°N, 103°45°E, 1985, D.H. Murphy. ZMG
191a, 222% (16.7, 21.8) 23d¢d (20.4, 219mm),
Mariveles, Aibuleit, Bohol, Philippines, coll. Semper,
1876. NTM Cr.003558, | juv. ¢ (7.8 7.2mm), Creek
‘H', East Arm, Darwin, N.T., mangrove lined creck,
mean low water, R. Hanley, 288.1985. NTM
Cr.003555, 1d (11.6X%10.7mm), same data as
Cr,003558. NTM Cr.003714, 1¢ (13,7*124mm),
Creek “H’, East Arm, Darwin, N.T., mangrove creek,
4m, R. Hanley, 31.10.1984. NTM Cr.003718, 19
(17.5 *16,lmm), same data as Cr.003714, NTM
Cr.001693, 1d (16.5X14.7mm), 12°34.2°S
130°56.3"E, Darwin, N.T., in mangroves, low water
spring, R. Hanley, 17.5.1984. AM P31821, 19
(13... 118mm), Andranangoo Ck, Melville 1, 1.5km
upstream, west bank, Rhizophora mudbank, D. Grace,
21.6.1975. SMF Unreg., 39 2 (21,7-25.8), Smith Ck,
Caims, Mangroves, M. Skinner, 5.61980. SMF Unreg.,
1¢ (27.1 X24.4mm), ar Australian Institute of Marine
Science, Cape Ferguson, NEQ, M. Tiirkay, 11,6,1980.
QM W12334, 3dd (21,6X20.0; 22,6%21,2;
22.2%20.6mm), 12 (21,819.4mm), Townsville,
NEQ, 19°16'S, 146°49°E,QM W8212, 12 (17.4mm),
Murray R., NEQ, 18°017S, 145°53°B, 23.5.1978,
P.Davie, at burrow entrance at night Om inland from
bank. QM W12333, 1d (15.6mm), Cardwell, NEQ,
18°16'S, 146°01°E, K.Broadbent, QM WL1100, Id
(16.Smm), Orpheus T., 18°4°S, 146°30' BE, in mangrove
area. QM W17485,5d cd (10.0-19.5mm), 32 2 (9,1-
15.2mm), Calliope R. and Auckland Ck, Gladstone.
MEQ, 23°51°S, 151°16°E, Queensland Electricity
Commision Survey. 1974-1983, P. Saenger. QM
W17486, 12 (12.2%11.0mm), smail branch of Ser
pentine Ck, SEQ, 27°24°S, 153°07’, Sept 1972, B.
Campbell et al.

DESCRIPTION

Carapace: ¢.1.1™ broader than long. Fronto-
orbital width c.0.8X carapace length. Carapace
deeply vaulted; evenly convex longitudinally, flat
from side to side. Depth c.0.9 x carapace width.
Regions moderately defined; mesogastric well
defined; cardiac distinct; intestinal indistinct,
defined laterally by branchio-intestinal grooves.
Lateral margins slighily divergent posteriorly;
slightly concave. Anterolateral margins regularly
convex; usually with two blunt, forwardly

REVISION OF THE GENUS SARMATIUM 89

FIG. 5. Sarmatium germaini (A. Milne Edwards), d (QM W14858). A, dorsal view. B, chela. C, upper surface

of palm of chela. Scale line in mm.

directed teeth behind exorbital angle, a third tooth
sometimes noticeable. Front c.0.4 X carapace
width; 0.5 Xfronto-orbital width; moderately
deflexed; with broad median concavity; lateral
angles obtuse; pre-orbital teeth obsolete; lateral
margins slightly diverging posteriorly. Post- fron-
tal lobes medially well defined, laterally indis-
tinct; without clumps of setae. Branchial ridges
moderately prominent; relatively short. Posterior
margin c.0.5 X carapace width. Carapace surface
smooth, shining; wrinkled and punctate poster-
iorly; setae arranged sparsely on branchial lines.
Upper orbital border evenly granular; with slight

median convexity; inner angle rounded. Lower
orbital border straight; evenly granular. Inner or-
bital tooth present; well developed; equilateral
triangular. Basal segment of antennal peduncle
with broad and rounded outer lobe. Inter-anten-
nular septum narrow; 0.25-0.3 X width of front.

Third maxilliped: Merus c.0.8 X length of is-
chium. Suture between merus and ischium
horizontal. Ischium sub- triangular; inner margin
smooth. Palp articulates near outer distal margin
of merus. Exopod narrow; 0.4 width of is-
chium; flagellum normal.

Chelipeds: Subequal; large and robust; merus

90 MEMOIRS OF THE QUEENSLAND MUSEUM

~Sss

SSS SSS:

——
ES

———

Imm

Ll mm

0.5 mm
0.5 mm

FIG. 6, Male first gonopods (setae removed, except for D). Sarmatium germaini (A. Milne Edwards) (QM
W14858). A, abdominal view. B, sternal view. C, lateral view of apex. S. unidentatus sp.nov. (holotype, AM
P31819). D, abdominal view. E, sternal view. F, lateral view of apex.

with posterior border minutely granulate; without
distinct subdistal spine; lower border granulate;
anterior border sparsely granulate, slightly coar-
ser medially; carpus with a broad tooth at inner
angle; tubercles present on inner face of carpus
just below inner angle, 1 or 2 ventrally on vertical
crest; outer margin smooth, wrinkled, upper
medial surface behind articulation with a broad
patch of small flattened vesicular tubercles. Palm
upper surface with a series of transverse grooves
separating swollen ridges: upper distal margin
behind articulation with a row of 14-17 blunt
chitinous tubercles, slightly larger towards inner

end; behind this a series of 8 transverse swollen
ridges, first 7 of similar shape evenly spaced, third
to seventh subequal in width, slightly narrower
than first and second; behind seventh, a much
broader sulcus separating a much broader,
proximal transverse swelling; ridges 3/4, 5/6, 7/8
narrowly connected top and bottom, more-or- less
obviously; ridges 3,5, and 7 with a line of puncta-
tions on distal edge, deeper on 5, very deep on 7
so as to form a series of gutters with a prominent
granule at the base of each gutter; behind
transverse ridges superior border with large
rounded granules. Outer surface of palm smooth

REVISION OF THE GENUS SARMATIUM 91

and punctate; without median longitudinal row.
Ventral ridge extends to tip of fixed finger, less
distinct distally, very prominent behind fixed
finger, extends less than halfway towards
proximal end. Outer surface of palm naked except
for fringe of short setae around dactyl articula-
tion; with a patch of ‘fur’ on supenor margin
behind transverse ridges. Inner surface of palm
smooth; without a vertical crest. Immovable
finger slightly flattened on outer surface;
moderately long. Length cutting edge c.0.4-
0,5 Xlength propodus, Ventral border of chela
convex, straight in small specimens. Dorsal sur-
face of dactyl bearing 3 large chitinous tubercles
in proximal half, set obliquely; followed immedi-
ately by 17-18 small acute chitinous tubercles.
Fingers pointed; slightly curved inwards; a wide
gape between cutting margins.

Walking legs: Medium length; cylindrical,
slender; second pair slightly the longest,
c.1.7 maximum carapace width. Third leg:
merus ¢.2.3-2.5* as long as wide; carpus c.2.6-
2.7 as long as wide; propodus c.2.2- 2.3 Xas
long as wide; dactylus c.0.95Xlength of
propodus. Dactyli slender and slightly recurved:
terminating in acute chitinous tips. Propodus
without short accessory carina on inferior
proximal portion of upper surface. Meri of legs
1-3 with a scattering of small sharp prickles on
outer surface; leg segments fringed with short
setae, setae arranged in thin tufts tn rows on
anterior and postenor faces,

Male abdomen: Relatively narrow; third seg-
ment the widest. First segment broad, only slight-
ly narrower than third segment. Segments
three-five slightly tapering, each with slightly
concave margins. Width segment three
¢.4.3length. Telson length subequal to sixth
segment; c.l.4Xlonger than wide; evenly
rounded_

Gonopods. G1 moderately stout; curved. Inner-
dorsal margin evenly curved onto palp. Dorsal
surface of stem flattened; completely calcified.
Palp poorly developed, not separated from stem,
large, narrow, rounded, calcified, Outer dorsal
margin of stem convex. Distal part of the stem
narrow. Apical ess present; corneous; strang-
ly produced; straight. Gonopore terminal. Setae
long, simple, lying around comeous tip and apical
part of stem obscuring structural detail. G2 short,
evenly tapering. moderately twisted, apically
pointed,

COLOURATION
Greenish brown with reddish tinge on legs (A.

Milne Edwards, 1869); palm of the chelipec yel-
lowish light brown (Seréne & Soh, 1971),

DISTRIBUTION

Poulo Condore Island, Vietnam, South China
Sea (type locality, A. Milne Edwards, 1869);
Singapore and Malaysia (Seréne & Soh, 1971,
and present record); Hong Kong (Soh, 1978);
Philippines (present record); northern Australia -
from Brisbane to Darwin (present records),

HABITAT

In the mangroves and nearly always out of the
water (A. Milne Edwards, 1869), Mangrove lined
creek at mean Jow water, Rhizophora mudbank;
at burrow entrance at night 10m inland from creek
bank (present records),

REMARKS

A, Milne Edwards (1869) did not specifically
designate a holotype but he gives the measure-
ments for only a single specimen of 17 16mm,
and does not mention its sex. From his descnption
of the ‘six ou sept trés-petits bourrelets obliques
et paralléltes’ on the chela, one might think that he
examined a male, but these ridges are also
present, albeit indistinctly, on the female, His
note on habitat - that this species was collected
almost always out of the water - does suggest that
he was given more than one specimen to examine.
De Man (1891) says, ‘I was enabled to study a
typical male specimen of Ses. Germani of the
Paris Museum’, but he does not give measure-
ments. Serene & Soh (1971) state, ‘The senior
author ... was able to re-examine the holotype of
S. germaini in the Paris Museum (April 1970); it
isa dry female and not a male as indicated by De:
Man (1891).’ The decision is taken here to desig-
nate the female as the lectotype - its measure-
ments do not precisely conform to those given by
A. Milne Edwards, but they are very near. There
are also two dry male specimens at the Paris
Museum, MP-B10472-3, and one of these was
presumably the specimen examined by De Man.
Arelatively modem label identifies them as being
collected by M. Germain from Poulo Condor, but
no original labelling has been preserved. The
pera {one is lithe more than a series of

gments) are indeed S$. germaini as defined
here. and by Seréne & Soh (1971), however they
are both much smaller than the measurements
given by A. Milne Edwards. They are considered,
along with the female discussed above, to have
been syntypes, and are thus designated here as

paralectotypes,

92 MEMOIRS OF THE QUEENSLAND MUSEUM

The spelling of the species name wus justifiably
emended by Seréne & Soh (1971) as the species
was clearly intended to be named after its collec-
tor, R. Germain, and can thus be considered an
“incorrect original spelling’ under Article 32(c) of
the Intemational Code of Zoological Nomencla-
ture (1989, Third Edition).

Sarmatiom hegerli sp.nov,
(Figs 1C.D. 3F-H, 7)

MATERIAL EXAMINED

HoLoryre: OM W698, 1d (19.6% 174mm), Base
Alligator R,, Kakadu National Park, N.T, burrow in
moist bank of drainage channel (K836), P. Davie, May
1980.

PaRATYres: QM W9699, 1d (9.98. 7mm), East Al-
ligator R,, Kakadu N.P., N.T.. edge of drainage channe)
in Ceriops/Avicennia sland with a duck Aegilaitiy un-
derstory, (T6Q2) P. Davie, 19.6,1982, OM W9700, 1g
(12.4% 10.8mm)}, East Alligator R., Kakadu N.P,.N,T,,
open Sonneratia forest, very soft sloppy mud, (T6Q4)
P. Davie 19.6.1982.QM W970), 1 juv. (6.66. lm),
East Alligator R, Mouth, Kakadu N.P., N.T., low open
AvicennialAegilains scrub, moist but finn substrate,
some succulent salimarsh plants, crab from shaded
raised burrow entrance very wear surface, (T5Q1), P
Davie, 16.6.1982. OM W9702, 12 (10,8 x 92mm},
East Alligator R,, Kakadu N.P., N.T., open Senneratia
forest, very soft sloppy mud {T6Q5), P, Davie,
19.6.1982, OM W9703, 492 (124% 10.4; 12.1*9.6
(bopynid infection); 10.9*9.0; 10.1% 8.5mm) Lt
(13.6 11,9mm), East Alligator R., Kakadu N.P., N.T-
(Site 27), P. Davie, May 1980. QM W9704, Id (73.2
*%12.0mm), Middle Ann, Darwin, N_T., in burrow
under rock rear boat ramp, firm mud, tidally inundated,
salinity 32 ppt at low water, P. Davic, 29.6.1982, NTM
Cr,001826, 1d (15.8 X 14.3rm), locality not supplied.
NTM Cr.003068 1 2 (24.6% 118mm), Creek 'H* East
Arm, Darwin Harbour, 12°33,2°S, 130°563°E.
mangroves, mean low water, R, Hienley, 1,7.1985

DESCRIPTION

Carapace: c.1.1 broader than long in males,
c.1.2 in females. Fronto-orbital width
¢.0.8Xcarapace length (0.9 in females).
Carapace deeply vaulted; convex antenorly, al
most flat from side to side. Depth c.0.8 X carapace
width. Regions moderately defined; mesogastric
well defined; cardiac distinct; intestinal indis-
tinct, defined laterally by branchio-intestinal
grooves. Lateral margins subparallel; slightly
concave. Anterolateral margins regularly convex;
with three blunt teeth behind the exorbital angle,
second small but obvious, third minute almost

obsolete. Exorbital angle bluni, rounded, not
projecting. First anterolateral similar in size to
exorbital angle but with longer margin. Front
c,0.36- 0.39% carapace width; c,0,5 x frento-
orbital width; moderately deflexed; with shallow
median emargination; lateral angles rounded:
pre-orbital teeth obsolete; lateral margins slightly
diverging posteriorly. Median post-frontal lobes
distinct, distinctly broader than laterals which are
not clearly differentiated. Branchial ridges
prominent as a series of short broken grmmnular
striations; last continuous, parallel] with
posterolateral margin. and finishing over coxa of
fourth walking leg, Posterior margin
o).S X carapace width. Carapace surface smooth,
shining, punctate. Setae in short tufts over entire
surface, less prominent on cardiac and intestinal
regions. Upper orbital border minutely granular;
straight; inner angle rounded. Lower orbital bor-
der straight; evenly granular. Inner orbital tooth
present; weil developed; acute equilateral tian-
gular. Basal segment of antennal peduncle with
well developed, elongate, rounded outer lobe.
{nter-anteonular septum narrow; 0.22-
0.24 width of front.

Third maxilliped: Meris c,1,4 length of is-
chum. Ischrum inner margin microscopically
granular. Palp articulates medially on distal mar-
gin of mems, Exopod narrow, barely visible in
frontal view; flagellum normal,

Chelipeds: Subequal; large and robust; merus
with posterior border minutely granulate; with
distinct subdistal spine; lower border minutely
granulate; anterior border sparsely granulate,
with a few large rounded granules medially on
convexity; carpus with a spine at inner angle;
innermargin minutely granular, rounded, ventral-
ly with minutely granulate crest and a long
proximal oblique ridge bearing row of long setae:
granules present on inner face of carpus just
below inner angle; outer margin striated, bearing
thick band of short setae: upper surface smooth
and sparsely granulate, with distinctive area of 9
promincntridges and 8 grooves, running transver-
sely along outer distal half behind joint. Palm
upper surface with a series of transverse grooyes
separating swollen ridges: distal margin of upper
surface of palm raised, with a series of 4-8 short,
broad, truncated, chitinous teeth; proximal to this
a series of 3 ridges separating 3 narrow smooth
grooves, or sometimes the first ridge low and
uneven and followed by area of small smooth
granules, and then the other two arced ridges; on
proximal slope of last ridge is a series of short
ridges and grooves, longest towards upper mar-

REVISION OF THE GENUS SARMATIUM 93

FIG. 7. Sarmatium hegerli sp.nov., 3 (holotype, QM W9698). A, dorsal view. B, chela. C, upper surface of palm
of chela. Scale line in mm.

gin; behind this a yery broad deep concavity, and
then a large swelling back to posterior margin, the
distal slope of which is long and covered in well
separated, medium sized, low round = granules;
superior margin with large rounded granules and
short setae. Outer surface of palm smooth;
without median longitudinal row; without a
ventral ridge. Outer surface of palm naked except
for fringe of setae at insertion of dactyl, and a
small patch ventro-proximally behind joint and as
a triangular patch proximally on ventral face.
Inner surface of palm smooth; without vertical
crest but with a few very low rounded tubercles,
almost indistinguishable. Immovable finger
rounded on outer surface; without ventral ridge;
moderately long. Length cutting edge c.0.45-
0.49 length propodus. Ventral border of chela

straight, or slightly convex. Dorsal surface of dac-
tyl with 1 large forwardly directed, bluntly
pointed, chitinous tubercle proximally near joint
on outer edge of superior margin; 5-6 additional
chitinous tubercles commencing c.two-fifths dis-
tance towards tip; first large, oblique, truncate;
second smaller of same form; third also similar
but very small, distal 2-3 minute and reducing
distally, last tubercle situated well before tip:
thick triangular patch of setae proximally on upper
surface near joint. Fingers pointed; slightly
curved inwards; a narrow gape between cutting
margins in largest male, no gape in smaller males.

Walking legs: Medium length; compressed;
second pair slightly the longest, c.1.3-1.5 * max-
imum carapace width. Third leg: merus c.2.4-
2.7 X as long as wide; carpus c.2.5- 2.7 Xas long

94 MEMOIRS OF THE QUEENSLAND MUSEUM

as wide; propodus c.2.3 X as long as wide. Dactyli
about equal to length of propodi; slender and
slightly recurved; terminating in acute chitinous
tips. Propodus with an indistinct accessory carina
on inferior proximal portion of upper surface.
Meri of legs 1-3 with scattering of small distally
directed prickles; dorsal and ventral borders of
meri granulate. Setae short, covering anterior and
posterior faces of carpi and propodi, and distal
upper margins of meri; continue on to proximal
half of dactyli in thin rows.

Male abdomen: Relatively narrow; third seg-
ment the widest, subequal with first. Segments
three-five tapering. Width segment three c.4.25-
4.9xXlength, relatively narrower in smaller
males. Segment six not elongated; 1.25-
1.35 wider than long. Telson longer than
preceding segments, only slightly longer than
segment 6; 1.4- 1.5 Xlonger than wide; evenly
rounded.

Gonopods: G1 moderately stout; slightly
curved. Inner-dorsal margin distally curved in-
ward. Dorsal surface of stem flattened; complete-
ly calcified. Palp poorly developed, not separated
from stem, large, broad, triangular, calcified.
Outer dorsal margin of stem convex. Distal part
of the stem broad but narrowing. G1 apical
process corneous; strongly produced; straight.
Gonopore slightly displaced towards the dorsal
surface. Setae short, simple, obscuring structural
detail. G2 short, relatively narrow, tapering,
twisted, tip blunt.

Sternum: Thick covering of setae between inser-
tion of telson and mouth frame.

HABITAT

Mangroves; burrows in riverbanks and banks of
drainage channels; moist to very sloppy mud; not
restricted to a particular mangrove zone. One
burrow, apparently belonging to this species had
a raised entrance, but this does not seem typical.

DISTRIBUTION
Only known from Northern Territory, Australia.

ETYMOLOGY

Named for Mr Ed Hegerl, director of the
Australian Littoral Society, and one of Australia’s
leading marine conservationists. Someone I have
followed into very many, very muddy places.

REMARKS

The key provided here separates Sarmatium
hegerli sp.nov. from all other known species of
Sarmatium. The pattern of grooves and ridges on

the upper surface of the palm of the cheliped; the
shape, number and position of the cheliped dac-
tylar tubercles; and the male first gonopod are
distinctive.

Sarmatium unidentatus sp.nov.
(Figs 1E, 6D-F, 8)

MATERIAL EXAMINED

HoLotyPe: AM P31822, 1¢é (12.5X10.9mm),
Nungbalgarri Ck, N.T., 2.5km upstream, west bank, D.
Grace, 27.7.1976.

PARATYPE: AM P31819,1¢ (11.7 10.4mm), Liver-
pool R., N.T., 14.3km upstream, amongst debris on
mud floor in Rhizophora mangrove forest, D. Grace,
30.1.1975.

DESCRIPTION

Carapace: c.1.15 X broader than long. Fronto-
orbital width c.0.85 X carapace length. Carapace
deeply vaulted; convex in both directions, only
slightly from side to side. Depth c.0.8 X carapace
width, Regions moderately defined; mesogastric
well defined; cardiac distinct; intestinal distinct.
Lateral margins slightly divergent posteriorly;
slightly concave. Anterolateral margins regularly
convex; with two blunt teeth behind the exorbital
angle. Exorbital angle blunt, rounded not project-
ing. First anterolateral tooth similar in size to
exorbital angle but with longer margin. Second
anterolateral tooth minute. Front c.0.4 X carapace
width; c.0.5 X fronto-orbital width; moderately
deflexed; with shallow median emargination;
lateral angles quadrate, rounded; pre-orbital teeth
obsolete; lateral margins sub-parallel. Median
post-frontal lobes distinct, distinctly broader than
laterals; lateral lobes not clearly differentiated. A
series of short broken granular branchial stria-
tions; last continuous, parallel with posterolateral
margin, and finishing over coxa of fourth walking
leg. Posterior margin c.0.5 X carapace width.
Carapace surface smooth, shining, punctate.
Setae arranged sparsely on branchial lines. Upper
orbital border minutely granular; straight; inner
angle rounded. Lower orbital border straight;
evenly granular. Inner orbital tooth present; well
developed; bluntly triangular. Basal segment of
antennal peduncle with a well developed elongate,
rounded outer lobe. Basal antennular segment
swollen. Inter- antennular septum narrow; 0.22-
0.24 X width of front.

Third maxilliped: Merus c.1.35 X length of is-
chium. Ischium inner margin granular. Palp ar-
ticulates medially on distal margin of merus.

REVISION OF THE GENUS SARMATIUM 95

FIG. 8. Sarmatium unidentatus sp.nov., 3 (holotype, AM P31819). A, dorsal view. B, chela. Scale line in mm.

Exopod narrow, visible proximally and distally in
frontal view but mostly hidden; flagellum normal.

Chelipeds: Subequal; large and robust; merus
with posterior border minutely striated; without
distinct subdistal spine; lower border granulate;
anterior border convex, coarsely granulate; car-
pus with inner angle granular; inner margin
rounded, ventrally with row of minute granules
and short proximal crest bearing row of long
setae; granules present on inner face of carpus just
below inner angle; outer margin striated, bearing
rows of short dark setae; upper surface smooth
and shiny distally, microscopically striated
proximally. Palm upper surface with a series of
transverse grooves separating swollen ridges: dis-
tal border of upper surface of palm with
prominent crest bearing 5-6 short, broad, trun-
cated, chitinous teeth; proximal to this a very

broad, deep, smooth, oblique depression; then a
very large medial transverse swelling; then a
series of seven very fine, long transverse ridges
ending near proximal margin of upper border;
first six ridges joined top and bottom to form 3
pairs; superior margin distinctly granular, and
lacking setation. Outer surface of palm smooth;
without median longitudinal row; without a
ventral ridge. Outer surface of palm naked except
for fringe of setae at insertion of dactyl. Inner
surface of palm sparsely granular, mostly smooth;
with a low vertical crest of c.3 granules. Immov-
able finger rounded on outer surface, or slightly
flattened on outer surface proximally; without
ventral ridge; moderately long. Length cutting
edge c.0.45 X length propodus. Ventral border of
chela concave at base of fixed finger, bearing a
shallow longitudinal depression. Dorsal surface

96 MEMOIRS OF THE QUEENSLAND MUSEUM

of dactyl tuberculate, bearing a pronounced slight-
ly oblique, elongate proximal tooth, truncated and
capped with chitin; a small patch of setae
proximal to this tooth; distally followed by a broad
gap and then a series of evenly spaced very small,
acute, forwardly directed, chitinous tubercles, be-
coming minute distally and finishing at about
three-quarters. length of finger. Fingers pointed:
curved inwards: a wide gape between cutting mar-
ins,

: Walking legs: Medium length; compressed;
second pair the longest, c.1.6-1,7 maximum
carapace width. Third leg: merus c,2,7 Xas long
as wide; carpus c.2.4- 2.5Xas long as wide:
propodus c.2.5% as long as wide. Dactyli about
equal to length of propodi: slender and recurved;
terminating in acute chitinous tips. Propodus
without distinct accessory carina on inferier
proximal portion of upper surface. Meri of legs
1-3 with scattering of small distally directed
prickles; meri granular on superior margins;
minutely striated over upper half, Setae very short
and sparse on anterior halves of meri; thicker on
upper halves of carpi, particularly along acces-
sory ridges; also present on propodi on superior
border and on both anterior and posterior faces,
but disappearing distally; most obvious on
anterior legs.

Male abdomen: First 3 sements subequal in
width in holotype but third segment the widest in
paratype. Segments three-five tapering, markedly
from 3-4, then moderately, Width segment three
c.4.6-4.8 * length. Segment six not elongated;
c.1.8% wider than long. Telson longer than
preceding segments; c.l.3% longer than wide:
evenly rounded.

Gonopeds: G1] moderately stout; slightly sig-
moid, Inner-dorsal margin distally curyed in-
ward. Dorsal surface of stem flattened;
completely calcified. Palp poorly developed, not
separated from stem, small, narrow, rounded, cal-
cified, situated c.two-thirds distance towards tip,
Outer dorsal margin of stem convex. Distal part
of the stem narrow. Gl apical process present:
comeous; strongly produced; stout not markedly
narrower than distal part of stem; straight.
Gonopore terminal. Setae long, simple, lie around
tip, mostly distal to palp. G2 shor, relatively
narrow, tapering, tip blunt.

Sternum: Thick covering of setae between inser-
tion of telson and mouth frame,

HABITAT
Amongst debris on mud floor in Rhizophora
tnangrove forest.

DISTRIBUTION
Only known from Nungbalgarri Ck and Liver-
pool R., Northern Territory, Australia.

ETYMOLOGY

Named for the single pronounced slightly obli-
que, elongate, proximal tooth on the dacty] of the
cheliped which is diagnostic.

REMARKS

The key provided here separates Sarmiectium
unidentatus sp.nov, from all other known species
of Sarmatium. The pattem of grooves and ridges
on the upper surface of the palm of the cheliped;
the shape, number and position of the cheliped
dactylar tubercles; and the male first gonopod are
distinctive,

ACKNOWLEDGEMENTS

Tam indebted to Dr Alain Crosnier of ORSTOM
for his assistance in helping me to undertake this
study, and for his encouragement; and the Board
of Trustees of the Queensland Museum for sup-
port and travel assistance. A large part of the work
was undertaken while at the Laboratoire de
Zoologie (Arthropodes), Muséum national d'-
Histoire Naturelle, Paris, under a grant from the
Institut francaise de Recherche Scientifique pour
le Développement en Coopération (ORSTOM).
Dr Danitle Guinot kindly arranged the loan of
comparative specimens, Dr Charles Fransen of
the Nationaal Natuurhistorisch Museum, Leiden,
and Dr Michael Tiirkay of the Forschungsinstitut
Senckenberg, Frankfurt, are both sincerely
thanked for their generosity and kindness whilst I
Was Visiting their laboratories. John Short is
thanked for his assistance with photography and
help with preparation of plates. M. Maurice Gail-
lard executed the excellent illustrations. } thank
too my wife Kathleen for her support and all
manner of practical assistance.

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Decapoda, Crustacea), Treubia. Buitenzorg 27(4):
387-416, pls 1-8.

1971. On the species of Sarmarium DANA, 1851
(Decapoda, Brachyura), Crustaceana. Internation-
al Journal of Crustacean Research 21 (3): 237-240,
figs 1-2, pls 1-2.

SOH, C.L. 1978, On a collection of sesarmine crabs
(Decapoda, Brachyura, Grapsidae) from Hong
Kong, Memoirs of the Hong Kong Natural History
Society 13: 9-22,

TESCH, J.J, 1917, Synopsis of the genera Sesarma,
Metasesarma, and Clistocoeloma witha key tothe
determination of the Indo-Pacific species.
Zoologische Mededeelingen, Leiden 3(2- 3): 127-
260, pis 15-7.

1918. The Decapoda Brachyura of the Siboga-Ex-
pedition. |. Hymenosomidaé, Retroplumidac,
Ocypodidac, Grapsidac, and Gecarcinidae,
Siboga-Expeditie, Leiden. Monogr, 39c: 1-148, 6
pls.

TWEEDIE, M.W.F. 1936, On the crabs of the family
Grapsidae in the collection of the Raffles
Museum. Bulletin of the Raffles Museum, Sin-
gapore 12: 44-70, text-fies 1-3, pls 14, 15.

98 MEMOIRS OF THE QUEENSLAND MUSEUM

THE RARE SHRIMP, PHYLLOGNATHIA CERATOPH-
THALMA (BALSS), DISCOVERED IN MORETON BAY.
Memoirs of the Queensland Museum 32(1): 98:- A collection
of Crustacea was made by SCUBA diving at Myora, North
Stradbroke Island, Moreton Bay, One particularly striking
species excited interest, and proved to be the rare shrimp
Phyllognathia ceratophthalma Balss. The Myora coral com-
munity occurs as a shallow (1-3m), narrow reef, with six coral
species (Lovell, 1989). It is unusual within Moreton Bay in
having large plates of Acropora digitifera (Dana), These grow
attached to loose rubble on a muddy-sand substrate, which
means they are casily overtumed to sample the associated
fauna. This small reef has a high diversity of invertebrates,
including species which are more typically expected on outside
reefs. This is possibly because during times of flooding when
the rest of the Bay is subject to severe freshwater dilution,
circulation patterns within the Bay cause a pooling of undiluted
seawater along the northern side of Stradbroke Island. The reef
also lies just to the side of the main channel leading in and out
of the Bay, so recruitment from open coastal waters, and the
warm southerly flowing East Australian Current, can be ex-
pected. See Bruce (1991) for full synonomy.

Phyllognathia ceratophthalma (Balss, 1913)

Hymenocera (7) ceratophthalma Balss, 1913: 236.
Phyllognathiaceratophthalma: Borradaile, 1915: 206; Bruce,
1991; 267-269, fig. 28.

Material Examined: QM W17487, 1d (8.0 mmcel (including
rostrum)) & 1? (11.0 mm cl), Myora, Moreton Bay, 2m, in
hole in base of Acropora digitifera plate, 14.4.1992, P. Davie.
Remarks:Bruce (1991) provided a modem figure and
description of this apparently rare shrimp. The present
specimens have an identical colour pattern to that described
by Bruce (1991) but differ in having a translucent

white/cream, rather than yellow-brown, background colour.
Despite being known since 1913, and despite widespread use
of SCUBA for underwater collecting over the past 25 years,
this strikingly coloured shrimp is known from only four
previous records: Satsuma, Japan, the type locality (Balss,
1913); Maldive Islands (Borradaile, 1915); Lizard Is., north-
em Great Barrier Reef; New Caledonia (Bruce, 1991), The
present record is a significant southerly range extension.

On the tenth day after capture, the female was seen to
release larvae from under her abdomen although her
ovigerous state had not been previously apparent from casual
observation, 25 first stage zoeae were released, and a further
16 were found in a ‘brood pouch’ formed by the broad,
overlapping pleopods. This species is reproductively a K-
Strategist, with a small number of large eggs. It probably can
maintain a breeding population at this southerly, subtropical
locality. The first zoeae will be described elsewhere. Facilities
were not available to rear them to later stages.

Literature Cited

Balss, H. 1913. Diagnosen neuer ostasialischer Macruren.
Zoologische Anzeiger 42: 234-9.

Borradaile, L.A. 1915. Notes on Carides. Annals and
Magazine of Natural History (8) 15: 205-13.

Bruce, A.J. 1991. Shallow-water Palaemorioid shrimps from
New Caledonia (Crustacea: Decapoda). Pp. 221-79, figs
1-31. In Richer de Forges, B, (ed.), ‘Le benthos des fonds
meubles des lagons de Nouvelle-Calédonie, Vol, 1.
Etudes et Théses.’ (ORSTOM: Paris).

Lovell, E.R. 1989. Coral assemblages of Moreton Bay,
Queensland, Australia, before and after a major flood.
Memoirs of the Queensland Museum 27(2): 535-50.

Peter J.F. Davie, Queensland Museum, PO Box 3300,
South Brisbane, Q. 4101, Australia; I May, 1992.

FIG. 1: Phyllognathia ceratophthalma (Balss, 1913), Queensland Museum W17487.

(CYMBASTELA, A NEW GENUS OF LAMELLATE CORAL REEF SPONGES
JOHN N.A, HOOPER AND PATRICIA R. BERGQUIST

Hooper, J.N.A. & Bergquist, P.R. 1992 06 29: Cymbastela, a new genus of lameliate coral
reef sponges. Memoirs of the Queensland Museum 32(1); 99-137. Brisbane. [ISSN 0079-8835.

Seven species of lamellate, stalked sponges from Australasian waters are referred to a new
genus, Cymbastela gen.noy. Six species from tropical, predominantly coral reef habitats: C.
stipitata (Bergquist & Tizard) (type species) and C. vespertina sp.nov., both from NW
Australia, C. concentrica (Lendenfeld) and C. coralliophila sp.nov. both from the Great Bamier
Reef, Queensland, C. marshae sp.nov. from the Houtman-Abrolhos Is, Western Australia, and
C.cantharella (Léyi) from New Caledonia), Another species is recorded from temperate South
Australian waters (C. notiaina sp.nov. from the Gulf of St Vincent). AL the species conformed
to the existing interpretation of Pseudaxinyssa Burton, but they are referred here to Cyrabastelia
since all are substantially different from the type species of Pseudaxinyssa, A, tethyoides
Kirkpatrick. Pseudaxinyssa is synonymized with Axinyssa Lendenfeld. The seven species of
Cymbastela ate primarily differentiated by their spicule length and geometry, axial and
extra-axial skeletal development, growth form, lamella thickness and live colouration. Four
species are sympatric over part of their range in tropical Australia, whereas three species have
more restricted, allopatric distributions in temperate Australia and the western Pacific Ocean.
CO] Pseudaxinyssa, Axinyssa, Cymbastela, new genus, new species, Porifera, Axinellidae,
taxonomy, biogeography, Australasia.

John N.A, Hooper, Queensland Museum, PO Box 3300, South Brisbane, Queenslund 410),
Australia; Patricia R. Bergquist, Department af Zoology, University of Auckland, Private Bag,

Auckland, New Zealand; 29 January, 1992.

No formal revision of the complete order
Axinellida has yet been published, even though
for many years it has been recognized these
marine sponges were polyphyletic (Bergquist &
Hartman, 1969). Recent work on several families
within the Axinellida, using a phylogenetic
framework, has gone some way towards
cedistributing some of these taxa amongst other
existing orders. Families considered so far (with
their affiniues as suggested by particular authors).
are Axinellidae (Halichondrida; Van Soest et al.,
1990), Desmacellidae (Poccilosclerida; Van
Soest, 1984), Hemiasterellidae (Hadromerida;
Hooper, 1986; Voultsiadou-Koukoura & Van
Scest, 1991), Rhabderemiidae (Poecilosclerida:
Hooper, 1990), and Raspailiidae
(Poecilosclerida; Hooper, 1991). However, it is
not yet clear whether the concept of an order
Axinellida should be abandoned completely, or
retained and restricted to one or few families.
Additional, non-morphological data is required to
confirm or refute these ideas (discussion in
Hooper et al., 1992), and until such is available
none of the above assignments can be completely
endorsed.

‘xinellidae is undoubtedly the most difficult of
all the families of Aximellida to interpret. The
original conception of the family, basically cor-
responding to the old subfamily Axinellinae of

Lendenfeld (1889), with major additions by
Hallmann (1916) and de Laubenfels (1936), in-
cluded genera with an ‘axinellid skeletal plan’
consisting of a condensed axial core with diverg-
ing plumose or plumo-reticulate extra-axial
branches. This definition stands today (e.g.
Hartman, 1982), even though nearly half of the 50
nominal genera included in the family have skele-
tal structures other than axinellid (e.g. evenly
reticulate - Reniochalina, sinuous - Rhaphoxya;
dendritic - Acanthella; thickly encrusting/lep-
tcelathrid - Fasubera), In such cases, genera with
atypical skeletal structures were included in
Axinellidae because their spicule geometnes were
similar to those with axinellid skeletons. How-
ever, recent biochemical evidence indicates that
the family contains several distinctly
heterogenous groups of genera (Bergquis! &
Hartman, 1969: Hooper et al., 1992),

Artecent revision of halichondnids by Van Suest
et al. (1990), based pnmarly on morphological
characters, suggested that genera of Axinellidac
could be redistributed into three existing families
(Axinellidae, Desmoxyidae and Halichondriidac)
and a new family (Dictyonellidae). Yan Soest et
al. (1990) referred these four families to the order
Halichondrida, and although we do not presently
support or refute this ordinal placement, we agree
that for the Axinellidae at least, there is a pressing

need for u re-evaluation of the major morphologi-
cal characters used te distinguish taxa within the
Axinellidae, including the relative importance of
these characters as reliable indicators of
phylogenetic affinity,

It is difficult in many cases to assign species of
Axinellidae to genera with complete assurance, as
there are few morphological characters with clear
importance at the supraspecific level, Unlike most
groups within Poecilosclerida, for example,
Axinellidae. lack any unique or diverse spicule
geometries, and lack microscleres which could
provide clues on affinities between genera. Thus,
the major problem preventing resolution of the
supraspecific taxonomy of Axinetlidae is the dif-
freulty or imability to define autapomorphies and,
furthermore, there is disagreement tn the inter-
pretation of the relative importance of the few
existing morphological characters that do occurin
the group - in particular spicule shape and skeletal
organization. Biochemical evidence may provide
more objective information on the affinities of the
numerous genera presently included in the family,
but the few data available are not entirely clear
(Mouper et al. 1992): there are, in many cases, no
obvious morphometric characters that correlate
consistently with generic groupings based on
chemotaxonomic data (see Hooper et al.. 1992).
The ‘axinellids’ are one group of Porifera that
require Migorous genetic study.

The existing subdivision witht Axinellidae,
resting solely on morphological features, concerns
the skeletal architecture, presence or absence of a
specialised eciosomal skeleton, and morphology
and diversity of structural megascleres. The cur
vature and geomeiry of megascleres, however, is
thought to vary considerably within a particular
species groups, and confuse the interpretation of
interrelationships between species even further.
This problem was first identified by Dendy
(1905), who noted that oxeas may become stron-
gyles or even styles, and curved or sinvous stron-
gyles, once considered apomorphic for genera
such as Acanthella, reappear throughout the fami-

y.

Consequently, it is not surprising that there are
many genera of Aninellidae that presently contain
heterogenous assemblages of species, particularly
the large groups of Phakellia, Acanthella, Axinel-
Ja, to each of which numerous species have been
assigned, A similar situation exists for
Pseudaxinyssa, and although it contains only etght
nominal species - prior to the present work - some
of these are obviously quite different from the type
species. Lévi (1983) first stated that Preudaxinys-

MEMOIRS OF THE QUEENSLAND MUSEUM

sa was probably polyphyletic, and this was recent-
ly eaerated by Van Soest et al. (1990). The
present paper re-evaluates Psendaxinyssa, returns
its type species and several other described species
to Axinyssa, and creates a new genus to receive
several described species and a number of new
species mainly from tropical Australian coral reef
habitats.

METHODS

Preparation of matemal for light microscopy and
scanning electron micrascopy (SEM) is described
elsewhere (Hooper, 1991). Abbreviations \sed in
the text areas follows: AM, Australian Museum,
Sydney; BMNH, The Natural History Museum,
London; CMNZ, Canterbury Museum, Marine In-
vertebrate collection, Christchurch; MNHN
LBIM, Muséum National d’ Histoire Naturelle,
Laboratoire de Biologie des Invertébrés Marins et
Malacologie, Paris; NCI Q66C-. US National
Cancer Institute shallow water collection program
(Australian Institute of Marine Science,
Townsville, contract) - primary material in
USNM, duplicates/fragments in NTM and QM;
NTM, Northern Territory Museum, Darwin;
ORSTOM, Institute Frangais de Recherche Scien-
tifique pour le Developpement en Cooperation,
Noumea; PIBOC, Pacific Institute of Bio-organic
Chemistry, Far Eastern Scientific Centre,
Academy of Sciences of the USSR, Vladivostok;
QM, Queensland Museum, Brisbane; SMF,
Natur-Museum und Forschungsinstitut Sencken-
berg, Frankfurt; USNM, United States National
Museum of Natural History, Smithsonian Institu-
lion, Washington; ZMB, Zoologisches Museum
fiir Naturkunde an der Humboldt-Universitit,
Berlin,

SYSTEMATICS

Family AXINELLIDAE Ridley & Dendy
Axinyssa Lendenfeld, 1897

Axinyssa Lendenfeld, 1897:116; Burton, 1931:349;
Van Soest etal,, 1990;25,

Axinomimus de Laubenfels, 1936:163; Van Soest et al.,
1990:27 (type species Axinella paradoxa Ridley &
Dendy, 1886:482 [by monotypy]; holotype BMNH
1887.5.2,68).

Psevdaxinyssa Burton, 1931:350; Van Soest et al.,
1990:27 (type species Axinyssa tethyoides
Kirkpatrick, 1903;245 [by original designation):
holotype BMNH 1902.11.16.25, schizorype MNHN
LBIM DCL298).

LAMELLATE CORAL REEF SPONGES

Not Pseudaxinyssa, sensu Bergquist & Tizard, 1967:
190; Léyi, 1983:719,

TYPE SPECIES

Axinyssa topsenti Lendenfeld, 1897:116 tby
munotypy) (holotype ZMB 2971, schizatype BMNH
1908.9.24.145),

DIAGNOSIS (emended)

Massive, frequently subspherical, sometimes
lobate or lobate-digitate growth forms. often with
in apical cloacal cavity or fistulose surface con-
ules, surface with prominent, regularly spaced
conules; choanosomal axial skeleton open
plumose, dendritic or vaguely radiating tracts of
larger oxeas, without axial compression, with
poorly developed spongin fibres; fibres cored by
larger oxeote megascleres; subectosomal extra-
axial skeleton plumose, plumo-reticulate, ascend-
ing to surface, without obvious spongin fibre
component; mesohyl usually with abundant col-
lagenous spongin; specialised ectosomal skeleton
may be present in the type species, or absent,
consisting of tufts of smaller oxeas. perched on
peripheral choancsomal skeleton, protruding
through surface, forming surface conules;
megascleres oxeas or modified oxeas, one or twa
Sizes.

REMARKS

Lendenfeld (1897:116) created the genus
Axinyssa for 4 single species, A. tapsenti Lenden-
feld, which had a vaguely plumose. ascending
choanosomal skeleton composed of multispicular
tracts of large oxeas. with smaller stylote oxeote
spicules in brushes on the surface. These
choanosomal spicule tracts and smaller ec-
tosomal brushes protrude through the surface for
a short distance, producing characteristic
microconules. There is no distinctive, compressed
core in the skeleton (termed ‘axial condensation’),
although there is differentiation between the axial
und extra-axial regions. Several other species
were subsequently described and included in
Axinyssa (Kirkpatrick, 1903; Topsent, 1906; Wil-
son, 1925).

Burton (1931) created Psewdaxinyssa tor two
species previously assigned to Axinyssa - the type
species A. tethyoides Kirkpatrick (1903), and A.
gravieri Topsent (1906). He also included a pre-
viously undescribed species, P. tenuispicula Bur-
ton (1931) from Natal, All three species differed
from Axinyssa sensu stricto in lacking a
specialised ectosomal skeleton of smaller styloid
oxeas. They were closer to Axielia in their skela-

101

tal structure, but had only oxeas as megascleres
(Axinelia usually has styles and oxeas, or only
styles, although at least one anomolous oxea-bear-
ing species is also known [see below]). Van Soest
et al. (1990) noted that a specialised ectosomal
skeleton, such as that seen in A. topsenti, common-
ly occurs within many axinellids and, as such, it
may not be a reliable or even valid taxonomic
character when used above the species level. We
provide further evidence to support this con-
clusion in the present work, (Le, Cymbustela
coralliophila sp.noy. having a specialised ec-
tosomal skeleton), supporting the proposal that
Axinyssa and Pseudaxinyssa (s.s,) should be
merged.

Pseudaxinryssa, as defined by Burton (1931),
were massive sponges, without strong differentia-
tion between the axial and extra-axtal skeletons,
and without any evidence of axial condensation,
The type species Axinyssa tethyoides
(Karkpatrick) ts subspherical, with fistulose,
usually blind surface papillae, with large oxeas
between 530-790m long scattered between
skeletal tracts and forming a branching reticulate,
nen-compressed, undifferentiated skeletal ar-
chitecture (i.e. no distinct axial and extra-axial
components). However, the subsequent addition
of two stalked, cup-shaped sponges from the Indo-
west Pacific. Pseudaxinyssa stipitata Berquist &
Tizard (1967) and P. cantharetla Lévi (1983). did
not fit at all with the original conception of the
genus, These two species had compressed-reticu-
late axial skeletons, with diverging plumose (or
plume-reticulate) extra-axial skeletons, and spon-
gin fibres were reasonably well developed. Both
groups of species were similar to A, tethyades
only in having diverging tracts of oxeas near the
periphery of the skeleton, whereas in other fea-
tures they were quite different. Therefore, it is
proposed here to include these anomolous stalked,
predominantly cup shaped species in anew genus,

Following from Van Soest et al’s (1990)
revision of Aximyssa and our own observations on
various museum specimens, 13 species may be
referred to the genus, most from the tropical
oceans {also indicating known geographic dis-
tributions, and material we have examined):

A. aculeata Wilson (1925-445) (Philippines;
none}.

A. ambrosia (de Laubenfels, 1934:14) (Puerto
Rico; none).

A, aplysinoides (Dendy, 1922:39) (Cargados
Carajos, Amirante, Seychelles; none).

A. digitata (Cabioch, 1968:229) (Roscoff;
none).

A. fenesiratus (Ridley, 1884:464) (Arafura Sea;
holotype BMNH 1882.2.23.255).

A, gravieri Topsent (1906:563) (Red Sea:
specimen of Léyi (1965:15), MNHN LBIM
DCL327L).

A. paradexa (Ridley & Dendy, 1886:482) (Tris-
tan de Cunha Is: none),

A. pitys (de Laubenfels, 1954:178) (Palau Is-
Jands; holotype USNM 23103).

A. radiata (Lévi & Lévi, 1983:968) (New
Caledonia; paratype MNHN LBIM DCL 2955).

A. tenuispicula (Burton, 1931:350) (Natal;
none).

A. tethyoides Kirkpatrick (Natal, holotype
BMNH 1902.1 1.16.25).

A. topsenti Lendenfeld (Adriatic; holotype
ZMB 2971).

A. valida (Thiele, 1899:12) (Moluccas; ?
paratype SMF 1821).

Another species exists in the MNHN collec-
tions, identified as anew species of Psendaxinyssa
by Claude Lévi (pers.comm.) (‘holotype’ MNHN
LBIM DCLI497) from Anborombe, Africa (M.
A Crosnier si CRO 69), and it too should be
included in Axtnyssa. The affinities of the remain-
ing species which have been included in
Pseudaxinyssa at one time or another are as fol-
lows:

P. cantharella Lévi (1983:719) (New
Caledonia) belongs to the new genus descnbed
below.

P. stipitata Bergquist and Tizard (1967:189)
(northern Australia) also belongs to the new genus
described below.

P. tricalyciformis (Bergquist, 1970:15)
(holotype CMNZ Mar. Inv, 3/63), a replacement
name for Axinella lamellata Bergquist (1961:188)
(preoccupied), from the Chatham Islands, was
referred to Pseudaxmyssa by Bergquist & Tizard
(1967:190). It has oxeas, some with styloid
modifications, forming a condensed axial skeleton
and plumose extra-axtal skeleton, and ts returned
to Axinella, Discrepancies in dates of referral and
description of that species are a consequence of
delayed publication of the 1970 paper, which was
completed in 1966 (Bergquist, 1970:4),

Following Van Soest et al.°s (1990) revision of
the *halichondrtd’ axinellids Axinyssa was
referred to the family Halichondriidae Vosmaer,
as emended by them, in having vague, ill-defined
spicule tracts, with a high ratio of spicules to
spongin. By comparison, ‘true’ members of the
Axinellidae have ordered, axially condensed,
extra-axially plumo-reuculate skeletons, The type
species of Pseudaxinyssa, A. tethyoides fits well

MEMOIRS OF THE QUEENSLAND MUSEUM

with the emended definition of Halichondnidae
(see Figs 1-2), and a redescription of A. tethyoides
is provided here for comparative purposes with the
new genus described below.

Axinyssa tethyoides Kirkpatrick
(Figs 1-2)

Axinyssa tethyoides Kirkpatrick, 1903: 245-6, pl. 5,
fig, 12, pl.6, fig,8a,b.; Van Soest et al., 1990:27,
fig. 18.

Pseudaxinyssa tethyoides; Burton, 193):350.

MATERIAL EXAMINED

HovLotyPe: BMNH 1902,11.16.25: Cone Point, Natal,
South Africa, 68m depth. SCHIZOTYPE: MNHN LBIM
DCL298.

SUBSTRATE AND DEPTH RANGE
Broken shell substrate.

GEOGRAPHICAL DISTRIBUTION
Known only from the type locality,

DESCRIPTION

Shape: Subspherical, massive growth form,
64mm diameter, with slightly flattened basal
region (point of attachment) (Fig. 1A).

Surface features; Detachable, relatively fragile
skin, covered with numerous small papillae, 2-
Smm long, 1-2.5mm diameter, most erect, fis-
tulose, but few open at the apex. When open,
diameter of oscule approximately 1mm.

Colour: Live colouration unknown, mottled
black and dark brown in ethanol.

Texture: Finn, barely compressible consistency.
easily crumbled.

Eclosome: Membraneous, without specialised
skeleton, with moderate quantities of detritus
within heavy organic matrix, occasionally with
spicules from peripheral choanosomal ‘skeleton
protruding through surface, corresponding to sur-
face papillae (Fig. 2B).

Choanosome; Skeleton branching reticulate,
open-meshed (mesh diameter [40-320um),
without trace of axial condensation or differentia-
tion between axial or extra-axial regions. Skeletal
tracts pauci- or multispicular, 80-170m wide,
composed of tightly bound oxeas, without spongin
fibres, bound together within heavy collagenous
mesohyl (Fig. 1B), Near periphery skeletal tracts
diverge to surface, protruding through ectosome
slightly or not at all.

Spicules: Oxeas of single size class, long,
robust, symmetrical, slightly curved at centre,

LAMELLATE CORAL REEF SPONGES

103

FIG, 1, Axinyssa tethyoides, holotype (BMNH 1902.11.16.25). A,whole specimen (scale=30mm). B,SEM of
skeleton (scale=I1mm). C,spicule (scale=250}.m).

tapering to sharp points (Fig. 2A) (length 533-
648.4-789).m, width 18-29.6-43,1m),

REMARKS

The large size of spicules, undifferentiated axial
and extra-axial skeletons, lack of a condensed
axial skeleton, absence of spongin fibres, presence
of heavy collagen in the mesohyl, and a spherical
fistulose growth form distinguish this species
from all the Australian ‘Pseudaxinyssa’ species.
For reasons discussed above we consider that this
distinction is at the generic Jevel.

Cymbastela gen.noy.

Pseudaxinyssa, in part, Bergquist & Tizard, 1967: 190;
Lévi, 1983:719.

TYPE SPECIES
Pseudaxinyssa stipitata Bergquist & Tizard, 1967:189
(holotype AM Z3101),

DIAGNOSIS
Typically stalked, cup-shaped, thinly lamellate
growth form, Ectosome with or without

MEMOIRS OF THE QUEENSLAND MUSEUM

specialised skeleton of smaller oxeas.
Choanosomal skeleton with compressed reticulate
axial region, in which major tracts run lon-
gitudinally through lamellae, and with gradually
ascending, diverging, radial, plumose or plumo-
reticulate extra-axial region, in which tracts be-
come plumose and project through surface.
Spongin fibres well developed, cored by oxeas,
frequently with telescoped points. Predominantly
autotrophic, most species containing symbiotic
cyanobacteria. Oviparous.

FIG. 2. Axinyssa tethyoides, holotype. A, spicules. B,
section through peripheral skeleton.

ETYMOLOGY

Latin, cymba - cup, stela - column (f.); Cymbas-
tela, for the predominantly stalked, cup-shape
growth forms.

REMARKS

Cymbastela is an unusual axinellid in its growth
form, although this morphology is also repeated
in several Haplosclerida, such as some Haliclona
and Amphimedon. Typically the genus has a
reticulated, compressed axial skeleton and a
plumose extra-axial skeleton, both composed of

LAMELLATE CORAL REEF SPONGES

TABLE 1. Comparison in spicule dimensions between
described species of Cymbastela. All measurements
given in micrometres, and expressed as minimum -
mean - maximum range of measurement. N=number
of specimens examined.

155-259.1-344 | 2.7-8.6-16.0
133-267.4-36) |2.0-i14-19.5

167-252.2-270 |5.2-11.9-15.0
152-238.8-283 |3.6-10,7-16.2

well formed spongin fibres. In the extra-axial
region there are always fewer paucispicular,
reticulate fibres interconnecting the multispicular,
radial or plumose, ascending fibres. This structure
shows some similarities to the skeletons found in
Axinella Schmidt (i.e. A. polypoides Schmidt,
sensu Vosmaer, 1912), in which there is a con-
densed, reticulate axis and plumose-branching.
and reticulated extra-axis, and Phakellia Bower-
bank (i.e. P. vertilabra Linnaeus, sensu Vosmaer,
1912), in which there is a condensed, reticulate
axis, and a plumose, radial, non-branching or
poorly-branching extra-axial skeleton. Both
Axinella and Phakellia haye spicules consisting of
styles, strongyles and oxeas, in all combinations,
whereas in Cymbastela spicules are only oxeas,
usually with telescoped ends. Cymbastela is su-
perficially most similar to Phakellia (sensu Vos-
maer), due to the emphasis of the extra-axial
skeleton over other structures, but it is probably
most closely related to Axinelia in all other fea-
tures.

Other specimens

105

It is difficult to define any particular character
which may serve as a synapomorphy for Cymbas-
iela, clearly differentiating it from its sister group
Axinella, in particular, Virtually no single feature
is unique; levels of homoplasy are comparatively
high within the Axinellidae, The lameliate growth
form is characteristic and consistent. but not uni-
que to Cymbasjela; two species of Axinella, A,
vaceletia (Pansini) and A. shegmakeri de Lauben-
fels, are also Jamellate, Similarly, Axinelia
polycapella de Laubenfels has a reduced spicule
complement, with mostly only oxea megascleres,
although in Cymbastela many of the oxeas have
telescoped ends, and itis probably this feature that
is most importantin defining the genus- Therefore,
all three characters in combination - growth form,
spiculation and skeletal architecture - are con-
sidered to be valid discriminatory characters
defining the new genus,

Seven species are referred here to Cynmibastela,
only three of which have been previously
described, The genus is most common in the tropi-
cal Australian - western Pacific region, alttiough
one temperate species is also known.

KEY TO THE SPECIES OF CYMBASTELA
Characters found in both preserved material
[and live specimens] are indicated.

|, Oxeas mostly with telescoped ends wu. ccuus 3.

Oxeas mostly with tapering CNdS voce de

2. Thicker /amellate, cup-shaped growth form, often
with external buttresses, concentric lamellae,
and lateral projections [typically pigmented
olive-green and mauve)) 2... C8
stipitata (Bergquist & Tizard).

Tihinly lamellate, usually simple, concentric or
asymmetrical cup-shaped growth forms.
wilhoul exterior projections [dark maroon-
brown, cvenly pigmented]
pyrtiyerrsnpareinenrateainenearneeree, C. Vespertinad sp.nov,
3. Oxeas Well stlicified oo... ee eseccnectsesseeecneaees 4.

Oxeas poorly silicified, vestigial, thin, with
strongyloid-rounded telescoped ends [small
symmetrical vase-shaped, thin lamellae, ever
surface. live colouration unknown]
a dosbebdeutchdysoiloruat dobcadetlatoéjebe .C. notiaina sp.nov.
4. Specialised ectosamal skeleton of smaller oxeas
present, overlaying projecting subdermal
skeleton of larger oxeas [wicle cups or thickly
Tamellate lobes attached directly to substrate,
usually with convoluted external surface, mot-

106

tled olive-green and olive-brown coloura-
PLOW: ..,cpvesssoessagsessencsts C. coralliophila sp.nov.

Without specialised ectosomal skeleton ..,... 5.
5. Axial skeleton not compressed, predominantly
plumo-reticulate, extra-axial skeleton radial,
not plumose, ectosomal skeleton with plumose
projecting spicules [concentric cups, usually
buttressed or with other sorts of external
projections, thin lamellae, convoluted margins,
with obvious differences between interior
(porous, inhalant) and exterior (smooth, ex-
halant) surfaces, pale orange]
a JpelpatldaUlngQayrsUrevesUaqdeUrpdebllac C. cantharella (Lévi).

Axial skeleton compressed, predominantly
reticulate, extra-axial skeleton predominantly
reticulate, ectosomal skeleton with plumose
projecting spicules [concentric or convoluted
cups, thin lamellae, evenly pigmented dark
brown or OliVe-DrOWN] ......c ce ceceepeeeeeseeecreeees

..C_concentrica (Lendenteld).

Axial skeleton compressed, predominantly
plumose or longitudinal, less obviously reticu-
late, extra-axial skeleton predominantly
plumose, ectosomal skeleton plumose [ir-
regular cups, thin lamellae, no basal stalk,
smooth surface on both interior and exterior
faces, evenly a olive-green/maroon|

devine peg UaeUen' ..C, marshae sp.nov,

Cymbastela stipitata (Bergquist and Tizard)
(Figs 3-5, Table 1)

Pseudaxinyssa stipitata Bergquist and Tizard,
1967:189-191, text-figs 3-4, p15, fig.3.

MATERIAL EXAMINED

HoLortyPe; AM Z3101: East Point Reef, Darwin, NT,
12°25.0'S, 130°49.1'E, intertidal, 13.x.1965, coll. B.
Pope and R.S, Williams.

OTHER MATERIAL: (All materia] collected by the author
unless otherwise indicated). DARWIN REGION, NT:
NTM Z163-165: East Point Aquatic Life Reserve,
12°25.0'S, 130°49.1°B, 0.5m depth, 13.xi.1981 (sm
EP6). NTM Z261 (4 specimens): same locality,
17.ix,198) (stn EP3), NTM 2294: same locality, 1m
depth, 18.viii,1981 (stn EP4). NTM Z309: same
locality, 0.5m depth, 20.ix.1981 (stn EPS). NTM
72145: same locality, 27.ix.1984 (sin EP17). NTM
22207: same locality, 23.xi.1984 (sin EP18). NTM
22225: same locality, 22.i,1985 (stn EP20). NTM
Z2240; same locality, 811.1985 (stn EP21), NTM
22250: same locality, 12.iv.1985 (stn EP22). NTM

MEMOIRS OF THE QUEENSLAND MUSEUM

Z2412: same locality, 7.x.1975 (stn Don.196, coll. A.J.
Dartnall), NTM 22422: same locality, 3m depth,
14,viii.1985 (stn EP24). NTM Z2542: same locality,
0,5m depth, 4,x.1985 (stn EP25). NTM 22547: same
locality, 15.xi.1985 (stn EP26; coll, A.M. Mussig).
NTM 22552: same locality, 12.xii.1985 (stn EP27; coll.
A.M. Mussig and C. Hood). NTM Z2250: same locality,
12°24,5°S, 130°48.0’E, 10m depth, 12.iv.1985, coll. C.
Hood and J.R. Hanley (stn EP22). QM G300001 (frag-
ment NTM Z2392): same locality, 8m depth,
29.vii.1985 (st EP23). NTM Z3179: same locality,
12°29,5’S, 130°48.0'E, 0.5m depth, 10.ix.1987, coll. N.
Smit, by hand (stn JH-87-017). NTM Z401; Vestey’s
Beach, 12°26.2’S, 130°49,9"E, intertidal, 11.xii.1981
(stn MB4), NTM 2479: Fannie Bay beach, 12°25.0°S,
130°50.0’E, intertidal, 9.11, 1932 (stn EP7). NTM Z2009

FIG. 3. Cynthastela stipitata. A, typical growth form
(specimen NTM Z2009). B, in situ (specimen NTM
22145).

LAMELLATE CORAL REEF SPONGES 107

FIG.4, Cymbastela stipitata. A, SEM of skeleton (specimen NTM 22009) (scale=500,.m; left magnified 40,8
times, right magnified 320 times). B, peripheral skeletal architecture (seale=1mm), C, oxea (scale=] 00m).

MEMOIRS OF THE QUEENSLAND MUSEUM

108

—

wi OOT

~S
=
—=

FIG. 5. Cymbastela stipitata, specimen NTM Z2009. A, spicules. B, section through peripheral skeleton. C,

enlarged portion showing the close-meshed reticulate near the axial skeleton. D, confirmed distribution.

LAMELLATE CORAL REEF SPONGES

(fragment QM G300006): W side of Weed Reef,
12°29.2’S, 130°47.1'E, 19m depth, 11.v.1984 (sto
WR2). NTM Z2190: ‘Bommics', Weed Reef,
12°29.2'S, 130°37.6'B, 6-10m depth, 16,xi,1984 (stn
WR6), SAM TS4023 (fragment NTM 2.1655): Old Man
Rock, Nightcliff, 12°21'S, 130°51'E, intertidal,
16.¥1.1962 (coll. R.N. Southcott and A.B. Adams, stn
AS73). NTM ZA424: Lec Point, 12°19,2'S, 130°53,1'E,
intertidal, 13.xi1,1981 (stn LP2). NTM Z779: W side of
Channel £, Middle Arm, 12°33.4'S, 130°52.5'E, 2m
depth, 7.vii. 1982 (sta ClL2). NTM 2796: 8 side of Chan-
nel L, Middle Atm, 12°32.3'S, 130°51.3"E, Lim depth,
16,¥ii, 1982, coll. S, Chidgey. NTM 22105: N side of
Fish Reef, Bynoe Harbour 12°26.2'S, 130°26.2'B, 9-
10m, 5.i%.1984 (stn FR2). CopourG PENINSULA
REOION, NT - NTM 22491: Coral Bay (now known as
Seven Spirit Bay), Port Essington, 11°09.4’S, 132°04"E,
4-7m depth, 13.ix,1986 (stn CP71), NTM 23301, 3302:
Inside outer barrier to Coral Bay, S side of Ray, Port
Essington, 11°09.4°S, 132°04.0’°E, 2-7m depth,
12,jx 1986, SCUBA (stn JH-85-020), NTM 23244; SW
side of chif face, Table Head, Port Essington,
11°13.5'S, 132°10-S°E, 2-5m depth, 11,ix.1986,
SCUBA (stn JH-85-019). NTM 2119: Sandy 1. No. 2,
Antfura Sea, 11°05.5'S, 132°17°E, 7m depth, 20x, 198!
(sty CP26), NTM 7536: EB of Point Edwards, Port
Bremer, CPMNP, 11°U8.5'S, 132°18.8"E, 6-7m depth,
1,v, 1982 (stn CP33).

SUBSTRATE AND DEPTH RANGE

Restricted to shallow coastal waters, ranging
from intertidal to 19m depth. Substrate
predominantly rock or coral rubble, and the
sponge is most abutdant in areas where there are
relatively light loads of sediment.

CIROGRAPHICAL DISTRIBUTION

Northwestern Australian waters (Fig. 5D). Lévi
(1983) suggested that C. stipitata also occurs in
the lagoon off Noumea, New Caledonia, but it is
possible that he was referring to the new species
described below from the Great Barner Reef,
Cymbasiela coralliophila sp.nov.

DESCRIPTION

Shape: Stalked, cup-shaped growth form, usual-
ly sith thick, uneven, crenellated margins, smooth
interior surface, mostly smooth exterior surface of
vase, but often with lateral growths and projec-
tions on exterior surface. Growth variants com-
mon, sometimes with 2-4 concentric, successively
smaller cups growing inside each other, with or
without buttressed exterior projections near base.
Margins and stalk typically relatively thick, up to
14mm and 32mm maximum widths, respectively.

109

Although growth variants may occur within
metres of each other, specimens tend to be thinner,
more elongate {more vasiform), and less con-
voluted with increasing depth (Figs 3A,B).

Surface features: Oscules not visible opncally,
although small, evenly distnbuted pores (0,4-
1.8mm diameter) seen under higher magnifica-
lion, usually situated on points of surface conules,
In some cases distribution of pores coincides with
distnbution of parasitic bamacles (Acasia sp.),
whereas other openings appear to be true intyalant
or exhalant pores.

Colour: Live colouration clearly dependent on
well-being of resident algal symbionts- vanations
in pigmentation related to degree of shading,
water depth and water clarity, Typical live
colouration of intertidal specimens olive-green on
margins (Munsell 2.SGY 5/4), olive-green with
maroon-brown tirige on inside of vase (SGY 4/4-
2.5R 4/2), maroon-brown on exterior of vase und
stalk (2,5R 4/4). Subtidal specimens typically
evenly pigmented olive-brown or olive-maroan
(2.5-IOR 3/2). Heavily shaded, damaged
specimens or material from waters with excep-
tionally high turbidity may be completely beige
(i.e. Without pigments),

Texture: Consistency always stiff, firm, but
flexible, with roughened, sand-paper texture
produced by spicules protruding slightly from der-
mal skeleton. Surface always optically smooth,
microscopically hispid and microconulase, fre-
quently raised into regular or irregular ridges or
Tounded digitale projections.

Ectosome: No specialized skeleton on surface,
but with prominent, relatively closely meshed,
diverging plumose brushes of choanosomal uxeas
protruding through ectosome for short distances
(upto 150j1m maximum), producing microscopic
surface conules. Subdermal skeleton more-or-less
plumose, with more closely meshed multisprcular
skeletal tracts than on ectosome. Tracts may or
may not be interconnected by paucispicular
transverse fibres. but usually hidden by diverging
spicules (Figs 4, SB-C).

Choanesome: Axial skeleton compressed, com-
pased of very close-meshed plumose or vaguely
plumo-reticulate multispicular fibres, mostly run-
ning longitudinally along core of lamellae, ov-
cupying majority of lamella diameter. Only few
interconnecting uni- or paucispicular tracts ob-
served at core, but these mostly obscured by close-
meshed reticulation. Disjunction between
Jongitudinally orientated axial skeleton and ver-
tically disposed extra-axial skeleton only notice-
able very close to periphery. Choanosomal fibres

110

lightly invested with spongm, with few extra-fibre
megascleres. Mesohyl matnx lightly reinforced
with collagen, with abundant microalgae scattered
throughout.

Spicules (refer to Table 1 for dimensions):
Megascleres unmodified oxeas, symmetrical,
thick, curved centrally, with mostly fusiform
sharply poimted ends, sometimes with very slight-
ly telescoped points but this feature never as well
developed as in other species. Structural
megascleres coring fibres slightly larger than
those scattered between fibres (Fig. 3A).

ASSOCIATIONS

With the exception of a few specimens from
very turbid waters, which may have shed their
algal symbionts, most specimens of C. stipitata
contained cyanobacteria (blue-green algae), dis-
persed evenly throughout the sponge mesohyl.
Only rarely were these symbionts seen inside
spongin fibres. The olive-green, olive-brown
sponge pigmentation, characteristic of this
species, may be a photosynthetic by-product of
these symbionts: the several shallow water
specimens tested were chlorophyll positive fas
demonstrated by acetone extracts of tissues).
Parasitic barnacles of the genus Acasta are also
common within this sponge.

REMARKS

This species is immediately differentiated from
other tropical Australasian Cymbastela species by
the emphasis of its plumose skeletal architecture
over other skeletal components, In other species
the reticulate connections within the skeleton are
more obvious, and overall skeletal architecture is
predominantly plumo-reticulate, but in C.
stipitata the reticulate component of the skeleton
is masked slightly by the dense plumose tracts. In
this regard the species is most similar to C. mar-
shae sp.nov. from the Houtman-Abrolhos Is, WA,
which has much smaller spicules (Table 1), In
some respects it also resembles the northwestem
Australian C. vespertina, which is found in
deeper, less: turbid waters. The latter species,
however, has much thinner lamellae and is usually
evenly pigmented maroon-brown to purple-grey
alive. Cymbastela stipitata is contrasted further
with other species below.

Cymbastela vespertina sp.noy.
(Figs 6-8, Table |)

Pseudaxinyssa n.sp. Hooper ci al., 1992: Table L, Fig
3b.

MEMOIRS OF THE QUEENSLAND MUSEUM

MATERIAL EXAMINED

HoLoTyPe: NTM 23939 (fragment QM G300010): N
side of Cumberland Strait, Wessel Is, NT, 11°27.6'S,
136°28.7'E, 32m depth, 15S.xi.1990, coll. J.N.A.
Hooper, SCUBA (stn JH-90-028). PARATYPES: NTM
Z2615: East Point Aquatic Life Reserve, Darwin, NT,
12°24.5°S, 130°48.0°E, 9-12m depth, 3.iv.1986, coll.
1.N.A. Hooper, SCUBA (stn EP28). OM G300002
(fragment NTM 73948): S of W headland, Rimbia I.,
Cape Wessel, Wessel Is, NT, 11°0.5'S, 136°43.8’E,
15m depth, [6.xi.1990, coll. J.N.A. Hooper, SCUBA
(stn JH-90-030).

OTHER MATERIAL: (All material collected by the author
unless otherwise indicated). NORTHWEST SHELF
REGION. WA: NCL Q66C-1323-F (fragment NTM
23470): Offshore from Norcape Lodge, Learmonth,
Exmouth Gulf 21°56.6°S, 114°09,1"E, 6m depth,
(7, vili.1988, coll, NCI, SCUBA. DARWIN REGION, NT:
NTM Z2696: Bast Point Aquatic Life Reserve.
12°24.5°S, 130°48.0°E, 9-12m depth, 3.iv.1986, coll,
1.N.A. Hooper, SCUBA (stn EP28), NTM 22699: East
Point Aquatic Life Reserve, 12°24.5°S, 130°48.0°E,
9-|2m depth, 3,1v.1986 (stn EP28). ARAFURA SEA, NT:
NTM Z3069, NCI-Q66-0519-F: Parry Shoals,
11°11,72'S, 129°43,26'E, 16m depth, 12. viii. 1987, coll.
A.M. Mussig, SCUBA (sm AM 87-2). Wesse. Is-
LANDS, NT: NCI Q66C-4828-0: S of W headland,
Rimbija 1. Cape Wessel. 11°0,5'S, 136°43,8°E, [5m
depth, 16.21.1990, call. NCT, SCUBA (stn JH-90-030),

SUBSTRATE AND DEFTH RANGE

Patch mock reef, fnnging coral reef, or rock
heads, sandy bottom over hard corals, gorgonians,
sponges and algae, in areas of strong current,
6-32m depth range.

GBOGRAPHICAL DISTRIBUTION
Northwest coast of Australia, from Exmouth
Gulf to Wessel Islands (Fig. 8C)-

ETYMOLOGY

Latin, vespertinis, Western, or of the evening;
for the west Australian coastal! distribution of this
species.

DESCRIPTION

Shape: Thin-walled, lamellate, cup shaped
sponges, 95-190mm high, 70-170mm maximum
diameter, lamellae 2-4.5mm thick. Lamellae typi-
cally do not rejom completely, usually producing
asymmetrical cups. Apical margins of lamellae
usually even, but larger specimens may consist of
2-4 concentric cups, with lamellae forming spiral
walls. Sponges attached to substrate by large, flat
basal plates, up to Simm maximum diameter, with

LAMELLATE CORAL REEF SPONGES

FIG. 6. Cymbastela vespertina sp.nov. A, holotype (NTM Z3939). B, paratype (NTM Z2615), in situ.

111

112

MEMOIRS OF THE QUEENSLAND MUSEUM

LAMELLATE CORAL REEF SPONGES 113

AY

SN
\ NN

7
oS
2
————————— ———-
===

ay }}
VY WI
| LG,

> YY
KOS
oe

, iy eg
<A

ty
4 a>

Lz, Ss
7 —

100 pum

be
Wass —S

—— ic
S

0

FIG. 8. Cymbastela vespertina sp.nov. (holotype NTM Z3939). A, spicules. B, section through peripheral skeleton.

C, known distribution.
FIG. 7. Cymbastela vespertina sp.nov. (paratype QM G300002). SEMs of skeleton showing: A, section through
lamella (scale=1mm); B, slightly compressed plumo-reticulate axial arrangement (scale=500.m); C, oxeas and

terminations (scale=100p.m).

114

short, thick stalk, 14-36mm long, 14-28mm
diameter (Fig. 6A,B).

Surface features: Smooth, even, uninterrupted
by any surface processes. Microscopically surface
with minute conules. Oscules not seen, but small
evenly distributed pores, up to 0.8mm diameter,
scattered over surface of lamellae.

Colour: Live colouration dark maroon brown
(Munsell 2.5R 4/4) to greyish purple (SR 5/2),
always evenly pigmented, whereas in ethanol
colour usually dark brown or dark grey.

Texture: Firm, slightly compressible texture,
sandpaper-like to touch.

Ectosome: Membraneous, without specialised
spiculation, but with regularly spaced plumose
brushes, composed of 3-8 oxeas, from ascending
extra-axial tracts, protruding through surface for
short distances, up to 100m from ectosome.
Position of plumose spicule brushes on surface
coincides with minute surface conules. Subec-
tosomal region predominantly plumose, slightly
plumo-reticulate, with ascending plumose portion
of skeletal tracts greatly emphasised over uni- or
paucispicular reticulate, transverse, connecting
tracts. Spongin fibre mesh size in peripheral
skeleton up to 140j.m diameter, more open than
in axial region (Figs 6,7).

Choanosome: Skeleton slightly axially com-
pressed, plumo-reticulate, with plumose com-
ponent emphasized only slightly over reticulate
component of skeleton: former consisting of mul-
tispicular ascending tracts, latter with
paucispicular, connecting tracts, both enclosed
within spongin fibres (Fig. 8B). Spongin fibres
light but well developed, with small mesh sizes,
up to 901m maximum diameter. Mesohy] with
moderate quantities of brown-pigmented col-
lagenous spongin, and only few spicules dispersed
outside fibres. Choanocyte chambers oval, 28-
90..m diameter.

Spicules (refer to Table 1 for dimensions):
Megascleres oxeas without any modifications,
relatively long, slender, slightly curved centrally,
tapering to sharp fusiform points, with very few
examples bearing telescoped points (Fig. 8A).

ASSOCIATIONS

This species is usually associated with sym-
biotic microalgae (cyanobacteria) in shallow
water (i.e. is chlorophyll positive), and the sym-
bionts are probably responsible for producing the
characteristic dark purple-mauve live pigmenta-
tion, although one specimen (Z3939) was pale
beige in situ and may have shed its algae. Parasitic
barnacles are rare.

MEMOIRS OF THE QUEENSLAND MUSEUM

REMARKS

This species was originally thought to be merely
a modified, deeper-water form of C. stipitata, as
there are similarities in skeletal architecture and
spicule dimensions. However, morphological dif-
ferences described above, particularly the asym-
metrical cup shape, the very thin lamellae and
darker pigmentation, and differences in free
amino acid profiles between C. vespertina and C.
Stipitata (Hooper et al., 1992), indicate that these
two sympatric populations are heterogenous and
sibling species.

Cymbastela concentrica (Lendenfeld)
comb.nov. (Figs 9-11, Table 1)

Antherochalina concentrica Lendenfeld, 1887:788,
pl.22, fig.42.

MATERIAL EXAMINED

LECTOTYPE (here designated): AM Z1993: Port Molle
(now Airlee Beach), Qld, 20°13’S, 148°49’E, no other
collection details known. (cf. published locality of St
Vincent Gulf, South Australia). PARALECTOTYPES:
BMNH 1886.8.27.451, 460: same locality.

OTHER MATERIAL. QUEENSLAND - NTM Z3169 (frag-
ment QM G300009): Outer slope of reef front, Blue
Lagoon, Lizard I., Great Barrier Reef, 14°40’S,
145°48’E, 10-20m depth, 1987, coll. A.W.D. Larkum
(ref.no. PC 348T, PC 346). QM G300003: Inner Gneer-
ings, Moreton Island, 26°38.5’S, 153°09.5’E, 14m depth,
01.v.1991, coll. Thorogood, JA, SCUBA (ref.JT-8).

SUBSTRATE AND DEPTH RANGE
Inshore fringing reefs and coral reef slope, up to
20m depth,

GEOGRAPHICAL DISTRIBUTION
Northern, central and southern Queensland (Fig.
11C).

DESCRIPTION

Shape: Cup- or vase-shaped growth forms, up
to 150mm high, 125mm maximum width, consist-
ing of relatively thin (approximately 2mm wide),
convoluted, conical lamellae or symmetrical cups
on short basal stalk. Stalk length up to 45mm long,
25mm diameter. Lamellae may produce con-
centric whorls inside cup, or merely contain a few
irregular bumps; margins of lamellae slightly con-
voluted (Fig. 9A-C).

Surface features: Lamellae smooth, slightly ir-
regular, with low conules and ridges on exterior
surface, more even interior surface; no buttresses
or external processes.

LAMELLATE CORAL REEF SPONGES 115

FIG. 9. Cymbastela concentrica. A, Paralectotype (BMNH 1886.8.27.451). B, specimen (NTM Z3169; photo A.
Larkum). C, field number PC 346, unaccompanied by voucher specimen (photo A. Larkum),

116 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 10. Cymbastela concentrica, A, SEM of skeleton (specimen NTM Z3169) (scale=500\.m; left magnified 21.2
times, nght magnified 186 times). B, peripheral skeletal architecture (scale=100p.m). C, spicule with telescoped
ends (lectotype AM Z1993) (scale=50j1m),

118

Colour; In life, dark brown (interior) (Munsell
5YR 3/2) or olive brown (exterior) (2.5 YR 4/4).
Chlorophyll pigments abundant (seen in acetone
extracts of freshly preserved material}, duc to
presence of large population of cyanobacteria
(microalgal) symbionts.

Texture: Smooth, suff and flexible consistency,
with a slightly velvet surface texture.

Ectosome: Optically smooth, lacking special-
ized skeleton, but microscopically villose, with
substantial portions of spicule from penpheral
skeleton protruding through surface and forming
plumose brushes. Spicules protrude up to 230um
from surface, and brushes may contain up to §
spicules (Figs 10, 11), Microalgae, seen in fresh
material, appear to be concentrated near periphery
of skeleton,

Cheanoseme: Skeletal architecture plumo-
reticulate, with poorly differennated axtal and
extra-axial skeletal regions - reticulate skeleton
emphasised over plumose portion (Figs 10, 11B).
Central region of choanosomal skeleton only
slightly axially condensed, predominantly reticu-
late, with plumose paucispicular ascending tracts
curving gradually towards surface, interconnected
by uni- or paucispicular transverse tracts, whereas
peripheral skeleton more clearly diverges into
plumose ascending multi- or paucispicular tracts,
interconnected by uni- or paucispicular transverse
tracts. Spongin fibres and spicule tracts, 40-60p.m
wide in axis, become less heavily cored and more
widely spaced towards periphery, 20-40..m wide,
ultimately forming distinctive plumose dermal
brushes, Spongin fibres (22-60j4m wide) lightly
developed, nearly fully cored by megascleres,
producing ovoid to eliptical meshes (42-77p.m,
60-185p.m wide in axial and extra-axial regions
respectively). Collagen abundant in mesohy] but
very lighlly pigmented. Choanacyte chambers
oval to elliptical, 45-150j.m diameter,

Spicules (refer to Table 1 for dimensions): Van-
able length. relatively, slender, fusiform oxews,
moderately curved at centre, usually with yery
faintly telescoped points (Fig. 11A).

ASSOCIATIONS

Cyanobacteria Were only sten in freshly
preserved material, not in the type material, and
appears to be concentrated mainly in the
peripheral skeleton,

REMARKS

Lendenfeld (1887) states that C. concentrica was
originally collected from St Vincent Gulf, South
Australia, Whereas both the AM und BMNH

MEMOIRS OP THE QUEENSLAND MUSEUM

registers, and the specimen labels with the type
material, indicate that the collection locality was
Port Molle, Queensland. Although the exact type
locality of this species will always be doubtful, the
new record of the Queensland material described
here from Lizard and Moreton Is suggests that the
Museum register notations may be correct. Ac-
cording to Burton (1934:558), and notes in the
BMNH register (in M. Burton’s handwniting), this
species has some similarities with Phakellia flabel-
Igta (Carter) from southem Australia. However,
the skeletal architecture and spiculation of C. cor-
centrice is quite different from that of Phakellia,
and the two species are regarded as convergent in
these two superficially prominent features,

The redescription presented above differs
slightly from the description given by Lendenfeld
(1887), who recorded oxea dimensions of
130x5jyum_ Lo this regard his descnptien conforms
more closely to the new specimens described
above than to the type material (Table 1). Apart
from their smaller spicule dimensions these two

specimens are identical to the type material in
growth form, including lamellae thickness and
surface features, skeletal architecture, spicule-
spongin fibre structure and spicule geometry, and
it is suggested here that all five specimens of C.
concentrica described above are conspecific,
However, the existence of these two anomalous
specimens casts some doubt over the accuracy of
absolute spicule dimensions as a useful diagnos-
tic character within this genus (sec Fig, 24), sug-
gesting that spicule size may vary with latitudinal
distribution,

Cyinbostela concentrica is similar to most of the
other lamellate Australasian Cymbasrela species
in a number of features. Many species contain
symbiotic algae (i.e. are chlorophyll positive),
they all have a thin, lamellar, plate-like, eup- or
vase-shaped growth form, with well developed
spongin fibres, and a more-or-less slightly con-
densed plumo-reticulate axial skeleton, plumose
of plumo-reticulate extra-axial skeleton, and if
present, a plumose ectosomal skeleton. Specics
appear to differ substantially only in spicule
dimensions, spicule geometry (presence or ab-
sence of telescoped points, and possession of has-
tate of fusiform points), and the degree to which
the skeleton is organized into axial and extra-axial
sections. For example, C. stipitara (from inshore
waters of northern Australia) has larger
megascleres. spicules are only rarely terminated
by telescoped points, and there is a more obvious
differentiation between the axial and extra-axial
regions of the skeleton, including a more con-

LAMELLATE CORAL REEF SPONGES

densed axial core, and an emphasis on the plumose
portion of the skeleton. Cymbastela cantharella
from the outer reefs of New Caledonia also has a
stalked lamellar growth form, slightly differen-
tiated axial and extra-axial skeletons, but it has
distinctive orange, yellow-orange live coloura-
tion. and relatively slender oxeas, In contrast,
Cymbastela concentrica has a relatively poorly
differentiated axial and extra-axial skeleton, only
slightly condensed choanosomal axis with em-
phasis on the reticulate extra-axial portion of the
skeleton. This takes the form of a plumo-reticu-
late extra-axial skeleton, a plumose ectosome, and
oxeas which are similar in size to those of C.
cantharella.

(Cymbastela cantharella (Lévi)
(Figs 12-14, Table 1}

Preudaxinyssa cantharella Lévi, 1983;719-722, fig. 1.
pl.l-

MATERIAL EXAMINED

Hovoryre: MNHN LBIM DCL3141: Onter reef, SW
coast New Caledonia, 40m depth, no other details
known.

OTHER MATERIAL: NTM 23869: Precise locality un-
known, Noumea lagoon, SW coast, New Caledonia,
22°20'S, 166°13"E, 25m depth, coll. ORstom, SCUBA
(ref. R1261). NTM 23872 (fragment OM G300004):
Exterior of Grand Recif Mbere, off Noumea, New
Caledonia, 22°20'S, 166°13’E, 30m depth, 21.ix.1990,
call, J.N.A. Hooper, SCUBA (sin JH-90-018),

SUBSTRATE AND DEPTH RANGE
Slope and ledges near base of barrier reef, dead
coral and silt substrates, 15- 75m depth.

GEOGRAPHICAL DISTRIBUTION
Known only from the New Caledonia region
{Fig. 14€).

DESCRIPTION

Shape: Short, erect cup-shaped or semi-
vasiform lamellae, 80-150mm high, up te 170mm
maximum diameter, on short stalk, 3040mm
long, 10-17mm diameter, with thin lamellate
walls, 3-6mm diameter, usually with convoluted
mnurgins, occasionally with secondary cups or
lamellae growing inside primary cup, often with
buttresses and secondary projections (Fig. 12A).

Surface features: Surface of lamellae
predominantly smooth, although distinct differen-
ces between interior (inhalant) and exterior faces
of cups exist. Exterior of lamellae microscopically

119

smooth, optically uneven, interior of cup wath
numerous small oscules, 0.5-2mm diameter, each
surrounded by slightly raised membraneous lip
(Fig. 12B). Oscules approximately 2mm apart,
throughout interior surface of lamellae producing
concentric striations within interior of cup.

Colour: Pale: orange alive (Munsell 2.5-7,5 YR
8/10), beige in ethanol.

Texture: Firm, flexible, slightly compressible;
exterior of lamellae with sandpaper like texture,
intezior more compressible.

Ectosome; Membraneous, without specialised
skeleton, with heavy layer of slightly darker col-
lagen. up to 80pm wide, through which
choanosomal oxeas protrude for about hald their
length. Projecting oxeas occur individually or in
paucispicular plumose bundles, regularly dis-
persed across surface, arising from ascending
radial tracts in peripheral skeleton (Figs 13, 14B).

Choanesome: Choanosomal skeleton plumo-
reticulate, wilhoul axial compressien or any
noticeable differentiation between axial and cxtra-
axial regions, apart from plumose ectosomal
spicules (Fig. 14B). Two components of skeleton
predominant: first, radial spongin fibres, cored by
muitispicular tracts of oxeas, run longitudinally
through lamellae, and gradually ascend to surface,
becoming plumose and diverging near periphery;
second, radial fibres interconnected by uniy or
paucispicular tracts of oxeas, only weakly or-
ganised as a reticulum. The skelcton in places is
almost halichondroid (Fig, 13). Spongin fibres
only lightly invested in spongin, up to 75pm
diameter, forming elongate orrectangular meshes,
up to 125m diameter, with small oval
choanocyte chambers, 25-40pm diameter,
Mesohyl! ‘heavily reinforced with collagen and
containing scattered small (possibly juvenile)
spicules.

Spicules (refer to Table 1 for dimensions):
Oxeas relatively short, slender, slightly curved at
centre, occasionally asymmetrical, tapering,
fusiform, sharply pointed but usually with very
slightly telescoped points at absolute apex of
spicule (Fig. 144).

ASSOCIATIONS
No symbiotic cyanobacteria were seen in any of
the three specimens examined.

REMARKS

Cymbastela cantharella is apparently unlike all
other species of the genus in that it contains u
number of biologically active metabolites (Lévi,
1983, and pers.comm.), although the nature of this

120

chemical activity is not yet published. It also dif-
fers from other tropical species in lacking any
evidence of symbiotic microalgae, and in this
respect it is similar to the two temperate species
described here. Cymbastela cantharella has other
features typical of the genus, especially growth
form - being thinly flabellate with irregular, but-
tressed surface processes similar to those seen on
C. stipitata, and the geometry of the oxeas is also
similar in these two species. However, the New
Caledonian species differs from other Cymbastela
in external colouration (orange alive, possibly due
to the lack of photosynthetic pigments found in
other tropical species), prominent surface
sculpturing on the oscular face of cups, skeletal
architecture (having a dense radial-plumose
skeleton, with a secondary paucispicular secon-
dary reticulate skeleton, with an overall nearly-
halichondroid appearance, and radial-plumose
ectosomal skeleton), and specific dimensions of
oxeas (Table 1).

MEMOIRS OF THE QUEENSLAND MUSEUM

Cymbastela coralliophila sp.nov.
(Figs 15-17, Table 1)

Pseudaxinyssa sp.; Wilkinson, 1983:411.
Pseudaxinyssa n.sp.; Bergquist et al., 1980:424, Table
1; Lévi, 1983:720.

MATERIAL EXAMINED

HOLOTYPE: NTM Z4038: S side of Blanchard Reef,
Qld, 13°23.4’S, 143°44.4’E, 14m depth, 12-xii-1990,
coll. J.N.A. Hooper, USSR RV ‘Akademik Oparin’,
SCUBA (stn JH-90-054).

OTHER MATERIAL: GREAT BARRIER REEF, QLD - FAR
NORTHERN SECTION: NTM Z4039: S side of Blanchard
Reef, 13°23.4’S, 143°44.4’E, 14m depth, 12-xii-1990,
coll. J.N.A. Hooper, USSR RV ‘Akademik Oparin’,
SCUBA (stn JH-90-054). NCI Q66B-2242-Y: W side
Canoe I., 10°20.6’S, 142°06.4’E, 4m depth, 6.v.1987,
coll. NCI, SCUBA. - CAIRNS SECTION: NTM Z3165,
3166, 3167, 3168 (fragments): Outer slope, reef front,
Blue Lagoon, Lizard I., 14°40’S, 145°28’E, 10-20m

FIG, 12. Cymbastela cantharella. A, specimen (NTM Z3872). B, close up view of interior surface of cup (specimen

NTM Z3869),

LAMELLATE CORAL REEF SPONGES 121

fs - f
= ws

FIG, 13. Cymbastela cantharella. A, SEM of skeleton (specimen NTM 23872) (scale=500j.m; left magnified 106
times, right magnified 566 times). B, peripheral skeletal architecture (scale=100j.m). C, oxeas, including juvenile
form (specimen ORSTOM RA1279) (scale=100.m).

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LAMELLATE CORAL REEF SPONGES

depth, 1987, coll. A.W.D. Larkum et al. (ref. PC341T,
PC342T, PC344T, PC345T, respectively). NCI Q66C-
0260-W: N end Bowden Reef, off Townsville,
19°2.06’S, 147°55.89°E, 14m depth, 3.ii.1987, coll.
NCI, SCUBA. - CENTRAL SECTION: AIMS RA3-PR
(fragments NTM Z2735, QM G300012): Davies Reef,
Townsville region, 18°50’S, 147°39’E, depth unknown,
17.viii.1982, coll. C.R. Wilkinson (stn Don.214), -
CAPRICORN SECTION: NCI Q66C-1766-L (fragments
NTM Z3509, QM G300008): 100m NW of small rocky
point at embayment edge, NW Brampton Island, Whit-
sunday Islands Group, Mackay region, 20°48.0’S,
149°15.0°E, 12m depth, 07.xi.1988, coll. NCI, SCUBA
(stn DON-0272). NCI Q66C-0816-A: W end Deloraine
I., Whitsunday Is Group, Mackay region, 20°09’S,
149°04’E, 20m depth, 10.xi.1987, coll. NCI, SCUBA.

SUBSTRATE AND DEPTH RANGE

Coral reef substrates, including lagoon, sloping
fringing reef, irregular coral boulders with muddy
sand between, gorgonian beds, and also areas of
high sedimentation. Depth range extends from
subtidal to 20m.

GEOGRAPHICAL DISTRIBUTION
Northern, central and southern sections of the
Great Barrier Reef, Queensland (Fig. 17C).

ETYMOLOGY

The species name refers to the restricted distibu-
tion of all material so far described, the sponge
being found exclusively on coral reefs.

DESCRIPTION

Shape: Large vase-shaped, cup-shaped or simp-
ly lamellate growth forms, usually with large mar-
ginal opening, or sometimes nearly flattened
completely against substrate, 80-170mm high, up
to 350mm maximum marginal diameter. Usually
attached to substrate by short stalk, up to 30mm
high, 25mm diameter, sometimes lamellae are
direct outgrowths of the basal attachment. Lamel-
lae relatively thick, lobate, 8-15mm thick exclud-
ing lateral projections (Fig. 15).

Surface features: External, exhalant, surface op-
tically smooth, relatively even, with or without
bumps, ridges and lateral growths. Internal, in-
halant, surface relatively uneven, often with con-
centric striations and ridges, or with irregular
bumps, lobate growths and depressions on lamella
walls, typically with small digitate or lobate
projections at centre of cup. Occurrence and
development of surface outgrowths, on both inte-
rior and exterior surfaces of lamellae is highly
variable. Oscules small, 0.5-2mm diameter, dis-

123

persed on tips of microconules or ridges on inte-
rior surface of cups, subdermal canals visible
below slightly translucent ectosome (Fig. 15C-D).

Colour: Live colouration typically mottled
olive-brown (Munsell 7.5YR 5/4), olive-green
(5Y 6/6) with maroon-brown tinge externally (SR
4/2), green (7.5GY 3/4) or olive-green internally
(2.5GY 5/4). Colour in ethanol white or greenish-
white.

Texture: Compressible, difficult to tear, rough
exterior texture.

Ectosome: Ectosome microscopically hispid,
with specialised skeleton. Subectosomal skeleton
ascending, plumose, produced by pauci- or multi-
spicular tracts of larger, choanosomal oxeas run-
ning at right angles to surface, overlain by
specialised ectosomal skeleton of tangential, mul-
tispicular brushes made up of smaller ectosomal
oxeas. Specialised dermal skeleton markedly
plumose, mostly orientated tangentially or
paratangentially to surface (Figs 16, 17B).

Choanosome: Choanosomal skeleton with only
slight axial condensation, with axial and extra-
axial components only slightly differentiated (Fig.
17B). Axial region predominantly radial-
plumose, less markedly plumo-reticulate, with
skeletal lines mostly directed longitudinally
through lamellae, but gradually ascending
towards peripheral skeleton; extra-axial skeleton
more-or-less radial-plumose, with multispicular
ascending spongin fibres interconnected by
paucispicular transverse fibres (Fig. 16). Spongin
fibres well formed but only very lightly invested
with spongin, 75-110,.m diameter. Fibre reticula-
tion forms elongate oval to elliptical meshes, 85-
350um diameter, relatively similar in diameter
between axial and extra-axial regions; choanocyte
chambers relatively large, oval, 90-210um
diameter. Ascending fibres cored by multispicular
tracts of larger choanosomal oxeas, whereas
transverse fibres uncored or with uni- or
paucispicular skeletal lines. Mesohy]l matrix rein-
forced with moderate quantities of collagen. Very
thin, smaller oxeas, similar in length to those on
surface skeleton are dispersed throughout the
mesohyl, possibly juvenile forms of larger
choanosomal oxeas.

Spicules (refer to Table 1 for dimensions):
Choanosomal oxeas thick, slightly curved at
centre, typically symmetrical, occasionally
anisoxeote, with hastate-rounded or slightly
pointed, usually very faintly telescoped ends, Ec-
tosomal oxeas thin, mostly straight, fusiform, with
sharply pointed, rarely telescoped ends (Fig.
17A).

MEMOIRS OF THE QUEENSLAND MUSEUM

124

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UL ‘aeT [aE] JO SaoRpANs ((q) JOUA\UT puE (5) JoLMalxa Jo MatA dn asoja Surmoys “(CELZ7Z WLLN) weuttoads ‘q-D ‘aleis AAT] UI av][aWE] Jo soRyins
JOUAUI SuTMOYs (991 €Z W.LN) Ueuttoads *g ‘(uosuTyTIAA ‘DO oloyd !¢EgLTZ WLLN) nus ur uaurtsads “y ‘aouds npydoijosos njaispquida ‘S| “OIA

LAMELLATE CORAL REEF SPONGES 125

FIG. 16. Cymbastela coralliophila sp.nov. A, SEM of skeleton (specimen NTM Z2735) (scale=500pm; left
magnified 19.5 times, right magnified 93 times). B, peripheral skeletal architecture (specimen NTM Z3509)
(scale=500p.m). C, two sizes of spicules (scale=100j.m).

126 MEMOIRS OF THE QUEENSLAND MUSEUM

100 pm

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eee ae Cymbastela coralliophila rope vet ay SNe a 735). A, choanosomal and ectosomal spicules. B,
n thro ust pe ciphers ral skelet mC kn tributi

LAMELLATE CORAL REEF SPONGES

REPRODUCTION
Oviparous, with synchronous release of eggs
and sperm (A. Ayling, pers. comm.).

ASSOCIATIONS

Sponges tested chlorophyll positive, and ir is
likely that the species’ live pigmentation is partial-
ly or entirely produced by cyanobacterial sym-
bionts. Several specimens also had numerous
zoanthids on upper and lower surfaces, and bar-
nacles (Acasta sp.) embedded within the mesohyl.

REMARKS

Cymbastela coralliophila has been recorded fre-
quently from the Great Barrier Reef. but until now
it has remained undescribed.

The species confonns with Burton's (1931) in-
terpretation of the genus Aximyssa, as distinct from
his nominal genus Pseudaxinyssa, in that it con-
tains two distinct size classes of oxeas, the smaller
found predominantly on the surface, the larger
forming the structural skeleton, On the othet hand,
the very close similarities in growth form, skeletal
architecture and spiculation that exist between this
species and the other Australasian lamellate Cym-
bastela suggests that this feature is not important
above the species level of classification, Cymbas-
tela coralliophila also differs from other members
of the genus, as construed here, in having an
almost flattened, usually relatively large cup-
shaped growth form, with flabellate lamellae. The
skeletal structure is most similar to C. cantharella,
with the cadial-plumose skeletal tracts
predominant over the reticulate component of the
skeleton, and with only minordifferences between
the axial and extra-amal regions. The species are
copirasted Further in Table 1.

Cymbastela marshae sp.nov.
(Figs 18-20, Table 1)

MATERIAL EXAMINED

HoLotype: NTM 22876: Northern edge of Pelsar Is-
lets, Houtman-Abrolhos Is, WA, 28° 47.6'S, 114°
00,7°R, 25m depth, 09,vii.1987, coll, J.N.A. Hooper,
USSR RV ‘Akademik Oparin’, SCUBA (stn JH-87-
OO!).

OTHER MATERIAL: HOUTMAN-ABROLHOS Is, WA -
NTM Z2901 (fragment QM G300005): Pelsar Isicts,
28°47.2°S, 113°58.5°E, 22m depth, 10.vii, 1987, coll.
1,N A. Hooper, USSR RV ‘Akademic Oparin', SCUBA
(stn JH-87-002). NCI Q66C-4198-F: 1,5 km N of north-
em point, West Wallabi I., Wallabi Group, 28°25,7°S.
113°42.0'E, 8m depth, 12.ix.1990, coll. NCI, SCUBA.

SUBSTRATE AND DEPTH RANGE

Growing at base and on reef slope of Acropora
and Seriatopora coral reef, in sand, silt or rock
substnvles, in Turbinaria zone, §-25m depth.

GEOGRAPHICAL DISTRIBUTION
Known only from the Houtman-Abrolhus Is-
jands, WA (Fig, 20C).

ETYMOLOGY

Named after Mrs Loisette Marsh, Curator of
Marine Invertebrates, Western Australian
Museum, Perth, in respect of her dedication ta the
study of the marine fauna of Wester Australia

DESCRIPTION

Shape: Thin, irregularly Jamellate lobes forming
asymmetrical vases, or symmetrical cup-shaped
sponges, 125-255mm high, 120-235mm wide at
apex of lamellae, with thin lamellae, 2-6mm max-
imum width, attached directly to substrate by a
basal attachment, without a basal stalk (Fig. 18)-

Surface features: Smooth, relatively even sur-
face. Exterior surface with few low, irregular
bumps, shallow depressions and concentric
ridges, with er without horizontal lamellate
projections growing on side of exterior surface.
Interior, oscular surface of lamellae smooth, with
concentric ridges, few low. irregular conules or
depressions, sometimes with secondary lamellac
or whole cups growing within interior surface.
Very small oscules scattered over interior of cups,
up lo 1.5mm diameter.

Colour: Olive-green and maroon alive (Mun-
sell SY 5/4 and 2.5R 4/8). evenly pigmented on
exterior and interior surfaces, beige in ethanol.

Texture: Firm, flexible, slightly compressible
texture, smooth to touch, difficult to tear.

Ectosome: Membraneous, without specialised
skeleton, with plumose brushes of choanosomal
oxeas protruding through surface for up fo
170m, consisting of 2-8 spicules per bundle.
Ectosame with thin layer of darker pigmented
granular collagen, 40-70um wide, with heavy
concentrations of microalgae, and also with a thin,
disorganised paratangential layer of choanosomal
oxeas scattered between erect, protruding ec-
tosomal brushes. Subectosomal skeleton cavem-
ous, predominantly plumose, slightly reticulate,
with reticulate component decreasing closer to
periphery (Figs 19, 20B).

Choanasome; Choanosomal skeleton plumo-
reticulate, with axial skeleton dominated by lon-
gitudinal tracts running through lameillac,
gradually ascending towards surface, becoming

FIG. 18. Cymbastela marshae sp.nov. A, holotype
(NTM Z2876), in sita. B-C, specimen (NTM 22901)
in situ (B) and preserved (C).

nearly plumose near peripheral skeleton (Fig.
20B). Axial skeleton slightly condensed at centre
of lamella with reasonably well differentiated
axial and extra-axial regions. Spongin fibres well
developed, with obvious differentiation between
multispicular primary (longitudinal and ascend-
ing) and uni- or paucispicular secondary
(transverse, connecting) elements (Fig. 19).
Primary fibres 30-80j.m diameter, rarely fully
cored with megascleres, secondary fibres much

MEMOIRS OF THE QUEENSLAND MUSEUM

smaller, 15-25j:m diameter, usually with greater
silica to spongin ratio. Fibre meshes elongate oval
or diamond-shaped in axis, 70-105,.2m maximum
diameter, generally more cavernous in periphery,
up to 150m maximum diameter. Choanocyte
chambers small, up to 454m diameter. Mesohy]
matrix only lightly invested with collagenous
spongin, with abundant microalgae scattered
throughout.

Spicules (refer to Table 1 for dimensions):
Oxeas relatively short, slender, slightly curved at
centre, with tapering, fusiform, usually telescoped
points (Fig. 20A).

ASSOCIATIONS

Although not analysed chemically (acetone ex-
tracted), the olive-green and maroon pigments of
this species are probably due to the presence of
cyanobacteria scattered within the mesohyl. Bach
of the specimens examined was also found to have
several barnacles boring within the lamella wall,
near the base of the sponge.

REMARKS

This species is most closely related to C.
stipitata in skeletal structure, as the plumose lon-
gitudinal and ascending primary tracts dominate
the skeletal architecture. Both species have con-
densed axial regions and differentiated axial and
extra-axial skeletons, with well developed
plumose ectosomal skeletons, Cymbastela mar-
shae also has a cavernous subdermal region, and
spicules are about half the dimensions of those in
C. stipitata (Table 1). The present species is close
to C. coralliophila in growth form, lamella thick-
ness and surface pigmentation

Cymbastela notiaina sp.noy.
(Figs 21-23, Table 1)

MATERIAL EXAMINED

HOLOTYPE; SAM TS4025 (fragments NTM Z1623a,
QM G300007): Precise locality unknown, Gulf of St
Vincent, SA, 34°33-45'S, 137°57'-138°10'E, depth and
date of collection unknown,

SUBSTRATE AND DEPTH RANGE
Unknown.

GEOGRAPHICAL DISTRIBUTION
South Australia (Fig. 23C).

ETYMOLOGY
Greek, notos, south; notiaina refers to the
southern distribution of this species.

LAMELLATE CORAL REEF SPONGES 129

FIG. 19. Cymbastela marshae sp.nov. A, SEM of skeleton (specimen (NTM 22901) (scale=lmm). B, peripheral
skeletal architecture (scale=!00j.m). C, oxeas (scale=100,.m).

100 pm

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 20, Cymbastela marshae sp.nov. A, spicules. B, section through peripheral skeleton, C, Known distribution,

DESCRIPTION

Shape: Small, symmetrical vasiform sponge,
96mm high, 125mm maximum diameter at apex
of cup, on short thick stalk with expanded basal
attachment, 12mm long, 35mm diameter, and thin
lamellae, 3-Smm thick (Fig. 21).

Surface features: Interior and exterior surfaces
optically even, without conules, buttresses. or
other lateral growth projections. Exterior surface

has longitudinal, concentrically striated ridges,
Oscules minute, less than Imm diameter, dis-
persed over interior surface of cup.

Colour; Live colouration unknown, olive-
brown in ethanol.

Texture: Firm, flexible, compressible in
preserved state, smooth to touch.

Ectosome: Ectosome optically smooth,
membraneous, slightly hispid due to protruding

LAMELLATE CORAL REEF SPONGES

spicule brushes, shghtly microconulose, with
mucroconules corresponding to surface brushes,
Ectosomal skeleton distinctly plumose, with
bundles of 2-15 choanosomal spicules protruding
up fo 90j.m from surface, perched on ends of
ascending extra-axial skeletal tracts (Figs 22,
23B).

Choanosome: Choanosomal skeleton with no
axial condensation, and only slightly differen-
uated axial and extra-axial regions (Fig. 23B).
Skeleton almost regularly reticulate at the core,
with oval meshes, 70-120|.m diameter, formed by
multispicular spongin fibres, both longitudinal
and connecting. Skeleton plumo-reticulale
towards periphery, with ascending multispicular
fibres predominant over uni- or paucispicular
transverse connecting fibres, Spicules form
plumose brushes close to, and protruding from,
surface, Fibre meshes in peripheral skeleton elon-
gate-rectangular, slightly larger than jn axis, 100-
140um diameter (Fig. 22). Spongin fibres
relatively large, 40-85j:m diameter, partially but
never fully occupied by spicules. Mesohyl matrix
lightly reinforced with collagen, choanocyte
chambers oval, 40-90,.m diameter.

Spicules (refer to Table | for dimensions):
Oxeas almost vestigial by comparison with other
specics, poorly silicified, very thin, small, only
slightly curved at centre, with prominently tele-
scoped ends and rounded tips (Fig, 253A).

ASSOCIATIONS
No symbiotic cyanobacteria were seen in the
holotype.

REMARKS

This species is presently known only from the
holotype described aboye, although a small frag-
ment from a second specimen exists in the SAM
collection (SAM TS4058 [section/fragments
NTM 21665, QM G300011), from Port Noarlun-
ga, SA), may also belong to the species. Cynthas-
tela notiaina is easily differentiated from other
taxa by its regular, nearly completely reticulate
choanosomal skeleton, the small size of
megascleres (Table 1), and the telescoped,
rounded (strongylote) pojnts of oxeas, unlike any
other species.

DISCUSSION

TAXONOMY

Small differences in field characteristics, such
us live colouration, growth form, surface features,
including outgrowths on the surface and lamellae

131

thickness, as descnbed above foreach species, can
serve as field identifiers for the seven known
species of Cymbestela, However, morphological
differences seen in preserved maternal, which cor-
relate with these specics boundaries are less ob-
vious. Certainly, in terms of general skeletal
structure and spicule geometry the genus is rela-
tively hatnogenous, and it is only through sniall
but significant differences in a small number of
skeletal characters that species can be reliably
differentiated, These characters, which are shown
to be most important, include spicule length,
spicule geometry, including the presence or ab-
sence of telescoped ends on spicules, and degree
of silicification, and the relative, proportional
development of the axial and extra-axial
skeletons. On this basis it was possible to construct
a key 16 species of Cymbastela (see above), with
emphasis given to the morphology of preserved
specimens, Thus, whereas the characters used 19
differentiate species in this key may secm trivial,
these differences correspond with more obyjous
field characteristics, many of which are no Jonger
visible in the preserved material.

It was also noted that, in general, species could
be separated by their mean spicule dimensions.
Generally, the relative size of oxea megascleres
decreases with increasing latitude (Fig. 24A-B),
such that species from low (tropical) latitudes (C.
stipuata, C. eeralliophila, C, vespertina) have
larger oxeas than those species from higher
latitudes (C. cantharella, C. marshae, C. notiana).
Cymbastela_ concentrica, which has a wide
latitudinal distribution (14-26°S; Fig. 24B), also
has a relatively wide range of spicule dimensions
(Fig. 244A, symbol 5).

BIOGEOGRAPHY

Although itis possible that more extensive sam-
pling along the coal temperate Australasian
coastline will reveal additional species, Cymhas-
tela is presently known to be predominantly 2
shallow waler, topical or subtropical genus,
usually associated with coral substrates. The
venus appears lo be endemic to the Indo-west
Pacific, extending only as far as New Caledonia
(longitude 166°) to the east, and to the Houtman-
Abrolhos Islands (113,5°E) to the west, and so far
there are no confinned records of the genus from
eller Indonesia (unpublished Siboga and Sneilius
IZ collections, R.W.M. Van Soest, pers.comm,) ov
New Zealand (personal observations}. However,
iLis possible that misidentified specimens of Cyr:
bastela exist In other mascum collections. The
genus is certainly not present in the published

132

collections of Hentschel (1912), from the Aru and
Kai Islands, southeast Indonesia (ZMB collec-
tions), in which close similarities with the
northwest Australian sponge fauna can be seen
(e.g. Hooper, 1991).

Three species of Cymbastela have restricted,
allopatric distributions (C. cantharella, C. mar-
shae, C. notiaina; Fig.25), whereas on the
northwest and northeast coasts of Australia there
are two groups of more-widely distributed, sym-
patric species. Within the northwestern area of
sympatry (Fig. 25), there is some evidence of
niche separation between the two populations, C.
stipitata and C. vespertina. The former species has
a predominantly shallow water distribution, found
mostly in the intertidal region and only exception-
ally down to 19m depth, whereas C. vespertina is
found mostly in deeper water, only rarely above
8m depth. Population characteristics of the north-
east Australian sympatric species, C. coral-
liophila and C. concentrica, have not yet been

MEMOIRS OF THE QUEENSLAND MUSEUM

determined, and the latter species, in particular, is
still poorly known.

PHYLOGENY

Under Van Soest et al.’s (1990) interpretation of
the existing family Axinellidae Cymbastela would
be placed with a group of genera having axially
condensed and extra-axially plumo-reticulate
skeletons, for example Auletta, Axinella,
Axinosia, Homaxinella, Phakellia, Pseudaxinella,
Reniochalina and Teichaxinella, which Van Soest
et al. (1990) retained in a newly defined concept
of family Axinellidae. Of these genera Cymbas-
tela is most closely related to Axinella. Converse-
ly, they placed the genus Axinyssa, previously
considered to be a close relative of
‘Pseudaxinyssa’ 8.1. (= Cymbastela), in a
redefined family Halichondriidae since it has
poorly developed spicule tracts. This group would
contain ‘axinellid’ genera such as Axinyssa and
Amorphinopsis, several genera previously in-

FIG.

21. Cymbastela notiaina sp.nov. A, holotype (SAM TS4025).

FIG. 22. Cymbastela notiaina sp.nov. A, SEM of peripheral skeleton (holotype, SAM TS4025) (scale=100,.m).
B, axial skeleton (scale=100j.m). C, spicules with telescoped ends (scale=50p.m).

LAMELLATE CORAL REEF SPONGES

133

134

"\

WNT WY

en

\

)

100 ym

YOAK vw
z ays

MEMOIRS OF THE QUEENSLAND MUSEUM

: N WH IAS © JA
E YS Yr A VAR
v : ENON OK IN’
= ARGH
XW VENI HNN
=~ \ a Vics Oe
a Oaag
5, AOC AE
iat a Ps a” LAY WY /} WO}
SoS ICONS
Se?” "GC ae a
C '

VY

FIG. 23. Cymbastela notiaina sp.nov. A, spicules, B, section through peripheral skeleton. C, known distribution.

cluded in Desmoxyidae (e.g. Myrmekioderma),
and other genera traditionally associated with the
halichondrids and hymentacidonids (e.g.
Ciocalypta, Hymeniacidon), A third group of
“axinellids’ with dendritic skeletons (¢.g. Acan-
thella, Dictyonella, Scopalina), were removed to
the new family Dictyonellidae, The fourth family,
Desmoxyidae, was also previously aligned with

the Axinellida, differing from other ‘axinellids’ in
having a reticulate-fasciculate choanosomal
skeleton and a fleshy corrugated surface (Van
Soest et al., 1990:18). It contained several des-
moxyid genera (e.g. Higginsia), but excluding
Myrmekioderma, as well as some Axinellidae
(e.g. Ptilocaulis). We consider that these new
family divisions proposed by Van Soest et al.

LAMELLATE CORAL REEF SPONGES

C. coralliophila 4

200 4

SPICULE LENGTH
3
it
=-—[_—id FC tipitata
No
coe ee

a {Tk C. marshae

pal
>
o
|
nN
s[-T] C. notiaina

= C. cantharella
ww ———_ [ }—— C- concentrrica

LATITUDE (°S)

FIG, 24. A, Comparison in spicule lengths (jzm) be-
tween seven species of Cymbastela, and B, latitudinal
differences in distribution for each species. Box-and-
whisker plots show the range of spicule dimensions
(i.e. whiskers’), the median values (i.e. 50% of data
values, between lower and upper quartiles - open
boxes), and the average length of oxeas for each
species (i.e. mid-range bars). Key to species codes: 1,
C. stipitata; 2, C. coralliophila (large and small
oxeas); 3, C. vespertina; 4, C. cantharella; 5, C. con-
centrica; 6, C. marshae; 7, C. notiaina.

(1990) have some merit over the existing, con-
fused system for Axinellidae - Halichondriidae -
Hymeniacidonidae, but we are uncertain whether
all these ‘axinellids’ and ‘halichondrids’ have a
monophyletic origin, and we leave open the ques-
tion of their ordinal classification for the time
being. Further evidence from molecular and
genetic nature may be useful in evaluating these
relationships in an objective context.

ACKNOWLEDGEMENTS

We thank Peter Murphy and associates (NCI,
Townsville), Clive Wilkinson (AIMS,
Townsville) and Cécile Debitus (ORSTOM,
Noumea), for providing access to their diverse

135

collections of marine sponges from Australia and
New Caledonia; Prof. Georgy Elyakov, scientific
staff and crew of the USSR RV ‘Akademik
Oparin’ (PIBOC) for their collaborative field
work at the Houtman-Abrolhos Islands (1987);
Avril Ayling (‘Sea Research’, Daintree) for un-
published information on reproductive observa-
tions, Tony Larkum (Sydney University) for other
information on several of the new species
described here, and Rob Van Soest (ZMA,
Amsterdam) for valuable unpublished informa-
tion and comments on the manuscript. Penny
Barents (AM, Sydney), Shane Parker (SAM,
Adelaide), Shirley Stone (BMNH, London),
Claude Lévi (MNHN, Paris), Dieter Kiihlmann
(ZMB, Berlin), Klaus Ruetzler (USNM,
Washington) and Manfred Grasshoff (SMF,
Frankfurt) are thanked for providing access to type
material from their respective collections.

LITERATURE CITED

BERGQUIST, P.R. 1961. Demospongiae (Porifera) of
the Chatham Islands and Chatham Rise, collected
by the Chatham Islands 1954 Expedition. New
Zealand Department of Scientific and Industrial
Research Bulletin 139. Biological Results of the
Chatham Islands 1954 Expedition (5): 169-206.

1970. The marine fauna of New Zealand: Porifera,
Demospongiae, Part 2. (Axinellida and
Halichondrida). New Zealand Department of
Scientific and Industrial Research Bulletin. New
Zealand Oceanographic Institute Memoir 197: 1-
85, pls 1-20.

BERGQUIST, P.R. & HARTMAN, W.D. 1969. Free
amino acid patterns and the classification of the
Demospongiae. Marine Biology 3(3): 247-268.

BERGQUIST, P.R., HOFHEINZ, W. & OESTER-
HELT, G. 1980. Sterol composition and the clas-
sification of the Demospongiae. Biochemical
Systematics and Ecology 8: 423-435.

BERGQUIST, P.R. & TIZARD, C.A. 1967. Australian
intertidal sponges from the Darwin area.
Micronesica 3: 175-202, pls 1-6.

BURTON, M. 1931. Ona collection of marine sponges
mostly from the Natal coast. Annals of the Natal
Museum 6(3): 337-358, pl. 23.

1934. Sponges. 513-621, pls 1-2. In ‘Scientific
Reports of the Great Barrier Reef Expedition
1928-29". Volume 4(14) (British Museum
(Natural History): London).

CABIOCH, L. 1968. Contribution a la connaissance de
la faune des spongiaires de la Manche occidentale.
démosponges de la région de Roscoff. (Travaux

136

MEMOIRS OF THE QUEENSLAND MUSEUM

area of sympatry

40°

FIG. 25. Distribution of Australasian Cymbastela species, indicating three groups of allopatric species (numbers
4,6,7), and areas of overlap between four sympatric species (numbers 1,2,3,5) on the northeast and northwest
coasts of Australia. 1, C. stipitata; 2, C. coralliophila; 3, C, vespertina; 4, C. cantharella; 5, C. concentrica; 6,

C. marshae; 7, C. notiaina.

de la Station Biologique de Roscoff. Nouvelle
Série 18.) Cahiers de Biologie Marine 9: 211-246,

DE LAUBENFELS, M.W. (see Laubenfels, M.W. de)

DENDY, A. 1905. Report on the sponges collected by
Professor Herdman, at Ceylon, in 1902. 57-246,
pls 1-16, In Herdman, W.A. (ed.), “Report to the
government of Ceylon on the pearl oyster fisheries
of the Gulf of Manaar’. Volume 3(18). (Royal
Society: London).

1922. Report on the Sigmatotetraxonida collected by
H,M.S. ‘Sealark’ in the Indian Ocean. Transac-
tions of the Linnean Society of London, Zoology
18: 1-164, pls 1-18. (In ‘Reports of the Percy
Sladen Trust Expedition to the Indian Ocean in
1905’. Volume 7).

HALLMANN, E.F. 1916. A revision of the genera with
microscleres included, or provisionally included,
in the family Axinellidae, with descriptions of
some Australian species. Part iii, (Porifera).

Proceedings of the Linnean Society of New South
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HARTMAN, W.D. 1982. Porifera. 640-666. In Parker,
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HENTSCHEL, E. 1912. Kiesel- und hornschwaimme
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295-448.

HOOPER, J.N.A. 1986, Revision of the marine sponge
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from northwest Australia. The Beagle, Oc-
casional Papers of the Northern Territory Museum
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1990. A new species of Rhabderemia Topsent
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199]. Revision of the family Raspailiidae (Porifera:

LAMELLATE CORAL REEF SPONGES

Demospongiae), with description of Australian
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HOOPER, J.N.A., CAPON, R.J., KEENAN, CP.,
PARRY, D.L. & SMIT, N. 1992.
Chemotaxonomy of marine sponges: families
Microcionidae, Raspailiidae and Axinellidae, and
their relationships with other families in the orders
Poecilosclerida and AxineJlida (Porifera:
Demospongiae). Invertebrate Taxonomy 6(2):
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KIRKPATRICK, R. 1903. Descriptions of South
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LAUBENFELS, M.W. DE 1934. New Sponges from
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1936. A discussion of the sponge fauna of the Dry
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general, with material fora revision of the families
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the Tortugas Laboratory 30: 1-225.

1954. The sponges of the west-central Pacific,
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1897. Die clavulina der Adria. Nova Acta Academy
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138

THE TADPOLE OF THE AUSTRALOPAPUAN FROG
RANA DAEMELI, Memoirs of the Queensland Museum
32(1): 138, 1992:- The ranid trog Rana daemeli (Steindach-
ner) is widespread in New Guinea and, in Australia, occurs in
northern Queensland north of Townsville (Menzies, 1987;
pers. obs.),

Little is known of the biology of this species. Zweifel
(1980) and Menzies (1987) analysed calls of R. daemeli from
Papua New Guinea, and Menzies (1987) described the form
of the spawn. In this note I describe the tadpole of R. daemeli
from northern Queensland,

Tadpoles were collected with a dip net and preserved imme-
diately in 10% formalin, orretumed tothe laboratory where they
were kept in glass aquaria and fed fresh algae. Measurements of
preserved specimens were made using a dissecting microscope
with an eyepiece micrometer. All measurements are in mil-
limetres; those involving apertures were taken from the centre
of the aperture. Terminology follows Altig (1970), and. the
staging system is that of Gosner (1960).

Tadpoles were collected in Melaleuca swamps and tem-
porary and permanent streams between Ollera Creek, 63km
north of Townsville, and Cape Tribulation. Specimens were
found primarily among debris and algae in the deepest sec-
tions of these habitats. Tadpoles and metamorphs have been
collected during March, May and September, indicating that
this species has an extended breeding seasan, or that larval
development is extended.

A stage 36 tadpole is illustrated in Figure 1. Details of the
oral disc are lustrated in Figure 2. A stage 36 tadpole has the
following measurements: 59.6 total length, 22.0 body length,
6.4 basul tail muscle height, 3.9 basal tai] muscle width, 4.0
maximun dorsal fin height located 14,8 trom boxy terminus,
3.0 maximum ventral fin height located 16.8 from body ter-
minus, 12.3 moxumum body width, 10,2 maximum body
lvight, 2.6 eye diameter, 7.3 interorbilal distance, 3.8 inter-
naral distance, 2.7 snout-naris, 7.0 snout-eye, 11.2 snout-
spiniclo, 4.2 naris-eye, 4.5 transverse oral disc diameter, Other
major charaetenstics include; oral disc emarginate, oriented
anieroventrally; marginal papillae large, ina single row lateral-
ly, tWo rows posteriorly, a wide gap anteriorly; papillae on
Posterior margin extremely elongated, forming a fringe wong
the postenor margin of the disc; submarginal papillae only at
the comer of the jaws; spiracle short, sinistral, focated
ventrolaterally and onented posterodorsally; vent tube dextral,
attached to yentral fin, opening 4.0 from tail-bedy junction.

The tooth row formula is 2(2)/3(1). The A2 gap is wide,
and the Pl gap extremely narrow, The keratinised jaw
sheaths are nacrow and serrated. The upper sheath is broadly
curved, and the lower sheath broadly V-shaped, The eyes are
dorsal. The nares are extremely small, and are covered with

FIG. |. Lateral view ofa stage 36 Rane daemeli tadpole. Scale
baer 1S.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 2 Oral dise of 4 atage 36 Rana daemeli tadpole. Scale
bar= lima.

wt
ate aust Paya

eanventymenie gant”
iat

r a
hy oe at
nen a anal — ans

skin in some specimens, The fing are low, tapering to a narrow
bul not sharp point.

In Jife tadpoles are heavily pigmented dorsally wath gold
and black, with a green tinge along the midline. Black pig-
mentation is more prominent laterally. Tadpoles are dark grey
on the anterior portion of the ventral surface, and light grey
with a purple Linge on the posterior portion of the ventral
surtace. Small clumps of gold pigment occur on the fins. A
tadpole reared to metamorphosis was olive green dorsally and
prey ventrally, weighed 1.42g and was 24.3mm in length.
These tadpoles belong to the Benthic (Type 2: Profundal)
ecomorphological guild of Altig & Johnston (1989). ‘The
typical ranid features of an emurginate oral disc and at fringe
of elongatcd papillae along the posterior margin of the disc
(Webb & Korky, 1977) should serveto distinguish this species
fron other Australian tadpoles so far described.

R. Alford commented on the manuscript. Field work was
funded by ARC grant AT8931875 to R. Alford.

Literature Cited

Altig, R.A, 1970, A key 10 the tadpoles of the continental
United States and Canada, Herpetologica 26; 180-207,

Altig, R. & Johnston, G.F. 1989. Guilds of anuran larvae:
relationships. among developmental modes, mor-
phologies and habitats, Herpetological Monographs 3:
81-108.

Gosner, KL, 1960. A simplified table for staging anuran
embryos and larvae with notes on identification, Her-
petologica 16; 183-190.

Menzies, Jl. 1987. A taxonomic revision of Papuan Rana
(Amphibia; Ranidae), Australian Journal of Zoology 35;
373-418.

Webb. R.G, & Korky. J.K.1977, Variation intadpoles of frovs
of the Rana farahumarac group in western Mexico
(Anura: Ranidoe). Herpetologica 33: 73-82.

Zweifel, R.G. 1980. Results of the Archbold Expeditions. No.
(03. Frogs wid lizards from the Huon Peninsula, Papua
New Guinea. Bulletin of the American Museum of
Natural History 165: 387-434.

Stephen J. Richards, Zoology Department, James Cook
University, Townsville, Queensland 481], Australia;
25 October, 1991.

TRACE FOSSILS FROM THE PHE FORMATION (LOWER CAMBRIAN), ZANSKAR
VALLEY, NORTHWESTERN INDIA

NIGEL C. HUGHES AND MARY L. DROSER

Hughes, N.C, & Droser, M.L. 1992 06 29: Trace fossils from the Phe Formation (Lower
Cambrian), Zanskar Valley, northwestern India. Memoirs Queensland Museum 32(1):139-

144, Brisbane, ISSN 0079-8835,

Trace fossils from the upper part of the Tsarap Member of the Phe Formation, Zanskar Valley,
northwest India, provide the first evidence for the age of these strata. Taphrhelminthopsis cl.
circularis and Helminthopsis sp. suggest an Early Cambrian age, T cf circularisin the absence
of body fossils may indicate that these traces predate the tnlobites and are correlative with the
Tommotian, (] trace fossil, Cambrian, Himalaya.

Nigel C. Hughes, Queensland Musexm, PO Box 3300, South Brisbane, Queensland 4101,
Australia; Mary L. Droser, Department of Earth Sciences, University of Callfornia, Riverside,

CA. 9252], USA; 10 April, 1992.

The lower Palaeozoic history of India is not well
known, despite much recent work. The Zanskar
region is particularly poorly understood, although
it seties of recent studies provide some under-
standing of the early Palaeozoic in this region
(Nanda & Singh, 1976; Srikantia et al., 1980;
Gaetani et al., 1986; Garzanti et al., 1986; Whit-
uington, 1986). Trace fossils studies are currently
of special significance because the Precambrian-
Cambrian boundary is formally based on the ap-
pearance of key ichnogenera (Narbonne et al.,
1987). This paper records trace fossils within the
Phe Formation and constrains the position of the
Precambrian-Cambrian boundary in this region,

TRACE FOSSILS AND THE
PRECAMBRIAN-CAMBRIAN BOUNDARY
IN INDIA

Following Seilacher’s (1955) pioneering work
on the trace fossils of the Salt Range in Pakistan
recent work in India has concentrated on the use
of trace fossils for recognising the Precambrian-
Cambrian boundary (Tandon & Bhatia, 1978;
Bhargava ct al., 1982; Bhargava & Snkantia,
1982; Raina et al., 1983; Kumar etal., 1983; Shah
& Sudan, 1983, 1984; Singh d& Rai, 1983: Kumar
ctal., 1984; Kumar, 1984). The diversity of trace
fossils increases at stratigraphically higher levels
in pre-trilobite bearing strata of India (Tandon &
Bhatia, 1978; Raina et al., 1983; Kumar, 1984),
which is consistent with pattems described from
many sections worldwide (e.g. Fedonkin, 1979;
Crimes & Anderson, 1985; Fritz & Crimes, 1985;
Crimes, 1987; Narbonne & Myrow, 1988; Nar-
bonne et al, 1987), The Precambrian-Cambrian
boundary is formally defined on the basis of trace

fossils and, while initial progess has been made,
much work is still needed before it can be ac-
curately determined in the Indian Himalaya.
Kumar ct al. (1984, fig. 2) reported that Phycades
first occurs at the top of member A of the Lolab
Formation in Kashmir, 750m above deposits con-
taining Upper Riphean microflora. If complete,
this section appears to be the best candidate fnr
recognising the Precambrian-Cambrian boundary
in India. A complete Precambrian-Cambrian
boundary section may be present in the Kro}-Tal
belt, from which Ediacaran fossils (Mathur &
Shanker, 1989; 1990), small shelly fossils (Bhatt
et al., 1983) and archaeocyathids (Singh & Rai,
1983) have been reported. However, the Echa-
caran faunas apparently occur above the small
shelly fossils (Kumar, 1990), and the accuracy of
some of these identifications is uncertain, M.
Wade (pers, comm.) suspects that many, but nol
all, are algal or cyanobacterial mat structures,

STRATIGRAPHY AND TRACE FOSSILS OF
THE PHE FORMATION

STRATIGRAPHY

Lowermost Palaeozoic rocks in the Zanskar
Valley are assigned to three formations; the basal,
silt/sandstone-dominated Phe Formation, the
dolomitic Karsha formation and the silt-
dominated Kargiakh Formation (Garzanti et al,,
1986). Although Garzanti et al. (1986) noted
bioturbation in the Phe Formation they did not
recognise any fossils. Trilobite fragments have
been reported (Garzanti et al., 1986) within the
Karsha Formation, and late Middle Cambrian
trilobites occur in the Kargiakh Formation (Gar-
zanti et al., 1986) (some 400 m above the top of

140

the Phe Formation). These relationships suggest
that the Phe Formation is older than late Middle
Cambrian; some of the Phe Formation might be
Precambrian (Garzanti et al., 1986), raising the
possibility of a Precambrian-Cambrian boundary
section in Zanskar.

The Phe Formation comprises at least 800 m of
siltstones and fine-grained sandstone (Garzanti et
al., 1986). Itis divided into a silt-rich lower Tsarap
Member, and an upper sandier Doda Member
(Nanda & Singh, 1976). Nanda & Singh (1976)
recognised an uppermost Thonde Member, from
which trilobites have been collected at Kargiakh
nulla (Dungrakoti et al., 1974). The tilobites
were assigned to Powhoparia sp and Asaphus sp
and, on this basis, it was suggested that the Phe
Formation is Upper Cambrian to Lower Or-
dovician (Gupta, 1978), However, further work
has shown that fossiliferous strata in Kargiakh
nulla are not part of the Phe Formation. Instead
they overlic the Karsha Formation. These deposits
are now assigned to the Sunchun Member of the
Kargiakh Formation (Garzanti et al,, 1986). Al-
though subsequent collection of the Kargiakh
nulla site yielded neither Plychoparia nor
Asaplus (these assignments are likely incorrect),
the annosid Lejopyge laevigata confirms a latest
Middle Cambrian age (Whittington, 1986). A
vanely of mid Middle Cambrian trilobites have
recently been collected from the Kargiakh For-
mation, some from limestones, by Mr, A. Dogra
and Dr. Sat Paul of Jammu University and are
currently being descnbed. As there are no major
unconformities or structural discontinuitics be-
tween the Kargiakh, Karsha and Phe Formations
the age of the Karsha/Phe Formations ts pre-mid
Middle Cambnan.

Gupta & Shaw (1981) reported and figured El-
Npsocephalus hoffi and Agraulos sp. which they
cluimed were collected from the Phe Formation
in the Kargiakh and Doda valleys. Reasons to
doubt the validity of this reportinclode: 1, the grid
co-ordinates given do pot coingide with the posi-
tions of these localities; 2, the three dimensional,
slightly distorted nature of their specimens con-
trasts with the condition of other Cambrian
trilobites collected from the Himalaya, which are
flattened and are strongly deformed; and 3, re-
collection of this area by M. Gaetani and N.

MEMOIKS OF THE QUEENSLAND MUSEUM

Hughes has not revealed any fossils similar to
those described by Gupta & Shaw (1981). Similar
doubts about the reliability of Gupta & Shaw's
(1981) paper has been recorded by Whitington
(1986) and their paper should be disregarded, in
line with the suggestion of Talent et al, (1990).

TRACE FOSSILS

The Phe Formation was examined at several
localities but trace fossils were observed only in
the section due north of the village of Purni
(33°8'24"N, 77°6’6"E) which corresponds to
Section I on the map of Garzanti et al. (1986).
These traces were described and photographed in
situ and were not collected.

Within this section, traces occur in the upper
part of the Tsarap Member of the Phe Formation,
which is predominately composed of hetcrolithic
dark grey siltstones and very fine grained light
grey/buff sandstones. Sandstones are thinly
bedded (1-Sem) and may be parallel, hammocky
or cross stratified, Siltstones range from thin
laminae to very thin beds (0,2-3.0cm). Where
ripple structures are well preserved, wavy bed-
ding is recognised (Fig, 1A). Wave, current, and
interference nipples are al] common on bedding
surfaces (Fig. 1H), Short, narrow tool marks indi-
cate palaeoflows towards the WSW. Slump feu-
lures are common, The Phe Formation was
deposited in an intertidal to shallow subtidal set-
ting (Garzanti et al., 1986; pers. obs. N.C.H.).

Trace fossils are common on several bedding
surfaces at the base of the section 100m north of
Purni village. Taphrhelminthopsis cf. circularis
(Fig. 1B-D) is most common occurring as a
bilobate structure with a well defined, broad
central furrow. Burrow width is 1.5-2.0cm. The
furrow is about (0).2-0.3 of the width of the trace
fossil. The trace fossils are sinuows to arcuate and
up to 50cm long. Striations are not present. No
complete circles were observed so specific as-
signment is questionable.

Helminthopsis (Fig. 1G) occurs as a groove,
0.2-D.4em wide consisting of a central furrow
with raised levees or as a full burrow. Traces are
up to 15cm long, forming irmegular meanders that
were not observed to cross and commonly occur
onnippled surfaces.

The tops of yertical burrows, 0.5-0.7em in

FIG. |, All photos except E fron Upper Tsarap Member, 10) metres north of Purni, Zanskar. A, Interbedded silt
and sand layers showing wavy bedding. B, Taphrhelminthopsiy cf, circularis on bedding surface, * 0,2. C,
Tophrhelminthopsis cf. cireuleris on bedding surface, x 0.5. D, detail of Taphrhelminthopsts cf. circularis, * 1.
E, bedding surface showing abundant Planolites and other traces, scree slope between Purni and Testa, probably
from Tsarap Member, % 0.3, P. back filled burrow, * 1. G, Helminthopsiy on bedding surface, ¥ 1. H, Bedding

surfaces showing wave, Current and interference ripples.

TRACE FOSSILS FROM THE PHE FORMATION 141

142

diameter, some upparently paired, are present on
bedding surfaces, but no cross sections of these
burrows were observed. Horizontal Planolites
burrows, up to 0.8cm wide und 7.0cm long, are
also present. One possible example of a back-
filled burrow (Fig. 1F) occurs in siltstones. The
burrow is infilled with very fine sand.

A slab of Phe Formation in scree on the track
between Purni and Testa villages (Fig. 1E) shows
abundant horizontal traces and vertical Skolithos
burrows, which are 0.5cm wide and unpaired. In
cross section thts slab shows an ichnofabric index
of 3 (Droser & Bottjer, 1986).

DISCUSSION

The trace fossils milicate that these deposits are
no older than Vendian. Taphrhelminthopsis is
characteristic of Cambrian and younger sirata
(Crimes, 1987, in press). Fritz & Crimes (1985)
reported Tapitrhelminthopsis from Precambrian
Strata from the Cassiar Mountains, British Colum-
bia but Cnmes (1987) suggested that Taphrhel-
minthopsts in these strata indicates that perhaps
the boundary in this section should be lower, In
the current Precambrian-Cambrian boundary
stratotype in Newfoundland Taphrhelminthopsis
first occurs in the second trace fossil zone,
(Rusophycus avalonensis) above the
Precambrian-Cambnan boundary (Narbonne &
Myrow, 1988).

Aelminthopsis similarly first appears in
Cambrian strata in well documented sections
(Crimes, 1987} and is also thought to be charac-
teristic of Phanerozoic strata. Both genera occur
throughout the Phanerozoic and therefore do not
further constrain the age. However, Taphirfielmin-
thepsis circularis, appears to range trom just
above the first Phanerozoic-type traces to just
below the first trilobites (Crimes, 1987, in press),

Skelithas, Planolites and simple back filled bur-
rows, all appear in the Yendian and range through
the Phanerozoic and thetefore do not aid in age
resolution,

Crimes (1992) recognised an evolutionary
development of simple sediment filled burrows
from the Precambrian to the Cambrian, The
straight to slightly curved Planolites first occurs
in Vendian deposits, Slightly more complex ir-
regularly meandering burrows which cross, such
as Gordia, occur later in the Vendian. Burrows
which spiral and have first-order meanders appear
subsequently. Helminthopsis, which consists of
irregular, but non-crossing first and second order
meanders normally first occur in Cambrian strata,

MEMOIRS OF THE QUEENSLAND MUSEUM

and are charactenstic of the Early Cambrian.
Crimes (1992) reported increasing complexity of
‘stuffed’ burrows, with the meandering bilobate
stuffed burrow Taphrhelminthopsis charac-
tenising Early Cambrian deposits.

We suggest that at least the upper part of the
Tsarap Member is of Early Cambrian age based
on Helminthopsis and Taphrhelminthopsis, and
particularly the close similarity to T. circularis. If
T, cireularis is restricted to the pre-trilobite
Cambrian (Crimes, 1987), it could explain the
absence of body fossils in the Phe Formation. The
Precambnan-Cambrian boundary may lie within
the lower, unfossiliferous 200m of the Tsarap
Member of the Phe Formation.

Like most other Early Cambrian trace fossil
assemblages (Crimes & Anderson, 1985; Crimes,
1987) the fossils from the Phe Formation are
found in shallow, nearshore deposits. Taphriel-
minthopsis is common in the Lower Cambrian of
India, occurring in the Himalaya in southeastern
Kashmir (Bhargava & Srikantia, 1982) and in the
Lesser Himalaya in the Tal Formation near
Kauriyala, Uttar Pradesh (Kumar ctal., 1983), and
possibly near Mussoorie (Singh & Rai, 1983),

ACKNOWLEDGEMENTS

Peter Jel] read the manuscript and offered help-
ful criticisms. Mary Wade kindly discussed
aspects of the paper, N.C.H. gratefully acknow-
ledges receipt of a Natural Environmental Re-
search Council Fellowship which enabled him to
visit the Himalaya. Rae and John Sheridan
provided skiliful assistance in the field and excel-
lent companionship. Rigsen Jora facilitated our
iravel within the region. Dawa Jora and Punzok
acted as guides and we are grateful to them and to
our horsemen for assisting our safe passage.
Isabel Montanez and Sue Turner kindly provided
essential literauire,

LITERATURE CITED

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BHARGAYVA, O.N. & SRIKANTIA, S.V. 1982.
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BHATT, D.K,, MAMGAIN, V,D., MISRA, RS. &
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TRACE FOSSILS FROM THE PHE FORMATION

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GAETANI, M., CASNEDI, R., FOIS, E., GARZANTI,
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GUPTA, VJ. 1978, A note on the Palaeozoic stratig-
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new species. of Retichonetes from Surichun La,
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GUPTA, V.J, & SHAW, P.C. 1981, Lower Palacozoic
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KUMAR, G, 1984, The Precambrian-Cambrian bound-
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1990, Late Proterozoic to Cambrian sequences and
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KUMAR, G., RAINA, B.K., BHATT, D.K. &
JANGPANGI, B.S, 1983, Lower Cambrian bixly
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KUMAR, G., RAINA, B.K., BHARGAVA, 0O.N.,
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1990, Bdiacaran medusoids from the Krol Fornva-
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NARBONNE, G.M & MYROW, P, 1988. Trace fossil
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SHARMA, V.P. 1983. ?Precambrian-Lower
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144

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SINGH, I.B. & RAI, V. 1983. Fauna and biogenic
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TALENT, J.A., BROCK, G.A., ENGELBRETSEN,
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MEMOIRS OF THE QUEENSLAND MUSEUM

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Himalayan palaeontologic database polluted:
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of the Tethyan Zone Kumaun Himalaya with
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Rivista Italiana di Paleontologia e Stratigrafia 92:
171-188.

SCALPELLID BARNACLES (CIRRIPEDIA: THORACICA)
FROM THE NORTHEASTERN AND CENTRAL EASTERN AUSTRALIAN
CONTINENTAL SHELF AND SLOPE

DIANA. JONES

Jones, D.S. 1992 06 29: Scalpellid barnacles (Cirripedia; Thoracica) from the northeastern and
central eastern Australian continental shelf and slope. Memoirs of the Queensland Museum
32(1); 145-178. Brisbane. ISSN 0079-8835.

Eleven species of scalpellid barnacles are described from continental. shelf and slope waters
off northeastern and central easter Australia. Nine of these species were collected by the
Cidaris | expedition which sampled barnacles mainly from the continental slope of north-
eastern Queensland and the adjacent Coral Sea. The additional two scalpellid species were
contained in the deep-water cirripede collections of the Queensland Museum, Brisbane, and
the Australian Museum, Sydney. The species are fully described and illustrated since previous
descriptions often have described conchology only and detailed descriptions, especially of
arthropodal structures and of dwarf or complemental males, are either absent or are brief and
inadequate. All the scalpellids collected by Cidaris I were first records from Queensland waters
and, with the exceptions of Verum aitstrali¢um Hoek and Anguloscalpellum pedunculaium
(Hoek), first records from Australian waters. They were listed, but not described, in the recent
catalogue of the Australian cirripede fauna (Jones et al,, 1990), In the present contribution
Trianguloscalpellum hirsutum sensu Jones et al. (1990) is regarded as a synonym of T, hamulus
(Hoek), 7. hantulus and A, persona are new records for Australian seas, and §. stearnsii is
recorded from Queensland for the first time. (] Crustacea, Thoracica, Lepadomorpha,
Sculpellidae, Australia, new records, taxonomy’.

Diana S. Jones, Western Australian Musewn, Francis 81, Perth, Western Australia 6000,

Australia; 14 November, 199],

Documentation of the littoral and shallow water
cirtipede fauna of Australia began during the
French expeditions of discovery at the beginning
of the nineteenth century (Jones, 1991), The first
contributions towards knowledge of the deeper
water Australian cirripede faunas were not made
until the late nineteenth and early twentieth cen-
turies and are contained in Hoek’s reports en the
Cirripedia collected during the voyages of
‘Challenger’, between 1873 and 1876, and
*Siboga’, from 1899 to 1900 (Hoek. 1883, 1907,
1Y13). Subsequently significant contributions
have been made through the works of Calman
(1918, 1919), Nilsson-Cantell (1927. 1928), New-
man (1960), Utmomi (1968), Foster (1981) and
Zevina (1981a). More recently, documentation of
Australian shelf and slope (as well as littoral)
cirnpede faunas has been completed (Jones et al.,
1990),

The Cidaris I expedition, which sampled the
deep-sea fauna of the Coral Sea, was organised by
the James Cook University of North Queensland,
Townsville, in 1986. Some samples were obtained
from the continental shelf and the majority from
ihe continental slope and adjacent Coral Sea off
the northeastern Queensland coast, owt from
Townsville (19° 16'S, 146°49’E), at depths of 287

to 1612m. The present report covers the nine
species of scalpellid barnacies collected by this
expedition and further reports will cover the
remaining lepadomorphs and the Verrucomarpha
and Balanomorpha collected during the expedi-
tion. On examination of the deep-water barnucle
collections housed in the Queensland Museuwrn.
Brisbane, and the Australian Museum, Sydney,
two further scalpellid species were discovered that
had been collected from the waters of the nuth-
easter and central eastern Australian continental
slope. For comparalive purposes, slope material
from northwestern Australia housed in the
Western Australian Museum, Perth, was utilized
to validate identifications, Where relevant this
miterial hes also been included in the present
contnbution,

Eleven continental shelf and slope scalpellids
are described and illustrated. Supplementary
descriptions of each species, especially of
arthropodal structures and dwarf and complenwn-
tal males, are given smmce these have either not
been deseribed previously, or existing descrip-
tions are brief or inadequate, The classification
and sequence used for the Scalpellidae is that of
Zevina (198la, 1982). All measurements are in
centimetres.

146

Abbreviations: AM, Australian Museum, Syd-
ney; QM, Queensland Museum, Brisbane; WAM,
Western Australian Museum, Perth; CH, capitular
height; CW, capitular width; PL, uncular
length; PW, peduncular width; R. range; (R) right;
(L) left. Registration numbers of AM material are
prefixed by P, QM material by W (with Cidaris I
material indicated by W*) and WAM matenal by
WAM,

LIST OF THE SPECIES

|, Scalpellum stearnsii Pilsbry, 1890
*2. Alcockianum alcackianum (Annandale,1905)
3. Alcockianum persona (Annandale.l916)

*4, Annandaleum lambda (Annandale,1910)

*5. Verum australicum (Hock, 1883)

*6. Anguloscalpellam peduncalaium (Hoek, 1883)

*7. Amigdoscalpellum daschae Zevina, 1981

*8. Amigdoscalpellum elegans (Hoek,1907)

*9. Triangulescalpellum hamulus (Hoek, 1907)
*10,
*11. Arcascalpellum moluccanum (Hoek,1883)
*indicates material collected by Cidaris 1.

Arcoscalpellum michelottianum (Soguenza,1876)

Trianguloscalpellum hirsutum sensu Jones et al.
(1990) is regarded as a synonym of 7. hamulus
(Hoek, 1907), T. hamulus and A. persona are new
records from Australian seas, and §. stearnsii is
recorded from Queensland waters for the first
lime.

SYSTEMATICS

Order THORACICA Darwin, 1854
Family SCALPELLIDAE Pilsbry, 1916
SCALPELLINAE Pilsbry, 1907
Scalpellum Leach, 1817

Scalpellum stearnsii Pilsbry, 1890
(Figs 1, 2)

Sealpellum magnum Darwin, 1851: 18, pl. 1, fig, 1.

Scalpellun stearnsi(i) Pilsbry, 1890a: 96; Pilsbry,
1890b: 441, fig.; Weltner, 1897: 250; Gruvel, 1905:
44, fig. 46; Pilsbry, 1907: 14, pl. 4, figs 1-5; Krtiger,
1911: 18, figs 26-35, pl. 2, figs 18-19; Pilsbry, 1911:
61; Annandale, 1916b: 293; Broch, 1922: 235, fig.
6; Broch, 1931; 16; Hiro, 1933: 22, fig. 4, pl. I figs
5-Sa; Nilsson-Cantell, 1933: 33; Newman & Ross,
1971: 123; Zeyina 198 1a; 98, hg. 68; Rosell, 1981:
279, pl. le; Ren, 1987; 189, fig. 1:1.

MEMOIRS OF THE QUEENSLAND MUSEUM

Scalpellum calcariferum Fischer, 1891: 116, fig.

Scalpellum inerme Annandale, 1905: 75, fig. 1, pl. 8,
figs I-1a; 1909: 270; Pilsbry, 1907; 14.

Scalpellum siearnsii var, inerme: Annandale, 191 6b:
293; Nilsson-Cantell, 1928: 2, fig. 1; Broch, 1931:
17; Nilsson-Cantell, 1933: 33; 1934: 44, pl. 5 fig. 1.

Scalpellum stearnsii yar. robusta and var. gemina Hoek,
1907; 69, 70, pl. 6, figs 1-12; Pilsbry, 1911: 62;
Weltner, 1922: 16.

Scalpellum stearnsii inerme: Nilsson-Cantell, 1938: 8:
Newman & Ross, 1971: 123.

TYPE LOCALITY
East coast of Japan, between the bay of Tokyo
and the Inland Sea.

DISTRIBUTION

Indian Ocean, Malay Arch., Indonesia,
northwestern Australia, South China Sea, Japan.
Depth range 182-2117m. Now known from
central eastern Australia (465-562m).

MATERIAL EXAMINED

W15533,S of Swain Reefs (23°28'S, 153°19"E), 562m,
1 spec.; W10196, 130km E of Capricom Gp (23°28'S,
153°19"B), 562m, 15 specs; W1097, 100km E of
Capricom Gp (23°30’S, 153°04'E), 540m, 6 specs;
W11303 (part), E of Bunker Gp (23°54'S, 153°01"E),
465m, | spec.; W14339 (27°20°S, 153°53’E), 600m, i
spec.; P35288, E of Gold Coast (28°02’-27°59'S,
153°59°B), 540m, 1 spec,

DESCRIPTION

Large individuals female. Capitulum laterally
compressed, fourteen capitular plates, calcareous,
partially reduced. Carina angularly flexed, umbo
at angle, Carinolatus with hom-like projection
which is very variable, almost wanting to strongly
developed, when free projecting portion can be
one-third to one- half length of plate. Peduncle
with prominent external rings of overlapping cal-
careous scales. Measurements of the 25
specimens; CH 1,75-4.93 (mean 3.28), CW 1.05-
3.90 (mean 2.17), PL 1.43-12.85 (mean 4.23), PW
0.55-1.98 (mean 23).

Cirrus [ with rami very unequal in length and
shape, anterior ramus much shorter than posterior
ramus, proximal and intermediate segments of
anterior ramus very protuberant especially to-
wards posterior, segments transversely oblong
and only united with one another over part of
breadth, segments of posterior ramus square. Cirri
If and Mil elongated, rami subequal, bearing three
to six pairs of long setae on anterior margins,
anterior ramus of cirrus Il more setose than

147

SCALPELLID BARNACLES

\\N f Hh ‘
NWN I

FIG, 1, Scalpellum stearnsii Pilsbry, 1890. a, lateral view. b, cirrus I, c, cirrus II. d, intermediate segments, antenor

ramus, cirrus III. e, intermediate segments, anterior ramus, cirrus V. f, maxillule. g, mandible.

148

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Scalpellum stearnsii Pilsbry, 1890. a, posterior view, caudal appendages and pedicels of posterior cirri. b,

lateral view of prosoma. c, dwarf males,

posterior ramus of cirrus I] and both rami of cirrus
lil. Cirri TV, V and VI very long, rami subequal,
segments elongated, bearing five to seven pairs
long setae on anterior margins, posterior margins
of proximal segments of cirri V and YI bearing
dense scales, scales more numerous on anterior
rami. Chaetotaxy ctenopod. Cirral formulae, see
Table 1.

Maxillule with sinuous culting edge bearing
many setae (28-34 pairs), seta at upper angle stout
and slightly larger than remainder. Mandible with
five main teeth, simple, second smallest, sub-
sidiary teeth may be present between main teeth,
lower angle pectinate with short, chitinous hairs.
Labrum strongly bullate, not cleft, teeth absent.
Caudal appendages small, multi-articulate (6-10
segments), slightly longer than height of pedicel
of cirrus VI, with tuft of long setae apically.

Dwarf males (8-10) at occludent margins of
both scuta of female, located just above adductor
muscle in deep pits in scutal membrane; sac-like,
boat-shaped when viewed laterally, peduncle and
capitulum not differentiated, four rudimentary,
calcareous plates around subterminal orifice, short
rudimentary tentacles between plates, body exter-

nally covered with minute erect spicules, two
small, prehensile antennae on ventral surface,

REMARKS

The species has been reported from
northwestern Australian waters (Jones et al,
1990) but not previously from eastern Australia.
S, stearnsii is similar to S. scalpellum (Linnaeus)
(= 8. vulgare Leach) but the form of the
carinolatera distinguishes S. stearnsit.

The species was first described from shallow
water (11-18m), from the east coast of Japan
(Pilsbry, 1890a,b). One year later Fischer (1891)
described S, calcariferum from Sagami Bay,
Japan, Which later authors considered to be
synonymous with S. stearnsii Pilsbry (Weltner,
1897; Grivel, 1902). Fischer (1891) and Annan-
dale (1909) also both pointed out the close
resemblance between S. stearnsii Pilsbry and the
fossil species S$, magnum Darwin of the Coralline
Crag, Sudbourne.

5. inerme, a species with highly degenerate
capitular plates, was described from a depth of
293m from the Bali Straits (Annandale, 1905).
Pilsbry (1907) considered §. inerme a distinct

SCALPELLID BARNACLES

149

TABLE 1. Measurements and cirral formulae of Scalpellum stearnsii Pilsbry, 1890.

cu | cw {et | pw

eee fei ee oe evi

eS
28 32 =

aaa

peso Tere Te Te

species. Hoek (1907) established S. stearnsii var.
gemina, which also had degenerate plates, and var.
robusta which had a broader and longer capitulum
than the typical form, from specimens collected
by ‘Siboga’ in the Malay Archipelago at depths of
204-450 m. Pilsbry (1911) synonymized S.
stearnsii var. gemina with S. inerme, but con-
sidered var. robusta identical with the typical
form. However, Annandale (1916b), after ex-
amining a good series of specimens from Japan
and the Malay Archipelago, considered S. inerme
to be an extreme form of Hoek’s var. gemina and
proposed the degenerate plate form as S. stearnsii
var. inerme, a view with which Nilsson- Cantell
(1928, 1934) agreed. Annandale (1916b) found
two Japanese specimens referable to var. robusta,
and a number of indeterminate forms between var.
robusta and var. gemina, and suggested that, as
there was no evidence for Japanese and Malayan
races, the two varieties should be united under S.
stearnsii.

The specimens listed here represent the first
records of S. stearnsii from eastern Australian
waters. The specimens showed conchological
characters which have been ascribed to the
degenerate plate forms S. stearnsii var. gemina
and S. inerme (tergum incompletly calcified in the
form of an inverted V, scutum and upper latus
reduced in size, peduncular scales smaller, ar-
ranged in more numerous rows). Soft-parts mor-
phology and the structure of the dwarf males,
however, agreed with descriptions of the more
perfectly calcified form S. stearnsii, thus support-
ing the inclusion of S. inerme and S. stearnsii var.
gemina in synonymy with the typical form.

MEROSCALPELLINAE Zevina, 1978
Alcockianum Zevina 1978

Alcockianum alcockianum (Annandale, 1905)
(Fig. 3)

Scalpellum alcockianum Annandale, 1905: 82; 1906a:
392; 1906b: 138; 1907: pl. 1, fig. 2, pl. 2, figs 2-2b;
1913: 228, 229; 1916a: 129, pl 6 fig. 5.

Scalpellum (Scalpellum) alcockianum: Calman, 1918:

115; Nilsson-Cantell, 1928: 6, fig. 3; 1931: 2; 1938:
7.
Arcoscalpellum alcockianum: Newman & Ross, 1971:
60; Lakshmana Rao & Newman, 1972: 70.
Alcockianum alcockianum: Zevina, 1978b:
198 1a: 149, fig. 106.

1345;

TYPE LOCALITY
S of Ceylon (5°48’15"N, 80°56’E);
‘Investigator’ station 277; 1750-1609m.

DISTRIBUTION

Indian Ocean, Malay Arch., Indonesia, between
Java and Australia (10°45’S, 120°50’E), between
southeastern Austalia and New Zealand (37°00’S,
165°00’E), New Zealand (46°16’S, 173°22’E),
mid-Pacific Ocean (18°31’N, 179°36’W). Depth
range 945-1800m. Now also recorded from north-
eastern and central eastern Australian waters
(945-1950m).

MATERIAL EXAMINED

W*13256 (16°54’S, 147°14’E), 1473-1590m, 3 specs;
P35286 E of Orient Pt, (34°56’-54’S, 151°15’-17’E),
1098- 1134m, 3 specs; P35287 E of Shoalhaven Heads,
(34°53-50'S, 151°14’-15’E), 945-1017m, 1 spec.;
P35294 E of Long Reef Pt, (33°43’-40’S, 152°03°-
05’E), 1026-1053m, 3 specs.

SUPPLEMENTARY DESCRIPTION

Large individuals hermaphrodite. Capitulum
oval, inflated in upper half, especially between
tergum and neighbouring valves. Fourteen
capitular plates, concealed by and embedded in
thick, opaque membrane, externally velvety;
some capitular plates imperfectly calcified, wide-
ly separated, paired plates relatively small.
Peduncle variable in length, depending on degree
of contraction, calcareous scales transversely
elongated, arranged in alternating rows. Measure-
ments of the 10 specimens: CH 3.15- 6.60 (mean
5.28), CW 2.47-5.81 (mean 4.31), PL 2.41-8.65
(mean 4.82), PW 1.14-2.90 (mean 2.36).

Cirrus I setose, well separated from cirrus II,
anterior ramus shorter than posterior ramus,
proximal segments of anterior ramus protuberant
posteriorly, inner surfaces of segments of both

150 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 3. Alcockianum alcackianum (Annandale, 1905), a, lateral view. b, cirrus I, c, cirrus II. d, distal segments,
posterior ramus, cirrus I], e, intermediate segments, anterior ramus, cirrus V. f, maxillule. g, mandible. h, distal
tip of caudal appendage. i, penis. j, lateral view of prosoma. k, complemental males.

SCALPELLID BARNACLES

TABLE 2, Measurements and cirral formulae of Alvockianum alcackianum (Annandale, 1905)

ca [aaa oe

rami extremely setose. Cirmus II long, rami sube-
qual, anterior ramus more densely setose than
posterior ramus, setae setulate, inner surfaces of
segments of both rami densely setose. Cirri II] to
VI very long, cirri increasing in length towards
posterior, rami subequal, segments elongate bear-
ing three to five pairs long, setulate setae on
anterior faces, bunch of long finely setulate setae
al postero-distal corner of proximal segments of
cirri I to V. Chactotaxy ctenopod. Cirral for-
mulae, see Table 2.

Maxillule with smuous cutting edge bearing
numerous paired setae (30-38 pairs), stout spine at
upper angle. Mandible with three main teeth, first
large, acute, widely separated from rest, lower
angle pectinate. Labrum prominent, bullate.
Caudal appendages multi-articulate (22-30 seg-
ments), one-third to one-half length of cirrus VI,
distal borders of segments with circlets of setae
bearing fine setules, terminal segment with dense
tuft of long setae with fine setules. Penis one-
quarter length of cirrus VI, slender, minutely an-
nulated, tapering distally, sparsely setose, setae
long, plumose. Complemental males (3-4) at oc-
cludent margins of both scuta of hermaphrodite,
located just above adductor muscle, in shallow
pocket formed by fold in scutal membrane; sac~
like, ovoid, calcareous plates and cirri absent,
orifice slit-like, lateral; externally with elongated
spicules arranged in transverse rows.

REMARKS

Material collected by Cidaris 1 is the first record
of A. alcockianum within Australian waters (Jones
ct al., 1990). Additional material examined in the
course of this study has extended the range of the
species from northeastern to central eastern
Australia. The specimens agreed in general with
published descriptions for this species, Some
variation in the form of the tergum has been
described - for example, the scutal margin of the
tergum may be less hollowed ont (Annandale,
1906a; Nilsson-Cantell, 1928) or more hollowed
out (Annandale, 1916a; Lakshmana Rao & New-
man, 1972). The Australian series (n = 10)
demonstrated ontogenetic changes of this plate,
which is less hollowed out in smaller specimens

Va tow ta [wl ~alalal~wloalefa|

jos [am fas fort @tetete ts tals |:
is 29 a8 40

epee perpefe teeta tele te

36 ad a2

Farseees
39

(e.g. CH 3.55) but more hollowed out as capitular
size increases (e.g. CH 6.60).

Nilsson-Cantell (1928) described a female
specimen of A. alceckianum from Mozambique
Channel at a depth of 1647m, All other deserip-
tions of this species are based on hermaphrodites,
including the Australian series.

Alcockianum persona (Annandale, 1916)
(Figs 4.5)

Scalpellum persana Annandale, 1916h; 295, pl. 4, fig.
3, pl. 5, figs 7-8, pl. 6, figs 3-5; Calman, 1918: 120:
Nilsson-Cantell, 1934: 44, pl, 5, fig, 2.

Alcuckianum persena: Zevina, 1978b; 1345; 19S )a:
150, fig. 107.

Graviscalpellam persona: Foster, 1980: 527, fig. Jo, 2.

TYPE LOCALITY
Timor Sea (10°22'30"S, 120°7°30"E);
366m; bollom temp 13.1°C.

109.

DISTRIBUTION

Indonesian seas, also New Zealand. Depth range
109-915m. Now also recorded from eastern
Australia (550m),

MATERIAL EXAMINED
W11327, B Bunker Gp (23°15'S, 153°07'E), 550 m, 7
specs,

SUPPLEMENTARY DESCRIPTION

Large individuals hermaphrodite. Capitulum
ovoid, inflated. Fourteen capitular plates em-
bedded and mostly concealed by opaque, thick
membrane, externally finely hirsute; umbos api-
cal, apices projecting through membrance (espe-
cially carinolatera); some capitular plates
imperfectly calcified (e.g. terga), all plates except
scuta relatively small, especially inframedian
Jatus, Peduncle as long as or longer than
capitulum, narrowing towards capitular junction,
large calcareous scales transversely elongated, ar-
ranged in alternating rows. Measurements of the
7 specimens: CH 1.55-4.05 (mean 3.20), CW
1.02- 2,79 (mean 2.07), PL 1.62-5.33 (mean 3.40),
PW 0.63-1.86 (mean 2.31).

152 MEMOIRS OF THE QUEENSLAND MUSEUM

ot diy :
' BAN’: f - Z :
PRI
a ¥Y WAZ im, :
y ey

FIG. 4. Aleockianum persona (Annandale, 1916). a, lateral view. b, cirrus I. c, cirrus IL. d, intermediate segments,
anterior ramus, Cirrus III, e, intermediate segments, anterior ramus, cirrus V. f, maxillule. g, mandible.

SCALPELLID BARNACLES 153

Cirrus I setose, set higher than and slightly segments of anterior ramus protuberant especially
separate from cirri II to VI, anterior ramus shorter towards posterior, inner surfaces of segments of
than posterior ramus, proximal and intermediate bothrami with dense, setulate setae. Cirrus I long,

wo ¢0

FIG. 5. Alcockianum persona (Annandale, 1916). a, posterior view, caudal appendages and pedicels of posterior
cirri. b, penis. c, lateral view of prosoma. d, complemental male.

154

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3. Measurements and cirral formula of Alcockianum persona (Annandale, 1911),

rami subequal, anterior ramus more setose than
posterior ramus, setae setulate, Cirri ITI to VI very
long, cirri increasing in length towards posterior,
rami subequal, segments square to elongate, bear-
ing three to six pairs long setulate setae on anterior
faces, bunch of long, finely setulate setae at
postero-distal corner of proximal segments of curti
Ito VI. Chaetotaxy ctenopod. Cirral formula, see
Table 3.

Maxilla with characteristic development of ol-
factory organ in the form of a blunt process. Max-

illule with slightly sinuous cutting edge bearing,

numerous paired setae (25-30 pairs), two stout
spines at upper angle, Mandible with 5 teeth, first
acute, large, widely separated from remainder,
lower angle molariform, Labrum extremely bul-
late, prominent. Caudal appendages multiarticu-
late (13-16 segments), twice length of pedicel of
cirrus VI, segments with circlets of sctulate setac,
distal segment with dense tuft of setac, proxirnal
segment expanded. Penis relatively long, one-
third to one-half length of cirrus VJ, annulated,
tapering distally. setose towards apex, apex blunt
with circlet of fine short setae, Complemental
males at occludent margin of both scuta of her-
maphrodite, located just above adductor muscle in
deep pit in scutal membrane; ovoid, sac-like, four
reduced calcareous plates around subterminal
orifice, externally with small spicules, two
prehensile antennae on ventral margin

REMARKS

A. persona is sitnilar to A, alceckianum but the
wo species are easily distinguished by characters
of the tergum, canmolatera, caudal appendages
and mandible,

The specimens ofA, persone described above are
the first collected from Australian seas. These
specimens agtee in general with published descrip-
tions of conchology for A. persana (Annandale,
1916). However, the form of the mandible of
Australian specimens differs from that previously
described in that they exhibit five definite teeth,
rather than four as figured by Annandale (1916b)
for Indonesia matenal and Foster (1980) for New
Zealand specimens. Australian specimens also do
not exhibit a notch on the maxillule, whereas New
Zealand specimens show a slight notch (Foster,
1980) and Indonesian material a broad shallow
notch(Annandale, 1916b). The characteristic blunt

pou | cw lo wm | wl [a [oa | oomfow lv [owl oe |
pe foe ee Dee eg Ce De ee
8 28 38 i+ 14+ 17+

conical olfactory lobe of the maxilla is, however,
exhibited by Indonesian, Australian and New
Zealand specimens.

Annandaleum Newman & Ross, 197]

Annandaleum lambda (Annandale, 1910)
(Fig. 6)

Scalpellum lambda Annandale, 1910: 115; 1913; 234;
191 6a; 130, pl.7, figs 6, 6a, pl, 8, figs 12-15,

Sealpellum longius Annandale, 1913: 234; 19164: 130,
pl, 7, figs 5, Sa; Stubbings, 1936; 27, fig. 11.

Annandaleum lambda: Newman & Ross, 1971: 122,
Zevina, [9Rla:s 16R, fig, 120,

TYPE LOCALITY
Eastern Indian Ocean (13°54'15"N, 94°02715"E);
‘Investigator’ station 372; 1176m.

DISTRIBUTION

Indian Ocean, Depth range 237-1960m, Now
also recorded from northeastern Australia (956-
469m).

MATERIAL EXAMINED
W* 13238 (18°08"S, 147°36'B), 956-969m, | spec.

SUPPLEMENTARY DESCRIPTION

Large individuals female, Capjtulum ivregularly
ovate, laterally compressed. Fourteen capitular
plates, smooth, thin, imperfectly calcified,
separate, eight in shape of Greck lambda (A).
Inframedian latus large, vase-shaped, Peduncle
shorter than capitulum, numerous calcareous
scales present, narrow, transversely elongated, ar-
ranged in alternating rows. Cirrus I long, placed at
a distance from citrus I, rami very unequal,
anterior ramus shorter and broader than posterior
ramus; inner surfaces of both rami densely setose.
Cir IL to VI very long, cirri increasing in Jength
to posterior, rami subequal, segments elongate,
bearing four to six pairs of long setae on anterior
faces, Chaetotaxy ctenopod. Cirral formula, see
Tafle 4.

Maxillule with definite notch, cutting edge
below notch bearing two to three pairs setae, lower
angle spinose, three stout subequal spines at upper
angle, Mandible with three main teeth, lower
angle pectinate. Labrum bullate, Caudal append-

SCALPELLID BARNACLES 155

ee.
OS,

“r “y Vf,
{lh sits
oe a

FIG. 6. Annandaleum lambda (Annandale, 1910), a, lateral view. b, cirrus I. c, distal segments, posterior ramus,
cirrus III. d, intermediate segments, anterior ramus, cirrus III. e, intermediate segments, anterior ramus, cirrus V.
f, maxillule. g, mandible. h, distal tip of caudal appendage. i, lateral view of prosoma. j, ova.

156

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 4. Measurements and cirral formula of Annandalewm lambda (Annandale, 1910)

ages moderately long, one-fifth length of cirrus
VI, multi-articulate (6-8 segments), long setae
with setules at distal tip, Penis absent, Dwarf
males unknown. Ova large, ovoid. yolky, lemon
yellow in preserved material.

REMARKS

This species was first described by Annandale
(1910) from material collected in the Andaman
Sea. Annandale (1913) also described S. longius
from the Andamans and commented that the spe-
cies may possibly be. *... merely a complete form
of S. lambda.,..’, differences being the degree of
reduction of the capitular plates, the structure of
the carinolatera and the relative lengths of the rami
of cirrus I and the number of segments in the
caudal appendages, Stubbings (1936) examined S.
longius from Zanzibar and found the specimens
agreed with the type description and ‘...showed no
tendency to vary towards S. lambda’ -

The sole Australian specimen descnbed here
shows characters attributable to both §. lambda
and §. Jongivs. The form of the carinolatera and
the degree of reduction of the capitular plates
agree in general with conchology described for S.
lambda but differ in the following details: the apex
of the carina almost touches the carinal margins of
the terga; the tergal branch of the scutum is much
shorter than that illustrated by Annandale (1910)
but not as short as shown by Zevina (198 1a);
sculum, rostrolatera and carinolatera are more
similar to the form illustrated by Annandale
(1910) than that of Zevina (198 1a); scutal arm of
upper latus exhibits a bifid tip. Mandibles and
maxillules of the Australian material resemble the
forms figured by Stubbings (1936) for S. longius,
In addition, both mandibles of the Australian
specimen have three small, well- spaced spines on
the upper margin of the third tooth and on the
upper margin of the lower angle, which have not
been figured previously for either §. lambda or S,
longus. Rami of cirrus L in the Australian
specimen are very unequal in Jength. These struc-
tures have been described as equal (Annandale,
1913) or subequal (Stubbings, 1936) in 8. longius
and unequal in §. fambda (Annandale, 1910).
Caudal appendages of the Australian spectmen are
eight segmented. This structure appears tw be
similar in S. lambda (8 segments; Annandale,

A TT
[re [om | om fom [ ml elelsis]e ls

1910) and S. fongius (6 segments; Annandale,
1913; 8 segments; Stubbings, 1936).

The similarity of conchological and arthropodal
characters support the conclusion that S. longius
is a junior subjective synonym of S. /ambda, as
originally suggested by Annandale (1913) and
later formalized - although not discussed - by
Newman & Ross (1971) and Zevina (1981a). The
Australian specimen collected by Cidaris 1 ex~
tends the geographical range of this species from
the Indian Ocean to the waters of northeastern
Queensland (Jones et al,, 1990),

ARCOSCALPELLINAE Zevina, 1978
Verum Zevina, 1978

Verum australicum (Hoek, 1883)
(Figs 7,8)

Sealpellum australicum Hoek, 1883: 118, pl. 5, fig. 11;
Weltner, 1922: 63.
Verum australicum: Zevina, 19§1a: 223, Fig. 160.

Type LOCALITY

Between New Guinea and Australia (12°08'S,
145°10°E); ‘Challenger’ station 184; 2561m, bot-
tom temp. 1.8°C; substrate grey ooze,

DISTRIBUTION J

Zanzibar: between Australia and New Guinea.
Depth range 463- 2561m_Now also recorded from
northeastern Australia (1187- 1590m),

MATERIAL EXAMINED

W* 13239 (16"54'S, 137°14"B), 1473-1590m, | spec.;
W*13240 (17° 19'S, 147°11E), 1402-1406m, 10 specs;
W" 13241 (17°20'S, 147°48’B), 1187-1200m, 1 spee.

SUPPLEMENTARY DESCRIPTION

Large individuals female. Capitulum oval, elon-
gate, laterally compressed. Thirteen capitular
plates, distinctly furrowed, closely approximate.
Inframedian latus elongate, vase- shaped.
Peduncle short, one-third to one-sixth capitular
height. calcareous scales distinct, larger towards
capitulom, arranged in five to six alternating rows.
Measurements of the 12 specimens: CH 1.24-2.40
(mean 2.00), CW 0.38-1.35 (mean 1.06), PL 0.20-
0.87 (mean 0.56), PW 0.09-0.68 (mean 0.52).

Cirrus [ set apart from cirrus I, rami unequal,

SCALPELLID BARNACLES 157

FIG. 7. Verum australicum (Hoek, 1883). a, lateral view. b, cirrus I. c, intermediate segments, anterior ramus, cirrus
IL. d, distal segments, posterior ramus, cirrus III. e, intermediate segments, posterior ramus, cirrus V. f, proximal
segments, posterior ramus, cirrus Y. g, maxillule. h, mandible. i, j, lower angle of mandible. k, distal tip of caudal
appendage.

158

0.56 cm

MEMOIRS OF THE QUEENSLAND MUSEUM

wo oO

WwW GOO

FIG. 8. Verum australicum (Hoek, 1883). a, lateral view of prosoma. b, dwarf males im scutal pocket. c, dwarf

males.

anterior ramus shorter and much broader than
posterior ramus, both rami densely setose. Cirri II
to VI very long, cirri increasing in length to
posterior, rami subequal, segments elongate, bear-
ing four to five pairs of long setulate setae on
anterior faces. Chaetotaxy ctenopod. Cirral for-
mulae, see Table 5.

Maxillule with slight notch, four to five spines
on cutting edge below notch, three stout subequal
spines at upper angle, lower angle spinose. Man-
dible with three main teeth, lower angle pectinate,
upper margin of third tooth with several small
spines. Labrum extremely bullate, teeth absent.
Caudal appendages short, marginally longer than
pedicel of cirrus VI, multiarticulate (nine seg-
ments), long setae bearing fine setules distally.
Penis absent. Dwarf males in pocket at occludent
margin, on inner surface of and towards apex of
right and left scuta of female, three to six males

per pocket; males sac-like, ovoid, externally with
chitinous spines, calcareous plates absent.

REMARKS

The material collected by Cidaris J extends the
range of V. australicum into the waters of north-
eastern Australia (Jones et al., 1990), The con-
chology of the specimens agrees in general with
the original description of Hoek (1883). However,
Hoek stated (p,118) that the 13 plates were ‘close-
ly locked together’ and that there were ‘no distinct
interspaces between the different valves’. In the
ontogenetic series of the Australian material an
interspace develops between the carinal margin of
the tergum and the upper portion of the carina in
larger specimens (CH 2.20+), Hoek's type
description was based on a smaller specimen (CH
1.25).

Hoek (1883) did not describe the internal struc-
ture of V. australicum and did not observe dwarf

TABLE 5. Measurements and cirral formulae of Verum australicum (Hoek, 1883).

cepa per pete le Ts

iG
aEaE

SCALPELLID BARNACLES 159

re Se
Wh,

Wit G2, | e-
a

FIG. 9. Anguloscalpellum pedunculatum (Hoek, 1883), a, lateral yiew. b, cirrus I, c, cirrus IL. d, cirrus IIL. e,
intermediate segments, posterior ramus, cirrus IV. f, maxillule. g, mandible. h, penis.

160

MEMOIRS OF THE QUEENSLAND MUSEUM

c= —_—
=o
je

ae

CELaaee

~ be T

|
{9
LOR

UP

DS

wa 90

FIG. 10. Anguloscalpellum pedunculatum (Hoek, 1883), a, lateral view of prosoma. b, complemental males.

males. Details of cirri, mouthparts, caudal ap-
pendages and the penis, and observations of com-

plemental males of the species are described here
for the first time.

Anguloscalpellum Zevina, 1978

Anguloscalpellum pedunculatum
(Hoek, 1883) (Figs 9,10)

Scalpellum pedunculatum Hoek, 1883: 99, pl. 6, figs

1-2.

Arcoscalpellum pedunculatum: Foster, 1978: 53, Pl.
TA, fig. 31.

Graviscalpellum peductulatum: Foster, 1980: 527, fig
3H

Anguloscalpellum pedunculatum: Zevina, 1981a: 266,
fig. 200.

TYPE LOCALITY

Pacific Ocean, near New Zealand (39°32’S,

171°48’E); ‘Challenger’ station 167; 274m, sub-
strate grey ooze.

DISTRIBUTION

Western Pacific Ocean, New Zealand, Ker-
madecs, Norfolk I., central eastern Australia.
Depth range 202-600m. Now also known from
northeastern Australia (300-540m).

MATERIAL EXAMINED

W11222, E of Murray I. (09°51'S, 144°26'E), 480m, 2
specs; W*13242 (17°35'S, 146°53'E), 458-500m, 1
spec.; W10193, SE of Saumarez Reef (22°36’S,
154°14'E), 522m, 13 specs; W10192, 50 km NE
Capricorn Gp (23°00°S, 153°00S, 153°18"E), 387m, 4
specs; W10191, 110 km E of Capricorn Gp (23°11’S,
153°00’E), 420m, 8 specs; W11324, E of Capricorn Gp
(23°21'S, 153°23’E), 410m, 2 specs; W10190, 100 km

SCALPELLID BARNACLES

161

TABLE 6. Measurements and cirral formulae of Anguloscalpellum pedunculatum (Hoek, 1883).
Peco | cw] om few | ot | co | co | cv | cv | cw | ce
Laer [ee [srl so ee] a Pee eS ee DS |

spate teletele terete [sts
Vv 27 33 EX)

E of Capricorn Gp (23°30°S, 153°04E), 540m, 29
specs; W11303 (part), E of Bunker Gp (23°54’S,
153°01"E), 465m, 2 specs; W11468, E of Bunker Gp
(23°59’S, 152°59'E), 380m, 12 nes W14334, off
Morton IL. (27°12'S, 153°52°E), 3 specs; W15351, off
Morton I. (27°13°S, 153°52°E), 590m, 1 spec.; P35288
E of Gold Coast (28°02"-27°59"S, 153°59'E), 540m, 20
specs; P35290 E of Gold Coast (28°02"- 27°59°S,
153°59'E), 300m, 26 specs; P35289 E of Gold Coast
(28°02°-D5'S, 153°57°E), 360m, 1 spec.; P35208, E of
Crowdy Head (32°06"-03'S, 153°04°-05’E), 360m, 1
spec,

SUPPLEMENTARY DESCRIPTION

Large individuals hermaphrodite. Capitulum
subtriangular, inflated towards peduncle. Four-
ieen capitular plates, concealed by chitinous, hir-
sute membrane, apices projecting through
membrane, plates of lower whorl short, small,
Carinolatus elongated, subtriangular, extending
outwards from capitulum laterally in larger
specimens. Peduncle longer than capitulum, cal-
cureous scales present, transversely elongated, ar-
ranged in alternating rows, covered by chitinous,
hirsute membrane. Measurements of the 49
specimens: CH 2.68-4.68 (mean 4.05), CW 1.91-
3.39 (mean 2.91), PL 4.60-12.97 (mean 8.19), PW
1.28-2.98 (mean 1.96).

Circus I shghtiy separated from cirrus Il, rami
unequal, proximal segments of anterior ramus
wider than those of posterior ramus, strongly
protuberant postenorly, distal segments antenni-
form, segments of posterior ramus not
protuberant. Cirri I to VI long, citi increasing in
length towards posterior, ramu subequal, segments
square, bearing four to seven parrs of long selac
on anterior margins. Chaetotaxy ctenopod. Cirral
formulae, see Table 6.

Maxillule with large spine at upper angle, cut-
ling edge slightly sinuous, bearing numerous pairs
of stout setae. Mandible with three large teeth,
lower angle molariform, pectinated, Labrum ex-
tremely bullate, cutting edge straight with minute
teeth. Caudal appendages multiarticulate, long (6-
37 segments), half to two-thirds length of cirrus
VI, Penis long, slender, tapering distally, sparsely
selose, circlel of short dense setae at distal tip.
Complemental males globulo-ovoid, onfice ter-

minal, surrounded by tour minute rudimentary
plates, externally with chitinous spines in regular
pattern, two prehensile antennae on ventral mar-
gin. extensile penis present; in shallow scutal
pocket of hermaphrodite, three to four males per
pocket, membrane of pocket with chitinous spines
in ‘ae pattern, membrane closely apposed ta
males,

REMARKS

Foster (1978, 1980) described material from
New Zealand waters, Norfolk Island and the Ker-
madecs and mentioned that the species was also
known to occur in the waters of eastern Australia
(29°S), at a depth of 500-600m, Cidaris |
speciemens of A, pedunculatum extended the
known geographic range of the species
northwards into the waters of north-eastern
Queensland (17°S) (Jones et al., 1990) and
material examined in the present study further
extends the total range of the species from 32'S to
Torres Strait (9°S).

Australian hermaphrodites agree well with pub-
lished descriptions. The presence of a penis in the
complemental male is confirmed and the orifice is
terminal, rather than lateral as illustrated by Foster
(1978) for New Zealand material, The close
proximity of the scutal pocket membrane to the
complemental males suggests the possibility of an
interlocking function between the chitinous spines
of the inner surface of the pocket and the external
surface of the males, thus maintaiming the males
securely in the pouch.

Amigdoscalpellum Zevina, 1978
Amigdoscalpellum daschae Zevina, 198!
(Fig. 11)

Amigdoscalpellum daschae Zevina, 198 ta:
219; 198 1h: 87, fig. 9.

290, fig.

TYPE LOCALITY
SE of Papua New Guinea (11°30.5'S,
152°11.7°E); depth 080m.

DISTRIBUTION
Type locality (3080m) and now also known
from north- eastern Australia (1517-1564),

162 MEMOIRS OF THE QUEENSLAND MUSEUM

i

: | AYYVIN \y\ & WIS
SS a dl \ | KX INNIS
LD ie

Pan LE fd 4

FIG, 11, Amigdoscalpellum daschae Zevina, 1981 .a, lateral view. b, cirrus I. c, cirrus IL. d, distal segments, posterior
ramus, cirrus III, e, median segments, posterior ramus, cirrus VY, f-h, maxillule. i, mandible. j, lower angle of
mandible. k, distal tip of caudal appendage. |, lateral view of prosoma. m, dwarf males.

SCALPELLID BARNACLES 163

TABLE 7, Measurements and cirral formulae of Amigdoscalpellum daschae Zevina, 1981.

_co | cwl om | wl | a | o | cm] cv | wl w

car [ow Pes Toe Poe ae be ee
B+ i+ 12+ 10+ 10+ 10+

sfeefesfe fers lets lelel sé.
13) 20 22, 26

TABLE 8. Measurements and cirral formulae of Amigdoscalpellum elegans (Hoek, 1907).

MATERIAL EXAMINED
W*13244 (16°59°S, 147°) 1B), 1545-1564m, | spec:
W*13243 (17°06S, 147°12°E), 1517-1539m, 2 specs.

SUPPLEMENTARY DESCRIPTION

Large individuals female. Capitulum irregularly
ovate, carinal margin more strongly curved than
occludent margin. Thirteen capitular plates par-
tially covered by thin, finely setose membrane,
fine growth lines visible externally on all plates.
Peduncle short, cylindrical, calcareous scales well
developed, overlapping, arranged in alternate
rows, covered with thin, finely setose membrane,
Cirrus I much shorter than and set apart from cirri
II to VI, anterior ramus shorter than and slightly
wider than posterior ramus, both rami setose,
especially inner surfaces. Cirti II to VI long, cir
increasing in length towards posterior, rami sube-
qual, segments elongate, bearing four to five pairs
long setae on anterior faces, Chaetotaxy ctenopod.,
Cirral formulae, see Table 7.

Maxillule with shallow median notch, two stout
subequal setae at upper angle, one long seta and
three pairs shorter setae below notch, Mandible
with three main teeth, lower angle molariform
with one or two small teeth on lower margin.
Labrum bullate, cutting edge straight, teeth ab-
sent, Caudal appendages multiarticulate (5-4 seg-
ments), short, extending just beyond pedicel of
cirrus VI, one twelfth length of cirrus VI, long
setae bearing setules apically. Penis absent, Dwarf
males in single row in shallow pocket at occludent
margin, on inner surface of both scuta, towards
apices; sac-like, four smal] rudimentary plates
around terminal orifice, externally with small
culicular spines.

REMARKS

Material collected by Cidaris I extended the
distribution of A. daschae into the waters of north-
castern Australia (Jones et al., 1990). The form of

a

0)

=
| 2

regenerating

the mandible of Australian specimens differs from
that figured by Zevina (1981), The pectination of
the molariform lower angle, especially of the
lower margin, is absent with only one or two small
teeth or residual teeth present on the lower man-
dibular margin in Australian material.

Amigdoscalpellum elegans (Hoek, 1907)
(Figs 12,13)

Scalpellum elegans Hoek, 1907;
Stubbings, 1936: 24.

Amigdoscalpellum elegans: Zevina 198la: 268, fig,
201.

107, pl, 8 fig. 9

TYPE LOCALITY
Sulawesi (5°39'S, 122°12°E), “Siboga’ Station
208, 1886m, substrate solid green mud.

DISTRIBUTION

Indo-west Pacific - South Arabian coust,
Sulawesi. Depth range 687-1886m. Now also
known from northeastern Australia (687- 122&8m),

MATERIAL EXAMINED

W*13245 (17°20'S, 147°48"E), 1187-1200m, 3 specs;
W*13246 (18°04'S, 147°19°E), 687-704m, 1 spec;
W* 13247 (17°45'S, 147°48'B), 1223-1228m, 2 specs;
W*13248 (18° 11'S, 148°22"E), 1109-1110m, 1 spec.

SUPPLEMENTARY DESCRIPTION

Large individuals female. Capitulum elongate,
uregularly ovate, laterally compressed, carinal
margin arched, occludent margin almost straight.
Thirteen capitular plates, closely approximate,
partially covered by thin, finely setose membrane,
all plates except carina externally finely striated.
Peduncle short, cylindrical, well-developed cal-
careous scales present, overlapping, arranged in
rows, covered with thin, finely setose membrane,
Measurements of the seven specimens: CH O.89-

164 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 12. Amigdoscalpellum elegans (Hoek, 1907). a, lateral view. b, cirrus I. c, distal segments, posterior ramus,
cirrus I. d, distal segments, anterior ramus, cirrus II. e, intermediate segments, posterior ramus, cirrus II. f, cirrus
II. g, intermediate segments, anterior ramus, cirrus V. h, intermediate segments, anterior ramus, cirrus VI. 1,
maxillule. j, mandible. k, lower angle of mandible.

SCALPELLID BARNACLES

165

FIG. 13. Amigdoscalpellum elegans (Hoek, 1907). a, lateral view of prosoma to show caudal appendages. b, lateral
view of prosoma. c, dwarf male in scutal pocket. d, dwarf male.

2.05 (mean 1.51); CW 0.39-2.05 (mean 0.69); PL
0.15-0.47 (mean 0.34); PW 0.14-0.48 (mean
0.35).

Cirias I set above and slightly apart from
remaining cirri, rami setose, anterior ramus
shorter than posterior ramus, with intermediate
segments broad but tending to antenniformy dis-
tally. Cirrus II with rami long, subequal, segments
becoming elongate distally, anterior ramus more
setose than posterior ramus. Cirri III to V1 very

long, rami subequal, segments elongate, bearing
three to four pairs long setae on anterior margins.
Chaetotaxy ctenopod. Cirral formulae, see Table
8.

Maxillule with small notch, definite, one stout
spine at upper angle and one pair shorter setae
above notch, three pairs of setae on cutting edge
below notch, lowest pair longest, three pairs small
setae towards lower angle. Mandible with three
main teeth, lower angle coarsely pectinate, upper

166 MEMOIRS OF THE QUEENSLAND MUSEUM

Ws -
ip oe —
) pA a
hy ih /S SPR st SS
Less g =
p

APR Ps
APP RRA PR
= iY VPS

Stl | INS .

‘SS ; aN
SE
SIO

FIG. 14. Trianguloscalpellum hamulus (Hoek, 1907). a, lateral view. b, rostrum, c, cirrus I. d, cirrus II. e, cirrus
Ill. f, proximal segments, anterior ramus, cirrus V. g, distal segments, posterior ramus, cirrus VI.

SCALPELLID BARNACLES

=="

=
aan IE
=

r\ 4
\|
l
t
[}

oY
‘
\|
a
a
1

|

wh
{ets
—Ttr” Ss
a

—=-
——

FIG. 15. Trianguloscalpellum hamulus (Hoek, 1907). a, maxillule. b, mandible. c, lower angle of mandible. d,

caudal appendage. e, penis. f, lateral view of prosoma.

margin of third tooth with two small spines.
Labrum bullate. Caudal appendages minute, less
than one-quarter length of basal segment of
pedicel of cirrus VI, with tuft of long setae apical-
ly. Penis absent. Dwarf male small, sac-like, four
small rudimentary plates around terminal orifice;
one specimen in shallow pocket at occludent mar-
gin towards apex of right scutum.

REMARKS
Material collected by Cidaris 1 extended the

distribution of A. elegans into the waters of north-
eastern Australia (Jones et al., 1990). This species
appears to be rare, being known previously from
only two specimens (Hoek, 1907; Stubbings,
1936). Conchological characters of the seven
Australian specimens agree fairly closely with the
original species description of Hoek (1907). The
appendages of the female, and the dwarf males of

A. elegans were undescribed prior to the present
report.

167

168

TABLE 9. Cirral formula for Trianguloscalpellum

hamulus (Hoek, 1907).

retele la D
26 21+ 25 27
Trianguloscalpellum Zevina, 1978

Trianguloscalpellum hamulus (Hoek, 1907)
(Figs 14,15)

Scalpellum hamulus Hoek, 1907: 86, pl. 7, fig. 14, 14a;
Annandale, 1916b: 291.

Arcoscalpellum hamulus: Rosell, 1981: 281, pl. 1, figs
a-d.

Trianguloscalpellum hamulus: Zevina, 1981a: 314, pl.
238.

Trianguloscalpellum hirsutum: Jones et al., 1990: 5
(non T, hirsutum (Hoek, 1883)).

TYPE LOCALITY
Banda Sea (5°26,6’S, 132°32.5’E); ‘Siboga’ sta-
tion 256, 397m, bottom greyish green mud.

DISTRIBUTION

Banda Sea, Johul Bank (198 km S of Timor),
Phillipines. Depth range 128-397m. Now also
recorded from northeastern Australia (490-
1119m).

MATERIAL EXAMINED
W*13249 (18°08’S, 148°15’E), 1115-1119m, 1 spec.;
W*13250 (18°04’S, 147°17’E), 490-512m, 1 spec.;
W10195, 130 km E of Capricorn Gp (23°28’S,
153°19’E), 562m, 1 spec.

SUPPLEMENTARY DESCRIPTION

Large individuals hermaphrodite. Capitulum
elongate, ovate, apex slightly produced, occludent
margin almost straight. Fourteen capitular plates,
closely approximate, covered by thin, finely
setose membrane. Peduncle short, cylindrical,
well developed calcareous scales present, ar-
ranged in rows, covered with thin, finely setose
membrane. Measurements as follows: CH 1.63,
0.83; CW 0.87, 0.35; PL 0.88, 0.25; PW 0.61,
0.30.

Cirrus I much shorter than cirri IT to VI and set
above and apart from remaining cirri, rami une-
qual, anterior ramus shorter and slightly wider
than posterior ramus, rami setose, setae long, bear-
ing setules. Cirri II to VI very long, cirri increasing
in length towards posterior. Cirrus II with rami
subequal, segments becoming elongate distally,
anterior ramus more setose than posterior ramus,
anterior faces of both rami bearing three to five

MEMOIRS OF THE QUEENSLAND MUSEUM

pairs of long, setulate setae on anterior margins.
Cirri II] to VI with rami subequal, segments elon-
gate, bearing three to four pairs of long setulate
setae on anterior margins. Chaetotaxy ctenopod.
Cirral formula, see Table 9.

Maxillule with slightly sinuous cutting edge
bearing numerous small spines (10 to 12 pairs),
upper angle with one stout spine, lower angle with
short spines. Mandible with three main teeth,
lower angle pectinate. Labrum extremely bullate.
Caudal appendages very thin, long, one-quarter to
one-third length of cirrus VI, multiarticulate (21
segments), circlet of fine setae apically. Penis
long, thin, tapering distally, sparsely setose but
setae more dense distally, circlet of fine setae
apically.

REMARKS

This species appears to be comparatively rare. It
was known previously from three specimens from
the Philippines (Rosell, 1981), one from Indonesia
(Hoek, 1907) and two from south of Timor (An-
nandale, 1916b). The three specimens collected
by the Cidaris I expedition extended the known
geographical range of T. hamulus to the waters of
northern Queensland (Jones et al., 1990). The
material agrees fairly closely with published con-
chological illustrations of the species (Hoek,
1907; Rosell, 1981). Only the larger of the two
Cidaris I specimens was dissected. Details of
arthropodal structures and mouthparts are
described here for the first time as only the man-
dible has been figured previously (Rosell, 1981).
Annandale (1916b) made brief mention of charac-
ters of the cirri and caudal appendages. The
smaller (and presumably juvenile) specimen col-
lected by the Cidaris I expedition is presently
attributed to T. hamulus, based on conchology.
However, the much deeper collection locality of
this specimen has to be reckoned with. Until more
specimens are collected and more information on
the ontological development of this species is
obtained, the identification cannot be confirmed.
As has been the case with previous specimens, no
complemental males were located in Australian
material and these are yet to be described.

Arcoscalpellum Hoek, 1907

Arcoscalpellum michelottianum
(Seguenza, 1876) (Fig. 16,17)

Scalpellum michelottianum Seguenza, 1876: 381, 464,
pl. 6, figs 15-25, pl. 10, fig. 26; de Alessandri, 1895:

SCALPELLID BARNACLES 169

FIG, 16, Arcoscalpellum michelottianum (Seguenza, 1876). a, lateral view. b, cirtus I. c, intermediate segments,
anterior ramus, cirrus II. d, intermediate segments, posterior ramus, cirrus II. e, intermediate segments, anterior
Tamus, cirrus ILI. f, intermediate segments, anterior ramus, cirrus V.

170

MEMOIRS OF THE QUEENSLAND MUSEUM

TS ee

,

FIG, 17. Arcoscalpellum michelottianum (Seguenza, 1876). a, maxillule. b, mandible. c, posterior view, caudal
appendages and pedicels of posterior cirri. d, dwarf male in scutal pocket. e, dwarf males.

263 (in part); 1897: 47; 1906: 251; Withers, 1953:
225.

Scalpellum velutinum Hoek, 1883: 96, pl. 4, figs 10-11,
pl. 9, figs 7-9; Gruvel, 1902: 56, 136, pl. 2, figs 3c,
10a-b, 14, pl. 3, figs 1, 27-31, pl. 4, figs 6, 11-22;
1905: 73, fig. 83; Annandale, 1906a: 389; Pilsbry,
1907: 26, pl. 3, figs 2-3; Annandale, 1908: pl. 4, fig.
7; 1911: 588; Gruvel, 1912; 2; Annandale, 1913:
229; Hoek, 1914: 4; Calman, 1918: 108; Gruvel,
1920: 27, pl. 1, figs 8-10, pl. 7, fig. 4; Weltner, 1922:
75; Nilsson-Cantell, 1927: 743, fig. 1; 1928: 4;
Broch, 1931: 18; Nilsson-Cantell, 1931: 7; Stub-
bings, 1936: 28, fig. 12; Tarasov & Zevina, 1957; 24,
fig. 9H; Stubbings, 1967: 234; Zevina, 1972: 43, fig.
9

Scalpellum eximium Hoek, 1883: 100, pl. 4, figs 6-7, pl.
9, fig. 10; Weltner, 1897: 247.

Scalpellum sordidum Aurivillius, 1898: 190.

Sealpellum erectum Aurivillius, 1898: 192; Gnivel,
1905: 74.

Scalpellum alatum Gruvel, 1900; 192, 1902: 57.

Arcoscalpellum michelottianum: Newman & Ross,
1971: 71, fig. 34, pl. 9B; Lakshmana Rao & New-
man, 1972:76, fig. 5; Zevina 198 1a: 343, fig. 263.

Trianguloscalpellum michelottianum. Liu & Ren,
1985: 207, fig. 14, pl. 4, figs 10-12; Ren, 1989; 442,
fig. 6.

TYPE LOCALITY
Unknown.

DISTRIBUTION

Atlantic Ocean; Indian Ocean; Malay Arch.,;
Pacific Ocean; Chile. Depth range 64-5190m.
Now also known from northeastern and north-
western Australia (458-736m).

MATERIAL EXAMINED
W*13251 (17°52’S, 147°08°E), 497-503m, 1 spec.;
W*13252 (17°35’S, 146°53°E), 458-500m, 1 spec.;

SCALPELLID BARNACLES

17]

TABLE 10. Measurements and cirral formulae of Arcoscalpellum michelottianum (Seguenza, 1876).

* segements not distinct

WAM 935-86 NW of Port Hedland (18°37’S,
116°40’E), 732- 736m, 1 spec.

SUPPLEMENTARY DESCRIPTION

Large individuals female. Capitulum subtrian-
gular, 14 capitular plates, approximate, fully cal-
cified, covered by thin, pale brown, chitinous
membrane, sparsely hairy, apices of plates
projecting freely through membrane, growth lines
faint. Peduncle one-quarter to one-third capitular
height, well-developed calcareous scales present,
arranged in alternating rows. Measurements of the
three specimens: CH 0.96, 1.68, 3.05; CW 0.56,
1.05, 2.43; PL 0.44, 0.88, broken; PW 0.31, 0.61,
broken.

Cirrus I with rami unequal, posterior ramus
longer than anterior ramus, anterior ramus with
intermediate segments protuberant posteriorly,
both rami with many long, finely setulate setae,
especially on inner surfaces. Cirrus II longer than
cirrus I, rami subequal, anterior ramus more den-
sely setose than posterior ramus, long setae finely
setulate. Cirri III to VI elongate, cirri increasing in
length towards posterior, rami subequal, segments
elongate bearing three to four pairs of long, setu-
late setae on anterior margins. Chaetotaxy
ctenopod. Cirral formulae, see Table 10.

Maxillule without definite notch but slight in-
dentation apparent below upper angle, one stout
spine at upper angle, cutting edge below indenta-
tion almost straight with many paired setae.
Mandible with three teeth, molariform lower
angle pectinate. Labrum extremely bullate.
Caudal appendages stout, approximately one- half
to two-thirds length of basal segment of pedicel of
cirrus VI, two- or three-segmented but segments
indistinct, setulate setae terminally. Penis absent.
Dwarf males globulo-ovoid, orifice terminal, cal-
careous plates absent, a pair of antennae at op-
posite pole to orifice, outer surface with muscle
striations, not covered with spicules arranged in
transverse rows but by minute spiniferous excres-
cences; in pocket at occludent margin, on inner

surfaces of right and left scuta of female just above
adductor muscle pit of scuta, one male per pocket.

REMARKS

The considerable range of variation exhibited by
this wide-ranging species has been noted by
various authors (e.g. Gruvel, 1902; Calman, 1918;
Newman & Ross, 1971). The two specimens col-
lected by Cidaris I were the first records of A.
michelottianum from the waters of eastern
Australia (Jones et al., 1990).

However the identification is provisional. Of the
three specimens examined, only the middle-sized
specimen (CH 1.68) was dissected. Differences
and similarities of this material to published de-
scriptions of A. michelottianum are discussed
below.

Australian material shows the maxillule without
a notch, but with a slight indentation below the
upper angle. Newman & Ross (1971), examining
slightly smaller specimens, interpreted a similar
but slightly more pronounced indentation as
evidence for a distinct notch in the maxillule.
However, for a larger specimen (CH 3.0) Nilsson-
Cantell (1927) described the cutting edge of the
maxillule as nearly straight, although Newman &
Ross (1971) suggested an indication of a slight
notch in Nilsson-Cantell’s illustration of this ap-
pendage. Zevina (1972) showed no distinct notch
but illustrated a more sinuous configuration to the
cutting edge than described previously.
Lakshmana Rao & Newman (1972) found no
supramedian notch on the cutting edge of the
maxillule in mid- Pacific material.

Segmentation of the caudal appendages of
young specimens of A. michelottianum is indis-
tinct but larger specimens show distinct segmen-
tation (e.g. Gruvel, 1902: 7-8 segments; Calman,
1918: 4-6 segments; Nilsson-Cantell, 1927: 7 seg-
ments; Zevina, 1972: 1-6 segments; Lakshmana
Rao & Newman, 1972: 2-4 segments). Australian
material exhibits indistinctly two- segmented (CH
1.68) or three-segmented (CA 3.05) caudal ap-
pendages. This character appears to vary with age.

The posterior cier) of the medium-sized
Australian specimen bear four pairs of long, setu-
late selae along the anterior margins of the inter-
mnediate segments. Newman & Ross (1971)
figured three pairs per segment for slightly smaller
animals and Nilsson-Cantell (1927) and
Lakshmana Rao & Newman (1972) desenbed
four pairs per segment for larger specimens; this
difference may, therefore, be attributable to age.

All three Australian specimens show
carinolatera projecting beyond the carinal edge of
the capitulum, the projection being strongest in the
larger specimen, This condition is not illustrated by
Newman & Ross (1971), whose material (CH 1, 1&-
0,32) covers the size range of the smallest
Australian specimen. The orginal illustration of §,
velutinum (Hoek, 1883) showed the carinolstera
projecting beyond the capitulum and this condition
is also illustrajed by Gruvel (1902, 1905) and
Zevina (1972). These plates, however, may only
slightly project from the capitulum (Hoek, 1883 as
S. eximiwm, Nilsson- Cantell, 1927). The degrec of
projection appears to be variable, reflecting dif
ferent stages in ontogenetic development.

Conchology of the two larger Australian
specimens is very similar to that figured for
Chilean specimens (CH 3.1-1.6) of 8. velutinwm
(Zevina, 1972), although the peduncular scales
appear more developed (i¢, larger in size and
closely approximate} in the Australian examples.
It must be noted thal the conchology of the smal-
lest, Australian specimen, presently assigned to A.
michelottianum, is also very similar to that il-
lustrated by Ultinormi (1968) for S, Aérsusun (CH
1.3, CW 0,9), This author commented that exter-
nally the female capitulum of §. hirsietum closely
resembled 8. veludinum but that churacters of the
maxillule (a distinct small notch below the upper
angle), mandible (pectination of the upper margin
of the third mandibular tooth) and the dwarf male
(minute spiniferous excrescences on surface) war-
ranted specific separation. However, the dwarf
males of the Australian material herein ascribed to
A. michelotianum exhibit similar surface
spiniferous excrescences rather than cuticular
huirs. S. hirsutum is only known from Lwo
specimens - the Arafura Sea, 390m (Utinomi,
1968) and the Moluccan Sea, 1485m (Hock,
]K83), Ren (1983) described A. ciliatum from the
East China Sea. It shows conchological characters
very similar to A. michelottianwn. He ulso
synonymized S. hirsutum (Utinomi, 1968) bubnot
§. hirsutum sensu Hock, 1883 with A. ciliatum,
Differences in the caudal appendages (uniarticu-
late, expanded oval-shaped) and the maxillule

MEMOIRS OF THE QUEENSLAND MUSEUM

(notched) distinguish A. cilingn from A, michelot-
tlanum, Dwart males of A. ciliatunr remain to be
desenbed.

Arcoscalpellum moluccanum (Hock, 1883)
(Figs 18,19)

Sealpetium moluccanum Hoek, 1883: 104, pl, 5, Figs
3-4; 1907: 56, 5, pl. 7, fig, 13; Gruvel, 1905; 76, Fig.
§5; Nifsson-Cantell, 1927: 747 fig. 3.

Arcoscalpellum moluccanum: Newman & Ross, 1971:

‘

TYPE LOCALITY

Banda Sea (4°21'S, 129°7'°E), ‘Challenger’ sta-
tion 195, 2606m, bottom temp. 3.0°C, substrate
Erey OOZe.

DISTRIBUTION

Indonesia, Tasman Sea. Depth range 788-
2745m. Now also known from northeastern
Australia (904-1491m),

MATERIAL EXAMINED

W* 13253 (17°20°S, 147°48'E), 1187-1200m, 7 specs:
W*13254 (17°51'S, 147°09°E), 904-916m, 1 spee.;
W*13255 (17°12'S, 147°11"E), 1489-1491m, 1 spec.

SUPPLEMENTARY DESCRIPTION

Large individuals female. Capitulum subtrian-
gular, inflated. Fourteen capitular plates, covered
by relatively thin chitinous membrane, sparsely
hirsute, apices of plates projecting freely through
membrane, growth lines fairly distinct. Peduncle
stout, narrowing towards base, shorter than
capitulum, well- developed calcareous scales
present, arranged in alternating rows. Measure-
ments of the seyen specimens: CH _ 1.78-2.09
(mean 1.91}; CW 1.10-1.33 (mean 1-22); PL 0.99
- 1.46 (mean 1.23); PW 0.87 - 1.17 (mean 1.02).

Cirrus I set apart from cirrus IT, stouter and much
shorter than cirri Il to VI, anterior ramus shorter
than posterior ramus, mtermediate segments of
anterior ramus strongly protuberant posteriorly,
segments of postetior ramus not protuberant, both
Tumi selose, especially posterior faces and medial
faces of anterior ramus, Cirn TI-VI elongate, rami
subequal, cirri increasing in length towards
posterior. Cirrus IE with three to four pairs of setac
along anterior border of segments, anterior ramus
more setose than posterior ramus, posterior sur-
faces of both rami setose. Cirri TH and IV with
anterior and posterior rami with two to four pairs
of setac along anterior margins of segments,
posterior surfaces of segments sparsely sctose-

SCALPELLID BARNACLES 173

FIG. 18. Arcoscalpellum moluccanum (Hoek, 1883). a, lateral view. b, cirrus |. c, proximal segments, cirrus I. d,
cirrus IT. e, intermediate segments, anterior ramus, cirrus II, f, cirrus III. g, intermediate segments, posterior ramus,
cirrus IIT.

174 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 19. Arcoscalpellum moluccanum (Hoek, 1883). a, intermediate segments, anterior ramus, cirrus IV. b,
intermediate segments, posterior ramus, cirrus V. c, cirrus VI. d, intermediate segments, posterior ramus, cirrus
VI. e, maxillule. f, mandible. g, lower angle of mandible. h, posterior view, caudal appendages and pedicels of
posterior cirri. i, lateral view of prosoma. j, dwarf male.

SCALPELLID BARNACLES

78

TABLE 11. Measurements and citral formulae of Arcoscalpellum moluccanum Hoek (1883).

| ch | cw] om | pw] | a | om | on | ew | ev | ow | G |

205 | 133 1.33 v = = _ 2 =
pa) | 29 > 31
23+ ef

13

Cirri V and VI with four pairs of setae along
anterior margins of segments, posterior surfaces
of both rami sparsely setose. Chaetotaxy
clenopod. Cirral formulae, see Table 11.

Maxillule without notch, large stout spine at
upper angle, cutting edge almost straight, bearing
seven to eight pairs of setae, lower angle slightly
protuberant, bearing five pairs of setae. Mandible
with three large teeth, lower angle pectinated.
Labrum extremely bullate, cutting edge straight,
teeth absent. Caudal appendages slightly longer
than basal segment of pedicel of cirrus VI, three-
segmented, several long, stiff setae distally, Penis
absent, Dwarf males small, elongate, ovoid, exter-
nally covered with chitinous, short, stout setae,
anfennae present, rudimentary plates around
onfice lacking; in pouch inside occludent margin
of right scutum of female, at base of occludent
onfice, just above adductor muscle.

REMARKS

The species was previously known from four
specimens (Hoek, 1883, 1907; Nilsson-Cantell,
1927) and only their conchology has been
described. The nine Australian specimens col-
lected by Cidaris I constitute the first records of A.
moluéecanum in Australian waters (Jones et al..
1990). They agree closely with the original
description of A. moluccanum (Hoek, 1883). The
conchology of both the Aulantic species T. regizen
(Thomson) and the Antarctic and eastern Pacific
A. darwinii (Hoek) is very similar to that of A.
moluecanum.,

The form of the carinolatera, especially where
they meet at the base of the carina, and the position
of the rostrum relative to the rostrolatera, separate
A. moluccanum, A. darwinii and T. regium. The
internal anatomy of A, moluccanum described
above is similar to previous brief descriptions of
these parts for T. regium (Hoek, 1883; Pilsbry,
1907). Descriptions of the dwarf males of 7.
regium (Hoek, 1884) are more similar fo those A.
moluccanum than to A. darwinii. The mouthparts
and the caudal appendages (three segmented in A.
moluecanum, 4 to 6 segmented in T. regium, four
lo five segmented in A. danwinit) are similar in all
three species. It is possible that A. moluccanum
may be a geographical ‘race’ as suggested by

a1 3
32

Hoek (1883). Until comparison of the type
Tatenal of A. regi and A. moluccanum can de
made, the identity of the Australian specimens is
tentative.

ACKNOWLEDGEMENTS

lamindebted to Dr M. Pichon, Dr P. Arnold and
Mr F, Hoedt, James Cook University, for the
opportunity to examine the cimipede matenal col-
lected by the Cidaris I expedition. Mr P. Davic,
QM, and Dr P, Berents and Mr R, Springthorpe,
AM, kindly loaned specimens from their respee-
tive collections, | thank Mr D, Neville for trans
lating Russian papers. In addition, ] thank Dr F.
Wells and Prof. B. Foster for critically reading the
manuscript and Mrs A. Nevin for typing the
THM ESCTTpL,

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1912, Note preliminarie sur les Cirrhipédes recueillis
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1884. Ibid. Anatomical Part: 1-47, Pls. I-VI.

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1913. Ibid. B. Cirripedia Sessilia, 31b: i-xxv, 129-
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178.

JONES, D.S., ANDERSON, J.T. & ANDERSON, D.T.

SCALPELLID BARNACLES

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_, the Australian Museum 3: 1-38.

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herausgegeben von F. Doflein, Kéniglich
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DISTRIBUTION OF THE LONG-NECKED FRESHWATER TURTLE CHELODINA

NOVAEGUINEAE AND NEW INFORMATION ON ITS ECOLOGY
RODNEY M. KENNETT, ARTHUR GEORGES, KEN THOMAS AND T.C. GEORGES

Kennel, R.M., Georges, A., Thomas, K. & Georges, T.C, 1992 06 29. Distribution of the
long-necked freshwater turtle Chelodina novaeguineae and new information on its ecology.
Memoirs of the Queensland Museurn 32(1): 179-182. Brisbane. ISSN 0079-8835,

Recent collections have extended the westerly distribution of Chelodina novaeguineae, The
species occupies semi-permanent and seasonally ephemeral waterholes from coastal northeast
Queensland to the Daly Waters region of the Northem Territory. C. novaeguineae, along with
C. longicollis and C. steindachneri, may have speciated trom a common ancestor whose
widespread distribution around the periphery of the continent covered the present distribution
of these species. C. noyaeguineae aestivates or migrates overland to permanent water when
the waterhole it occupies dries, Its reproductive pattern of dry season nesting, small egg size
and Jong incubation period conflicts with a current interpretation of the phylogeny of
Australian chelid turtles. (]Chelidae, Chelodina novaeguineue, ecology, distribution.

Reiney M. Kennett, Wildlife Research, Conservation Commission of the Northern Territory,
PO Box 496, Palmerston, Northern Territory O831, Australia, and Faculty of Science,
Northern Territory University, PO Box 40146, Casuarina, Northern Territory 0811,
Australia; Arthur Georges, Ken Thomas and 7.C. Georges, Applied Ecology Research Group,
University of Canberra, PO Box I, Belconnen, Australian Capital Territory 26/6, Australia;

7 November, 199).

Chelodina novaeguineae was first described
from the Binaturi River in New Guinea
{Boulenger, 1888) and later found to occur also
in north-eastern Queensland, Australia (Goode,
1467). Its distribution in Australia is poorly
known, Covacevich and Couper (1991) provide a
map showing its Queensland distribution based
on specimens held in the Queensland Museum
and these data together with anecdotal reports
(Goode, 1967; Cann, 1978) suggest it is dis-
tributed not only on Cape York Penninsula but
throughout the Gulf of Carpentaria, in Amhem
Land, and as far south as Newcastle Waters in the
Northern Territory. More recent collections
(King & Horner, 1987; Covacevich et al., 1990)
confirm that the species" distribution extends into
the Gulf country of western Queensland and the
Northern Territory,

In this note, we provide a range extension for
C, novaeguineae, teview Muscum specimens to
provide an updated map of the distribution of the
species, and provide new information on its
ecology,

RESULTS

On 29 April, 1990, we collected two live
specimens and three shells of C. novaeguineae
from Malogie Waterhole, a small permanent
waterhole in the Roper River catchment near Scar-

let Hill on Kalala Station (16°8’S 133°36'E) in the
Northern Territory. Another three shells were
collected from Stuart Swamp near the township of
Daly Waters (16°14’S 133°27'E) (Table 1), The
specimens were determined to be C. nevae-
guineae on the basis of the following characters:

* The plastronis greatly expanded anteriorly,
less than 1.9 limes longer than broad {un-
like Chelodina rugosa, C. expansa and C.
oblonga), but extends barely as far as the
inner edges of the overlying marginal plates
of the carapace (unlike C, longicellis}.

* Theintergular shield is markedly more than
twice as long as the suture between the
pectoral shields (unlike C. rugosa, C. ex-
pansa and C. oblonga).

* The commen boundary of the 12th mar-
ginals is not raised to form a distinct in-
vented “V' when viewed from the rear
(unlike C_ longicollis).

* The carapace is broadly oval (unlike C.
steindachinert).

Stuart Swamp drains into Daly Waters Creek in
the upper catchment of the Roper River. When
full, the swamp covers several square kilometres
but poor wet scason rainfall over the previous
three years had reduced the waterhole {0 a rela-
tively small muddy hole about 75min diameter at
the time of our visit, The waterhole dricd com-
pletely a few months later for the first Lime in many

180

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE L. Details of Chelodina novoeguinene collected from Kalala Station, Norther Temtory, April 1990,

years (Bill Cook, Kalala Station, pers, comm.),
Malogie Waterhole was the only waterhole in the
area lo retain water in 1990, while many others
dried for the first time in living memory (Bill
Cook, pers, comm).

The turtles were transported to Canberra Where
abdominal palpation on 26 May, 1990 revealed
the female to be gravid. Nine hard-shelled (cal-
careous) eggs Were obtained on 28 May, 1990 by
inducing oviposition with an injection of synthetic
oxytocin (Ewert & Legler, 1978), A tenth egg was
found in the holding tank weeks Jater but was not
measured. Mean egg length was 31,40.2mm,
egg width was 21.8+0.]1mm and egg weight was
8.6+0.1g2 (mean, standard error, n=9). The eggs
were incubated at 30°C in a plastic container on a
bed of vermiculite moistened to a constant ratio of
4e vermiculite to 3g water. An opaque while
patch appeared on each of the nine eggs within 36
hours indicating development had commenced
(Thompson, 1985) and the eggs hatched on 25
September, 1990, 120 days later, Mean hatchling
carapace length was 31.3+1.0mm, mean carapace
width was 26.620,4mm and mean weight was
§.4=0.3g (mean, standard error, 1=3),

DISCUSSION

Recent collections of C. nevaeguinewe (King &
Homer, 1987; Coyaceyich etal,, 1990; this paper)
have substantially itcreased the known Australian
range of the species. Formerly only known from
northeast Queensland (Cogger, 1988), C
novaeguinege has now been recorded from the
Gulf country near the Queensland and Northern
Terntory border (Covacevich et al, 1990) and
from Maria Island in the Gulf of Carpentaria (King
& Horner, 1987). King & Homer (1987) suggest
that the population on Maria Isjand in the Gulf of
Carpentaria may have been established by turtles
swept out of rivers during flooding in cither the
Roper, Towns or Limmen-Bight Rivers, Our find-

ing of C. hovaeguineae in the upper reaches of the
Roper River, is consistent with this suggestion and
conficms Cann’s (1978) reports of C.
novaéguineae in the Daly Waters area and his
reference to their presence in the lower reaches of
the Roper River (Cann, 1972), A list of current
holdings in Australian Museums (AM, Australian
Museum; QM, Queensland Museum; SAM, South
Australian Museum; NTM, Northem Territory
Museum), excluding those of Table 1, is as fol-
lows:

AM R40700, O8°55°S 143°03°E, Kuru [Village],
Western District, PNG; AM R129346, Greta Creek,
near Bowen; QM J5269, |8°04'S 146°33°E, Palm Is.,
NEQ; QM J10265, 20°24’S 148°35’E, Proserpine,
NEQ; QM J13326, 19°16'S 146°49°E, Townsville,
Thornley Pk, NEQ; QM J15560, 315900, 19°13'S
146°38"E, Townsville, Black R., NEQ; QM
¥20627/28/30/3 1/33/35, 14°35"S 144°03"E, Wakooka
Outstation, lagoon 6.4km W, NEQ, 29 Nov, 1970; QM
J36751, 15°25'S 141°58°B, Mitchell R., NWQ, late
1978; QM J37566, 14°07'S 143°16'B, StewartR, 23km
NNW Mt Croll NEQ, 6 Aug, 1979; QM J37819,
17°S3'S 142°34’E, Cape York Peninsula, Gilbert R.,
NEQ, 10 Aug, 1979; QM J45005, 19°14'S 144"02'E,
Lynd Hwy, NEQ, | Dee 198}; QM 147923, 180°39'S
46°01 E, Ingham Q. 19 Mar, 1988; QM J50730,
1916'S 146°49'B, Townsville Town Common , NEQ,
2 Jul, 1990; QM 550731, J50732, 150997, J50998,
16°58'S 138°05'E, Skin SW of Old Doomadgee, NWQ,
24 Jun 1990; SAM R14361, 17°S6°S 138°49°E,
Doomadgee Mission, NWQ, Jun, 1963; NTM R5753 ,
14°53'S 135°43'E, Maria Is., NT, |1 Jul, 1972; NTM
R5898, 14°53"S 135°43°E. Maria Is., NT, 26 Jul, 1972,

Chelodina longicollis, C. novaeguineae and C.
steindachneri are closely related, Their distribu-
tions (Fig. 1) suggest that their common ancestor
may have had a distribution extending around
much of the periphery of the Australian continent.
This widespread species may have exhibited
clinal variation in morpholegy and reproductive
parameters consistent with differences we observe

DISTRIBUTION GF CHELODIA NOY AEGUINEAE

“tS
| Chetogina
“) steinaachaort

Chelodina
fongicallis a

FIG. 1. A map of Australia showing the locations of
collection of specimens held in Australian museums
(@) and the likely distribution of Chelodina
nevaeguineae in stipple. Shown also are the distribu-
tions of C. longicollis and C, steindachneri (atter
Cogger, 1988). The exact boundary between C, lon-
xicollis and C. novaeguineae, while in the Burdekin
drainage (John Cann, pers. comm.), is uncertain.

in these characters among the three species today.
Progressive aridity in the Pleistocene, around
25,000 years ago, or earlier during similar periods
of expanding aridity in the past 15 million years
may have led to the separation of three isolated
populations, the one in the west speciating to
Chelodina steindachneri, the one in the north and
New Guinea speciating into C. novaeguineae and
related forms and the one in the southeast speciat-
ing into C. longicollis. Studies of populations of
C. longicollis and C. novaeguineae in the Bur-
dekin drainage where they are in sympatry or
parapatry with possible hybridization (John Cann,
pers. comm.) are worthy of further investigation.
Chelodina novaeguineae occupies semi-per-
manent and seasonally ephemeral waterholes
throughout its range and occurs sympatrically
with C. rugosa in the Gulf country (Covacevich et
al,, 1990), Unlike C. megosa in these waterholes,
C. novaeguineae will migrate many kilometres
overland to permanent water rather than aestivate
(Covacevich et al., 1990); later expanded in pers.
comm.). This seems to be a hazardous strategy in
waterholes that dry annually. C. novaeguineae in
the Daly Waters area, on the other hand, appear to
acstivate and can be dug out of the mud when the
waterhole dries up (Bill Cook, pers. comm.)

The presence of C, novaeguineae in the Roper
River was known to Aboriginal people at Jinduck-
in near Mataranka, who hint for ‘Nganymalin’,
the ‘smelly annpit turtle’, in small crecks well
upstream (Jesse Roberts, Mangarray) language
group, pers. comm.). A pungent secretion exuded
from Mn ds in the auxillary and inguinal pockets
is a distinctive feature of C. novweguineae and its
southern relative, C. longicollis, Four other
species of freshwater turtle also inhabit the Roper
River, Jarnamanjarr (C. rugosa), “Gorronani’
(Blseya dentata), ‘Marndirrin' (&ntydura
‘australis’) and ‘Jabada’ (Jesse Roberts, pers-
comm,). Jabada is described as a flat-headed
short-necked turtle and may possibly be Elseya
latisternum, although &. latisternum has not been
officially recorded from the Roper. Interestingly,
Aboriginal people at Jinduckin claim that, con-
trary to other reports (e.g. John Bywater, cited in
Georges & Kennett, 1989), the Pig-Nosed Turtle
(Carettochelys insculpta) does not occur in the
Roper, the nearest place they caught them was in
the Victoria River at Timber Creek. The distribu-
tion of C. inseulpta may be more restricted in the
Roper River than in other drainages where it is
found.

The eggs and hatchlings of C. newieguineae
(this study, Grossman, 1988) are similar in size
and appearance to those of C, longieallix
(Vestjens, 1969; Kennett & Georges, 1990). The
bright orange stripes on the plastron of hatchlings
may be aposematic which, allied with their pun-
gent secretion, may act as a deterrent to predators,
The colouration fades with age (Cann, 1981) and
may not be as marked in captive raised juveniles
(Grossman, 1988).

Our gravid specimen and a gravid female col-
lected by John Cann (10 August, 1979) from the
Gilbert River on the Cape York Peninsula north
east Queensland (QM J37819) indicate that nest-
ing may occur throughout the tropical dry season,
equivalent to the temperate autumn and winter. [fF
our observed incubation period of 120 days ap-
proximates the natural incubation periad, then
hatching would occur in the late dry or carly wet
season, equivalent to the temperate spring and
summer. These results are in conflict with the
conclusions of Legler (1981, 1985) who describes
C. novaeguineae as a temperate breeder, nesting
in spring with hatching before autumn. Legler
(1981, 1985) recognizes two groups within the
genus Chelodina and characterizes the groups as
exhibiting a tropical reproductive pattern and a
temperate reproductive pattern, C nevaeguinede
effectively straddles both groups, with the small

182

egg size of the temperate pattern and the nesting
and long incubation period of the tropical pattern.
These data suggest that the reproductive pattern of
C. novaeguineae may have changed as it has
moved north from its hypothesised evolution in
southern Australia (Legler, 1981, 1985) and that
phylogenetic conclusions based on reproductive
patterns may be unsound,

LITERATURE CITED

BOULENGER, G.A. 1888. On the chelydoid
chelonians of New Guinea. Annali del Musco
civico di storia naturale di Genova (2)6:449-452

CANN, J. 1972. Notes on some tortoises collected in
Northern Australia. Victorian Naturalist 89:165-
168.

1978. ‘Tortoises of Australia’,
Robertson: Sydney).

1981. Confessions of a tortoise freak. Geo 3(2):50-
69,

COGGER, H.G. 1988. ‘Reptiles and Amphibians of
Australia’. (A, H. and A, W. Reed; Sydney),

COVACEVICH, J. & COUPER, P. 1991. The reptile
records. Pp. 49. In Ingram, G.J. and Raven, R.J.
(eds), ‘An Atlas of Queensland’s Frogs, Reptiles,
Birds and Mammals’, (Board of Trustees,
Queensland Museum: Brisbane).

COVACEVICH, J., COUPER, P., MCDONALD, K, &
TRIGGER, D. 1990. Walunarra, Bungarra mali,
and the Gangalidda at Old Doomadgee. Memoirs
of the Queensland Museum 29(2):322

EWERT, M. A. & LEGLER, J.M, 1978. Hormonal
induction of oviposition in turtles. Herpetologica
34:314-318.

GEORGES, A. & KENNETT, R. M. 1989, Dry-season
distribution and ecology of Carettochelys in-

(Angus and

MEMOIRS OF THE QUEENSLAND MUSEUM

sculpta (Chelonia: Carettochelydidae) in Kakadu
National Park, Northern Australia. Australian
Wildlife Research 16:323-35

GOODE, J. 1967, ‘Freshwater Tortoises of Australia
and New Guinea’, (Lansdowne Press: Mel-
bourne).

GROSSMAN, P. 1988. Beobachtungen an und Nach-
zucht von Chelodina novaeguineae (Boulenger,
1988). Sauria LO(2):7-11-

KENNETT, R. M, & GEORGES, A. 1990. Habitat
utilization and its relationship to growth and
reproduction of ihe Eastern Long-Necked Turtle,
Chelodina longicollis (Testudinata: Chelidae),
from Australia. Herpetologica 46:22-33.

KING, M. & HORNER, P, 1987. An additional fresh-
water turtle species and the confirmation of
Chelodina novaeguineae in the Northem Ter-
ritory, Australia, The Beagle 4(1):83-84.

LEGLER, J, M. 1981. The taxonomy, distribution, and
ecology of Australian freshwater turtles (Tes-
tudines: Pleurodira: Chelidae). National
Geographic Society Research Reports 13:391-
404.

1985. Australian chelid turtles: Reproductive pat-
terns in wide-ranging taxa. Pp. 117-123. In Grigg,
G,, Shine, R. and Ehmann, H. (eds), ‘Biology of
Australasian Frogs and Reptiles’. (Royal Society
of New South Wales: Sydney).

THOMPSON, M. B. 1985. Functional significance of
the opaque white patch in eggs of Emydura mac-
quarii. Pp, 387-395. In Gngg, G., Shine, R. and
Ehmann, H. (eds), ‘Biology of Australasian Frogs
and Reptiles’. (Royal Society of New South
Wales: Sydney).

VESTIJENS, W.J.M. 1969. Nesting, egg-laying and
hatching of the Snake-Necked Tortoise at Canber-
ra, A.C,T. The Australian Zoologist 15:141-149.

RE-EXAMINATION OF THE VENATION OF OSMYLOPSYCHOPS SPILLERAE
TILLYARD FROM THE TRIASSIC OF QUEENSLAND

KEVIN J, LAMBKIN

Lambkin, K.J. 1992 06 29: Re-examination of the venation of Osmylopsychops spillerae
Tillyard from the Triassic of Queensland. Memoirs of the Queensland Museum: 32(1):

183-188, Brisbane. ISSN 0079-8835.

The finding of new material has permitted re-examination of the venation of the Triassic
neuropteran Osmylopsycheps spillerae Tillyard, Many of the features of Tillyard’s
reconstruction of the forewing of the species are shown to be erroneous. [JOsmylopyychops
spillerae, Neuroptera, Insecta, Triassic, Queensland.

Kevin J, Lambkin, Honorary Research Fellow, Queensland Museum, PO Box 3300, South
Brisbane, Queensland 4101; Correspondence ta; 75 Alexandra Si, Bardon, Quevnslanel,

Australia 4065; 22 November, 1991.

In the last of his 10 part series on the Mesozoic
insects of Queensland, Tillyard (1923) described
u broad-winged psychopsid-like lacewing from
the Triassic of Denmark Hill in south-eastern
Queensland (27°37'S, 152°47’E) as Osmylop-
sychops spillerae Tillyard. Osmylopsyehops Til-
lyard was ascribed to the new family
Osmylopsychopsidae by Martynoya (1949) and,
in 1953, Ellenberger et al. augmented the genus
with their description of O. radialis from the
Upper Triassic of France.

O. spillerae was based on two wing fragments
(Tillyard, 1923, pl. XLII, figs 39,40), from
which Tillyard ‘reconstructed’ the forewing
venation (Fig.1). Because of the fragmentary na-
lure of this type material (Figs3,4), much of the
reconstruction was notional and, as often happens
with such “best guess* reconstructions, its publi-
cation and subsequent reproduction (e.g. Riek,
1970) entrenched in the literature an assumed to
be accurate ‘complete’ venational scheme that
became a baseline for subsequent comparisons
with similar species.

The purpose of the present work is to provide a
more accurate view of the venation of QO. spillerae
and in so doing to exclude Tillyard’s reconstruc-
lion, visually appealing though it may be, from
the data set of fossil Neuroptera. The re-examina-
tion of the venation of the species has been made
possible by the collection in more recent years of
several specimens of O. spillerae from the Trias-
sic of Mount Crosby in south-eastern Queensland
(27°32°S, 152°48"E), from where the species was
first recorded by Riek (1955), Examination of this
new material as well as of Tillyard’s original
Denmark Hill specimens provides an almost
complete picture of the forewing venation of the
species (see Fig.2).

Tillyard’s matenal was collected in grey shales
of the Blackstone Formation that were exposed
in asmall quarry at Denmark Hill, a suburb of the
town of Ipswich (Dunstan, 1916). Mount Crosby
fossil insects occur in green shales of the Mount
Crosby Formation and have been collected at five
separale exposures designated as Fossil Insect
Localities A, B, C, D and E (details in Allen,
1961), Both Formations are Late Triassic (Kar-
nian) and form part of the Ipswich Coal
Measures, with the Mount Crosby Formation
lower in the sequence (Cranfield et al., 1976).

With one exception, the determination of
whether the isolated wing fragments described
herein are of fore- or hindwings has not been
possible. The closest living species for compara
live purposes is the psychopsid Megapsychops
illidgei (Froggatt), In this species the basic vena-
tional patterns of the fore- and hindwings are very
similar (see Tillyard, 1919, pl. LAX VII, fig.4;
New, 1988, figs}96,197) and, because of this
similarity, 1 believe it is valid, for the sake of
convenience of discussion, to regard all wing
fragments of O. spillerae considered herein as
probable forewings. The exception to this uncer-
tainty is specimen UQDE CB 14 (Fig.6). Ithas the
proximal regions of AAj4 and AP widely
separated and the stem of AAi+ clearly
developed (both forewing characters of M, il-
lidgei and other living psychopsids), and is there-
fore almost certainly a forewing.

The venational terminology used herein is that
of Kukalové-Peck (1983, 1991; see also
Lawrence et al., 1991). The following muscum
abbreviations apply: GSQ, Geological Survey of
Queensland; QM, Queensland Museum; UQDE,
University of Queensland, Department of En-
tomology.

(iy 2
(\ /

cr
aA

Nn, >

HE

: yg

= , ,

p 4 :
S) \ e &
ja

ye

SX Va

=

Lt |

2

t,
é

REXAMINATION OF THE VENATION OF OSMYOPSYCHOPS SPILLERAE

MATERIAL EXAMINED

Holotype GSQ 314 (central region of ?forew-
ing), “Osmylopsychops / reevei spillerae Till. /
Holotype / Denmark Hili / Ipswich’, ‘3149.9';
Heautotype [= autotype] GSQ 283a (apical */s of
?forewing}, ‘G’, ‘Osmylopsychops / reevet spil-
lerae Till. { Heautorype/ Denmark Hill / Ipswich
( Trias.’; QM F18921 (central region of ?forew-
ing), Mount Crosby Fossil Insect Locality D (col-
lected by K J Lambkin, 20.viii.1972); UQDE
CB14 (base of forewing), Mount Crosby Fossil
Insect Locality By QM F18922 (apex of ?forew-
ing), ‘L114 Mt Crosby shale / (910805-911805
Ipswich / 1 mile military map - upper bed) /
10.vii.1969 ¢ E. Dahms’, [Mount Crosby Fossil
Insect Locality Cj; QM F18923 (?forewing frag-
ment), Mount Crosby Fossil Insect Locality B
(collected by K ] Lambkin, 24.viti.1974),

DESCRIPTION OF FOREWING VENATION
(Figs 2-8)

Broad Wing of psychopsid type, length c.
138.5mm (Fig.2); costal space quite broad basally,
tapering apically (Figs4,5); costal crossveins ir-
regular, mostly deeply forked and extensively
branched, almost hpright basally, becoming
gradually more inclined apically, more closely
spaced in ‘pterostigmal’ area, with a limited
series of interconnecting crossveins basally just
above ScP (Figs3-7); ScP converging and either
Tfused of Toonnected by a very short crossvein
with RA apically (Figs4,7), entering margin
before wing apex; 1 upright scp-ra just beyond
tadial fork (Figs5,6); ScP. RA and stem of RP
curved apically (Fig.2), 1 inclined fa-rp at c.
midlength (Fig.5); RP with at least 18 pectinate
branches, a few of these forked basally or c.
midlength, all with apical branches (Figs4,5,7,8);
crossveins of RP field apparently few in number,
only 1 or 4 detected (Figs 5,8); MA simple or
deeply forked (Figs3,5,6,8); MP occupying a4
large central area of the wing, its primary forking
very close to the base (before that of R - Fig.6),
with 6 or 7 branches at midlength (Figs3,5,6), and
extensively dichotomously forked apically
(Figs5,7); MP)\42 dichotomous in holotype, butin
some specimens tending to pectination, in UQDE
CB14 this pectination mirroring that of RP
(Fig.6); MPss4 forking a litle later than MPy42
(Figs3,6,8), MPs of holotype with an enclosed
cell at c. midiength (Fig,3): medial, cubital and
anal fields witha few randomly placed crossveils
as illustrated in Figs3.5.6,8 (possibly preserya-

185

tion dependent); CuA extensively dichotomously
forked from c.7/3 length with branches curved
anteriorly (Figs3,5}, CuP occupying only a nar-
row area of the wing, more deeply dichotomously
forked than CuA, CuP1 more deeply forked than
CuP2 (Figs3,5,6); AAr2 developed as a shor
spur directed obliquely tawards Cu (Fig.6);
AAgeq extensively dichotomously forked from
cJ¥q length (Fig.6); AP1+2 dichotomously
forked from c.'/2 length, less extensive than
AAass oF AP3s4 (Fig.6); APs joined to AP2 hy a
short crossycin basally, simple for almost entire
length, with 2 small branches apically (Fig.6);
AP, extensive, occupying as much of the margin
as APi, APz and APa combined, extensively
dichotomously forked with branches directed
apically (Fig,6); iichosors present, at least along
fore margin (Figs5,6); 1 nygma detected, placed
centrally between 2nd and 3rd branches of RP
(Fig.5).

NorTes

Figure 2 is a composite of UQDE CB 14 (Fig,6),
QM F1892] (Fig.S), QM F18922 (Fig.7), ws well
as a small part of GSQ 283a (Fig.4)
(‘pterostigmal’ region only). It Wlustrates the
known features of the presumed forewing of O,
spillerae, The major differences between this
composite and Tillyard’s reconstruction (Fig.1)
are listed below. Certain features, such as the
wing shape and the venation of the humeral
region and paris of the fore, apical and hind
margins, are not preserved in the material cx-
amined and remain unknown,

1. Structure of MP (= M of Tillyard’s notation)

Because Tillyard's specimens did not show the
important basal structures of the main veins (as
now provided by UQDE CB14 - Fig,6}, he con-
jectured that MP was of minor extent with its
primary forking well beyond that of R (as is ap-
parently the case in recent psychopsids). The new
material indicates that MP forks very early (before
the primary radial fork), is multibranched, und
occupies a large central area of the wing. The veins
shown by Tillyard as the first two branches of RP
(Rs of his notation) are indeed part of MP.

The structure of MP in O. spillerae is especially
interesting in providing a clue (o the homology of
‘MP’ in the wings of living species of the
Psychopsidae. In the forewing of these species
there is a variably developed weak posterior
branch leaying MP before the primary forking of
R, In exactly the same position as MPa in 0.
spillerag, Iw larger species fe.g, Mf. ilidgei,

186 MEMOIRS OF THE QUEENSLAND MUSEUM

FIGS 3-8, Osmylopsychops spillerae (proximal sections of ScP, RA, RP, CuA (thicker veins) not drawn to scale).
3, GSQ 314 (holotype). 4, GSQ 283a (*heautotype’). 5, QM F18921. 6, UQDE CB14. 7, QM F18922. 8, QM
F18923.

REXAMINATION OF THE VENATION OF OSMYGPSYCHOPS SPILLERAE

Psychopsis barnardi Tillyard) this branch exists
asa very weak vein which peters out after running
for a short distance along a fairly deep furrow in
the membrane. In species of smaller size the
furrow and a kink in the stem at the point of
forking (e.g. P. insolens McLachlan), or simply
the suggestion of a fold in the membrane (e.g. P.
coelivagus (Walker)). is all that remains. Based
on the structure of MP in O. spillerae, the much
reduced extreme basal posterior branch of MP
described above is the rermant of MP3.4 and the
vein system usually labelled as ‘MP’ in living
psychopsids is MPi42 only.

2. Structure of Cu

In Tillyard’s reconstruction the primary forking
of Cu is a considerable distance from the base,
and CuP forks at the same level as CuA and has
only three branches. Specimen UQDE CB14
(Fig.6) shows that the primary fork of Cu is
placed extreme basally (as oceurs in the Psychop-
sidae and most Neuroptera), and that CuP forks
much earlier than CuA and has at least six major
branches

3. Structure of A

The anal area in Tillyard's reconstruction is
almost entirely conjectural and bears only mar-
ginal resemblance to that described herein (see
especially Fig.6). The new information shows
that the basal sections of Cu-CuP, AAz+4 and
APj4z are much more widely spaced than in
Tillyard’s reconstruction; the basal section of
AAaiais curved anteriorly, not posteriorly; AA3+4
and AP}42 are not fused at the base; and AP344 is
smaller in area and has fewer major branches
which are directed apically rather than posterior-
ly.

4. Primary forking of R

Tillyard’s reconstruction shows the primary
forking of R very close to the base of the wing.
UQDE CB14 (Fig.6) shows that R forks much
further distally than this, well beyond the pnmary
forkings of Cu and AA3.4.

5. Marginal end-twigging of veins

End-twigging of Veins is much more
pronounced along the fore and hind margins (see
Fig.2) than indicated by Tillyard.

6, Wing proportions

Comparison of Figures | and 2 indicates that
the presumed forewing of O. spillerae is not us
broad as suggested hy Tillyard.

187

The relationships of QO. spillerae (and O.
radialis which, based on the figure of Ellenberger
el al. (1953, fig.}), appears to be congeneric)
remain problematical, Osmrylopsychops is just
one of a large number of diverse Mesozoic
psychopsid-like wings (see Martynova, 1949,
and numerous subsequent descriptions of aclkdi-
tional species) that, as a group, requires critical
re-examination and reappraisal. Whatcan be said
al this stage, however, is that Osmylopsychops
does not possess the set of apomorphic wing
features characteristic of the Psychopsidae (viz
ScP, RA and stem of RP evenly spaced and
braced by numerous crossveins; MP3.4 much
reduced; CuA and CuP strongly posteriorly pec-
tinate; crossveins mostly aligned into two or three
egradate series} and is thus not attributable to that
family {see also Riek, 1955),

ACKNOWLEDGEMENT

Jam grateful for the assistance of Susan Partrey
of the Queensland Department of Resource In-
dustrics who arranged the loan of Tillyard's

types,
LITERATURE CITED

ALLEN, R.J, 1961, The Kholo Sub-group al the
Ipswich Coal Measures. Geological Survey of
Queensland Publication 300; 1-14,

CRANFIELD, L.C., SCHWARZBOCK, H. & DAY.
RW, 1976. Geology of the Ipswich and Brisbane
1:250000 Sheet areas, Geological Survey of
Queensland Report 95; 1-176,

DUNSTAN., B. 1916. Mesozoic and Tertiary insects of
Queensland and New South Wales Stratigraphi-
cal features, Geological Survey of Queensland
Publication 253: 1-10.

ELLENBERGER, F, & P., LAURENTIAUX, D. &
RICOUR, J. 1953. Note préliminaire sur la fauna
et un niveau insectifére des lentilles de grés ct
schistes noirs des gypses de la Vanoise (Trias
Supérieur), Bulletin de la Société Géologique de
France (6)2: 269-274.

KUKALOVA-PECK, J. 1983. Origin of the insect
wing and wing articulation from the arthropodun
leg. Canadian Journal of Zoology 61: 1618-1669,

1991], Fossil history and the evolution of hexapod
structures. Pp. 141-179, In CSIRO (ed,), “The in-
sects of Australis A textbook for students and
research workers’, 2nd ed. Vol.J. (Melbourne
University Press; Carlton), S42pp.

LAWRENCE. L.F., NIELSEN, E.S, & MACKER-
RAS, LM. 1991. Skeletal anatomy ond key to

188 MEMOIRS OF THE QUEENSLAND MUSEUM

Orders. Pp.3-32. In CSIRO (ed.), ‘The insects of 1970. Fossil history. Pp.168-186. In CSIRO (ed.),

Australia: A textbook for students and research ‘The insects of Australia: A textbook for students
workers.’ 2nd ed. Vol.1. (Melbourne University and research workers.’ (Melbourne University
Press: Carlton). 542pp. Press: Carlton). 1029pp.

MARTYNOVA, O.M. 1949. Mesozoic lacewings TILLYARD,R.J. 1919. Studies in Australian Neurop-
(Neuroptera) and their bearing on concepts of tera, No.6. The family Psychopsidae, with
phylogeny and systematics of the Order. Trudy descriptions of new genera and species. Proceed-
Paleontologicheskogo Instituta 20: 150-170. [In ings of the Linnean Society of New South Wales
Russian] 43: 750-786.

NEW, T.R. 1988. The Psychopsidae (Insecta:Neurop- 1923. Mesozoic insects of Queensland. No.10. Sum-
tera) of Australia and the Oriental Region. Inver- mary of the Upper Triassic insect fauna of
tebrate Taxonomy 2: 841-883. Ipswich, Q. (With an appendix describing new

RIEK, E.F. 1955. Fossil insects from the Triassic beds Hemiptera and Planipennia). Proceedings of the
at Mt. Crosby, Queensland. Australian Journal of Linnean Society of New South Wales 48: 481-

Zoology 3: 654-691. 498.

NOTES ON SPONDYLUS LINNAEUS WITH DESCRIPTIONS
OF TWO NEW SPECIES
FROM WESTERN AUSTRALIA (MOLLUSCA: BIVALVIA: SPONDYLIDAE)

KEVIN LAMPRELL

Lamprell, K. 1992 06 29: Notes on Spondylus Linnaeus with descriptions of two new species
from Wester Australia (Mollusca: Bivalvia; manele Memoirs of the Queensland
Museum 32(1): 189-194. Brisbane. ISSN 0079-8835.

Two new species of Spondylidae, Spondylus clarksoni spmov. and S. eastae sp.nov., are
described from Western Australia. Additional taxonomic notes are presented on ather Recent
Spondylidae. Spondylus asperrimus Sowerby, 1847 is elevated to specific rank; 5. buileri
Reeve, 1856 is removed from the synonymy of S. longitudinalis; 8. erinaceus Reeve, 1856 is
relegated to the synonymy of S. gilvus Reeve, 1856; §. pratri Parth, 1990 is placed in the
synonymy of §. pictorwn Schreibers, 1793; areliable locality is established for S. gravis Fulton,
1915; S. raoulensis Oliver, 1915 (July) is given priority over S. iredalei Fulton, 1915 (October);

and S. prionifer Iredale, 1931 is redescribed.
new species, taxonomy, Western Australia.

‘ollusca, Bivalvia, Spondylidae, Spondylus,

Kevin Lamprell, Queensland Museum, PQ Box 3300, South Brisbane, Queensland 4/01,

Australia; 3 February, 1992.

Since publication of Spondylus: Spiny Oyster
Shells of the World (Lamprell, 1987), I have
studied type specimens in overseas museums and
a range of new material from the Australian
region, Two undescribed species of Spondylus
from Western Australia were ftecognised and
taxonomic information on several previously
described species was updated.

Abbreviations used in text: AM = Australian
Museum; QM = Queensland Museum; WAM =
Western Australian Museum; MV = Museum of
Victoria; py = paired valves.

SYSTEMATICS

Family SPONDYLIDAE Gray, 1826
Spondylus Linnaeus, 1758

Spondylus clarksoni sp.nov.
(Fig. la-d)

ETYMOLOGY
For Mr Peter Clarkson who donated specimens
for study.

MATERIAL EXAMINED

HOLOTYPE: WAMS562-91, 1 pv, exposed on dead coral
faces, 9-13m, Bennett Shoal, central Exmouth Guif,
W.A., 1991, P. Clarkson. Height 100mm, length 86mm.
width 48mm.

PARATYPES: AMC167693, MVF6552, QMMO032909,
3 py, exposed on sponge covered limestone reef in tide
np, 15-22m, S end South Muiron 1., Exmouth Gulf,
W.A., 1991, P. Clarkson.

OTHER MATERIAL: Four specimens from Pt Lefroy,
southern end Exmouth Gulf, north W.A., P. Clarkson.

DIAGNOSIS

Shell pecten-shaped, slightly oblique, solid.
Hinge typical for the genus. Top (left) valve
moderately convex, more so umbonally; internal-
ly, slightly excavated under the hinge plate with a
strong, wide, crenulated margin; hinge-line
moderately wide, Sculpture of five, strong,
rounded, principal radial ribs ornamented by
strong, upright, spatulate spines which are chan-
nelled on the underside, narrow and strong at their
bases, wide and flattened at their tips. Rib inter-
stices with five to twelve minor radial ribs, central
longer and stronger ornamented with dense, sharp,
overlapping spines, and whose interstices contain
smaller ribs which are ornamented with even
smaller and finer spines or nodules. Lower (right)
valve equally as convex as the top valve; cardinal
area triangular and moderately deep; internally,
moderately deep under the hinge plate with a
strong, wide, crenulated margin. Sculpture (where
not affixed) of thin concentric foliations with few,
irregularly placed, strong spines. Area of attach-
ment varying from 50-100% of the lower surface.
Colouration pure white with pink-rose umbonally
extending approximately one-third down the shell
surface. Height 70-100mm. length 70-87mm,
width 38-54mim.

HABITAT AND DISTRIBUTION
Covered in red sponge attached to low-profile
limestone reefs, on a sandy bottom, among spon-

190

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. |. Spondylus clarksoni sp.nov. a-d, Bennett Shoal, central Exmouth Gulf, Western Australia. WAM562-91,
holotype. a, top valve. b, sculpture on top valve. c, side view. d, interior of lower valve.

ges and gorgonian corals. Apparently restricted to
north Western Australia (Shark Bay to Broome).

REMARKS

Compared with Spondylus clarksoni, S. sinensis
Schreibers, 1793 has fewer, stronger, interstitial
ribs devoid of ornamentation, strong principal ribs
on the lower valve (absent in S. clarksoni) and a
very small attachment area. Spondylus victoriae
Sowerby, 1843, which occurs sympatrically, has
long, sharp to spatulate principal spines on both
valves and fewer, stronger interstitial spines.Com-
pared with S. tenellus Reeve, 1856, which occurs
in southern Australia, S. clarksoni is larger and
heavier, has stronger ribs, wide, spatulate spines
and weaker interstitial spining. The spining and

ribs in both valves of S. tenellus are always orange
or pink whereas those of S. clarksoni are white.

Spondylus eastae sp.nov.
(Fig. 2a-d)

ETYMOLOGY
For Aileen East who has donated material for
study.

MATERIAL EXAMINED

HOLoTyPE: WAMS563-91, 1 pv, 7-l1m, Schofield
Shoal, SW Exmouth Gulf, W.A., 1991, P. Clarkson.
Height 82mm, length 64mm, width 37mm.
PARATYPES: QM MOQ32910, AM C167694, MY
F6553, 3 pv, exposed on sponge covered limestone reef

NEW SPECIES OF SPINY OYSTERS 191

FIG. 2. Spondylus eastae sp.nov. a. Schofield Shoal, South West Exmouth Gulf, Western Australia, WAM563-91,
holotype, b-d, Muiron I,, Exmouth Gulf, Western Australia, paratype. a, top valve. b, sculpture of top valve. c,

interior of lower valve. d, side view.

in tide rip, 15-22m, S end South Muiron I., Exmouth
Gulf, W.A., 1991, P, Clarkson.

OTHER MATERIAL: Cape Keraudren, W.A., attached to
rock at low tide, Sept. 1987, K. Lamprell; Port Hedland,
W.A., attached to coral, low tide, Sept. 1987, K.
Lamprell; Exmouth Gulf, Town Beach, 6m, attached to
rocks, Sept. 1987, K, Lamprell and P. Spoor; Cape
Keraudren, W.A,, Sept. 1982, L. Marrow.

DIAGNOSIS

Shell ovate to pear-shaped; solid, Hinge typical
for the genus. Top (left) valve moderately convex
umbonally, remainder of surface depressed to
moderately concave; internally smooth, slightly
excavated under the hinge plate, with a strong,
wide, moderately crenulated margin; sculpture of
numerous, weak radial ribs, ornamented by short,

192

dense, strong, depressed, sharp spines and a few
longer, irregular, flat, spatulate spines which are
stronger at their bases. Rib interstices with a few
irregular, obsolete radial ridges which are orna-
mented with dense, nodules or small spines,
Lower (right) valve convex, moderately shallow
to deep; cardinal area narrowly triangular, shal-
low; internally moderately excavate with wide.
moderately crenulated margin; sculpture (where
not affixed) of thin concentric foliations covering
most of the surface, or otherwise with similar ribs
and ornamentation to upper valve. Area of attach-
ment varying from 10-70% of lower surface-
Colouration typically brown/maroon to brick- red,
orange or purple; umbones white with small flecks
of brown; lower valve with similar colouration to
top valve; internally blue-white centrally with
deep purple-brown, red or orange margins and a
dark brown margin towards the hinge. Height 72-
93mm, length 57-79mm, width 30-42mm.

HABITAT AND DISTRIBUTION

Embedded in dead coral boulders and depres-
sions on reef, among sponges and gorgonian
corals; usually covered in red sponge. Restricted
to north Western Australia.

REMARKS

Compared with Spondylus eastae, S. barbatus
Reeve, 1856 has five or six strong, principal ribs
with alternating, dense, sharp and spatulate spines,
and colour bars or bands at the auricles. Spondylus
vielascens Lamarck, 1819 has been confused with
5. eastae but has much stronger and denser spining
on both valves, much less affixation and spatulate
spines which are hollow rather than flat. The
north-west Australian distribution given by
Lamprell (1987) for 8. violascens represents the
range of S. castae, Spondylus multiniuricatus
Reeve, 1856 is similar in shape and size to S.
eastae but has stronger radial nbs omamented
with hollow, blunted spines (not spatulate) and
with interstices that have few, irregular, smaller
stout spines,

ADDITIONAL NOTES
Spondylus asperrimus Sowerby, 1847

REMARKS

Lamprell (1987: pl.3, fig. la-c) considered this
species to be a form of §. linguafelis Sowerby,
1847. However it can be separated from that
species by its more elongate shape, shorter spines:
and rows of fine nodulose spines in the interstices.

MEMOIRS OF THE QUEENSLAND MUSEUM

It is usually affixed by almost the entire lower
valve (or surface) whereas S. linguafelis is usually
affixed by the cardinal area.

DISTRIBUTION
Philippines, north Western Australia and central
Queensland coastal areas.

Spondylus botleri Reeve, 1856

REMARKS

Lamprell (1987: p14, fig. 2a-c) considered 5.
longitudinalis Lamarck, 1819 to be a prior name
for the more widely used S. burleri, The type of 8.
longitudinalis (Museum of Natural History,
Geneva) was examined and considered to be the
same as S. zonalis Lamarck, 1819. Another
specimen figured and referred to by Chenu (1845:
pl.9, fig 2-2a) as S. longitudinalis (but not marked
asa type), was even more similar to the type of S.
zonalis . Fulton (1915:360, species 86) remarked,
*Lhave received a photograph of the shell figured
by Chenu (S. longitudinalis), which is now in the
Geneva Museum; the authorities there cannot say
with any certainty that it is the actual type
specimen’. Spondylus butleri: should be used for
the species figured in Lamprell (1987: pl.4, fig
Qa-c),

DISTRIBUTION ‘
Philippines, Madagascar, Micronesia and the
Great Barrier Reef, Queensland.

Spondylus gilvus Reeve, 1856

§. erinaceus Reeve, (856.

REMARKS

Although figured separately by Lamprell
(1987: pl. 17, tig. 2a, b; pl. 17, fig. 1), types
(British Museum Natural History) of 8. gilvus and
5. erinaceus revealed insufficient variation to
maintain their separation.

Spondylus pictorum Schreibers, 1793

S. crassisquamatus Lamarck, 1819.
3. prant Parth, 1990,

REMARKS

Three specimens of S$. pratt? from the type
locality (Somalia) were compared with S. pic-
torum. Two were very mature but retained most
of their sculpture; the third was younger with full
sculpture. In describing §. prasi, Parth referred to

NEW SPECIES OF SPINY OYSTERS

the two valves as unequal, the top valve larger and
more gibbose. In all specimens examined, each
valve was equally convex and gibbose. Parth also
referred to both valves of S. pratti as equally
sculptured with six principal ribs. In two of the
specimens the top valve had six principal ribs
while in the lower valve, one specimen had six and
the other seven principal ribs.

While Parth (1990:6) only compared S. prarti
with S. americanus ‘because of its large size’,
numerous other Spondylus are equally as large or
larger than this species: in particular, S. pictorum
Schreibers, 1793 (Lamprell, 1987: pl. 23, fig, la-
d), which shares a similar distribution to $. pratti,
viz Somalia. Spondylus pictorum is equally as
convex as S, pratfi, has similar sculpture, nb
count, interstitial sculpture and spining; and both
have stout, upright principal spines that are chan-
nelled beneath, extending the full length of the
spine. Colouration in specimens examined was
similar to $. pictorum: white in base colour with
bands of pink, red or brown and pink in the um-
bonal area. The juvenile specimen had white in the
interstices and pink spines whereas, in some
specimens of S. pictorwnt, the interstices are white
and red or orange-brown. I am unable to find any
features, apart from this interstitial colour varia-
lion, to separate S. pratti from §. pictorwm.

Spondylus gravis Fulton, [915

REMARKS

The type is in the British Museum (Natural
History) without a specified locality, Recent
trawling of several specimens (Lamprell, 1987: pl.
28, fig. 1) ih 200-300m off Capo Ras Hafun,
Somalia, gives reliable locality data.

Spondylus raoulensis Oliver, 1915
S. fredale? Fulton, 1915.

REMARKS

Lamprell (1987; pl. 29, fig, 4) incorrectly listed
this species as §. fredalei. Fulton’s work was
published in two parts and the section naming 3,
iredalei was included in the second part printed in
October 1915, The descnption of §. raeulensis by
Oliver was published on the 12 July 1915 and thus
has priority over S. fredalet.

Spondylus prionifer Iredale, 1931

REMARKS
Lamprell (1987: pl.25, fig.1) moorrectly listed

193

Spondylus prionifer as a synonym of S. tenellus
Reeve, 1856. The holotype (Australian Museum,
Sydney) is immature and superficially similar to
§_fenellus, However, numerous specimens recent-
ly collected on the Gneering Shoals,
Mooloclooba, south Queensland, differed fram 3.
lenellus. On the basis of this material, 2 more
detailed description of S. prionifer follows, The
species was figured by Lamprell (1987: pl. 25, fig.
3b; pl, 30, figs 1, 2).

DESCRIPTION

Shell height to 85mm, width to 65mm and depth
of conjoined yalves to 45mm, pecten-shaped. Top
(left) valve slightly convex to depressed; lower
(right) valve moderately deep, cup-like, cardinal
area trigonal, moderately extended; internally
with the margins strongly crenulated; area under
the lunule and hinge plate, deep and extended;
sculpture of both valves with 35-40 strong radial
Tibs with weaker intermediate ribs in the inter-
stices; ribs ormamented with numerous depressed
or hooked spines which vary from sharp to spaiu-
late, stronger marginally and with a longitudinal
furrow beneath and finely denticulate edges; in-
terstitial ribs with two or three rows of fine spines
or striae. Area of attachment of the lower valve
variable but usually restricted to the umbonal area,
Colouration variable. usually white with con-
centric bands of red and white ribs marginally,
rarely all orange or red-brown; internally bluc-
White with margin crenulations variegated white
and red or orange.

HABITAT AND DISTRIBUTION

Atiached to corals or rock to 33m. Central
Qucensland (Capricorn Channel), south
Queensland (Gneering Shoals) and Sydney Har-
bour, N.S.W,

ACKNOWLEDGEMENTS

Thanks are due to the Curators and staff of the
Britsh Museum (Natural History), Museum Na-
uional d'Histoire Naturelle, Paris and the Museum
of Natural History, Geneva for their assistance in
examining essential study material; to Dr John
Stanisic, Queensland Museum and Dr John Healy,
Queensland University for critically reading the
manuscript and for advice on manuscript prepara-
tion; to Mr lan Loch, Australian Museum, for his
research on Sponaylus iredalei/raoulensis; and to
Mr Luigi Bozzeiti, Italy, for the loan of specimens.

This work was carned out through a grant from
the CSIRO, Science and Industry Endowmeni

194 MEMOIRS OF THE QUEENSLAND MUSEUM

Fund. Overseas photographic expenses were par-
tially met by funds received from the Keith
Sutherland Award, Australian Museum, Sydney.

LITERATURE CITED
CHENU, J.C. 1845. ‘Illustrations conchyliologiques’.

Vol.3. Bivalves. Part 2. (Paris).
FULTON, H.C. 1915. List of the Recent species of

Spondylus Linné with some notes and descriptions
of six new forms. Journal of Conchology
14(11):331-360.

LAMPRELL, K.L. 1987. ‘Spondylus: spiny oyster
shells of the world’. (Robert Brown and As-
sociates: Bathurst).

PARTH, M. 1990. Spondylus pratti, spec.nov., eine
neue Art aus Somalia (Mollusca, Bivalvia, Spon-
dylidae). Spixiana 13(1): 5-7.

THE SUBTRIBE HICKSBEACHINAE (PROTEACBAB)
IN THE AUSTRALIAN TERTIARY

ANDREW C_ ROZEFELDS

Rozefelds, A.C. 1992 06 29: The subwibe Hicksbeachiinae (Proteaceae) in the Australian
Tertiary, Memoirs of the Queensland Muséurn 32(1): 195-202. Brisbane. ISSN 0079-8835.

Wilkinsonia bilaminata F. Muell,, a fossil fruit from deep leads at Gulgong, New South Wales
is similar to fruits of extant genera Athertonia and Heliciopsis (subtribe Hicksbeachiinae:
Proteaceae), Athertonivides Rozefelds is considered a synonym of Wilkinsonia.. Fossil
evidence of Wilkinsortia in the mid-Tertiary demonstrates that the subtribe was more
widespread than at present. Wilkinsonia is more closely related to Athertonia, the aorth
Queensland rainforest endemic, than to the South East Asian genus Heliciopsis.C] Proteaceae,
Aicksbeachiinae, Athertonia, Heliciopsis, Wilkinsonia, Athertonivides, fossil fruit, Australia.
Tertiary.

Andrew C, Rozefelds, Queensland Museum, PO Bax 3300, South Brisbane, Queensland, 4301,
Australia; present address: School of Botany, University of Melbourne. Parkville, Victoria

3052, Australia; 30 January, 1991,

Mueller (1879, 1883) described Wilkinsonia
bilaminata on the basis of woody fruits from deep
leads (buried placer deposits) below basalts in the
Orange and Gulgong gold mining areas of New
South Wales. Mueller (1879, 1883) recorded W.
bilaminata from Beneree, near Orange at a depth
of about 110-130ft and in deep leads at Gulgong
at a depth of 140-175ft (Fig. 1). He was unsure of
the affinities of W. bilaminata but suggested
(1879: 170) that it ‘may perhaps have belonged to
the order of Sapindaceae; but with the material at
present available it is impossible to find its exact
ordinal position, as no extemal resemblance indi-
cates any close relationship to existing genera’.
The only surviving specimen located is in the
Museum of Victoria (NMVP53092) from the Gul-
gang locality in New South Wales.

Palynological studies of the Home Rule Kaolin
Deposit which is part of one of the deep Jeads in
the Gulgong area indicates they are of late early to
middle Miocene age (McMinn, 1981). Potassium-
Argon dates of 14.8=1.2 and 13,8+1.1my BP,
for basalts that overlie deep leads in this area
(Dulhunty, 1971) indicate a minimum mid-
Miocene age for this Flora.

Fruits from the Oligocene Glencoe locality in
central Queensland were placed in a new form
genus Athertonioides which closely resembles
those of the north Queensland endemic Ather-
tania, and South East Asian genus Heliciopsis,
(both in subtribe Hicksbeachiinae) (Rozefelds,
1990a). Re-examination of topotypic material of
W, bilaminata P. Mueil. from the Gulgong locality
also suggests inclusion in Hicksbeachtinae (Fig. 2).

In this paper W. bilaminata is redescribed and

it’s affinities discussed. The status of Afher-
tonioides Rozefelds is also re-examined. Fruit
anatomy of extant genera Heliciopsis, Athertonia
and Hicksbeachia 1s also figured and the fossil
record of Hicksbeachiinae in Australia js
reviewed.

METHODS AND MATERIALS

To permit comparison with the fossil material,
the soft outer exocarp and mesocarp layers of
modern fruits were removed. Persistent fibres at-
tached to the stony layer (endocarp) of the fruit
were removed with a nylon brush, The endocarp
was sectioned with a coping saw and the
transverse sections polished with 1200 grit “wer
and dry sandpapers’. The sections were
photographed with a Carl Zeiss DRC Stereo
Microscope with MC63 camera attachment. Com
parative modern Australian material figured in
this paper will be donated to the Queensland Her-
barium (BRI).

The term endocarp is used here in a descriptive
sense and follows that of Sleumer (1955a,b),
(Smith & Haas, 1975) and Filla (1926). Johnson
& Briggs (1975), however, have concluded that
this inner stony layer is part of the mesocarp. The
mesocarp consisting of an outer zone of
parenchymatous tissue with radial fibres and inner
dense hard zone of tangentially oriented fibres.

Silicified fossil material from Gleneoe
(Queensland Museum locality QML511) is
preserved as moulds with little or no infernal ce),
lular preservation, Gulgeng and other deep Jeads
fruits approach fursanites in appearance, which

196

suggests they have been subjected to considerable
heat and pressure. During fossilization, the more:
volatile endocarp wall constituents are lost, leay-
ing only the ligneous structure of the fruit. In
modern fruits the volatile endocarp wall con-
stituents are still present making comparisons dif-
ficult.

SYSTEMATIC PALAEOBOTANY

Family PROTEACEAE
Tribe MACADAMIEAE
Subtribe HICKSBEACHIINAE

Rozefelds (1990a) erected Athertonioides glen-
coensis for fossil fruits collected from the Glencoe
locality in central Queensland. Re-examination of
Wilkinsonia bilaminata also indicates close af-
finities with Athertonia. It is not warranted to
maintain two form genera for fossil Athertonia-
like fruits and so Athertonioides is here reduced to
a synonym of Wilkinsonia. In his diagnosis of
Wilkinsonia, Mueller (1879, 1883) drew
prominent attention to the flattened nature of the
endocarp and the ridging which are features, that
are in part, due to compression during fossiliza-
tion. An emended diagnosis of Wilkinsonia
bilaminata is provided. The original specimen
figured by Mueller (1879: pl. 3, fig. 4a,b) has not
been located in the Museum of Victoria, Geologi-
cal Survey of New South Wales or Australian
Museum collections. It seems likely that as the
original material was obtained by C.S. Wilkinson
(Geological Survey, New South Wales), the
figured specimen would have been Jodged in their
collections, but may have been subsequently lost
in the Garden Palace fire, Sydney, 1882 (J. Pickett,
pers comm. 1992). A neotype is erected here,
which is probably the specimen figured by
Mueller (1883: pl. 3).

Geological Survey of New South Wales
specimens (MMF31337-31343) from the
Newstead Mine, Elsmore, northern NSW have
been tentatively compared with Wilkinsonia
(Pickett et al., 1990). These compressed fruits
however lack the reticulate lacunose omamenta-
tion and longitudinal ridges characteristic of this
genus. The affinities of these fruits remain un-
known.

Wilkinsonia (FP. Muell., 1879) emend.

TYPE SPECIES
Wilkinsonia bilaminata F. Muell,, 1879.

MEMOIRS OF THE QUEENSLAND MUSEUM

Hicksbeachia pilosa

Glencoe @®

Brisbane

Hicksbeachia

pinnatifolia
Gulgonge

Beneree @

FIG. 1. Localities mentioned in text and distribution of
modern genera and species of Australian Hicks-
beachiinae.

EMENDED DIAGNOSIS

Endocarp indehiscent, variable in shape and
size, with vaniable development of longitudinal
ridges, reticulate lacunose surface consisting of
irregular depressions. Depressions variable in
size, Base of depressions confluent with internal
wall of endocarp. Incipient curved longitudinal
ridge continuous from apex to base of endocarp.
Irregular rows of small depressions between in-
cipient ridge and edge of endocarp. Monolocular.

HICKSBEACHIINAE IN THE AUSTRALIAN TERTIARY

197

FIG. 2. Reproduction of Mueller’s (1883) original figure of Wilkinsonia bilaminata, X 0.7. The specimens were

originally illustrated by Mueller at natural size.

s

Wilkinsonia bilaminata (F. Muell., 1879)
emend. (Figs 2, 3A-D).

1879. Wilkinsonia bilaminata F. Muell., p. 170, pl. 3,
fig. 4a,b.

1883. Wilkinsonia bilaminata F. Muell., p. 7, pl. 3
(refigured herein).

MATERIAL EXAMINED
NEOTYPE: NMVP53092 (here designated), Black Lead,
140-170ft below basalt, Gulgong, NSW (Fig. 3A-D).

EMENDED DIAGNOSIS
Large round monolocular endocarp, with reticu-

late lacunose ornamentation consisting of lon-
gitudinal ridges with ramifying supporting lateral
ridges. Approximate size of endocarp (due to fray-
ing of edges), length 46.3mm, width 45.9mm. A
prominent incipient lateral ridge extends from the
apex to the base of the endocarp. A broad lateral
field occurs between incipient ridge and edge of
endocarp with minor accessory lateral ridges and
irregular rows of small depressions. Endocarp
thinly elliptical in lateral view, although this is due
to lateral compression. Multiple irregular rows of
depressions and secondary ridges occur between
the lateral ridge and the edge of the endocarp. The
endocarp wall which is 2-3 mm in thickness, con-

198

sists of tangentially oriented interweaving fibres.
Evidence of radial fibres around lacunae is miss-
ing but as these fibres are not strongly lignified
they are unlikely to be preserved in fossil material.

Wilkinsonia glencoensis (Rozefelds) comb.
nov. (Fig. 4A-C).

1990a. Athertonioides glencoensis Rozefelds, pp. 123-
4, fig. 3,A- C,D(left figure), E,H.

MATERIAL EXAMINED

Holotype, QMF17212 and additional specimens
from the mid-Tertiary Glencoe locality
(QMLS11), near Capella, CQ, figured by
Rozefelds (1990a).

DIAGNOSIS

Endocarp variable in size, broad to suboval or
rounded in outline, with prominent reticulate
lacunose ornamentation, ridging poorly
developed, length 36-43mm; width 25-34mm,
height 14-21mm. A narrow lateral field with mul-
tiple irregular rows of depressions occurring be-
tween incipient ridge and edge of endocarp.
Endocarp wall thick, but ligneous structure not
preserved.

REMARKS

The two Wilkinsonia species differ in the width
of the lateral field between the incipient ridge and
edge of the endocarp. In W. bilaminata this zone
is broad while in W. glencoensis and closely re-
lated modern genera of the Hicksbeachiinae, it is
narrow. This broad field in W. bilaminata is
reflected by the presence of multiple irregular
rows of small depressions with accessory ridges
between the incipient ridge and edge of the en-
docarp. W. bilaminata also appears to possess
more pronounced longitudinal ridges than W.
glencoensis, but these ridges along with the flat-
tened appearance of the endocarp are in part, due
to compression.

ENDOCARP MORPHOLOGY OF EXTANT
GENERA IN THE HICKSBEACHIINAE

The endocarps of Wilkinsonia are strongly lig-
nified which helped facilitate fossilization. The
endocarp wall in Heliciopsis and Athertonia con-

MEMOIRS OF THE QUEENSLAND MUSEUM

sists of tangentially and radially oriented fibres. In
Athertonia the tangentially oriented fibre bands
are numerous and diffuse while in Heliciopsis
artocarpoides, the endocarp wall is thin and there
are areas of lignified parenchyma interspersed
around the tangentially oriented fibres (Fig. 4F, J).
The radial fibres in Athertonia and Heliciopsis
tend to be perpendicular to the outside edge of the
endocarp. In Heliciopsis the radial fibres around
the lacunae are variously oriented, and sometimes
oblique to the endocarp wall (Fig. 4F-G,J-K).

In H. artocarpoides (Elm.) Sleumer the reticu-
late lacunose ornamentation is less developed and
the ridging is less prominent than in Athertonia
(Fig. 4H-I). Most species of Heliciopsis conform
to the endocarp morphology in H. artocarpoides
although material from Sumatra, figured as
Heliciopsis sp. and tentatively compared with H.
incisa by Sleumer (1955a:191,194) resembles
Athertonia in having more thickly walled and
more strongly ornamented endocarps. The en-
docarp wall of Heliciopsis sp. also lacks the inter-
nested lacunae typical of H. artocarpoides
(Sleumer, 1955a).

Hicksbeachia is the only other genus of this
subtribe in Australia with two allopatric species in
eastern Australia (Weston, 1988). The endocarp
of Hicksbeachia pinnatifolia lacks reticulate
lacunose ornamentation. In the specimens of
Hicksbeachia available there is not a clear separa-
tion between the inner ligneous (endocarp) and
outer fleshy mesocarp. The endocarp in Hick-
sbeachia, in transverse section has prominent vas-
cular bundles that are oriented longitudinally,
(absent in Heliciopsis and Athertonia) and poorly
lignified radial fibres extend from the outer edge
of the endocarp into the fleshy mesocarp (Filla,
1926; Fig. 4L, M). Similarly in Athertonia and
Heliciopsis radial fibres extend from the endocarp
into the mesocarp, with persistent radial fibres
evident around the lacunae. The endocarp (inner
mesocarp of Weston, 1988) in Hicksbeachia con-
sists of tangentially oriented fibres and lignified
parenchyma.

The remaining genera in the Hicksbeachiinae;
Virotia from New Caledonia lacks reticulate
lacunose ornamentation of the endocarp while the
fruit morphology of Malagasia from Madagascar
is unknown (Johnson & Briggs, 1975; Virot,
1968).

FIG. 3. A-D, Wilkinsonia bilaminata. F. Muell. (NMVP53092). A,C, Internal view of endocarp wall x 1.2. B,D,
External views of endocarp wall. Note prominent lacunae and the position of the incipient ridge indicated by
arrows and wide lateral field x 1.3. E-F, Athertonia diversifolia (C.T. White) L. Johnson & B. Briggs X 1.5.

E, Internal view and F, external view of endocarp wall.

HICKSBEACHIINAE IN THE AUSTRALIAN TERTIARY 199

200

CONCLUSION

The large endocarp, incipient ridge extending
from the apex to the base and prominent reticulate
ornamentation and arrangement of tangential
fibres in Wilkinsonia strongly suggest affinities
with genera in the subtribe Hicksbeachiinae
(Sleumer, 1955 a,b; Rozefelds, 1990a). These ex-
ternal endocarp features are shared with Ather-
tonia and Helictopsis but are absent From
Hicksbeachia and Virotia.

Wilkinsonia is more closely related to Adter-
tonia than to Heliciopsis. In Heliciopsiy the
depressions on the endocarp wall fend to consist
of internested small lacunae while in Alkertonia
and Wilkinsonia they tend to be separate and not
internested, The endocarp wall in W, bilaminata
is strongly ornamented, consists of separate
lacunae surrounded by tangentially oriented
fibres. The endocarp wall is relatively thin and the
simple perpendicularly oriented lacunae in
Wilkinsonia are similar to Athertenia. The internal
ligneous structure of W, glencoensis is not
preserved although the endocarps are thick and
woody, have perpendicularly oriented lacunae as
in Athertonia and differ from the relatively thin
walled endocarps of Heliciopsis (Rozefelds,
1990a) (Fig, 4).

The fossil distribution of Wilkinsonia
demonstrates that the subtnbe Hicksbeachiinae
was more widespread than at present (Fig. 1).
Wilkinsonia is most closely related to Alhertonia
which is presently restnicted to rainforest com-
munities on the Atherton Tablelands of north east-
ern Queensland. Similarly the allopatric
distribution of Hicksbeachia, and restriction to
widely separated mesic rainforest communities is
suggestive of a more widespread range for this
genus during the Tertiary, which may have en-
compassed much of eastern Austrulia (Fig. 1). The
close relationship of Wilkinsonia to Atherfonia,
and the presence of Elaeocurpus (Rozefelds
19906) in this flora suggests that the Gulgong area

MEMOIRS OF THE QUEENSLAND MUSEUM

during the mid-Miocene was dominated by rain-
forest. The taxonomic diversity and structure of
these mid-Tertiary rainforests will be more fully
understood when the affinities of the remaining
fruits and seeds described by Mueller (1879,
1883) are recognized. This will only be achieved
by the concurrent study and description of fruits
and seeds of the contemporary Australian flora.

ACKNOWLEDGEMENTS

The Jabatan Perhutanan Herbarium, Sandakan,
Sabuh, Malaysia provided material of Hellclopsis
artocarpoides (SAN 28114) for study. Athertenia
and Hicksbeachia fruits were supplied by T, Irvine
and P. Weston respectively. P. Jelland A. Drinnan
gave helpful advice and comments on this paper.
The Museum of Victoria provided funding for my
visit, allowing access to collections and
photographed the onginal plates from Mueller’s
papers. Most of the remaining photographs were
taken by the pholography section at the
Queensland Museum. Information and/ or access
to collections was provided by R. Jones
(Australian Museum), E. Thompson (Museum of
Victoria) and J. Pickett (Geological Survey of
New South Wales).

LITERATURE CITED

DULHUNTY, J.A. 1971. Potassium-Argon dates and
their significance in the Ilford-Mudgee-Gulgong
region, Journal and Proceedings of the Royal
Society of New South Wales 104: 39-44.

FILLA.P, 1926, Das pericarp der, Proteaceac. Flora 120:
99-142,

JOHNSON, L.A.S. & BRIGGS B.G. 1975. On the
Proteaceae - the evolution and classification of a
southern family. Botanical Journal of the Linnean
Society of Londen 70: 83-182,

McMINN, A. 1981, A Miocene microflora from the
Home Rule Kaolin Deposit. Quarterly notes of the
Geological Survey of New South Wales 43: 1-4,

FIG. 4, A-C, Wilkinsonia glencoensis (Rozefelds), A, QMF17212, externa) view of endocarp note prominent
simple lacunae & 1,2. B, lateral view of same specimen showing narrow lateral field between edge of endocarp
and lateral ridge * 1.2.C,QMF1721 1, natural transverse section ofexoocarp * 1.7.D-G, Athertoniadiversifolia.
D. Extemal views of endocarp * 1.1. E, Lateral view of same specimen showing narrow lateral field X 1.1.
F- G, Transverse section of endocarp wall showing strongly lignified tangentially onented fibres (TF) and poorly
lignified radially oriented fibres (RF) F & 2.3, G & 6.7. H- K, Heliciopsis artocarpoides (SAN 28114). H,
External view of endocarp showing interested lacunae X 1.7. J, Jateral view of same specimen showin
internested Jacunae of the endocarp wall and fine radial fibres that extend into mesocarp, the thin exocarp wall
has flaked away from the mesocarp, * 1.5. J-K, transverse section of endocarp wall showing radial fibres (RF),
transverse fibres (TF) and figneous parenchyma (LP), J x 5.8, K * 20.5. L-M, Transverse sections of
Hicksbeachia pinnatifolia F. Muell., endocarps showing vascular bundles (VC), radial fibres extending from
near the mid-region of the endocarp extending into the mesocamp; tangential fibres occur along inner wall of
endocarp, L & 4.3,M % 17.0.

HICKSBEACHIINAE IN THE AUSTRALIAN TERTIARY

01

202

MUELLER, F. Von 1879. Appendix D. Descriptive
notes on the Tertiary flora of New South Wales.
Pp. 169-72. (Annual Report, 1878, Mines Depart-
ment, New South Wales).

1883, ‘Observations on new vegetable fruits of the
auriferous drifts Dec.2.’ (Geological Survey of
Victoria; John Ferres, Government Printer: Mel-
bourne). 23pp.

PICKETT, J.W., SMITH, N., HILL, R.S., MACPHAIL,
M.K. & HOLMES, W.B.K. 1990. A stratigraphic
evaluation of Ettingshausen’s New England Ter-
tiary localities. Australian Journal of Earth Scien-
ces 37; 293-303.

ROZEFELDS, A.C. 1990a. A mid Tertiary rainforest
flora from Capella, central Queensland. Pp. 123-
136. InDouglas, J.G. & Christophel, D.C. (eds),
‘Proceedings, Third International Organization of
Palaeobotany, 1988’. (A-Z Publishers: Mel-
bourne).

MEMOIRS OF THE QUEENSLAND MUSEUM

1990b. ‘A taxonomic study of extant and fossil fruits
of the genus Elaeocarpus (Elaeocarpaceae) in
Australia and New Zealand’. (Unpublished
Honours Thesis, Botany Department, University
of Adelaide).

SLEUMER, H. 195Sa. Proteaceae. Flora Malesiana
(1)5: 147-206.

1955b. Studies in Old World Proteaceae. Blumea 8:
2-95.

SMITH, A.C. & HAAS, J.E. 1975. Studies of pacific
island plants. XXIX. Bleasdalea and related
genera of Proteaceae. American Journal of Botany
62(2):133-147.

VIROT, R. 1968. Proteacées. In Aubréville, A. (ed.),
‘Flora de la Nouvelle Calédonie et depéndances.’
2: 1-254. (Muséum National d’ Histoire Naturelle:
Paris).

WESTON, P.H. 1988. A revision of Hicksbeachia
(Proteaceae). Telopea 3(2): 231-239.

TERTIARY OCCURRENCE OF THE FERN LYGODIUM (SCHIZAEACEAE)
IN AUSTRALIA AND NEW ZEALAND

A.C. ROZEFELDS, D.C. CHRISTOPHEL AND N.F, ALLEY

Rozetelds, A.C., Christophel, D.C. & Alley, N.F. 1992 06 29: Tertary occurrence of the fem
Lygodium (Schizacaceae) in Australia and New Zealand, Memoirs of the Queensland Museum
32(1): 203-222. Brisbane. ISSN 01079-8835.

Lygodium is ubiquitous in early Tertiary floras of Australia, Sterile and fertile leaves of the
climbing fern Lygedium are found i Eocene deposits from Anglesea (Victoria), Maslin Bay
and Golden Grove (South Australia) and in the early Tertiary from Dinmore in Queensland
(L, dinmerphylium Churchill, 1969), Sterile material is known fromm Laté Eocene/Early
Oligocene sediments at Cethana (Tasmania) and the Middle Eocene at Temuka, in New
Zealand.

New matenal from the Golden Grove, Maslin Bay and Anglesea localities including sterile
and fertile pinnules, spores, and cuticular structure is referred to L, dinmorphyllum. The fertile
pinnae contain spores of Cyathidites splendens Harris, 1965. This palynomorph has a Late
Cretaceous (Maastrichtian) to Pliocene range in southern Australia. The Australian species is
very similar to the Eocene Chilean L. skotisbergii Halle, 1940, in sterile and fertile pinnule
morphology. Heterophylly presents problems in interpreting variation in fossil Lygodium.
LLygedium, Schizaeaceae, Cyathidites splendens, Tertiary, palynology, palaeebotany,
biogeography, Australia, New Zealand.

Andrew C, Rozefelds, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101,
Australia; present address: School ef Botany, University of Melhaurne, Parkville 3052,
Victoria, Australia; D, C. Christophel, Botany Department, University of Adelaide, PO Box
496, Adelaide 5001, South Australia, Australia; N, F. Alley, South Australian Department of
Mines and Energy, PO Box 151, Eastwood 5053, South Australia, Australia; 25 June, 1991.

Extant Lygodium is a pantropical climbing fern
with temperate outliers occurring in New Zealand
(i, articularum A. Rich.}, eastern United States (L,
palmatum (Bernh,) Swartz and Japan [LE
japonicum (Thunb.) Sw.]. About 40 species are
known world wide with four occurring in
Australia (Holttum, 1959). The Australian species
L. japonicum (Thunb,) Sw., L. flexuosum (L.) Sw.,
L. microphyllum (Cav) R. Br and L. reticulatum
Schkuhr are tropical occurring in northern and
eastern Australia, In this paper, fossil Lygediun
leaves including sterile and fertile pinnules are
described from Dinmore (Queensland), Maslin
Bay and Golden Grove (South Australia) and
Anglesea (Victoria) and a number of other
localities in southern Australia and New Zealand
(Fig. 1). Previous records of Lygodium or
Lygodium-like ferns are reviewed. Problems in-
herent in classifying and describing variable taxa
are discussed.

PREVIOUS RECORDS OF FOSSIL
LYGODIUM IN AUSTRALIA
AND NEW ZEALAND

Lygodium or Lygoediuni-like fossils have been
peenied by Johnston (1894), Ettingshausen
(1886, 1888), Shirley (1898), Churchill (1969),

Hill et al. (1970), Douglas (1978), Peters & Chris-
tophel (1978) and Christophel & Greenwood
(1987). Chorchill’s material was the first une-
quivocal fossil evidence of the genus in Australia,
The taxa described in the 19th century were based
on fragmentary sterile pinnules but none can be
confidently assigned to Lygodium (Churchill,
1969). Of these taxa the type of Osmunda tas-
manice Johnston from Glenora, could not be lo-
cated in collections in Tasmania (R. S, Hill & N.
Kemp, pers.comm.). The holotype of L. strzeleck-
if Ettingshausen, 1886, collected from the
Vegetable Creek locality was restudied and
prepared further (Fig. 3A) but referral to
Lygodium is dubious because it is based upon
small pinnule fragments with abraded margins.
The Late Triassic Lygedium antiquorum Shirley.
1898 from the Ipswich Coal Measures is based
upon a trilobate seed fern or fern pinnule (Fig. 3F)
with no evidence of sporangia, and is not referable
to Lygodium as Walkom (1917) indicated.
Churchill (1969) described L. dinmorphyllum
based upon two fertile pinnules from the early
Tertiary Redbank Plains Formation at Dinmore in
south eastern Queensland. Hill et al. (1970)
recorded sterile Lygodiuwn pinnules from Din-
more, which they referred to L. skottshergii.
Douglas (1978) figured a fertile Lygedium pinna

204

from Anglesea in Victoria. A fern frond tentative-
ly compared with Lygodium has also been figured
from the Cenomanian Winton flora of Western
Queensland (Peters & Christophel, 1978). Chris-
tophel & Greenwood (1987) recorded Lygodium
from the Golden Grove locality.

Additional material was collected from Din-
more and Anglesea by ACR and DCC respective-
ly over anumber of years. Lygodium has also been
recently collected from the Middle Eocene Maslin
Bay and Golden Grove floras (South Australia)
and the Late Eocene/Early Oligocene Cethana
flora of Tasmania (R. Carpenter, pers. comm.).
Lygodium pinnae have also been recognized from
Temuka in New Zealand (D. Mildenhall, pers.
comm. 1986).

LOCATION OF SITES AND AGES
OF FLORAS

The Dinmore site in the Redbank Plains Forma-

Golden Grove
Maslin Bay

Anglesea

Cethana
Glenora

MEMOIRS OF THE QUEENSLAND MUSEUM

tion, south eastern Queensland is of
Paleocene/Eocene age (Day et al., 1983). This
largely undocumented flora includes ferns,
Podocarpaceae and dicotyledonous leaves (Hill et
al., 1970; Selling, 1950).

The diverse Anglesea flora occurs in the Middle
Eocene Eastern View Coal Measures, and while
Lygodium occurs in most lenses the majority come
from the Gymnostoma lens (Christophel et al.,
1987). Elements of the Maslin Bay flora were
described by Blackburn (1981) from a car-
bonaceous clay lens in the Middle Eocene North
Maslin Sands (McGowran et al., 1970; Alley,
1987). A well preserved flora from the North
Maslin Sands at Golden Grove, north of Adelaide
is also Middle Eocene in age. A recent overview
of the flora has been provided by Christophel &
Greenwood (1987).

The Late Eocene/Early Oligocene Cethana flora
has yielded Nothofagus leaves (Hill, 1984), and
the remaining flora, which includes ‘tropical’

LYGODIUM LOCALITIES

e(Dinmore

egetable Creek

FIG. 1. Fossil Lygodium localities in Australia and New Zealand.

TERTIARY OCCURRENCE OF FERN LYGODIUM 205

FIG. 2. Lygodiumarticularum A. Rich. from Fairy Falls Track, Waitakere Range, South of Auckland, New Zealand.
DSIR, Christchurch CHR153015. Note abscission nodes at base of sterile pinnules and the varying development
of lamina between sorophores. [Photograph supplied by Bryony MacMillan, DSIR, Christchurch, New Zealand].

206 MEMOIRS OF THE QUEENSLAND MUSEUM

TERTIARY OCCURRENCE OF FERN LYGODIUM

rainforest elements ts presently being studied (R.
Carpenter, pers. comm., 1987). The geology and
stratigraphic information on the Temuka potteries
clay pits in New Zealand indicates an Eocene age
for this flora (Wellman, 1953; Gair. 1967).
Specimens from these two localities were not
available for study.

METHODS AND PRESERVATION
OF FOSSILS

The Dinmore, Cethana and Vegetable Creek
materials. are preserved as impressions in fine
prained mudstones lacking cuticle. Latex peels of
the surface of sterile Lygodium pinnules from
Dinmore, examined with a SEM, yielded no cel-
lujar information (D_ Dilcher, pers. comm.)}. How-
ever, SEM examination of fragments of the
original specimen do show some cellular informa-
tion (Fig. 11A.B). The Maslin Bay and Anglesea
specimens are preserved as compressions or im-
pressions while Lygodium leaves and sorophores
from Golden Grove are ‘mummified’, The method
for preparing ‘mummified’ leaves follows that of
Christophel (1984). The Golden Grove Lygadium
leaves are usually opaque and could not be studied
using the fluorescence technique of Friedrich &
Schaarschmidt (1977). Small areas of venation in
these specimens were illuminated using narrow
beams of light from cold light sources.

The mummified sorophores and small frag-
ments of leaf cuticle from Golden Grove were
cleaned with concentrated HF acid fora few days
and the sporangia were gently macerated to
release spores. Spores were studied and photo-
graphed with a Zeiss Photomicroscope 111. Frag-
ments of cuticle from the sorophores and leaves
were washed in distilled water, airdried and sput-
ler coated with platinum. Material was examined
with 4 JEOL ISM 840 scanning electron micro-
scope with lanthanum hexaboride filament at 5kv.

Lygoedium examined inthis paper (Appendix A)
includes material from the Queensland Museum
(QMF), Geological Survey of Queensland
(GSQF), University of Queensland (UQF),
Museum of Victona (NMVP), University of Tas-
nyania (UTC), University of Adelaide (Maslin
Bay, UAS; Golden Groye, UAG), and Geological
Survey of New Zealand (GSNZ), Modern

207

Lygodium material collected by ACR for this
study, will be donated to the Queensland Her-
barium (BRI),

SYSTEMATIC PALAEOBOTANY
Family SCHIZAEACEAE
Lygodium Sw.

The new Lygodiun material from the various
Australian localities is considered to be con-
specific with L. dimmorphyllum Churchill from
Dinmore in south eastern Queensland, An
emended diagnosis of this taxon including infor-
mation on spores, cuticle and foliar heteromor-
phism is given, as Churchill's description only
covered the gross morphology of the fertile pin-
nules.

Lygodium dinmorphyllum Churchill, 1969
(Figs 4-10)

1969 Lygodium dinmorphyllum Churchill, pp. 257-25,
fig,2,

1970 Lygedium skotisbergii Halle, Hill et al., pl. Cell,
fig. L.

EMENDED DIAGNOSIS

Sterile pinnules asymmetrical, usually 2-3
lobed, although unilobate, quadrilebate and pen-
talobate pinnules occur, Lateral lobes tend to he
longer than central lobes, Pinnule base is cuncate.
to truncate, apex rounded to acute, Pinnules petio-
late, lacking abscisston node at pinnule base.
Venation palmate, dichotomously branching trom
the central vein, on each lube, two to three times
before reaching Jeaf margin. Leaf length variable,
maximum lobe lengihabout J 10mm. Pinnule mar-
gin variable, undulose in larger specimens,
smooth in small pinnae, rarely lobed. Pinnules
probably hypostomatic, Anticlinal walls of
abaxial epidermis strongly sinuous, stomata
anomocytic, Stomatal apparatus 30m long, (29-
32,2.m) and 27pm wide (25-32y.m).

Fertile pinnules in branched axes often lack
lamina, althouzh lamina sometimes present as
narrow wing or flange at the base of sozophores,
Individual sorophores variable in size, ranging

FIG. 3. A, Holotype, Lygodium strzeleckii Ettingshausen, from Vegetable Ck, GNSWF8820. x 3. B,C. Cethana
ferns. B, fern aff. Lygedium, sterile pinnule UTC235. C, Bilobate sterile Lygodium pinnule UTC232, x 2, D.B.

Temuka Lygodium, New

Zealand. D, GSNZ25/90, * 1.7. E, GSNZ25/92, & 2. F. Holotype, Lygoulitunp

anigquorum Shirley, GSQF143, * 3.5, [D, Mildenhall (DSIR, New Zealand Geological Survey, Lower Mutt)
and R. Carpenter (University of Tasmania) supplied photographs of the Temuka and Cethana specimens

respectively].

208

from about 4.0-14.0mm in length, often united in
groups of 2-4, Spores referable to Cyathidites
splendens Harns, 1965. Sinuosity of anticlinal
walls of epidermis on sorophore lamin not as
pronounced as in Sterile pinnules.

REMARKS

The Golden Grove and Anglesea localities have
yielded Lygodium sorophores containing C. splen-
dens spores. C, splendens js also 4 common
palynomorph in the Maslin Bay palynoflora.
While spores have not been isolated from the
Dinmore sorophores and the cuticle is poorly
known, it scems preferable to refer the new
material to L, dinmerphyllam than erect a doubt-
fully distinct taxon, Based upon the widespread
cistribution of C. splendens in the carly Ternary
and Late Cretaceous in Australia il seems likely
that the Dinmore sorophoresalso had C, splendens
spores. Lygodium dinmorphyllum is closely com-
parable with L skotesbergii but is considered dis-
tinct based upon spores and cuticular information.
Halle (1940; 259) noted that the spores of L.
skonsbergii ate finely punctate or almost smooth
and hence differ from C. splendens.

The only contraindication that the Lygodium
material from Australia may include a number of
taxa is the variation in the angle of Jower order
branching in the fertile axes at different localities.
In a relatively complete non-distorted specimen
from Anglesea (Fig. 8D), the angle of branching
is (130-150°), which is similar to that of modem
L. articulatunt (Fig. 2). The Dinmore maternal
typically has acutely angled lower order branch-
ing in the fertile pinnae, i.e. (75-85°) (Fig. 44,
G,H), Relatively complete and undistorted fertile
axes have not been collected from Golden Grove
or Maslin Bay so the angle of branching in the
fertile axes from these localities is unknown. The
systematic implications are difficult to assess be-
cause insufficient materia] has been collected
from snough localities lo assess yariation.

MEMOIRS OF THE QUEENSLAND MUSEUM

FOLIAR HETEROMORPHISM IN EXTANT
LYGODIUM

Foliar heteromorphism is the production of mor-
phologically variable leaf types on the same in-
dividual (Eckenwalder, 1980). Changes in the size
and shape of successive leaves can be due to
heteroblastic development (Wareing & Phillips,
1982). This may occur in the transition from
juvenile to adult leaves or may result from en-
vironmental factors. In Lygedium, transition from
sterile to fertile pinnae results in morphologically
distinct pinnae. Pinnae intermediate in character
between sterile and fertile pinnules occur in L.
articulatum and L. palmatum. The lamina in the
fertile pinnae of the extant species L. articulatum
and L. palmatum is very reduced and the pinnules
are strongly dimorphic (Fig. 2; Manchester &
Zavada, 1987, figs 9,11). The degree of lamina
reduction varics in fertile leaflets of L. articulatum
(Fig. 2).

In Lygodium, there is great plasticity of form in
sterile and fertile pinnae, and fusion of pinnules
occurs, Sterile leaflets in extant L articulatum are
characteristically unilobate although fused bilob-
ate pinnules also occur. Fusion of pinnules may
explain some variation in fossi] material from
Australia.

Aberrant pinnules with accessory lobing were
collected from Dinmore, Maslin Bay and Golden
Grove (Fig. 9G,H) and were also figured for L.
kaulfussi Heer, 1861 by Manchester & Zavada
(1987). In modern L. rericulatiwm from north east-
erm Queensland similarly deformed leaves also
occur.

FOSSIL LYGOQDIUM MORPHOLOGY

Taxonomic treatments of extant Lygodium are
based upon the form of sterile and fertile pinnules,
Spares, venation and mode of branching. Holttum
(1959: 39) commented that ‘there is so much
variation in leaflet form due to the age of the plant,

FIG, 4. Fertile Lygedium dinmorphyllum pinnules from Dinmore. A, QMF14465, * 1.9, sporangial head free of
Jamina. B, QMF14530, x 2, lamina around sporangial heads. C,D, GSQF12973, C, X2.8. D, * 1.0. E,F,
QMFI4361, E, ¥3.1. F, X 1.G,H, QMF12544. G, * 3.3. H, x 1,

FIG. 5, Stele Lygodjum dinmorphyllum pinnules trom Dinmore, illustrating variation in pinnules, A, QMF18035.
B, QMF15320. C, QMFI4365. D, QMF2858. E, QMF15319. F, QMF14974. G, QMF14363. H, UQF32006. 1,
GSQF12974. J, OMF14366. All natural size.

FIG, 6, Sterile Lygodiuwn dinmorphyllum pinnules from Dinmore, illustrating variation. A, QMF 18035, x 1, B,
OMF14809, x 2. C, UQF32006, * 1.4. D, QMFI4363, * 3.5.

FIG, 7, Fertile and sterile Lygedium dinmorphyllam pinnules from Maslin Bay. A,B, Sterile trilobate pinoule,
UAS7I5.A, % 2,B, ¥ 1.C, Fertile pimnifeUAS3524, x 5, D,E, Sterile unilobate pinnule with poorly developed
secondary lobe, UAS2274. D, 2.3. EB, &% 1.0, F,G, Bilobate stenle pmnule, UAS1211. F, & 1,9.G, % 10.

TERTIARY OCCURRENCE OF FERN LYGODIUM 209

MEMOIRS OF THE QUEENSLAND MUSEUM

210

Figure 5

TERTIARY OCCURRENCE OF FERN LYGODIUM 211

12 MEMOIRS OF THE QUEENSLAND MUSEUM

ay
Seta

TERTIARY OCCURRENCE OF FERN LYGODIUM

213

214 MEMOIRS OF THE QUEENSLAND MUSEUM

TERTIARY OCCURRENCE OF FERN LYGODIUM 215

Figure 10

+

MEMOIRS OF THE QUEENSLAND MUSEUM

TERTIARY OCCURRENCE OF FERN LYGODIUM

height sbove ground and perhaps polyploidy and
hybridization that even with ample material it is
not easy to give specific limits’. Fossil Lygodium
presents further problems due to the incomplete
und fragmentary material available for study, The
overlap jn morphology and variation in sterile
Lygodium pinnules from sites in Australia and
New Zealand is presented i) Table |, As Dinmare
and Golden Grove have yielded the largest
amount of vegetative and fertile material, discus-
ston centres on maternal frum These two localities,
The strongly dimorphic sterile and fertile pinnules
occur in association at Dinmore, Golden Grove,
Anglesea and Maslin Bay. Malenal from Dinmore
indicates that the stevile and fertile Lyeedisen pin-
nules can be referred to the same plant (Fig. 4E-F),

STERILE PINNULE MORPHOLOGY

Fossil Lygodium pinnules from the various siles
are asymmetrical and palmately lobed, The num-
ber of lobes in slenile Lygodium: was used by Berry
(1930) to define new fossil taxa, although it is an
extremely variable feature. Lygodium pinnules it
most sites, but particularly at Dinmore and Golden

217

Grove, exhibit considerable variation in the num-
ber and degree of lobation (Table 1). Ai Dinmore,
unilobate and trilobate and bilobate and trilobate
pinnules have been found in organic connection
(Figs 5E,H,6C). In the extant New Zealand
species L. articularm, unilobate pinnules
predominate but fused bilobalé pinniiles occuy,
Similarly, in fossil Lygodium from Dinmore,
bilobate and trilobate pinnules are [he musi com-
mon, although rare quadrilobate and pentalobate
pinnules occur (Fig. 5B,1). These are interpreted
as resulting from the fusion of trilobate with
unilobate and bilobate pinnules. Bilobate and
trilobate pinnules also predormnate m the Golden
Grove, and Maslin Bay localities (Table }), The
Australian fossil Lygodimm pinnules resemble ex-
tant L. palmartun: in having multilobale pinnules
but like L, kaulfussit Heer, they differ in their
larger, more elongate pinnules,

AlLthe major localities in Australia have yielded
sterile pinnules with petiolules. In most specimens
the petiulules are short (<3.0mm long}, except a
few specimens from Dinmore which have a
preserved petiolule length of up to 15,0 nim (Fig

FIG 8, Sterile und fertile Lygodiam dimmorphyllam Irons Anglesea. A, Quadrilobate sterile pinnule, NMVP178150),
* 2.3. B, Hilobate sterile pinnule, NMYPIT7S148, & 2. C, Bilobate stenle pinnule, NMVPI73149, * 2,2. D,
Sorophore, arrow indicates prominent obtuse branching of sorophore, NMVP165681, x 4.6, E, Fragmentary
single lobe of large sterile pinnule, NMVP178145, x 2. [Photographs of NMVP1635681 provided by J. Douglas}

FIG. 9. Sterile and fertile Lygodium dintsorphyliam from Golden Grove. 4-H, Sterile pinnules. A, Small pinaule,
UAG3024, * 2.5, B, Bilobate pinnule, UAG3022, * 1.5. C, Bilobate pinrule, VAG2604, * 1.6. D, Unilobate
pinnule, UAG3004, X 1.2. E, Trilobate pinnule, UAG1379, * 1,2. F, Small unilobale pinnule, UAG3003. x
1_G, Aberrant trilobate pinnule with multiple accessory lobing, UAG2718, * L.7. H, Aberrant bilobate pinnule
with multiple accessory lobes, UAG3018, * 1.5. 1-O, Terminal sorophores, all ¥ 4, 1, UAG2037. J, UAG3036,
K, UAG3039, L, UAG3038, M, UAG3042, X L7.N, UAG3O40, * 1.7.0, UAGI041, & L.5.

FIG. 10. Spores macerated from Lygodium dinmerphyllwn sporophores fram Golden Grove and Anglesea, and
late Paleocene Cyathiditey splendens trom the Lake Eyre Basin. A, Ornamentation of distal surface of
dinmorphyllum spore, Golden Grove. Original magnification * 40, Zeiss location 20,8/87.4; England Finder
Location V 17/0, B, Equatorial view of L, dinmorphyllum spore, Golden Grove, showing thickened exine and
reduced ornamentation. Original magnification x 40. Zeiss location 19.6/99,0; England Finder Location U 29/0.
C, Proximal view of L. dinmorphyllum spore, Golden Grove, showing slightly thickened radials and gaping
lgesura, Original magnification * 40. Zeiss Jocation 19.6/122.5; England Finder Location U 53/0, D, Proximal
view of L. dinmerphyllum spore, Golden Grove, showing thickened labra and reduced ornamentation. Original
magnification x 40, Zeiss location 14.8/101.3; England Finder Location P 31/0, E, Oblique equatorial view af
L. dinmorphyllum spore, Anglesca, PB5304, with strongly developed rugulate ormamentation. Original mag-
nification, ¥ 40. Zeiss location 18.8/104.9; England Finder Location T 35/0, F, Distal surface of C. splendens-
Muloorina 2 well, 34 metres depth, showing rugulate and verricate omamentation, Original magnification *
16. Zeiss location 10.5/95.8; England Finder Location L.26/1, G, Proximal surface of sanve specimen with reduced
ornamentation, gaping lacsura and slightly thickened radials.

FIG. 11. SEM of fossil and modern Lygadium cuticle. A,B, Anticlinal walls of epidermis in sterile L. dinnerphyllin
pinnules from Dinmore, QMFI8904, A, Venation and general SERA NOE of Jeaf impression, note venation *
50. B, Sinuous anticlinal walls of epidermis * 140. C, SEM of adaxial surface of L. articelarum cuticle, showing
‘wax covered cuticle and fungal hyphae, * 270, D.E, SEM of abaxial surface of L. articulatum cuticle. D,
showing arrangement of stomata * 260), E, Detail of stomata * 700; P-H, Cuticle from L, dinmorphyllum leaves
und sorophores from Golden Grove. F, Abaxial surface, showing arrangement of stomata, UAG3021, x 320.6,
Cuticle from fertile sorophore of UAGS041, * 430. H, ?Adaxial surface of Ieaf cuticle, UAG3N21, % 350,
[Photographs wl! Figs 11 4,4 supplied by MLE. Detimann and HT, Clifford],

218

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1. Comparison of the morphology of sterile Lygodium pinnules from localities in Australia and New

Zealand.

L. dinmorphyllum
Dinmore, Queensland

asymmetrical/palmate;
bi., tri., quad.,
pentalobate

asymmetrical/palmate;
bilobate, quadrilobate

Anglesea, Victoria

Maslin Bay, S. Aust asymmetrical/palmate;
uni., bi., trilobate
asymmetrical/palmate;

uni.,bi., trilobate

Golden Grove, S. Aust

Lygodium sp.

Cethana, Tasmania

Temuka, New Zealand | asymmetrical/palmate?;

uni., trilobate
Vegetable Creek, N.S.W | unilobate
(L.strzeleckii)

LOCALITY PINNULE SHAPE VENATION |PINNULE PINNULE REFERENCES
APEX BASE

dichotomous

dichotomous

dichotomous

dichotomous

asymmetrical/palmate?; |dichotomous |acute?, obtuse | cuneate
uni., bilobate

dichotomous

dichotomous | obtuse

Churchill, 1969;
Hill et al., 1970

acute, obtuse, | cuneate, truncate

retuse

acute, obtuse, | cuneate Douglas, 1978

retuse

acute, obtuse

acute, obtuse, | cuneate, truncate
retuse

cuneate, truncate

Christophel and
Greenwood, 1987

obtuse, acute truncate

Ettingshausen,
1886

L. skottsbergit

Chile (fossil) asymmetrical/palmate;

bi., trilobate lL

dichotomous pe obtuse

truncate, broady _| Halle, 1940
cuneate

L. articulatum

usually unilobate, rarely
bilobate

New Zealand (extant)

6B). The end of the petiolule in the Golden Grove
material is frequently enlarged indicating a point
of attachment and probably also dehisence (Fig.
9B-E,H). In extant L. articulatum, sterile and fer-
tile pinnules lack a petiolule, and the abscission
node occurs at the leaf base (Fig. 2). This feature
also occurs in the fossil L. kaulfussi and was used
by Manchester and Zavada (1987) to differentiate
between L. kaulfussi and L. skottsbergii.
Lygodium skottsbergii and the fossil Australian
material resemble extant L. palmatum in that their
leaflets are firmly affixed to short petiolules
(Manchester & Zavada, 1987).

The pinnule margins in L. dinmorphyllum are
entire to undulose with some specimens showing
irregular lobation (Fig. 9G,H). One of the Cethana
specimens (Fig. 3B) is irregularly serrate and is
tentatively referred to Lygodium. The serration is
accentuated by tearing of the leaf lamina and
margins during fossilization. The other specimen
is asymmetrical and bilobate (Fig. 3C) and com-
parable to other Lygodium material. The concave-
ly curved basal margin of the pinnule is similar in
both specimens. This feature occurs consistently

dichotomous

acute cuneate

|

in most of the Tertiary Lygodium material in
Australia. The base of the pinnule is typically
cuneate or truncate while in L. kaulfussi it is
rounded to obtuse or cordate (Manchester &
Zavada, 1987).

The venation in the fossil Australian and New
Zealand material is dichotomous, branching 2-3
times before reaching the margin. The venation of
the Australian Lygodium material appears indis-
tinguishable from L. kaulfussi from the Eocene
material from Wyoming figured by Manchester &
Zavada (1987). Dichotomous venation is typical
of most Lygodium species except those with
reticulate venation, e.g. L. reticulatum.

Preservation of the mode of branching and
evidence of the climbing habit which is charac-
teristic of modern species in this genus is rare in the
fossil record and is unlikely to occur in those
species whose pinnules dehisce at senescence. The
only evidence of the mode of branching in fossil
Lygodium comes from impressions at Dinmore.

FERTILE PINNULE MORPHOLOGY
The sorophores of L. dinmorphyllum, like those

TERTIARY OCCURRENCE OF FERN LYGODIUM

of L. kaulfussi and the extant species L, palma
and L. articulate, are produced terminally on
branched axes. The sorophores vary in size,
reflecting their position on the plant and stage of
development. Lygodiunt skottsbergii, L. ar-
ficulatum and L. dinmorphylium all show varia-
tion in reduction of lamina in ferule pinnae, The
fertile pinnae of L. articulatum vary in the free
lamina between sorophores (Fig, 2) and L. skot-
tsbergii shows similar vanation in the amount of
jeaf lamina between sorophores (Halle, 1940: pl.
1, figs 4-8). Lygedium dinmorphyllwn pinnae
from Dinmore exhibit similar ranges of variation
(Fig. 4) with some fertile pinnae ‘free’ of lamina,
and others beanng extensive laminae between the
sorophores (Churchill, 1969: fig, 2), In most of the
Golden Grove, Maslin Bay and Anglesea material
the fertile pinnae are nonlaminar. Fertile pinnae of
L. kaulfussiare also nonlaminar although pinnules
intermediate in character between sterile and fer-
tile pinnules are preserved (Manchester & Zayada,
1987: fig. 8). The sorophores attached to the sterile
pinnule from Dinmore can similarly be explained
ws a ‘leaflet’ intermediate in character between
sterile and fertile pinnules (Fig. 4E-F). The an-
nulus on the sporangia is not preserved in material
from any site. Halle (1940) and Manchester &
Zavada (1987) noted that the annulus could not be
discerned in £ sketrsbergii and L. kaulfiessi
respectively.

SPORE MORPHOLOGY

Spores from complete sorophores of L. dikmor-
phyllum are known from Anglesea (Douglas,
1978) and have been extracted from Golden
Grove material, The spores broadly resemble
those from fossil Lygedium (excluding L. poalen-
sis Chandler, 1955) described from the Nesthem
Hemisphere (Buzek & Konzalova, 1983;
Chandler, 1955; Manchester & Zavada, 1987).
Although spores were recovered from L. dinmur-
phyllum from a number of sites the best preserva-
tion is in the Golden Grove material and
descriptions are based on this material, In general,
the spores possess a subtriangular amb with con-
vex to straight (occasionally concave) interradials
and rounded radials. The laesura are slightly
sinuous, offen open and gaping, and extend two
thirds of spore radius towards the amb. The laesura
are bordered by labra 3-7pm wide and raised
f-Bum. Spore exine is approximately 2j.m thick
und thickened in the radial areas to Sum, Or-
namentation 1s bestdeveloped on the distal surface
and comprises |ow verruca, short rugula and low
sinuous to bifurcating ridges enclosing short ir-

219

regular lumina; omamentation is reduced on the
proximal surface, particularly in the proximal
polar area (Fig. 10). In optical section the amb is
gently undulating. Some specimens appear to bear
remnants of an irregular perisperum. Based upon
measurements from 15 specimens the equatonal
diameter of the spores is 67 (82) 95j.m and polar
diameter is 59 (75) 90pm.

The spores from L. dinmorphyllum sorophores
are identical to the form species Cyathidites splen-
dens Harris, 1965. The holotype of C. splendens
was examined but the mounting medium and
Spore are too dehydrated to provide useful mor-
phological information compansons were
limited to photographs and published descrip-
tions, However free spores of C. splendens oecur
consistently in the Golden Grove and Maslin Bay
carbonaceous lenses as well as other Australian
early Tertiary sediments (Fig. 10). Cyathidites
splendens has also recently being recognized in
Late Cretaceous (Maastrichtian) sediments from
the offshore Bight and Duntroon Basins, off South
and Western Australia (NA, pers. obs). Spores of
the form species C, gigantis (Cookson) Harns,
1965 also bear some similarity to the spores ex-
tracted from the L dinmerphyllum sorophores.
However the holotype of C. gigantis and other free
spores from carly Tertiary sediments were cx-
amined and found to be smaller, possess a much
thickened exine with prominently thickened
radials, and coarse ormamentation in the proximal
polar area, diminishing greatly in the equatorial
interradial areas. The L. dinmorphyllum spores
therefore correlate with the form species C, splen-
dens Harvis, 1965.

CUTICLE MORPHOLOGY

Atlempts to prepare cuticle fromthe mummified
Golden Grove leaves have proved difficult but
small fragments were obtained, Lygodium din-
morphyllum pinnules from Golden Grove are
probably hypostomatic as is modern L. ar-
ticulatum. The stomata of £, dinmorphyllum from
Golden Grove are anomocytic, as in extant
Lygodium (Clifford & Constantine, 1980) and the
anti¢linal epidermal walls are strongly sinuous
(Fig. 11 F,H). The stomata appear to be irregularly
oriented on the leaf surface as in modem L. ar-
ticulatum (Fig. 11D). SEM micrographs of im-
pressions of Dinmore material also demonstrate
sinuous anticlinal epidermal walls but stomata are
not evident (Fig. 1) A,B). Lycodium Kaulfussi hus
sinuous anticlinal epidermal walls and stomata
comparable in size to those of L. dinmarphylluan.
Unlike L, dinmerphyllum the stomata in £. kaul-

220 MEMOIRS OF THE QUEENSLAND MUSEUM

fiessi are diacytio, arranged in rows and the an-
ticlinal epidermal walls are not as strongly sinuous
(Krausel & Weyland, 1950). The stomata in £.
kaulfussit are onented between two epidennal
cells, while in L. dinmorplhylium the stomata are
surrounded by four or five cells.

The cuticle of L. dimmorphyllum sorophores in-
dicates that the anticlinal epidermal walls are
sinuous but not to the same extent as the leaves
(Fig. 11G). No stomata were seen.

CONCLUSION

Even with ample material as provided by the
Golden Grove and Dinmore localities, species
assessmentis difficult, as the material is extremely
variable. The limited material from Cethana or
Temuka is therefore difficult to interpret par-
ticularly with only photographic evidence avail-
able, Similarly early records of Lygodium (such as
Ettingshausen, 1886, 1888}, which are based sole-
ly on sterile foliage are also not presently resolv-
able, Northern Hemisphere spectes based on
limited material present similar problems as
Manchester & Zavada (1987) have shown.

Palaeobiogeographical speculation on the sadia-
tion and evolution of Lygodiwn is difficult be-
cause of the cosmopolitan distribution and
extensive Tertiary fossil record (Kréiuse] &
Weyland, 1950; Reed, 1947). The extensive fossil
record of Lygodium in Australia suggests that the
extant Australian species may be derived from
these early Tertiary forms. However, as fossil
Lygadiain is also known from the Miocene
Notonakajima flora of Japan and Oligocene floras
of Manchuria (Matsuo, 1963; Florin, 1922) it is
equally possible that the modern northern
Australian species L. japonicum, L. flexuosum, L.
microphyllum and L. reticulatwr are recent ar-
rivals from Asia as Page & Clifford (1981) specu-
lated, These species are all widely dismbuted
within the Malesian region (Holttum, 1959). The
weakly dimorphic fertile foliage of most of the
Malesian species, Le. the fertile pinnulestesemble
the sterile pinnules, suggests a closer affinity to L
mioscandens Matsuo{ 1963) from Japan than with
the fossil material from the early Tertiary of
Australia. Manchester & Zavada (1987) suggested
that species with lamina surrounding the
sorophores, e.g. L. japonicum and L. flexuasum
are celatively recent in origin. The high diversity
of Lygodium species in the Malesian area may be
due toa relatively recent and rapid speciation and
Tadiation io this region. This may explain, in part,

Holttum's (1959: 39) difficnilies in defining
modern species within the genus.

Cyathidites splendens ranges from the Late
Cretaceous through to the Phocene in southern
Australia. In the early Tertiary Eyre Formation of
the Lake Eyre Basin C. splendens is common and
can be locally more frequent in parts of the
Paleocene section. The frequency of the spore in
palynomorph assemblages, however, decreases
through the Tertiary and C. splendens is rare in
Pliocene assemblages. Cyathidites splendens is
commonly associated with palynofleral as-
semblages that are indicative of rainforest com-
munities. The gradual decrease in the frequency
of C. splendens through the Tertiary in southern
Australia may be related to the overall decline in
temperature and precipitation during this interval.

ACKNOWLEDGEMENTS

We thank the following institutions and staff for
access fo collections, loan of material or
photographs: Museum of Victoria, Geological
Survey af Victoria, Geological Survey of
Queensland, University of Queensland, Geologi-
cal and Mining Museum, Sydney, University of
Tasmania, New Zealand Geological Survey,
Dominion Museum, Auckland, Department of

Scientific and Industrial Research Organization,
Christchurch, New Zealand, and Swedish
Museum of Natural History. J. Baker
photographed and drafted Figs 1 and 9. The
remaining photographs were taken by staff at the
Queensland Museum and the South Australian
Department of Mines and Energy.

Our special thanks to G. Atherton, J. Webb, T.
Poole and KR. Naumann who donated specimens to
the Queensland Museum. We also thank Alcaa
(Australia) and Monier Ltd for access to the
Anglesea and Golden Grove localities respective-
ty and for grants which enabled early collections
to be made. M.E. Dettmann and HLT. Clifford
provided Figs L1A,B. A. Drinnan and J, Conran
provided advice on this paper, N.F. Alley publish-
es with the permission of the Director-General,
South Australian Department of Mines and Energy.

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WAREING, P.P. & PHILLIPS, I.D.J. 1982. ‘Growth
and differentiation in plants’. (Pergamon Interna-
tional Library: Oxford). 343pp.

WELLMAN, H.H. 1953. The geology of Geraldine
Subdivision. Bulletin New Zealand Geological
Survey 50: 1-72.

APPENDIX A
LYGODIUM MATERIAL EXAMINED
(F=fertile. S=sterile)

Dinmore, Redbank Plains Formation, SE Qld

QMF2857-8 (S); QMF12544 (F); QMF13464 (S);
QMF14096 (S); QMF14360 (S); QMF14361 (S/F);
QMF14362-3 (S); QMF14365-7 (S); QMF14464-7 (F);
QMF14530 (F); QMF15809 (S); QMF14924 (F);
QMF14974 (S); QMF15314 (S); QMF15317 (F);
QMF15318-9 (S); QMF15320-1 (S); QMF18035 (S);
QMF18904 (S). UQF10620 (F); UQF10720 (F);

MEMOIRS OF THE QUEENSLAND MUSEUM

UQF10726 (S); UQF12450-1 (F); UQF32006 (S).
GSQF12973 (F); GSQF12974 (S); NMVP26688 (F)
Holotype, L. dinmorphyllum; NMVP26689 (F)
Paratype, L. dinmorphyllum.

Cethana, Tasmania
UTC232 (S); UTC235 (S).

Vegetable Creek, N.S.W.
GNSWF8820 (S), Holotype, L. strzeleckii.

Maslin Bay, North Maslin Sands, S.A.
UAS715-6 (S); UAS1211 (S); UAS1868 (S);
UAS2274 (S); UAS2513-4 (S); UAS2691-2 (S);
UAS3502 (S); UAS3524 (F); UAS3537-8 (S);
UAS3546-7 (S).

Golden Grove, North Maslin Sands, S.A.
UAG1379 (S); UAG1398 (S); UAG1400 (S);
UAG2604-5 (S); UAG2607-8 (S); UAG2718 (S);
UAG2720 (S); UAG3001-26(S); UAG3036-42 (F).

Anglesea, Eastern View Coal Measures, Victoria
NMVP165681 (F); NMVP178144-9 (S);
NMVP178150 (F).

Temuka, New Zealand
NZGSB25/90-92.

BRYOZOA FROM HERON ISLAND, GREAT BARRIER REEF
J.S, RYLAND AND P.J. HAYWARD

Ryland, 3.8. & Hayward, PJ. 1992 06 29: Bryozoa from Heron Island, Great Barrier Reef.
Memoirs of the Queensland Museum 32(1): 223-301. Brisbane. ISSN 0079-8835.

Eighty-one species of Bryozoa are described from shallow reef habitats around Heron Island,
Queensland. Fourteen new species are described: Puellina decipiens, P. egretta, P. vulgaris,
Celléporaria desperabilis, Robertsonidra novella, Parasmittina galerite, P. onychor-
rhyncha, P. turbula, Pleurocodonellina clavicula, Microporelia fimbriata, Celleporina
reginae, Rhynchozoon ardeolum, R. limatulum and R. scopulorum. Twenty four species are
recorded for the first time from Australian waters, [lHeron Island, Great Barrier Reef,
Bryozoa, taxonomy, new species.

JS. Ryland and P.J, Hayward, Marine, Environmental and Evolutionary Research Group,
School of Biological Sciences, University of Wales, Swansea, Singleton Park, Swanzea SA2

8PP, United Kingdom; 10 June, 1991),

The coral-associated bryozoan fauna of the
Great Barrier Reef (GBR) is undoubtedly very
diverse, probably comprising several hundred
species. It is, however, poorly known. Thus, in

¢ late 19th century, when the rich bryozoan
fauna of Victoria and, to a slightly lesser degree,
of New South Wales and South Australia, was
being described, the GBR bryozoans were known
only by a few specimens from HMS Rattlesnake
{Busk, 1852a), from Port Denison and Holbourne
Island (Haswell, 1881) and from A.C. Haddon's
collections in Torres Strait (Kirkpatrick, 1890a).
This lack of study continued through most of the
20th century, with coastal Queensland or GBR
bryozoans being described in only a few short
papers (Livingstone, 1926; Hastings, 1932; Silén,
1942; Ross, 1974; Ryland, 1974, 1984) and lists
(Livingstone, 1927; Dall & Stephenson, 1953;
Hall, 1984; Winston, 1986), Consequently, the
view prevailed that bryozoans were poorly repre-
sented in coral reefs generally and in the GBR
particularly (Ryland, 1974), That this must have
been erroneous was evident from Maxwell's
(1968) study of sediments in which bryozoan
fragments, representing a wide diversity of
species, formed an important constituent. Recent
semi-popular descriptions of the GBR have been
given by Bennett (1971), Endean (1982) and Tal-
bot (1984), providing usefu) background infor-
mation on the fauna generally but litle on the
Bryozoa specifically.

The GBR ts extensive, stretching some 1900km
from Lady Elliot Reef (24°07'S), slightly south
of the Tropic of Capricorn, to the Warrior Reefs
and Bramble Cay in the Gulf of Papua (9°09°S).
The reefs are dispersed over, and along the edge
of, the continental shelf and occupy an area of

some 215,000km7?. It is to be expected that the
bryozoan fauna will vary with latitude and, most
species being essentially subtidal, would be ap-
propriately studied and collected by scuba. The
present contribution, in which we describe 81
species, is manifestly preliminary being based on
a stil] incomplete examination of specimens col-
lected intertidally on Heron Island reef, one of the
southemmostin the GBR, mainly overa few days
in 1972.

Heron Reef is one of 16 comprising the
Capricom Group which, together with four of the
Bunker Group, lic between latitudes 23° and
24°S, at the southern end of the GBR. These reefs
are separated from Breaksea Spit (Sandy Cape)
to the southeast and from the mainland coast in
the neighbourhood of Gladstone by Curtis Chan-
nel, while the Swain Reefs lie about 150km
northwards across the Capricorn Channel. The
200m isobath lies some 16-25km east of the
Capricom Group, whilst the mainland is about
70km to the southwest, A map of the group, with
bathymetry, is given by Jell & Flood (1977) and
Mather & Bennett (1984).

Mean water temperatures are 26-27°C in sum-
mer dropping to 20-21°C in winter, but tempera-
tures over the reef flat may exceed 32°C or drop
to 17°C, or even lower (Endezan et al.. 1956).
Prevailing winds are southeasterly, or from be-
tween southeast and east during summer. Low-
amplitude ocean swells thus approach
predominantly from roughly the southeast. The
East Australian Current outside the shelf edge
flows polewards but there is little information on
the current patterns within the southern GBR. In
the central GBR the residual flows are Geter-
mined by prevailing winds, setting northerly

224

during periods of steady wadewinds but southerly
during light or variable breezes (Wolanski &
Pickard, 1985), Releases of surface drifters in the
Capricorn Channel suggested a nett southerly
movement (Woodhead, 1970). Tides are semi-
diurnal, though successive tides frequently differ
in height: mean amplitudes are 2m on springs and
Im on neaps. Tidal currents set westerly on the
flood, easterly on the ebb.

Heron Island, the cay (23°27°S, 151°55"E), is
sttuated at the leeward (western) end of an egg-
shaped lagoonal platform reef (Jell & Flood,
1977), roughly 8S by 4km with the Jong axis
orientated ESE-WNW (Fig. 1). Atits caster end,
Heron Reef is separated from the smaller Sykes
Reef, with which it shares a recful platform, by a
3km wide channel of about 15m depth (map in
Jell & Flood, 1977). The southwestern quarter of
Heron Reef ts separated by achanne], about 700m
wide and 30m deep, from Wistari Reef. The
disposition of the two reefs ensures that the
western extremity of Heron Reef, and especially
the northem aspect of it, is well sheltered from
direct impact of the tradewind swells, although
waves from the southeast are refracted around the
reef. Waye-transported particles are deposited in
the vicinity of the cay (Flood, 1974). During the
ebb, water from the reef drains i¢ a radial manner
(except near the boat channel, see below) until the
rim becomes exposed, A shallow lagoon occupies
the east-central area of the reef. At spring tides
there is an extensive peripheral reef flat, especial-
ly at the western end, covered by 0.5m of water
or less. The edge of the reef is then accessible
from the cay by wading- Access to other parts of
the reef edge at low tide, and to the drop off for
scuba, must be by boat,

Most descnptions of Heron Island refer more
to the cay than to the reef (e.g, Steers, 1938) but
brief accounts of the reef have been given by
Endean et al. (1956), Cribb (1966), Jell & Flood
(1977) and Mather & Bennett (1984). Important
man-made features are the boat channel and the
cays western retaining wall. Manne engineering
works up to 1980 were summanzed by Gourlay
& Flood (1981), with particular reference to the
stability of the cay, although there have been
consequences on the reef flat to the south of the
cay, In 1945 a gap was blasted in the recf rim,
opposite the northern tp of Wistan Reef, to pro-
vide access for small boats. A wreck (still there)
was positioned to provide shelter. The gap was
extended In 1966-67 to provide a dredged boat
channe) and harbour, The channel has swb-
sequently been filled with sand mobilized by

MEMOIRS OF THE QUEENSLAND MUSEUM

vyclones, e.g. ‘Emily’ (Apr 1972), ‘David’ Jan
1976) and ‘Simon’ (Feb 1980). cach time being
dredged clear afterwards. The boat channel was
extended in 1987-88 (Flood, 1989) and is now
associated with along and substantial jetty. These
developments have affected sediment deposition
(Gourlay & Flood, 1981), a symptom of a sub-
stantially altered flow regime over the southern
cay-side flat. Most of the Bryozoa descnbed in
this paper were collected from this area in 1972.
Water no longer drains radially from the reef here
but (1988) flows strongly along the flat towards
the channel. The reef flat provided a rich co}lect-
ing arca in 1972 but seemed to have deteriorated
markedly by 1988, even before the consequences
of the 1986-87 developments (already apparent
on water flow) could have affected the hving
coral framework.

The reef flat extends seawards from the cay to
the inner side of the reefrim. Atlow water springs,
away from the boat harbour, water drains through
gaps and channels in the rim, but the flat never
wholly dries. The inner part of the flat is largely
sundy, with patches of coral rubble and some
living coral, €,2. Porites cylindrica. The outer flat
is characterized by living coral, extensive
development of crustose coralline algae, and drifts
of coral shingle; sind is restricted to small pools
and narrow channels. In 1972 the flat at the
western end (beyonil the boat harbour} was exten-
sively carpeted with a level forest of Acropera but
shingle has increasingly accumulated since then,

The reef rim is higher than the upper level of
live coral on the outer flat. Its surface is cemented
with crustose coralline algae, and zoanthids
(especially Palythoa spp.) are conspicuous. Coral
shingle, coral boulders (derived from Acropora
hyacinthus heads), and sometimes large blocks
derived from bommies (plate 30B in Saville-
Kent, 1893), accumulate on and just inside the
rim. While the nm pavement ts devoid of habitats
suitable for bryozoans, the boulders constitute the
richest intertidal habitat for them,

The reef slope is the sublittoral extension of the
reef im, On the windward (southeastern to south-
westem) side of the reef, the upper slope has spur
and groove form and the gradient varies from 1
in 20 in the southeast to | in 4 southwest of the
cay (Jeli & Flood, 1977). When accessible in
calm weather the coral growing on the spars
would undoubtedly prove a rich habitat for
Bryozoa, On the lee side of the reef the spur and
groove formation does not develop; rather there
is extensive growth of tabular corals, such as
Acropora hyacinthus, In 1972 some bryozoans

BRYOZOA FROM HERON ISLAND

225

Lagoon

+ 239°30's

152900'E

Spur and groove

200 0

Profiles

FIG. 1. Heron Island and Reef, showing location of samples. Profiles from Jell & Flood (1977).

were collected by snorkelling and scuba from
under such tables, and photographs show the
richness there of sessile fauna. It is such tables
that, when struck by tropical cyclones, are broken
free and hurled onto the edge of the reef. They
ultimately settle in the only stable position, with
the former attachment facing upwards. The cryp-
tic surface is usually somewhat concave, such
that the boulder rests on its rim; irregularities
ensure that water circulates through the space,
keeping the boulder surface clear of sediment. In
this dark, protected habitat ascidians, bryozoans
and sponges develop in profusion. Thus the
boulders in the Blue Pools area, north of the cay,
which were deposited by cyclone ‘Emily’ (Apr

1972), by 1988 had developed an abundant and
colourful sessile fauna of great diversity.

In 1988 occasional pieces of drift Sargassum
heavily colonized with bryozoans such as Aetea,
Crisia, Lichenopora and Membranipora and
hydroids Eudendrium and Plumularia were col-
lected on the beach of the cay. At the time it was
not known where the Sargassum was growing.
Subsequently, a bed of Sargassum has been lo-
cated by Lin Baldock and Myriam Preker in
10-15m water in the channel between Heron and
Sykes reefs. The species here differ from those of
boulders and discovery of the Sargassum bed
should greatly benefit future studies on the
Bryozoa.

226 MEMOIRS OF THE QUEENSLAND MUSEUM

MATERIALS

With the exception of two species of Bugula
found on boulders near Blue Pools in 1988, the
bryozoans described in this paper are from 14 of
the 28 samples obtained by JSR in 1972. These
are identified by sample numbers (Fig. 1) as
follows:

2. 9 Apr, 1972, Gn the reef flat south of the cay,
Bryozoa were collected on pieces chipped off
tabular Acrapera, in situ, or from dislodged
heads resting on the sand. Bryozoans were
present on under surfaces, though rarely abun-
dant, and less common than sponges,

5. 11 Apr, 1972. On the reef flat WNW of the cay.
Flat plates of coral, the outer parts spread as
horizontal laminae, the inner were convoluted,
forming recesses and clefis sheltering Margaret-
ta, lace corals (Phidolopandae), and cellularines.

6. 11 Apr, 1972. Near 5 on the reef flat WNW of
the cay. Plates of Echinopora with encrosting
bryozoans on the lower surface.

8. 11 Apr, 1972. Near 5 and 6. The flat here was a
level platform of abraded Acropora with a
maze of cavities underneath, and coral
‘fingers’ rising from the surface, The encrust-
ing fauna seemed poorer than under individual
tables,

11. 12 Apr, 1972. On the reef flat west of the cay,
to the north of the boat channel. Part of a
well-developed Acropora head which had
tipped when stepped on, The outer parts were
coated with a thin, filmy prussian blue sponge.
Several Reteporella graeffei and incrusting
species were present.

12. 12 Apr, 1972. On the reef flat, near 11, Below
the actively growing Acropora head, sup-
ported by a slender pedestal, was a layer rep-
resenting an earlier growth level, 3-Scm thick,
with sediment lying on its upper surface. The
lower surface was while, lacking encrusting
coralline algae, but bearing Tubastrea, spon-
ges. Filograna, a brachiopod, and bryozoans.
Below this was coral rubble.

14. 12 Apr, 1972. Near 12. Pieces broken from a
large Acropora head which had become free
and turned upside down, though continuing to
grow,

16, 13 Apr, 1972, Crest to the west of the cleft in
the reef edge, northeast of the cay, The crest
here was strewn with boulders deposited (or
redistributed) by cyclone ‘Emily’ (1-2 Apr).
The sessile fauna on the Acropora was
protected by the knobbly surface of the pedes-

tal. Fragments of typically subtidal erect
Bryozoa, such as Triphyllozoon and violet
fodictyum, were also picked up here, (These
boulders were examined in 1988, having been
on the reef for 16years. The faunais still being
studied).

2). 14 Apr, 1972, Reef crest south of the cay. Most
of the flat approaching the crest at this point
was level, easy to walk on, rather than being
a forest of heads. Several coral boulders
derived from Acropora heads were present.
The living coral of the crest tended to form
layers (almost shelves). While water on the
flat appeared to ebb over the coral, on the crest
it noticeably percolated through the coral. The
space matrix would seem ideal for bryozoans
being fairly dark and with long periods of
water flow, The coral heads typically, though
not invariably, displayed a lower surface
zonation, with the peripheral fingers covered
with lithothammion and the centre region
white, with Filograna tubes, sponges, com-
pound ascidians and Bryozoa. The sample
was a piece of boulder showing a
corul/bryozoan interface,

22.15 Apr, 1972, Reef crest at the cleft, northeast
of the cay. The crest here is more complex than
at 2). There is the actual edge, marked by surf,
then a moat; the coral then rises onto the crest
which merges into the zone of coral boulders
and rubble (see profile D, fig. 1, from Jell &
Flood, 1977). Along the crest coral was dying
back, apparently because cyclone “Emily* had
altered the drainage pattern, leaving the coral
heads dry and dying; or live coral had been
flattened by pounding with the Acropora
boulders produced by the cyclone. Sample
was from these boulders, though most /odic-
iyum fragments were unattached.

23.16 Apr, 1972. Reef crest south of the cay;
collection made by turning boulders. Tide
poor; reefedge nol exposed; water running off
the reef as a sheet.

24.17 Apr, 1972, Near 23; from well-incrusted
boulders.

25.18 Apr, 1972. Collection from 4.5-9m at bom-
mic made by D.R, Robertson. Included
Rugula dentata, Nellia simplex and
Phidoloporidae.

26. 18 Apr, 1972. Near 25, abont 3m depth on the
slope.

Holotypes and paratypes of the new species
described here are in the Queensland Museum,

BRYOZOA FROM HERON ISLAND 227

ACKNOWLEDGEMENTS

We are grateful to the Nuffield Foundation for
the provision of a Travelling Fellowship to JSR
in 1971/72, to the Royal Society for a travel grant
in 1988, and to the Australian Biological Resour-
ces Survey for generous financial support for our
research. Thanks are also due to the Director of
the Heron Island Research Station, Dr Ian Lawn,
and to Miriam Preker, who kindly supplied data
on the distribution of Sargassum, and Dr P.G.
Flood for freely given advice and information on
the physical environment of Heron Island. P.J.
Chimonides and Mary E Spencer-Jones (Natural
History Museum, London) were most helpful in
facilitating the loan of type specimens. Dr MLR.
Fordy (School of Biological Sciences, University
of Wales, Swansea) is especially thanked for his
valued expertise in scanning electron microg-
raphy.

SYSTEMATIC ACCOUNTS

The taxonomic order adopted here follows that
established by Gordon (1984, 1986, 1989b).
Taxonomic diagnoses are provided at family and
genus level, and the species descriptions and
measurements are based on Heron Island
specimens. The synonymies given for each
species are deliberately selective, being limited
to the original description and the most useful,
recent accounts. Unverified synonymies have
been avoided. The occurrence of each species at
Heron Island, and its wider geographical distribu-
tion, are briefly noted. It is intended that a more
precise account of the ecological distribution of
the bryozoan fauna will be prepared after the
completion of the present, contmuing research
project.

Class STENOLAEMATA Borg, 1926
Order CYCLOSTOMIDA Busk, 1852a
Family LICHENOPORIDAE Smitt, [867

Colony encrusting. regularly circular or oval.
occasionally developing as a cylinder around
erect substrata: concave or domed, nodular or
mamillate, often lobed; bordered by a peripheral
lamina of common basal wall. Autozooids-radiat-
ing from centre of colony, sometimes in irregular
quincuncial series, more usually in connate,
uniserial or multiserial rows. Extrazooidal space
between zooid rows divided by calcified struts,
defining polygonal spaces referred to as alycoli.
Brood chambers onginating in centre of colony,

with lobes extending between zooid rows, roofed
by finely granular calcification; ooeciostomes
situated at edges of brood chamber. between
zooid rows.

Lichenopora Defrance, 1823

With the characters of the family.
Type species: Lichénopora turbinata Detrance,
1823.

Lichenopora novaezelandiae (Busk)
(Fig. 2a,b)

Discoporella novaezelandiae Busk, 1875: 32, pi. 0,
fig. 2.

Lichenopont novaezelandiae: Brood. 1976: 299. figs
17A-C (not figs 17G-1,= Disporella sibogae Burg);
Hayward & Cook, 1983: 137.

DESCRIPTION

L. nevaezelandiae, as described and figured by
Busk (1875) and Harmer (1915), has uniseri!
zooid rows and a simple, rounded ooeciostome.
Brood (1976) appears to have confused his
figures with those of Disporella sibogae, a super-
ficially similar species in which the zooid rows
are always biserial. Colonies are typically oval
and domed, and may reach 10mm in length. This
species Js probably quite common in shallow rect
environments, but it is generally inconspicucus,
Several large colonies were present in the inter-
stices of Acropora in sample 2.

DISTRIBUTION

It is distributed throughout the western Pacific,
from New Zealand to Japan, and westwards to the
east coast of Africa.

Class GYMNOLAEMATA Allman, 1856
Order CHEILOSTOMIDA Busk, 1852a
Suborder ANASCINA Levinsen, 1909
Superfamily MEMBRANIPOROIDEA Busk,
1854
Family CALLOPORIDAE Norman, 1903

Colony encrusting, forming coherent or dis-
junct sheets, or reticulate to uniserial runners; oF
erect, developing bilaminar plates or arborescent,
vinculariiform growths, attached by an encrust-
ing base. Frontal wall of autozooids with exten-
sive membrane, sometimes overarched by spines
but always conspicuous; gymnocysial and/or
cryptocystal calvificatton usually present, often
well developed, Spines present or absent-

228 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2, a,b, Lichenopora novaezelandiae; a, whole colony, * 9; b, detail showing ooeciostome between two
zooid rows, X30. c, Antropora granulifera, X80. d, Parellisina curvirastris, ¥ 60. ¢, Cranosina coronata,
»¢50. £, Micrepora variperforata, X75.

BRYOZOA FROM HERON ISLAND

Avicularia adventitious and/or vicarious, some-
times lacking. Ovicells generally present, hyper-
stomial, or partly immersed, reduced. Vertical
walls with large basal pore chambers, or mural
septula,

Antropora Norman, 1903

Colony encrusting. Autozooids with well
developed cryptocyst; gymnocyst negligible or
absent, No spines. Avicularia interzooidal, small,
sometimes Jacking. Ovicell endozooidal, its
presence indicated by a slight thickening at the
distal end of the autozooid. Basal pore chambers
present,

Type species: Membranipora granulifera
Hincks, L880b.

Antropora granulifera (Hincks)
(Fig. 2c)

Membranipora granulifera Hincks, |880b: 72, pl. 9,
fig, 4.

Antropora granulifera: Harmer, 1926: 232, pl. 14, figs
11-14; Cook 1968s: 138, text-fig, 9.

DESCRIPTION

Colony forming flat, unilaminar sheets.
Autozooids irregularly oval to hexagonal, each
with a raised crenellated rim; 0.4-0.5*0.2-
0.3mm. Cryptocyst occupying about half total
autozooid length, flat or slightly concave, coarse-
ly beaded; opesia subtriangular, At the distal end
of each autozooid is a pair (rarely one) of small,
interzooidal avicularia, each budded from a disto-
lateral pore chamber; rostrum acute to frontal
plane, directed distally or disto-medially,
0.07mm long, with a short tnangular mandible.

DISTRIBUTION

Antropora granulifera is possibly circumtropi-
cal in distribution. It has been reported from
Madeira, West Africa, Sri Lanka, Indonesia and
the tropical eastern Pacific, but has not been
recorded previously from the Great Barrier Reef.
It was found at two stations on Heron Island
(samples 2 and 16), on the reef flat to the south of
the cay, and on the reef edge to the northeast.

Parellisina Osbum, 1940

Colony encrusting. Autozooid with narrow
cryptocystal rim, and minimal area of gymnocyst,
Spines reduced or absent. Avicularia vicarious,
large, each associated with a small kenozooid.

hw
hm
<Z

Ovicell prominent, hyperstomial, Basal pore
chambers present.

Type species: Membranipora curvirastris
Hincks, 1862.

Parellisina curvirostris (Hincks)
(Fig. 2d)

Membranipora curvirostris Hincks, 1862: 29: 188th
153, pl. 20, figs 5,6.

Ellisina curvirestris: Harmer, 1926: 228, pl.14, fig. 7.

Parellisina curvirosiris; Cook, 1968a: 156, text-fig.
16; Winston, 1984: 7, fig. 14; Winston & Heimberg,
1986: 6, figs 7,8.

DESCRIPTION

Colony forming small, unilaminar patches-
Autozooids broadly oval or pear-shaped, com-
monly 0.5X0.3mm, the extensive frontal
membrane bordered by a raised mural im; a pair
of delicate, evanescent, distal oral spines present
in early ontogeny. Avicularia frequent; distine-
tive: rostrum acute to frontal plane, 0.25mm long,
laterally curved: associated with a kenozooidal
polymorph, with a rounded opesia, visible distal
to the rostrum. Ovicell relatively small, the ec-
looecium membranous frontally, exposing
granular entooecium,

DISTRIBUTION

This familiarspecies has a pantropical distribu-
tion, fanging into temperate regions in the north»
east Atlantic and perhaps elsewhere. It forms
small, and generally inconspicuous, colonies on
hard substrata, particularly biogenic carbonates.
itis common at Heron Island, occurring in seven
of the sample stations reported here.

Crassimarginatella Canu, 1900

Colony encrusting; or erect, attached by an
encrusting base, bilaminar or vinculariiform.
Autozooids with extensive opesia, bordered by a
narrow, granular cryptocyst: gymnocyst variably
developed. Spines present or absent. Avicularia
vicarious. Ovicell prominent, hyperstomial, or
small and cap-like. Vertical walls with basal pore
chambers or mural septula.

Type species: Membranipora crassimarginata
Hincks, 1880b.

Subgenus Corbulella Gordon, 19%4

Autozooids with well developed gymnocyst,
and numerous marginal spines. Avicularia

vicarious, with proximal opesia bordered by
spines. Ovicell prominent, hyperstomial, with a
broad frontal fenestra,

Type species: Membranipora corbula Hincks,
1880c.

Crassimarginatella (Corbulella) corbula
(Hincks) (Fig. 3)

Membranipora corbula Hincks, 1880c: 378, pl. 17, fig,
6

Crassimargtnatella (Carbulella) corbula: Gordon,
1984: 29, pl. 3, figs DE; 1986: 32, pl. 5, fig. A.

DESCRIPTION

Colony encrusting, forming a small, flat,
unilaminar sheet, Autozooids oval, 0.5%
0,35mm, with large oval opesia, a narrow,
granular cryptocyst and a reduced, smooth gym-
nocyst. Twelve to 18 pairs of spines distributed
around the opesia, the distal one or two pairs
stout, erect, the rest thin, incurved over the frontal
membrane, Ovicell spherical. with a transversely
oval frontal fenestra, its distal edge often
developing a short umbo. Avicularium slightly
larger than autozooids, its distal half comprising
a raised, smooth-edged rostrum, supporting a
semielliptical mandible; proximal half consisting
of a shortened opesia bordered by three or four
pairs of curved spines.

DISTRIBUTION

This species is widely distributed in the westem
Pacific from Japan to New Zealand. A single
colony was present in sample 16, from the reef
edge northeast of the cay.

Cranosina Canu & Bassler, 1933

Colony encrusting, Frontal membrane of
autozooid underlain by a broad, peripheral cryp-
tocyst; gymnocyst absent, No spines, Avicularia
interzooidal, typically one budded from each
autozooid; mandible setiform, Ovicell reduced,
endozooidal, or lacking. Large basal pore-cham-
bers present.

Typespecies: Membraniporacoronata Hincks,
188 la.

Cranosina coronata (Hincks)
(Fig. 2e)

Membranipora coronaia Hincks, 1881a: 147, pl. 10,
fig. 1.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. Crassimarginatella (Corbulella) corbula,

Cranosina coronata: Winston & Heimberg, 1986: 6,
figs 3-6; Hayward, 1988; 281,

DESCRIPTION

Colony forming flat, unilaminar sheets.
Autozooids irregularly oval to hexagonal,
separated by distinct grooves; 0.65-
0.7 Xc.0.4mm. Cryptocyst coarsely beaded, the
beads aligned in ridges, directed medially and
imparting acrenulate edge to the cryptocyst. Dis-
tal end of autozooid raised, with a hood-like
border of smooth, gymnocystal calcification sup-
porting the operculum. Avicularium developed
from the distal pore chamber of each autozooid,
its cystid not clearly distinguishable from the
distal wall of the autozooid; rostrum transversely
orientated, at a slight angle to the frontal plane,
constricted medially, its distal end flared; man-
ens setiform, very finely toothed, about 0.4mm
ong.

DISTRIBUTION

A single small colony of this species was
present in sample 16, It is distributed throughout
the Indo-West-Pacific region, from the Philip-
pines to Sri Lanka and Mauritius. Also reported

BRYOZOA FROM HERON ISLAND

from Puerto Rico (Osburn, 1940) and Venezuela
(Winston, 1986),

Family QUADRICELLARIIDAE Gordon, 1984

Colony erect, branching, jointed. Autozooids
elongate, opesiae extensive; cryptocyst
moderately to well developed; gymnocyst
reduced. Avicularia adventitious, or absent.
Ovicells present or absent. Ancestrula resem-
bling later autozooids, but with a tubular
proximal portion rising from an uncalcified stem,

Nellia Busk, 1852b

Colony erect, branching, jointed: internodes
square-sectioned, consisting of four longitudinal
series of autozooids, arranged in alternating,
back-to-back pairs. Autozooids with well-
developed pymnocyst; opesia with broad,
proximal border of cryptocyst and surrounded by
raiséd mural rim, No spines, Avicularia adyen-
titious. Ovicells small, partly immersed.

Type species: Nei/ia oculata Busk, 1852b-

Nellia tenuis Harmer
(Fig. 10c)

Netlia tenuis Harmer, 1926: 245, pl. 14, figs 16-17.
Nellia tenuis Hayward, 1988; 286, pl. 1, fig. a; Gordon,
1989a: 450, fig. 3.

DESCRIPTION

Colony delicate, inconspicuons, formed from
intemodes 1-4mm long, 0.25mm wide; dividing
dichotomously atinfrequent intervals, most often
developing simply an undivided series of two or
three internodes. Joints between internodes con-
sisting of slender, cuticular tubes: each intemode
finely tapered proximally, Autozooids about
0.5 X*0.25mm, the narrow, oval opesia compris-
ing about three-fifths of total autozooid length. A
pair of ayicularia at the distal end of each
a\itozooid; rostra pointed, hooked, at right angles
1o long axis of internode.

DISTRIBUTION

Nellia tenuis is probably quite common in shal-
low reefal habitats, At Heron Island, il was
present in samples 14 and 24, from the reef flat
west of the cay, and the reef crest jo the south.
Harmer (1926) had fragmentary specimens from
three localities in the Indo-Malaysian region, and
listed a fourth from the South China Sea. Ryland
(1984) illustrated specimens from Fiji. It was

231

abundant on Mauritian reefs. as part of a

bryozoan sward developed beneath coral
boulders (Hayward, 19838).

Superfamily BUGULOIDEA Gray, 1848
Family BUGULIDAE Gray, 1848

Colony erect and branched, unjointed,
unilaminar, attached by rhizoids. Zooids long,
parallel-sided, with almost all of the frontal sur-
face membranous; lateral walls lightly calcified:
marginal spines usually present. Pedunculate
‘bird’s head’ avicularia characteristically
present, Ovicells independent and hyperstomial,
with the ectooecium membranous.

Bugula Oken, 1815

Colony erect. growing from an upright an-
cestrula, branching; attached by rhizoids which
issue from frontal, lateral and basal surfaces of
the autozooids- The latter arranged in two or more
series, alternating; boat-shaped, with the
proximal end forked and the distal end of the next
lowest zooid wedged into the fork, extending
beyond it on the frontal side; with a row of
uniporous, or one to two muluporous, septula.
Secn from the front the zooids are usually trun-
cate distally and slightly attenuate proximally,
Basal and lateral walls lightly calcified, the
membrane occupying most of the front: orifice
closed by a sphincter and no operculum can be
distinguished. One orf more spines may be
present, usually confined to the distal angles of
the zooid. In most species, a pedunculate
avicularium shaped like a bird's head present on
the side of all or many zooids. Ovicell hyper-
stomial. with calcified endooecium and
membranous ectooecium; typically globular, but
sometimes reduced to a hemisphere or less,
closed by the inner vesicle which sometimes
forms the major part of the ovicell.

In biserial species the arrangement of zooids at
a bifurcation usually conforms to one of three
palterns, onginally described by Harmer (1923)
as types 3, 4 and 5. The bifurcations are best
studied from the basal side in stained and cleared
preparations, but can gencrally be observed in a
piece of a clean specimen viewed by transmitted
light. The pattem of consecutive bifurcations is
laterally reversed, so that the bifurcations imme-~
diately above and below any given one are mirror
images of il. For descriptive purposes the zooids
involved in the bifurcation are distinguished by
letters (Fig. 4). The first zooid from which two

232

zooids. are budded is referred to as A, the one
beside it (which also buds two) as B. The outer
zooids above A and B are C and D respectively
while the inner zooids are E and F. In bifurcations
of type 3 the axil is formed by zooids E and F. In
type 4 the axil is formed by F and G (G being
immediately distal to Ein the same series), Zooid
Eis thus excluded from the axil, and the proximal
part of G connects with F. In type 5 the enclosing
process has been further extended so that both E
and F are completely surrounded, and the axil is
formed by G and H (Fig. 4). It should be noted,
however, that the connecting portion between G
and H may be very slender. In a further develop-
ment of the enclosing process, several zooids may
be surrounded; the branch then becomes quad-
riserial below the bifurcations,

Type species: Sertularia neritina Linnaeus,

758.

Bugula dentata (Lamouroux)
(Fig. 4)

Acamarchis dentata Lamouroux, 1816; 135, pl, 3, figs
3a,b.

Bugula dentata: Busk, 1852b: 46, pl. 35, figs 1-5;
Harmer, 1926; 439 (cum syn,), pl. 30, figs, 5,6; pl.
32, figs 21-25; Ryland 1974: 343; 1984: 68, fig. 4;
Winston, 1986; 6,

Bugula sp.: Doak, 1971, colour plate 31 and rear cover;
Coleman, 1977; 80 (colour plate),

1mm

MEMOIRS OF THE QUEENSLAND MUSEUM

DESCRIPTION

Colony tufted, not on a stalk, with some tenden-
cy to spiralling of branches; to 10cm. Branches
narrow, of uniform width, the zooids biserial.
Bifurcations of type 5, i.e., with two zooids whol-
ly enclosed, most obvious in basal view. Zooids
0.6% 0.25mm (basal), in Heron Island material.
Opesia occupying three-fifths or more of the fron-
tal surface; the external margin of the zooid in-
rolling proximally, narrowing the opesia. Inner
distal angle with one spine, outer with two, some-
times long, and jointed at the base; a third outer
spine some way below the other two. The spines
of equal size on both sides of the branch, or the
outer ones (with reference to the proximal bifur-
cation) much larger. Avicularia of two kinds:
ordinary marginal avicularia attached about two-
thirds way down the zooid, a little above or just
below the vortex of the opesia; giant avicularia
present or not. Ordinary avicularia about 0.2mm
in length, a little less than the width of the bearing
zooid at the point of attachment; dorsal profile of
rostrum straight, its tip abruptly hooked (rectan-
gular), Giant avicularia, when present, most often
replacing normal avicularia on the two outer
zooids (C, D) at bifurcations; much longer than
the width of the bearing zooid; its entire upper
profile convex, the rostral tip sometimes with
lateral cusps; the mandible clavate, with a weak
terminal point (Harmer, 1926, pl. 32, figs 21,23,
25). Ovicells globose, attached somewhat obli-
quely over the inner distal angle (Harmer, 1926,

FIG. 4, Bugula dentata. A, Group of autozooids, with ovicells and avicularia. B, Autozooids in lateral view. C,

Avicularium, D, Basal view of a dichotomy.

BRYOZOA FROM HERON ISLAND

pl. 32, fig. 21) or wider than long, occupying the
full width of the distal margin; surface calcifica-
tion rippled; the closing membrane cuticularized.

REMARKS

The living colony is deep bottle green in colour,
though acquiring a bluish sheen in flash
photographs (Doak, 1971, pl. 31, & back cover;
Coleman, 1977:80); on preservation the colour
disappears leaving the colony of leaden ap-
pearance. The variability of this species was men-
tioned by Ryland (1974). In particular, the
obliquely orientated ovicells, emphasized as
diagnostic by Harmer (1926), seem to have been
observed only in material from Indonesia. Giant
avicularia have been recorded only from Sin-
gapore, the Banda Sea and Torres Strait. The
normal, marginal avicularia are attached about
two-thirds way down the outer wall of the zooid
but, since the opesiae vary in length, the
avicularian attachment may be either lateral or
proximal relative to the opesia, The unusual and
distinctively cuticularized ooecial closure, un-
remarked until 1974, characterizes all specimens
from Madeira to New Zealand.

o.smm

DISTRIBUTION

Widely distributed in the tropics and subtropics
from Madeira and Cape Verde Is., to southern
Africa, Indian Ocean (Amirante Island), In-
donesia, Japan, Lifu, and northern New Zealand.
In Australia, Coleman (1977) gives the depth
range as 2-40 m; the habitat as rocky reefs, coral
reefs and wharf piles; and its distribution to all
States except N.T.

Bugula vectifera Harmer
(Fig. 5)

Bugula vectifera Harmer, 1926: 442, pl. 31, figs 1-6,

DESCRIPTION

Colony a loose spreading tuft, delicate, pale
translucent straw-brown; to about 15mm in
height; anchored proximally by rhizoids; the
branches also attached to the substratum by mter-
mittent rhizoids. Branches with zooids in two
series, with type 4 bifurcations (Fig. 5D). Zooids
weakly calcified; elongate, narrowed proximally,
0.55 (SD 0,03) 0.13 (SD 0.01)mm basally;
opesia occupying about three-quarters of the
frontal surface. No marginal spines. Avicularia
attached on pedicels, proximal to the opesia,

FIG. 5. Bugula vectifera, A, Group of autozooids, with ovicells and avicularia. B, Antozooids in lateral view,
showing attachment of ovicells and avicularia, C, Part of a branch in basal view. D, A dichotomy in basal view.

E, Avicularium. F, Palatal view of avicularium.

234

close to the distal margin of the preceding zooid;
their length approximately equal to the terminal
width of the preceding zooid; rostrum as long as
the head, its tip sharply downcurved (Fig. 5E);
width about the same as in B. subglobosa but, the
rostrum being longer, lacking the swollen ap-
pearance of that species (Fig. 5F, cf. Fig. 6D).
Ovicells prominent, a little longer than broad,
narrowly attached to the inner distal angle of the
bearing zooid (Fig. 5A,B), transversely orien-
tated; the opening directed downwards and across
the end of the bearing zooid, invisible from most
viewpoints, close to the avicularium of the suc-
ceeding zooid. Embryos pale mahogany or pale
purple-brown. Tentacle number 17-18, ‘about
12’ in Harmer’s material.

DISCUSSION

Some differences are apparent between the
Heron Island material and that described by
Harmer. These probably relate mainly to the nar-
rowness of the zooids in the Indonesian
specimens (0.53 (SD 0.05) x 0.085 (SD 0.01)mm
in British Museum specimen 1928.3.6.287, Bor-
neo Bank), with ‘their frontal surfaces sloping
obliquely away from the median raised part of the
branch’. In the Heron Island specimens this is
slightly evident in the proximal part of the colony,
but not elsewhere where the distal angles of the
zooids tend to be raised well above the surface of
the next distal zooid. Zooids tend to be uniserial
proximally, though not to the extent described by
Harmer in one of the Borneo Bank specimens;
neither is there any longitudinal thickening here.
Indeed, the habit of the colonies is semi-repent
(Scrupocellaria-like) rather than arborescent,
with a supporting stem. In other respects the
agreement between the two groups of specimens
is very close.

OCCURRENCE

Found, usually with B. subglobosa, among the
under fauna of several boulders in the tract at Blue
Pools, 27-31 Jul 1988. First record from
Australia.

DISTRIBUTION
Great Barrier Reef, Java, Borneo Bank (Strait

of Makassar), eastern Honshu, Japan (Harmer,
1926).

MEMOIRS OF THE QUEENSLAND MUSEUM

Bugula subglobosa Harmer
(Fig. 6)

Bugula subglobosa Harmer, 1926: 444, pl. 32, figs
15-18.

DESCRIPTION

Colony a loose spreading tuft, attached
proximally by rhizoids, delicate, pale straw
coloured, up to about 15mm; the branches also
attached to the substratum by intermittent
rhizoids. Branches with zooids in two series, with
type 4 bifurcations (Fig. 6B). Zooids elongate,
narrowing proximally, 0.59 (SD 0.04) x 0.13 (SD
0.01)mm basally; weakly calcified. Opesia oc-
cupying about three-quarters of the frontal sur-
face. No marginal spines. Avicularia attached on
pedicels, proximal to the opesia, almost under the
distal edge of the preceding zooid; short (much
shorter than the terminal width of the preceding
zooid), very convex dorsally; the rostrum com-
prising less than half the total length, sharply
downcurved (Fig. 6C); the head broad in relation
to overall length, appearing swollen (hence sub-
globosa, Fig. 6D). Ovicells prominent, longer
than broad, narrowly attached to the inner distal
angle of the bearing zooid, transversely orien-
tated (Fig. 6A); the opening directed downwards
and across the end of the bearing zooid, concealed
from most viewpoints, opposite the avicularium
of the succeeding zooid. Embryos pale milky
yellow. Tentacle number about 18.

The Siboga specimen (British Museum
1928.3.6.91) from North Celebes has larger
zooids, 0.7 (SD 0.03) X 0.15 (SD 0.04)mm, but is
of the same delicate colony form.

OCCURRENCE

Found, usually with B. vectifera, among the
under fauna of several boulders in the tract at Blue
Pools, 27-31 Jul 1988. First record from
Australia.

DISTRIBUTION

Great Barrier Reef, Flores, Sumbawa, Timor,
North Celebes and eastern Honshu, Japan
(Harmer, 1926).

Family BEANIIDAE Canu & Bassler, 1927

Colony encrusting, loosely adherent,
unilaminar; or erect, bilaminar, foliaceous; at-
tached to substratum by rhizoids. Autozooids
contiguous to widely disjunct, linked by long or
short tubular extrusions of the mural septula.

BRYOZOA FROM HERON ISLAND

FIG. 6, Bugula subglobosa. A, Part of a branch, with autozooids bearing ovicells and avicularia. B, A dichotomy
in basal view, C, Avicularium, D, Avicularium in distal view.

Frontal surface entirely membranous, with or
without a border of marginal spines. Avicularia
adventitious, pedunculate, of bird’s head form.
Ovicells present or absent.

Beania Johnston, 1840

Colony encrusting, unilaminar, ramifying or
reticulate, attached to the substratum by rhizoids
and generally loosely adherent. Autozooids
closely packed, or widely spaced apart, but not
contiguous, linked to each other by tubular exten-
sions of mural pore chambers; recumbent, or
erect, sometimes with an elongate, narrowly
tubular, proximal portion. Frontal membrane oc-
cupying whole of broad distal portion of
autozooid, bordered by thin spines in many
species; distal spines or spinous processes present
or absent. A vicularia present or absent; peduncu-

late, with a hooked rostrum. Ovicells reduced,
vestigial or absent.
Type species: Beania mirabilis Johnston, 1840.

Beania magellanica (Busk)

Diachoris magellanica Busk, 1852b: 54, pl.67, figs
1-3,

Beania magellanica: Harmer, 1926: 412, pl. 28, figs
1-4, text- fig.1; Gordon, 1984: 46, pl. 11, fig. H.

DESCRIPTION

This species is distinguished from all other
species of Beania by the six, regularly-arranged
connecting tubes linking each autozooid with its
neighbours, by a complete absence of spines,
except for two to four tiny processes at the distal
end of the autozooid, and by its greatly enlarged,
paired avicularia. It is probably common in reef

236

habitats, but its filmy, transparent colonies are
quite inconspicuous in the field, and only become
readily apparent once specimens begin to dry.

DISTRIBUTION

Beania magellanica seems to be practically
ubiquitous in tropical, subtropical and temperate
seas. It has a circumpolar distribution in the
Southern Ocean, though not extending into An-
tarctic waters, and in the western Pacific ranges
northwards to Japan, and in the Atlantic region as
far as the Mediterranean. It is certainly present in
the eastern Pacific (Osburn, 1950) and probably
occurs throughout the Indian Ocean.

Family CABEREIDAE Busk, 1852a

Colony erect, branching, jointed, attached to
substratum by rhizoids; branches dividing
dichotomously, unilaminar, with frontal and
basal surfaces, autozooids in two, sometimes
more, alternating longitudinal series. Autozooids
well calcified. with moderately sized opesia;
cryptocyst generally narrow, gymnocyst well
developed, Spines present, or rarely, absent, one
often modified asa flat plate, the scutum, overar-
ching the frontal membrane. Avicularia adven-
titious, sessile, sometimes absent, vibracula
generally present on basal surface, Ovicell hyper-
stomial, prominent.

Scrupocellaria van Beneden, 1845

Colony branching, erect and bushy, or repent
and spreading; attached to the substratum by
rhizoids. Branches biserial, with defined frontal
and abfrontal faces, dividing dichotomously at
regular intervals, each jointed at its base.
Autozooids with well developed gymnocyst and
oval frontal membrane; spines usually present at
distal end of autozooid; a specially modified
lateral spine, or scutum, present in many species,
flattened and arching over the frontal membrane,
sometimes hiding it completely. Heterozooids
polymorphic: lateral avicularia typically present
at puter distal angles of autozooids, often dimor-
phic; frontal avicularia often present on gym-
nocyst, also often dimorphic; vibracula typically
present on abfrontal surfaces. Ovicell hyper-
stomial, globular, with or without pores, or larger
fenestrae.

Type species: Sertularia scruposa Linnaeus,
1758.

Species of Scrupocellaria may be quite com-
mon in shallow reefal habitats, forming swards

MEMOIRS. OF THE QUEENSLAND MUSEUM

on the undersides of boulders, in association with
other erect Cheilostomata, such as Margaretta,
Nellia and Vasignyella. Three common tropi-
cal/subtropical species occur on the reef flats of
Heron Island.

Scrupocellaria diadema Busk
(Fig. 7)

Scrupocellaria diadema Busk, 1852a: 370; 1852b: 24,
pl. 28, figs 1-3,

Scrupocellaria diadema: Harmer, 1926; 375, pl. 25,
figs 20-25; Ryland, 1984: 68, fig. 3; Hayward, 1985:
285, fig. 3C.

DESCRIPTION

This species forms stout, bushy, erect colonies.
Autozooids 0.35—0).5mm long, with the opesia
comprising about three-quarters total length; the

FIG, 7. Serupocellaria diadema.

BRYOZOA FROM HERON ISLAND

opesiae of the two series of autozooids overlap-
ping along the branch. Each autozooid with two
pairs of short, disto-lateral spines, and a single
median distal spine; scutum narrowly oval,
covering just the central area of the frontal
membrane, but frequently absent. Disto-lateral
avicularia small, monomorphic, a single, abfron-
tal vibraculum just visible distal to each; frontal
aVicularia shortly columnar, with the rostrum al-
most perpendicular to frontal plane, dimorphic.
those borne by autozooids proximal to
dichotomies greatly enlarged; a single
vibraculum occurs in the axil of each dichotomy.
Ovicell characteristic: wider than long, with con-
Spicuous tubiform pores.

DISTRIBUTION

S, diadema was described from Moreton Bay,
Queensland. It ranges northwards to Japan,
throughout the Indo-Malaysian region, and
westwards to India and Mauritius.

Scrupocellaria maderensis Busk
(Fig. 8)

Serupocellaria maderensis Busk, 1860: 280; 1861: 77,
pl. 32, fig. 1,

Serupocellaria maderensis: Harmer, 1926: 372, pl. 25,
figs 16-19; Gordon, 1984: 47, pl. 12, figs F,G;
Ryland, 1984, fig. 2; Hayward, 1988: 283,

DESCRIPTION

Colony an erect, freely branching tuft, with the
internodes between dichotomies often exceeding
2mm. Autozooids commonly about 0.35mm
long; opesia small and round, comprising less
than half total autozooid length, and those of the
two autozooid series do not overlap along the
branch. Up to four short spines on the outer distal
corner of the autozooid, one or two on the inner;
the scutum, invariably present, is rounded, with a
straight distal edge, and almost completely hides
the frontal membrane proximal to the operculum.
Disto-lateral ayicularia monomorphic, very
large, with hooked rostrum, giving a serrated
outline to the branch edge; frontal avicularia
usually absent; vibraculum present on the abfron-
tal surface of each autozooid, and two in the axil
of the dichotomy. Ovicell longer than wide, tilted
towards axis of branch; flattened frontally, with
# Single fenestra.

DISTRIBUTION
Serupocellaria maderensis is widely dis-
tributed in the warm temperate and tropical

237

waters of the eastern Atlanuc and Indo-West-
Pacific regions. In the western Pacific it ranges
from southern New Zealand to Japan.

Scrupocellaria spatulata (d’Orbigny)
(Fig. 9)

Cellularia spatulata d’ Orbigny 1851; 50,

Scrupocellaria spatulata: Harmer, 1926: 382, pl. 26,
figs 1-10; Winston & Heimberg, 1986: 7, figs 11-12;
Hayward, 1988: 283.

DESCRIPTION

Colony forming delicate, spreading tufts up to
10mm high. Autozooids slender, 0.4-0.5mm
long, the opesia comprising slightly less than half
total frontal length; two series of opesiae just
overlapping along the branch. Two or three
spines on the outer distal comer of the autozooid,
and usually two on the inner; non-ovicellate
autozooids may have, in addition, one or two
median distal spines; scutum narrowly oval,
covering only a small central portion of the fron-
tal membrane. Disto-lateral avicularia dimor-
phic: most usually tiny, with a laterally
orientated, triangular mandible, occasionally

FIG. 8. Serupecellaria maderensis

FIG. 9. Scrupocellaria spatulata,

replaced by a grossly enlarged type with slender,
digitate rostrum orientated parallel to the long
axis of the branch: frontal avicularia sparse,
monomorphic, columnar, situated close to the
base of the scutum of the adjacent autozooid; a
single vibraculum on the abfrontal surface of
each autozooid, and one in the axi] of each
dichotomy. Ovicell as wide as long, globular,
with small, regular perforations.

DISTRIBUTION

This species is distributed throughout the Indo-
West-Pacific region, from Queensland to East
Africa, and from Victoria northwards to the
Philippines.

Superfamily MICROPOROIDEA Gray, 1848
Family MICROPORIDAE Gray, 1848

Colony encrusting or erect. Autozooids with
membranous frontal wall underlain by an exten-
sive cryplocyst; opesia reduced, often scarcely
exceeding area of operculum; gymnocyst usually
little developed. Spines present or absent.
Avicularia present, adventitious, interzooidal or
vicarious. Ovicells present or absent. Vertical
walls with basal pore chambers or mural septula.

MEMOIRS OF THE QUEENSLAND MUSEUM

Micropora Gray, 1848

Colony encrusting. Autozooids typically flat,
with lateral walls raised and thickened, and often
developed as prominent umbones lateral to the
opesia; frontal membrane entirely underlain by a
thickly calcified, perforate cryptocyst, with
paired opesiules situated close to its distal
corners; opesia small, exactly coincident with
orifice, with a thickened rim; no gymnocyst.
Spines present or absent. Avicularia vicarious,
smaller than autozooids; lacking in some species,
Ovicells partly immersed, prominent, with small,
scattered pores; closed by autozooid operculum.
Vertical walls with large mural septula, which
may extend to the basal wall.

Type species: Flustra coriacea Johnston, 1847.

Micropora angusta MacGillivray
(Fig. 10a)

Micropora voriacea var. angusta MacGillivray, 1887:
67.
Micropora sagitiata a’ Hondt, 1986; 712, pl. 6, figs 1,2.

DESCRIPTION

Colony loosely encrusting, forming flat. multi-
Jarninar sheets. Autozooids narrowly rectangular,
0,6-0.75 * 0.25-0.3mm; cryptocyst flat. finely
granular, densely perforated, bounded by a thick,
crenulate mural rim. Opesia raised above plane
of eryptocyst, its lower lip almost perpendicular,
thick, with a crenulate rim; more or less semicir-
cular, with rounded corners. No oral spines.
Opesiules indistinct. Avicularia relatively large;
rostrum 0.16mm long, with broadly triangular
mandible, proximo-laterally directed. Ovicell
elongate oval, surface coarsely granular, with
numerous tiny perforations; opesia of ovicellate
autezooids longer than in non-ovicellate
autozooids.

REMARKS

MacGillivray (1887) described the morphol-
ogy of this species quite precisely although,
regarding it simply as an interesting variant of M.
cortacea Johnston, he unfortunately did not fig-
ure it or give. locality for his material. D'Hondt’ s
maternal, which is unquestionably the same
species as that from Heron Island, was from New
Caledonia. P.L. Cook (in litt.) has re-examined
MacGillivray’s specimens of M. coriacea var.
angusta, in the Victoria Museum, Melbourne,
and reports that it is the same species as recorded
here from Heron Island; specimen F45537A, Port

BRYOZOA FROM HERON ISLAND 239

FIG, 10. a, Micropora angusta, * 110. b, Steginoporella magnilabris, *~50. c, Nellia tenuis, x70. de
Thalamoporella stapifera; d, autozooids and an avicularium, x70; e, group of calipers, * 800.

240

Phillip Heads, is here designated as lectotype of
Micropora angusta MacGillivray. The loosely
encrusting, multilaminar colony form, and the
finely punctate ovicell are unusual features for a
species of Micropora.

Micropora variperforata Waters
(Fig. 2f)

Micropora variperforata Waters, 1887:51, p18, fig. 27.
Microporavariperforata: Gordon, 1986: 69, pl. 27, fig.
B.

DESCRIPTION

Colony forming an irregular, spreading
unilaminar sheet. Autozooids oval to hexagonal,
0.45-0.6 X 0.35-0.4mm; cryptocyst flat, nodular,
with about 20-35 small, randomly distributed
pores; opesiules recessed but distinct, proximo-
lateral to opesia; an irregular series of compara-
tively large pores developed around the margins
of the cryptocyst; mural rim crenulate, thickened,
in later ontogeny continuous with a conspicuous
proximal ridge of cryptocystal calcification.
Opesia wider than long, 0.12-0.14 x 0.06-
0.08mm; no oral spines. Avicularium scarcely
larger than opesia; rostrum triangular, acute to
frontal plane and laterally directed. Ovicell about
as wide as long, the surface finely nodular, except
for a smooth or faintly striated frontal band im-
mediately above the aperture.

DISTRIBUTION

Several colonies occurred in sample 22, from
the reef crest, northeast of the cay. The Heron
Island specimens are strikingly similar to the
Recent New Zealand material figured by Gordon
(1986), so that M. variperforata is known from
the western South Island of New Zealand as well
as from New Zealand Tertiary deposits.

Family STEGINOPORELLIDAE Hincks, 1884

Colony encrusting; or erect, frondose or vin-
culariiform, attached by an encrusting base.
Autozooids with well developed cryptocyst,
which descends to basal wall in distal half of
zooid, delimiting a tube for the exsertion of the
polypide; gymnocyst reduced or lacking.
Autozooids often dimorphic: unmodified A
zooids represent majority; B zooids, perhaps
primitive avicularia, are characterized by en-
larged, toothed opercula. No ovicells. Vertical
walls with multiporous mural septula.

MEMOIRS OF THE QUEENSLAND MUSEUM

Steginoporella Smitt 1873

Colony encrusting, or erect, frondose or narrow
and branched. Autozooids typically dimorphic,
both types containing polypides, distinguished
principally by relative size of operculum. Frontal
wall membranous, lacking a gymnocyst; under-
lain by a concave cryptocyst, which descends to
the distal wall or to the basal wall in the distal half
of the zooid, thus dividing its cavity into two
chambers; withdrawn polypide located in
proximal chamber, exserting via a tubular
foramen (‘polypide tube’) in the distal part of the
cryptocyst. Vertical walls of autozooids with
large multiporous septula. No ovicells; embryos
brooded within maternal autozooids. No spines.
B zooids, characterized by enlarged and thickly
sclerotized opercula, perhaps represent primitive
vicarious avicularia.

Type species: Membranipora magnilabris
Busk, 1854.

Steginoporella magnilabris (Busk)
(Fig. 10b)

Membranipora magnilabris Busk, 1854: 62, pl. 65, fig.
4.

Steganoporella magnilabris; Harmer, 1926; 277, pl.
17, figs 1-3, 7, 9, 12, text-fig. 10; Cook, 1964a: 53,
pl. 1, fig. 4, text-fig. 2.

Steginoporella magnilabris: Winston, 1984: 10, figs
17,18.

DESCRIPTION

Colony forming extensive, flat, encrusting
sheets, or occasionally erect, bilaminar or uni-
laminar plates; deep reddish-brown or horn-
coloured. Autozooids rounded distally, straight
proximally, commonly 0.9-1.0 X0.6-0.7mm.
Frontal membrane thick, opaque, bordered by a
raised, crenulate mural rim; A zooids with distinc-
tive semicircular operculum, as wide as zooid and
about half as long, with a dark brown arched
sclerite, and a submarginal series of small teeth
on its inner face; B zooids, having proportionately
much larger operculum, armed with stout, con-
spicuous, marginal teeth on its inner face, varying
in frequency, from 0.2-30% in different
geographical populations (Cook, 1964a). Cryp-
tocyst occupying about half total length of
autozooid, depressed, finely nodular, with
numerous fine perforations; descending abruptly
to basal wall of autozooid, with rounded polypide
tube situated centrally in vertical distal part of
cryptocyst and not visible in frontal view. Upper

BRYOZOA FROM HERON ISLAND

distal edge of cryptocyst produced as a concave,
or deeply cupped, process immediately above
polypide tube. Distal half of autozooid with a
raised rim supporting the operculum, below
which is a broad oral shelf.

DISTRIBUTION

5S. magnilabris occurs in shallow tropical
waters, from the Caribbean, West Africa, East
Africa and the Indian Ocean, Malaysia, Indonesia
and the Philippines, and northwards to southem
Japan.

Family THALAMOPORELLIDAE Levinsen,
1909

Colony encrusting or erect. Autozooids with
frontal membrane entirely underlain by a
granular, concave cryptocyst; opesia usually
coincident with operculum, single or paired
opesiules present, large. No gymnocyst,
Avicularia vicarious. Ovicell large, bivalved; or
lacking. Vertical walls with multiporous mural
septula.

Thalamoporella Hincks, 1887

Colony encrusting or erect. Autozooids with
membranous frontal wall underlain by extensive
cryptocyst; no gymnocystal calcification. Cryp-
tocyst concave or depressed, with a pair of large
opesiules bounded by basally deflected cryp-
tocyst, which may extend to the basal wall; opesia
small, usually almost coincident with operculum,
Nospines, Avicularia vicarious, Ovicell bulbous;
bivalved, with a longitudinal median suture,
borne by a modified maternal zooid. Vertical
walls with multiporous sepiula. Calecareous
spicules, of unknown significance, present in
cavity of autozooid.

Type species: Flustra rezieri Audouin, 1826.

Thalamoporella granulata Levinsen
(Fig. 1 1a,b)

Thalamoporella granulata Levinsen, 1909; 789 (part),
pL. 6a, figs la-f.
Thalamoporella granulata: Harmer, 1926; 297.

DESCRIPTION

Colony forming a flat, encrusting sheet.
Autozooids rectangular to polygonal, bounded by
low, crenulate, mural rims; 0,6-0,7 < 0,3-0,4rnin,
Cryptocyst with finely beaded surface, flat or
slightly convex proximally, concave distally;

741

opesiules irregularly oval, of unequal size,
bounded proximally and Jaterally by descending
cryptocyst which reaches to the basal wall of the
autozooid; opesia with semicircular distal rim, its
proximal edge shallowly concave. Avicularium
as long as autozooid; rostrum constituting half
total length, narrowly spatulate, curved laterally
to left or right, the autozooid embraced by the
curve consequently with an asymmetrically
placed opesia. Brooding zooid shorter than
autozooid, with proportionately broader opesia;
capped by a thin-walled, bivalved ovicell, with
triangular aperture closed by a modified oper-
culum. Spicules of two types: widely opened
compasses, of two sizes, and small, closed
calipers,

DISTRIBUTION

The type locality for this species is Tomes
Strait; Harmer (1926) redescribed the material on
which Levinsen (1909) founded T. granulara, but
the species does not seem to have been reporicd
subsequently from elsewhere. ltappears common
at Heron Island; large colonies were present in
three of the samples reported on here.

Thalamoporella stapifera Leyinsen
(Fig. 10d,e)

Thalamoporella granulate var, stapifera Levinsen,
1909; 188, pl. 6, figs Sa-e.

Thalamoporella stapifera: Harmer, 1926; 297, pl, 19,
figs 17,20,25; Soule & Soule, 1970; 10, figs 4C-G;
Winston & Heimberg, 1986: §, figs 13-16,

DESCRIPTION

Colony forming a broad, flat, encrusting shect.
Autozooids elongate, reclangular to uregularly
hexagonal, bounded by thick, crenulate mural
rims; 0.6-0,8% 0,25-0.5mm, Cryptocyst more or
Jess flat, slightly concave proximal to opesia;
opesiules irregularly oval, of equal or unequal
size, cryptocyst descending to basal wall, to form
complete tube; opesia with semicircular distal
rim, its proximal edge deeply concave.
Avicularium shorter than autozooid; rostrum
constituting less than half total length, symmetri-
cal, distally rounded. Brooding zooid shorter than
autozooid, with proportionately broader opesia,
and especially pronounced opesiular tubes;
ovicell thin-walled, bulbous, with triangular
aperture closed by a modified operculum,
Spicules of two types: widely open compasses,
and large, wishbone-shaped calipers.

i)
>
rm

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 11. a,b, Thalamoporella granulata; a, autozooids, an ovicell and an avicularium, *70; b, group of
compasses, X 250. c, Labioporella bursaria, * 60, d,e,. Puellina egretia; d, X65; e, * 400.

BRYOZOA FROM HERON ISLAND

DISTRIBUTION

This species has been reported fram Hawaii,
Timor, Lombok and Mauritius, and is perhaps
widely distributed through the Indo-West-Pacific
realm. It has not been previously recorded from
Australia, It was present in two of the Heron
Island samples, numbers 24 and 26.

Labioporetla Harmer, 1926

Colony encrusting or erect. Autozooids with
membranous frontal wall underlain by extensive,
porous cryptocyst; opesia constituting one-third
or less of total length; no gymnocyst. Distal edge
of eryptocyst deflected to meet basal wall, divid-
ing cavity of cystid into two parts, forming a
prominent tube through which the polypide pas-
ses on eversion, No spines. Avicularia vicarious,
with extensive palatal cryptocyst; cystid divided
by transverse septum, rostrum with proximo-
lateral condyles but no crossbar. No ovicells.
Vertical walls with multiporous septula.

Type species: Lebiopora crennfata Levinsen,
1909.

Labioporella bursaria (MacGillivray)
(Fig. Llc)

Amphiblestrum bursarium MacGillivray, 1887; 66, pl.
2, fig. 2.

Labioporella bursaria: Harmer, 1926; 285, pl 21, figs
12, 15.

DESCRIPTIGN

Colony forming flat, unilaminar, encrusting
sheets. Avtozooids elongate, rectangular,
separated by distinct sutures; 0.6-0.85 <c.
0.35mm. Mural rim raised and crenulate in distal
half of autozooid, smoothly rounded along mid-
dlc portion of distal terminal wall for reception of
operculum, Cryptocyst occupying three-quarters
of autozooid length, finely granular, with a group
of about 30 small pores in its middle region; flat,
dipping sharply adjacent to polypide tube, which
is markedly conyex frontally, with a transversely
oval aperture. Avicularia as large as autozooids;
éryplocyst finely granular, imperforate, with an
oval foramen through whicha transverse, medial-
ly-thickened septum is visible, dividing the cystid
cavity in half; rostrum slightly less than half
length of avicularium, smoothly rounded, semi-
elliptical, with thickened, proxtmo-lateral con-
dyics.

243

DISTRIBUTION

Described by Harmer (1926) from Queensland
and the Celebes, L. bursarvia is also known from
Victoria and South Australia. At Heron Island it
occurred in two samples, from the reef flat and
the reef crest south of the cay.

Suborder ASCOPHORINA Levinsen, 1909
Superfamily CRIBRILINOIDEA Hincks, 1879
Family CRIBRILINIDAE Hincks, 1879

Colony encrusting, sheet-like; or erect, at-
tached by an encrusting base. Frontal surface of
autozooids consisting of a shield of fused, flat-
tened spines (costae) overarching the frontal
membrane; costae hollow, often with one or mare
uncalcified spots (pelmatidia) and with intercos-
tal spaces between the points of fusion permitting
the passage of water to the space between the
shield and the membrane. Distal oral spines
present or absent. Avicularia adventitious or
vicarious, or lacking. Ovicell hyperstomial. Ver-
tical walls with basal pore chambers or mural
septula.

Puellina Jullien, 1886

Colony encrusting, unilaminar. Autozooids
with reduced proximal gymnocyst, and costate
frontal shield covering whole of frontal
membrane. Frontal shield consisting of radiating
series of costae, each fused at intervals along
lateral margins, with small intercostal pores be-
tween points of fusion; cach costa with one or
more uncalcified frontal pores {pelmatidia),
Operculum D-shaped, closely fitting a calcified
secondary orifice, proximal edge of which (aper-
tural bar) is formed by the first pair of costac.
Short, slender spines present around distal and
lateral borders of orifice. A pair of slender. non-
calcified papillae present, proximo-lateral to
orifice, each protruding through a proximal inter-
costal pore. Avicularia interzooidal, with well-
developed gymnocystal cystid. Ovicell hyper-
stomial, imperforate, closed by awlozooid oper-
culum, Basal pore chambers present. Ancestrula
latiform.

Type species: Lepralia gattyae Landsborough,
1852.

Species of Puellina are frequently charac-
terized by the small size of both autozooids and
colony, They are abundant in all temperate and
tropical seas, apparently favouring biogenic car-
bonate substrata. They are al] confusingly similar
and many more species may exist than has been

previously recognized. Bishop & Househam
(1987) reviewed species of Puellina in British
coastal waters, and redescribed and figured P.
innominata (Couch), which, together with P-
radiata (Moll), has been reported from almost all
the world’s seas, All records of these, and other,
species of Puellina (formerly Cribrilaria) need to
be examined in the light of Bishop & Househam’s
(1987) work. The three species here described
from Heron Island have probably been reported
previously from Queensland and adjacent areas
as P. innominata or P. radiata. Features impor-
tant in identifying species of Puellina are: size
and shape of orifice, number of spines, form of
the suborificial lacuna, and morphology of the
avicularium,

Puellina decipiens sp. nov,
(Fig. 12d,e)

TYPE MATERIAL
Horotyre: GH5225. Heron Island, Stn 22, 15 Apr
1972,

ETYMOLOGY
Latin decipio, -ere, to beguile or cheat.

DESCRIPTION

Autozooids broadly oval, flat, separated by
shallow grooves. Gymnocyst limited to a narrow,
proximal band; frontal shield consisting of about
16 fused costae, each with a pronounced tubercle
basally (i.e., at periphery) bearing the single pel-
matidium, A single, large, transversely oval
lacuna Immediately proximal to the apertural bar.
Secondary orifice wider than long, with five,
evenly spaced disto-lateral spines, four persisting
in ovicelled autozooids. Avicularia infrequent;
cystid small, inconspicuous, rostrum elongate tri-
angular. Ovicell slightly wider than long, with
indistinct longitudinal carina.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.29+0.02;
width 0.2140.02; orifice length 0.05+0.002;
width 0.060.003; avicularium length 0.10£0.01
(n= 14),

REMARKS

The short avicularium is very similar to that of
the European P. innominata (Couch), but P.
decipiens differs from that, and other, species of
Puellina in its very large subapertural lacuna and
flat, non-umbonate frontal shield.

MEMOIRS OF THE QUEENSLAND MUSEUM

Puellina egretta sp. nov.
(Fig. 1}d,e)

Cribrilaria aff. radiata: Ristedt, 1985; 20, fig. 3e-h .

TYPE MATERIAL
HoLotyee: GH5226, Heron Island, Stn 12, 12 Apr
1972.

ETYMOLOGY
French aigrette, a heron.

DESCRIPTION

Autozooids broadly oval to irregular, flat,
separated by deep grooves. Gymnocyst minimal-
ly developed, inconspicuous; frontal shield
formed from about 19 fused costae, each
broadened but scarcely tuberculate proximally,
pelmatidia indistinct. Apertural bar with a small
median thickening; proximal to which is a small,
round lacuna, clearly distinct from the first series
of intercostal pores. Secondary orifice wider than
long; five closely spaced oral spines present.
Avicularia frequent; cystid well developed, with
broad, smooth gymnocyst; rostrum slender, elon-
gate, directed disto-medially along the border of
the adjoining autozoojd, Ovicell not found.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.34+0.03;
width 0.2740.03; orifice length 0.05+0.003;
width 0.07+0.005; avicularium rostrum length
0.24+0.05,

REMARKS

This species develops rather larger colonies
than the other two described here, and is thus
more immediately obvious. The broad, flat
autozooids and numerous elongate avicularia are
especially distinctive.

DISTRIBUTION

Ristedt (1985) figured specimens of a Puellina
from the Seychelles and the Philippine Islands
which are clearly referable to P. egretta.

Puellina vulgaris sp. nov.
(Fig. 12b,c)

TYPE MATERIAL
Ho.Lotyre: GH5227, Heron Island, Stn 23, 16 Apr
1972.

ETYMOLOGY
Latin vulgaris, commonplace or ordinary.

BRYOZOA FROM HERON ISLAND 245

FIG. 12. a, Figularia fissa, ¥ 75. b,c, Puellina vulgaris; b, X 130; c, X 450. de, Puellina decipiens; d, * 220;
e, *500.

DESCRIPTION

Colony forming smal] unilaminar patches-
Autozooids oval, convex, separated by deep
grooves. Gymnocyst clearly visible as a smooth
border around the autozooid; frontal shield
formed from 15-16 costae, each broadened
proximally but not tuberculate; pelmatidia indis-
tinct; mtercostal pores relatively large, particular-
ly in distal half of shield. A single, moderately
sized subapertural lacuna present. Secondary:
onofice wider than long, with seven closely spaced
spines, four persisting in ovice)led autozooids.
Aviculatia infrequent; cystid small, indistinct;
rostrum narrowly tiangular, relatively short,
directed distally or disto-laterally. Ovicell spheri-
cal, as wide as long, with a small frontal umbo.
Measurements (means and standard deviation of
20 values, mm): autezooid length 0.27+0.02;
width 0.2140.02; orifice length 0.04+0.003:
width 0.0540.005; avicularium rostrum length
0,12+0,02 (n =5).

REMARKS

The seven oral spines, conspicuous subaper-
tural Jacuna, and rather small avicularium
together distinguish this species from other tropi-
cal species of Puellina,

Figularia Jullien, 1886

Colony encrusting. Autozooids with well-
developed gymnocyst and, often extensive, cos-
tate frontal shield; costae fused medially and
along their length, with a few irregular pores
between the points of fusion, and with one or
more pseudopores on each. Aperture rounded-
quadrangular; operculum pivoting on lateral con-
dyles, frequently compound, with a transverse
line of flexure level with the condyles. Oral
spines present or absent. Avicularia vicarious or
interzooidal, generally sparse. Ovicell hyper-
stomial, prominent, | by autozooidal oper-
culum; typically with a longitudinal, median
suture, and one or more large pores, or fenestrae,
on each side. Vertical walls with uniporous or
multiporous septula. Ancestrula tatiform.

Type species: Lepralia figularis Johnston,
1847.

Figularia fissa (Hincks)
(Fig. 12a)

Cribrilina figularis var. fissa Hincks, 1880a: 196, pl.
26, fig, 8.
F ieularla fissa: Harmer, 1926: 478, pl. 34, figs 19-23,

MEMOIRS OF THE QUEENSLAND MUSEUM

DESCRIPTION

Colony forming extensive, unilaminar sheets;
brownish-coloured, with the opercula of the
autozooids conspicuously dark mahogany.
Autozooids elongate, convex, with costate frontal
shield comprising about one-third total length;
0.55-0.8 %c.0.35mm. Costae flat. 8-10 in total:
widely spaced, with elongate gaps between, cach
with a single, distal, pseudopore. Avicularia rare,
about as long as autozooid, witha striking, spoon-
shaped rostrum. Ovicell wider than long, the ec-
tooecium on each side of the median suture with
an clongate, curved, frontal fenestra.

REMARKS

This distinctive species is immediately recog-
nized by its costate frontal shield, which has
relatively few, widely spaced costae, and by its
characteristic avicularia. F. fissa is evidently
quite common at Heron Island, and was present
in four of the samples described here.

DISTRIBUTION

F. fissa does not seem to have been recorded
previously from Australia; it is known to occur in
the Indo-Malayan region, from Singapore to New
Guinea, and perhaps has a limited distribution in
the western Pacific.

Superfamily CATENICELLOIDEA Busk,
1852a
Family SAVIGNYELLIDAE Leyvinsen, 1909

Colony erect, consisting of branching, uniserial
chains of autozooids: jointed, each internode
formed from a single clavate autozooid. Frontal
wall of autozooids a porous gymnocyst, with or
without mural septula; orifice with or without a
sinus, its proximal rim sometimes formed from a
pair of fused costae; oral spines present or absent.
Avicularia adventitious, adjacent to orifice, or
lacking. Ovicell present or absent.

Vasignyella Gordon, 1989a

Colony erect to repent, attached by rhizoids.
Autozooids elongate, clavate, in branching
uniserial chains. Frontal wall largely gymnocys-
tal, with scattered pores; proximal side of orifice
bordered by a pair of small, fused costae, No
spines. Avicularia paired, lateral to orifice.
Ovicell unknown. Large uniporous septula in
lateral walls of autozooid.

; gu pe species: Catenaria otophora Kirkpatrick,

BRYOZOA FROM HERON ISLAND

Vasignyella otophora (Kirkpatrick)
(Fig, 14a)

Catenaria otephora Kirkpatrick, 189(b; 17, pl. 5, figs
la-c,

Sevignyella otephora: Harmer, 1957: 763, pl. 31, igs
19-21.

Vasignyella otophora: Gordon, 1989a: 453, figs 13-15.

DESCRIPTION

Colony erect or repent, attached to the sub-
stratum by delicate, chitinous rhizoids; composed
of uniserial chains of slender, clavate autozooids,
branching at irregular intervals. Autozooids very
lightly calcified, clavate or fusiform, broadest
distally, tapered proximally; 1.0-1.3X 0.25mm.
Primary orifice terminal, about as wide as long.
the proximal edge shallowly concave between
indistinct condyles; proximal border of orifice
formed by a pair of short, fused costae. Frontal
wall evenly perforated by large. round pores; a
row of about five large, uniporous septula in each
lateral wall. Avicularia lateral to orifice. paired;
the rostrum triangular, distally directed, project-
ing conspicuously. Each autozooid buds another
from its disto-basal surface, and one or more
additional autozooids may be budded from the
lateral septula; the proximal part of each
autozooid, immediately above its poin{ of origin,
is uncalcified, and appears as a brown, cuticular
node

DISTRIBUTION

V. otophora is probably widely distributed
throughout the Indo-West-Pacific realm, but is
particularly inconspicuous and has been reported
on very few occasions. Most records are centred
on the Indo-Malayan region, but it has also been
recorded as far west as the Sudanese Red Sea
(Dumont, 1981) and, to the east, from Samoa
(Gordon, 1989a),

Superfamily HIPPOTHOOIDEA Busk, 1859
Family HIPPOTHOIDAE Busk, 1859

Colony encrusting or erect, Warious zootd
polymorphs occur, including female zooids and
dwarf zooeciules. Autozovids with gymnocystal
frontal wall; imperforate, or with small punctae
and/or tubercles. Primary orifice rounded or
sinuate; condyles present: oral spines generally
lacking. Avicularia interzooidal or vicarious, or
absent. Ovicell hyperstomial, closed by zooidal
operculum; or absent. Vertical walls. with basal
pore chambers and small mural septula

247

Hippothoa Lamovroux, 182!

Colony encrusting, consisting of branching,
uniserial chains of polymorphic zooids.
Autozooids clavate, with a slender, stolon- like
proximal portion; ovicellate female zooids often
smaller than autozooids, ovicell with frontal
umbo or apical perforation, closed by zooidal
operculum; zooeciules minute, cylindrical, with
simple orifice, of unknown function. Frontal wall
imperforate, smooth, or ridged. Primary orifice
with sinus and distinct condyles; no oral spines,
No avicularia, Tubular pore chambers in lateral
walls. Ancestrula kenozooidal or tatiform,

Type species: Hippothea divaricata
Lamouroux, 1821.

Hippothoa calciophilia Gordo
(Fig. 133.b)

Hipporhoa calciophilia Gordon, 1984; 108, text-fig.
10, pL 42, figs A-C.

DESCRIPTION

Colony diffuse, spreading Widely over the sub-
stratum; zooid chains branching in a cruciform
manner, each zooid producing one distal and
paired lateral buds, Distal, clavate portion of
autozooid about 0.3 ¥ 0.15mm; filiform proximal
portion 0.02mm wide, 1-15 length of distal
portion. Primary orifice slightly longer than wide,
commonly 0,06 0.05mm, with a deep, broad,
U-shaped sinus, and small, rounded condyles.
Frontal wall steeply convex, often with fine,
transverse corrugations, and a longitudinal ndge.
Female zooids most usually laterally budded
from autozooids, the distal portion of similar size
to autozooids, the filiform proximal portion
usually shorter; ovicell with semicircular aper-
ture, and a single, small, apical perforation,
Zooeciules sporadic, budded from either or both
of the two basal pore chambers on each side of an
autozooid; (.05-0.1mm long, with minute distal
orifice.

REMARKS

H. caiciophilia has only recently been distin-
guished from other species of Aippothoa (Gor-
don, 1984). It is probably widely distribyted
throughout the Great Barrier Reef province, but
its small size and cryptic habits render it very
inconspicuous.

248 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 13. a,b, Hippothoa calciophilia; a, part of a colony, X50; b, a female zooid, with zooeciules, 215. c,
Chorizopora brongniartii, X85. d, Exochella conjuncta, X70. e, Tremogasterina spathulata, < 65.

BRYOZOA FROM HERON ISLAND

Family CHORIZOPORIDAE Vigneaux, 1949

Colony encrusting, unilaminar. Frontal wall of
nulozooids imperforate, or with minute punctae.
Primary orifice semicircular, without a sinus.
Avicularia interzooidal. Ovicell hyperstomial,
imperforate, Small kenozooids present. Vertical
walls with tubular basal pore chambers.

Chorizopora Hincks, 1879

Colony encrusting, forming thin unilaminar
sheets, Autozooids slightly disjunct, linked by
tubular extensions of the basal pore chambers.
Frontal wall gymnocystal, thinly calcified, im-
perforate. Primary orifice semicircular or D-
shaped; oral spines present or absent. Avicularia
vicarious, smaller than autozooids; small
kenozooids with rounded membranous frontal
fenestrae interspersed among autozooids. Ovicell
hyperstomial, imperforate, closed by autozooid
operculum,

Type species: Flustra brongniartii Audouin,
1826.

Chorizopora brongniartii (Audouin)
(Fig. 13c)

Flastra brongniartii Audouin, 1826; 240; Savigny pl.
10, fig. 6.

Chorizopora brongniartii: Hayward & Ryland, 1979:
240, fig. 103; Gordon, 1984: 113, pl. 44, fig. C;
Hayward, 1988; 327.

REMARKS

The thin, hyaline colonies of C. brongniartii are
inconspicuous on the carbonate substrata it ap-
pears to favour and are easily overlooked in wet
material. At least three other species of
Chorizopora are known from New Zealand (Gor-
don, 1984), but C. brongniartii is readily recog-
nized by the smooth or transversely wrinkled, but
non-spinous, frontal walls of its autozooids.

DISTRIBUTION
C. brongniartii is widely distributed in all
temperate and tropical shelf seas.

Superfamily ARACHNOPUSIOIDEA Jullien,
1888
Family EXECHONELLIDAE Harmer, 1957
Colony encrusting, Avtozooids with per-

forated, umbonuloid frontal wall, Orifice with
well marked peristome; lacking spines.

249

Aviculana adventitious and/or vicarious or inter-
zooidal. Ovicells present or absent.

Tremogasterina Cana, 1911

Colony encrusting or erect, multilaminar,
Autozooids with umbonuloid frontal wall, per-
forated by few, large, frontal foramina, and with
small marginal pores. Calcified orifice rim with
lateral condyles; no oral spines. Aviculana inter-
zooidal and/or vicarious. Ovicells prominent, hy-
perstomial, imperforate; not closed by autozuoicd
operculum. Autozooids linked by large, multi-
porous septila. Ancestrula tatiform,

Type species: Tremogasterina problematica
Canu, 1911.

Tremogasterina has a worldwide distribution,
in shallow coastal waters, between the 30° north
and south parallels. The genus was studied in
detail by Cook (1977).

Tremogasterina spathulata (Canu & Bassler)
(Fig. 13e)

Hiantopora spathulata Canu & Bassler, 1929: 116, pl.
11, figs 13, 14.

Arachnopusia spathulata: Harmer, 1957: 657, pl. 68,
figs 22, 23, 25, 31.

Tremogasterina spathulata: Cook, 1977: 136, pl. 5, fiz
A; text- figs 1B, 3A.

DESCRIPTION

Colony forming small, unilaminar patches.
generally inconspicuous. Autozooids oval, con-
vex, separated by deep grooves: 0,5-0.6 x 0,3-
0.4mm. Orifice longer than wide, narrowest
proximally; proximal edge more or less straight
thickened, and frequently with a short, suboral
mucro, Four or five delicate, distal spines present,
Frontal wall with a variable number (2-20) of
irregular foramina, each with a broad, thick nm,
the sutures between giving a tessellated effect;
marginal pores inconspicuous, Avicularia
sporadic, developed on disto-lateral borders of
autozooids, 0.2mm long, typically distally
directed; rostrum spatulate, with straight distal
edge, upeurved. Ovicell wider than long.

DISTRIBUTION

Widely distributed in the Indo-West-Pactfic
region, from the Red Sea to the Philippines. 7,
spathulata is essentially a shallow water species,
and its small colonies occur on coral rubble and
other hard substrata; on Mauritian reefs it is espe-
cially common on the inner reef flat (Hayward,

250

1988). At Heron Island it was present if half of
the samples studied to date.

Superfamily UMBONULOIDEA Canu, 1904
Family EXOCHELLIDAE Bassler, 1935

Colony encrusting or erect. Autozooids with
umbonuloid frontal shield bordered by con-
spicuous marginal pores. Orifice with well
developed peristome, incorporating one or more
denticles on its inner, proximal edge; oral spines
present. AVicularia adventitious, Ovicell present.
Vertical walls with basal pore chambers or mural
septula,

Exochella Jullien, 1888

Colony encrusting, unilaminar. Frontal wall of
autozooids umbonuloid, imperforate other than
fora single series of large, marginal pores, Orifice
obscured in early ontogeny by a complex peris-
tome, its lobed proximal edge corresponding with
ridges on its inner face which delimit one or two
narrow channels; at the lip of the peristome, these
channels may close distally to form one or two
distinct spiramina. Oral spines typically present,
Aviculana adventitious, budded from marginal
pores (frontal septula) proximo-lateral to orifice;
one, two, Or more per autozooid, usually with
acuminate rostrum. Ovicell spherical, imper-
forate. sometimes partly immersed, not closed by
autozooid operculum, Basal or mural pore cham-
bers present.

Type species: Mucronella tricuspis Hincks,
188 le,

Exochella is distributed almost exclusively in
the southern hemisphere. There are several An-
taretic species, and a number of others from cold
temperate or subantarctic waters of South
America, the Southem Indian Ocean and New
Zealand (Hayward, 1991). E. conjuncta Brown is
the only species known to range into tropical
waters.

Exochella conjuncta Brown
(Fig. 13d)

Exochella vonjuncia Brown, 1952: 294, figs 220-223.
Exochella conjuneta: Gordon, 1989b;30, pl.14, fig. A.

DESCRIPTION

Colony forming small, rounded patches,
Autozooids oval to hexagonal, canvex, separated
by thickened ridges which become indistinct in
later ontogeny; small, 0.35-0.4 by about 0.25mm,

MEMOIRS OF THE QUEENSLAND MUSEUM

Frontal wall with few, large, marginal pores;
thickening steadily, becoming vitreous and
smooth, with the boundaries of each autozooid
marked only by the pronounced, rounded, mar-
ginal pores. Peristome with paired inner ridges,
delimiting a single median channel, opening ex-
ternally as an oval spiramen. Two short, distal,
oral spines present in newly budded autozooids,
obscured by development of peristome. One or
two avicularia per autozooid, 0.13mm long, with
slender, laterally-directed rostrum. Ovicell
spherical, smooth, largely immersed and hidden
in later ontogeny.

DISTRIBUTION

E. cenjuncta was first described from Pliocene
deposits of New Zealand (Brown, 1952), and
subsequently shown to have a Recent distribution
extending from the South Island of New Zealand
to New South Wales (Gordon, 1989b). Its
presence a{ Heron Island marks a considerable
extension in its known range.

Family CELLEPORARIIDAE Harmer, ]957

Colony encrusting, multilaminar, massive or
nodular, often erect, developing stout, irregular
branched or lobed forms; or pisiform, encrusting
erect substrata; or discoidal and unattached,
Autozooids recumbent to erect, typically without
regular orientation; frontal wall umbonuloid,
with marginal pores. Onfice without a sinus, with
or without one or more proximal denticles; oral
spines present or absent. Avicularia adventitious,
associated with orifice, and vicarious; Lypically
polymorphic, Ovicells present.

Celleporaria Lamouroux, 1821

Colony encrusting, or erect and irregularly
branched, or massive; multilaminar, developed
by frontal budding of successive autozooid
generations. Autozooids recumbent at growing
margins, elsewhere suberect, with varying orien-
tation. Frontal wall umbonuloid, with indistinct
marginal pores. Oral spines typically present,
Avicularia adventitious and vicanous, frequently
polymorphic, Ovicell hyperstomial, small and
spherical, imperforate.

Type species: Cellepora cristata Lamarck,
1816,

This genus is Widely distributed in tropical and
subtropical seas, with many species reported
from shallow feef habitats, Colonies are often
large, and conspicuously pigmented. Unfor-

BRYOZOA FROM HERON ISLAND 251

tunately, identification is often difficult. The
taxonomy of Celleporaria is confused, at least
partly because many species have still to be
named, and a majority of descnbed species are
poorly known and have never been adequately
illustrated.

Celleporaria tridenticulata (Busk)
(Fig. 14b,c)

Cellepora tridenticulata Busk, 1881: 343,344,347, pl.
26, fig. 9; 1884: 198, pl. 29, fig. 3; pl. 35, fig. 17.
Celleporaria tridenticylata; Harmer, 1957: 670, pl. 42,

figs 5-10; Gordon, 1984: 115, pl, 45, figs C-E.

DESCRIPTION

The aperture of C. tridenticulata bears three
conspicuous denticles on its concave proximal
border, which may be pointed, truncate, bifid, or
even trifid. It has two (rarely three or four)
slender, distal oral spines, and a median,
proximal, suboral avicularium on a short um-
bonate cystid, with the oval rostrum acute to the
onfice plane and facing laterally. Vicarious
aviculana vary from 0.3 to 0.5mm Jong, with a
slender rostrum, slightly spatulate distally, with a
finely serrate edge, and a stout crossbar, which
lacks a columella.

DISTRIBUTION

Described originally from Torres Strait. this
species has been recorded from throughout the
Great Barrier Reef province, from New South
Wales, Victoria and New Zealand. It is dis-
tributed through the Indo-Malaysian Ar-
chipelago, westwards to India, Sn Lanka and
Maunitius, Cany & Bassler’s (1930) record from
the Galapagos Islands should perhaps be re-ex-
amined.

REMARKS

C. vagans (q.v.) also bears three, thin, pointed
denticles on the proximal border of the orifice. It
is distinguished from C. tridenticulata by its lack
of oral spines; by its deeply notched peristome;
and by its vicarious avicularium, which is more
slender than that of C. tridenticulata, with a
smooth edge to the rostrum and a distinct
columella.

Celleporaria aperta ((Hincks)
(Fig. 14d,e)

Schizoporella aperta Hincks, 1882: 126, pl. 3, fig. 3.

Celleporaria aperta: Harmer, 1957; 673 (in party;
Winston & Heimberg, 1986: 32, figs 79-84.

DESCRIPTION

Colony multilaminar and nodular, but rather
small, rarely occupying more than a few cm’ of
substratum. Recumbent autozooids commonly
0.5 0.3mm, with convex, nodular frontal wall;
marginal pores few, small and indistinct. Primary
orifice wider than long, about 0.1 X 0,15mm, the
proximal border slightly concave, witha shallow,
uregularly U-shaped sinus medially; condyles
small, pointed, inconspicuous; typically with
two, widely spaced, disto-lateral spines, and oc-
casionally with a third, distally situated, Suboral
avicularium characteristic: proximal to sious,
small, with oval rostram, finely denticulate dis-
tally, 0.0Smm long; perpendicular to plane of
orifice and facing laterally, Vicarious avicularia
sporadic, rostrum commonly 0.2-0.3mm long,
parallel-sided, and rounded distally, or rather lan-
ceolate with a tapered distal tip; crossbar stout,
flattened. Ovicell shallow, imperforate, with
finely nodular calcification,

REMARKS

Winston & Heimberg (1986) re-examined
some of the material studied by Harmer (1957),
and considered that he had confused several
species of Celleporaria in his account of C. aper-
ta.

DISTRIBUTION

C. aperia seems to be widely distributed in the
western Pacific, from the South China Sea to
Queensland, and throughout the East Indies,
ranging to Sri Lanka and East Africa. Its reported
occurrence from West Africa, Cape Verde Is-
lands (Cook, 1968b, 1985) and the Caribbean
should be regarded with caution until specimens
from those regions have been examined by SEM.

Celleporaria desperabilis sp. nov.
(Fig. t5e,f)

TYPE MATERIAL

HototypPe: GH5228, Heron Island, Sin 14, 12 Apr
1972.

PARATYPE: GH5229, as for holotype,

ETYMOLOGY
Latin despero, déspéerabilis, hopeless.

DESCRIPTION ..
Colony forming a multilaminar, encrusting

252 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 14. a, Vasignyella otophora, * 120. b, c, Celleporaria tridenticulata; b, X50; c, detail to show primary
orifice and a vicarious avicularia, X 120, d,e, Celleporaria aperta; d, X 85; e, vicarious avicularia, < 95.

BRYOZOA FROM HERON JSLAND

sheet. Autozooids 0,6-0.7 x 0,4-0.5mm,; frontal
wall convex, finely nodular or wrinkled, with
few, relatively large, marginal pores. Primary
onfice wider than long, about 0,12 0.7mm, the
proximal border gently concave, bearing three
denticles within its edge: lateral denticles stout.
pointed; median denticle vanable, quadrate, or
anvil-shaped, its free edge often notched or bifid,
sometimes completely divided to give two
separate, slender denticles, No oral spines, Peris-
tome developing early in ontogeny, enclosing
and obscuring proximal and lateral borders of
orifice: a single median suboral avicularium,
columnar, the rostrum perpendicular to plane of
orifice, elongate triangular, about 0.15mm long,
with finely toothed dista] edge; crossbar slender,
with a conspicuous columella; apically, the
avicularium is produced as a tapered umbo, Ad-
jacent to the proximal end of the avicularium, the
peristome is deeply notched. Tiny adventitious
aviculana, with oval rostrum, often frequent
along margins of autozooids- Vicarious
avicularia rare, apparently absent from many
colonies; small, rostrum 0.15-0.25mm long,
parallel-sided, deeply cupped distally, Ovicell a
simple hemisphere, widely open frontally, lightly
calcified, but thickening slightly in later on-
togeny.

REMARKS

This species is distinguished from C. triden-
ticulata by its large, columnar suboral
avicularium, deep peristome, and lack of oral
spines. C. yagans (q.v.) has three thin, pointed
denticles within the proximal edge of the orifice:
its suboral avicularium is proportionately much
smaller than that of C. desperabilis, and the
ovicell larger, with a granular surface, and an
asymmetrical aperture which may bear a small
aVicularium on its rim.

Celleporaria fusca (Busk)
(Fig. 15a-d)

Cellepora fusca Busk, 1854: 88. pl. 119, fig. 2: pl. 120,

fig. &.
Ceileporaria fusca: Harmer, 1957: 680, pl. 43, figs 1-7.

DESCRIPTION

Colony forming massive, nodulated, multi-
laminar sheets, often exceeding 25cm”; less fre-
quently, developing erect, irregularly branching
growths; deep purplish-brown in colour. Aper-
ture aboutas wide as long, more or less D-shaped,
the proximal border slightly concaye, and arched

2353

frontally. No oral spines. Penstomial thickening
most pronounced proximally, incorporating a
single medio-proximal aviculatium, Vertically
orientated with respect to the aperture, its cystid
produced apically as a stout, pointed umbo;
rostrum facing laterally, small and oval (0.05mm
long), or longer (0.15-0.30mm), with a strongly
denticulate distal edge. Vicarious avicularia
generally infrequent, but characteristic: rostrum
slender, lanceolate, rounded distally and often
with a slight lateral curve; 0.2-0.6mm long.
Ovicell spherical, imperforate, its orifice rim with
an asymmetrical thickening which may include a
small adventitious avicularium. Immediately
below the avicularium the peristome rim bears a
natrow notch, on the other side of which the nm
is slightly thickened.

REMARKS

Winston & Heimberg (1986) indicated that
Hurmer’s (1957) account of C. fusca was founded
on a number of similar, deeply pigmented
species, Harmer's figures, however, appear to
represent true C. fusca. The type specimen
(BMNH reg. no. 1854.11.15.180)} 1s labelled ‘pl.
119. Cape Capricom' in Busk's hand, although
the published type locality was given as Bass”
Strait. A second Busk collection specimen
(BMNH 1899.7.1.4867A) is labelled ‘Bass’
Strait’; both this and the type specimen are
labelled 'R’. for HMS Rattlesnake. A Hincks
collection specimen from Bass’ Strait (BMNH
1899.5,.1,1316), Torres Strait specimen figured
by Harmer (1957)(BMNH 1963.9.8.11), and a
Siboga specimen from Makassar (BMNH
1963.9.8.14 part) have all been compared with
the two Busk collection specimens, and with the
present series from Heron Island, and it js clear
that they all represent a single species. On the
basis of these specimens, C. fusca can be said to
range along the entire east coast of Australia,
from Bass Strait to Torres Strait, and into the
Indo-Malaysian Archipelago, In view of Winston
& Heimberg’s (1986) account, the synonymy
constructed by Hanmer should be viewed with
caution, until records from the Indian Ocean can
be re-examined

Celleporaria pigmentaria (Waters)
(Fig, 16d,e)

Heloporella piymentarta Waters, 1909: 163, pl. 15,
figs 16-19; pl. 16, figs 9-16; pl. 17, figs 22, 23,

Cellaporaria pigmentaria: Harmer, 1957; 684, pl, $3,
fig. 8; text- fig. 58.

254 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 15. a-d, Celleporaria fusca; a, primary orifice, x 290; b, part of colony, with elongate vicarious avicularia,
* 45; c, part of colony, with spatulate vicarious avicularia, X70; d, detail to show ovicell, x 120. e,f,
Celleporaria desperabilis; e, * 55; f, * 100.

BRYOZOA FROM HERON ISLAND 255

FIG. 16, a-c, Celleporaria vagans; a, X 45;b, X50; c, X45, de, Celleporaria pigmentaria; d, * 45: e, * 60. f,
Arthropoma cecilti, X50.

256

DESCRIPTION

Colony multilaminar, nodular, occasionally
developing stout, erect, branching forms, deep
brown to black. Autozooids oval, strongly convex,
thickly calcified; commonly 0.5-0.6X0.4-
0.5mm. Frontal wall coarsely nodular, with small,
indistinct marginal pores. Aperture orbicular,
slightly wider than long, its rim thickened but not
especially prominent; a very small avicularium
present mid-proximally, vertically orientated,
with oval rostrum 0.05mm long; adjacent to the
avicularium the peristomial rim clearly notched.
No oral spines. Vicarious avicularia rare, usually
absent; rostrum narrowly scaphoid, 0.25mm long,
with incomplete crossbar. Peristomial
avicularium occasionally giving rise to a stout,
spiked mucro, which may be particularly accen-
tuated among the oldest autozooids. Ovicell
hemispherical, thickly calcified, inconspicuous.

REMARKS

This species was originally described from the
Red Sea (Waters, 1909), and reported from a
single Indonesian locality by Harmer (1957). The
present material has been compared with the
specimens from Ghardaqa and Sumbawa listed
by Harmer, and with a specimen collected pre-
viously from Heron Island by L. Vail, and all
seem to represent a single species.

Celleporaria vagans (Busk)
(Fig. 16a-c)

Cellepora vagans Busk, 1881: 343; 1884: 198, pl. 29,
fig. 10; pl. 35, fig. 11.

Celleporaria vagans: Harmer, 1957: 671, pl. 42, fig. 4;
text-fig. 55.

DESCRIPTION

Colony forming a multilaminar, encrusting
sheet. Autozooids 0.6-0.8 Xc.0.4mm; frontal
wall convex, nodular and rugose, bordered by
small but distinct marginal pores. Primary orifice
wider than long, commonly 0.17 X0.12mm;
proximal edge shallowly concave, with three
thin, pointed denticles evenly spaced just within
its rim. No oral spines. Peristome deep, develop-
ing in early ontogeny, enveloping lateral and
proximal borders of orifice; a single median sub-
oral avicularium, small, columnar, its apex
produced as a short, pointed umbo; rostrum 0.1-
0.2mm long, oval, orientated vertically to orifice
plane and directed laterally. Peristome rim deeply
notched adjacent to the proximal end of the
avicularium; in later ontogeny a process extend-

MEMOIRS OF THE QUEENSLAND MUSEUM

ing from the peristome rim eventually fuses with
the opposite rim, below the avicularium, to
delimit a rounded spiramen. Vicarious avicularia
infrequent, variable in length, but reaching
0.6mm; elongate, narrowest medially, spatulate
and slightly cupped distally. Ovicell more than a
hemisphere, overarching the primary orifice, sur-
face finely granular, with a few indistinct perfora-
tions; its aperture irregularly triangular, the rim
often peaked medially, and bearing a small
avicularium in later ontogeny.

DISTRIBUTION

C. vagans is known from Hawaii, Torres Strait,
Indonesia, and the eastern Indian Ocean, but from
relatively few specimens. Busk’s (1881, 1884)
material from Crozet Island perhaps represents a
different species, and requires re-examination.

Superfamily SCHIZOPORELLOIDEA Jullien,
1883
Family HIPPOPODINIDAE Levinsen, 1909

Colony encrusting. Autozooids with cryptocys-
tidean frontal wall, evenly perforated. Orifice
more or less bell-shaped: arched distally, shal-
lowly concave proximally, with prominent lateral
condyles. No oral spines. Avicularia adven-
titious, typically associated with orifice. Ovicell
partly or wholly immersed, perforated, closed by
autozooid operculum.

Hippopodina Levinsen, 1909

Colony encrusting. Frontal wall of autozooid
with cryptocystidean development, perforated by
numerous, closely spaced, small pores. Primary
orifice bell-shaped; proximal border shallowly
concave between distinct lateral condyles; no oral
spines. Avicularia adventitious. Ovicell partially
immersed in next autozooid, finely perforate,
closed by autozooidal operculum. Vertical walls
of autozooids with numerous, small, uniporous
septula.

Type species: Lepralia feegeensis Busk, 1884.

Hippopodina feegeensis (Busk)
(Fig. 17a)

Lepralia feegeensis Busk, 1884: 144, pl. 22, figs 9, 9a,
9b.

Hippopodina feegeensis: Harmer, 1957: 974, pl. 67,
figs 7-9; Winston & Heimberg, 1986: 16, figs 28-30.

BRYOZOA FROM HERON ISLAND

257

FIG. 17. a, Hippopodina feegeensis, * 55. b, Cosciniopsis lonchaea, X 50. c, d, Tharnelya perarmata;, c, * 60,

d, *85

DESCRIPTION

Colonies forming thin, very extensive, light
yellowish-brown, unilaminar sheets, glistening
when dried. Autozooids oval to hexagonal, con-
vex, separated by distinct grooves, lightly cal-
cified; commonly 0.75-0.9X0,5-0.6mm,
Primary orifice longer than wide, bell-shaped,
with sharp, down-curved condyles. Frontal wall

pearly or nodular, with numerous, densely-
spaced, small pores, giving a speckled ap-
pearance at low magnification. Avicularia single
or paired, disto-lateral to orifice, slender,
acuminate rostrum directed medially, close to the
distal border of the orifice.

Ovicell elongate oval, very prominent, densely
perforated, with arched aperture.

REMARKS

H. feegeensis has a pan-tropical distribution in
shallow waters. It forms broad flat sheets, often
exceeding 100cm*, on all kinds of hard substrata,
and may be the most abundant bryozoan occur-
ring on coral rubble in reef flat habitats. It is also
often the primary colonising species on artificial
substrata. It was common in 11 of the samples
from Heron Island reported here.

Cosciniopsis Canu & Bassler, 1927

Colony encrusting, often strikingly pigmented.
Antozooids large, robust, with thickly calcified,
tuberculate frontal wall, pierced by numerous
small pores. Primary onfice elongate-oval or
bell- shaped, the proximal border broadly con-
cave, between distinct lateral condyles. No oral
spines. A well developed peristome typically
present. Adventitious avicularia developed
lateral to orifice. single or paired; lacking in some
species. Ovicell hyperstomial, globular, with
tuberculate calcification pierced by numerous
small pores; closed by autozooid operculum.

Type species: Coseiniopsis coelatus Canu &
Bassler, 1927.

Cosciniopsis lonchaea (Busk)
(Fig. 17b)

Lepralia lonchaea Busk, 1884: 146, teat-fig. 43.
Cosciniopsis lonchaea: Harmer, 1957: 1083, pl. 72,
figs 16, 17, 19, 20.

DESCRIPTION

Colony forming small, unilaminar patches,
Autozooids oval to hexagonal. strongly convex,
separated by deep grooves; 0.7-1.0%¢.0.6mm.
Primary orifice elongate-oyal, the proximal edge
with thickened, concave nm; condyles small and
rounded; enveloped distally and laterally by a
thin, raised peristome. Frontal calcification strik-
ingly tuberculate, the tiny pores just visible be-
tween the tubercles. Most autozocids with a
single, lateral-oral, aviculanum; its blister-like
cystid closely applied to the peristome, the
acuminate rostrum acute to frontal plane, directed
disto-medially towards the orifice. In some
autozooids the lateral-cral avicularium is paired,
one each side of the orifice; in others an identical
avicularium occurs on the distal edge of the peris-
tome, with the rostrum perpendicular to the fron-
tal plane. directed towards the orifice of the
autozooid. Ovicell slightly wider than long,

MEMOIRS OF THE QUEENSLAND MUSEUM

globular, with tuberculate calcification and fine
perforation identical to that of the autozooid.

DISTRIBUTION

Described originally from New Guinea, C.
lenchaea has been reported from Torres Strait
and South China Sea by Kirkpatrick (1890a,b)
and from Indonesia by Harmer (1957). Waters
(1909) recorded it from Sudan, while Harmer's
(1957) synonymy of Lepralia vestita Hincks ex-
tends its range to Tahiti and Hawaii.

Thornelya Hanmer, 1957

Colony encrusting, unilaminar. Primary orifice
of autozooid bell-shaped, with broad, concave
proximal border; condyles distinct; oral spines
present. Frontal wall with large marginal pores,
and small, scattered frontal pores. Adventitious
avicularia present, often numerous, one to four
per autozooid, most usually associated with the
orifice or the ovicell, generally rather small.
Oyicell recumbent on next autozooid, prominent,
irregularly rounded, surface densely perforated,
sutured, often bearing avicularia; closed by
autozooid operculum.

Type species: Lepralia ceylonica Thornely,

905,

Thornelya perarmata Harmer, 1957
(Fig. 17c.d)

Thornelya perarmata Harmer, 1957: 1105, pl. 73, figs
6-8.

DESCRIPTION

Colony forming very small, unilaminar
patches. Autozooids imegularly rectangular to
hexagonal, rather flat, separated by thickened
sutures; 0.4-0.5 X0.3-0.35mm. Primary orifice
0.12mm long, with six spines in a distal arc;
condyles rounded, distinct. Frontal wall smoothly
calcified, nodular; with 10-30 small pores
centrally, and a single series of larger, elongate
oval marginal pores. Avicularium single or
paired, Jateral to orifice, rostrum triangular,
O.lmm long, acute to frontal plane, directed
towards orifice; one or two, occasionally more,
avicularia developing in later ontogeny on distal
edge of orifice, with rostrum directed towards it;
additional avicularia developed along lateral
margins of autozooids, Ovicell about as wide as
long, prominent, with densely punctured, nodular
surface: a single avicularium present on frontal
surface, with rostrum proximally directed,

BRYOZOA FROM HERON ISLAND

DISTRIBUTION

This species was described by Harmer (1957)
from 141m off Waigeo Island, northwest New
Guinea. It has not been recorded previously from
the Great Barrier Reef, or from reef flat habitats,
it seems to be common at Heron Island, and
occurred in seven of the samples studied.

Family HIPPOPORINIDAE Brown, 1952

Colony encrusting; or erect with an encrusting
base. Autozooids with regularly perforated fron-
tal wall. Primary orifice shallowly concave
proximally, or distinctly sinuate, with con-
spicuous condyles:.no oral spines. Avicularia ad-
ventitious. Ovicell typically large, inflated, often
sutured; in some geneta brooding autozooids
with large. dimorphic orifice.

Calyptotheca Harmer, 1957

Colony encrusting: sheet-like; or erect.
bilaminate, plate-like. Frontal wall of autozooids
densely perforated. Primary orifice sinuate, or
with evenly concave proximal edge; dimorphic,
typically larger and broader in brooding
autozooids. Avyicularia typically small, adven-
titious, lateral oral or sutural; rarely, vicarious? or
absent. Ovicell large and conspicuous, per-
forated, closed by autozeoid operculum.

Type species: Schizeporella nivea yar.
wasinensis Waters, 1913.

Calyptotheca tenuata Harner
(Fig, 18b,c)

Calyptotheca tenuata Harmer, 1957: 1016, pl. 68, figs
16, 17.

DESCRIPTION

Colony developing extensive, multilaminar
sheets. Autozooids oyal to hexagonal or ir-
regularly polygonal, convex, separated by thin,
raised sutures; commonly 0.6-0.7*0.Smm.
Primary orifice slightly wider than long, proximal
border shallowly concave between knob-like
proximo-lateral condyles; with a low, nodular
peristome developing in later ontogeny. Frontal
wall thick, densely and evenly perforated by
regular, round pores, with nodular calcification
between. Ovicell partly immersed in distally suc-
ceeding autozooid, wider than long,, rather flat-
tened frontally, with a coarsely nodular surface
perforated by imregular pores. Avicularia sparse,
developed on lateral margin of autozooid,

259

proximal to orifice, directed medially and reach-
ing midpoint of frontal wall; rostrum slender.
acuminate, 0.35mm long.

DISTRIBUTION

C. tenuata was known only from Indonesia, the
Loyalty Islands and Sri Lanka; Heron Islancl
provides the first Australian record.

Calyptotheca thornelyae Dumont
(Fig. 18a)

Lepralia subimmersa: Thomely, 1903; 122,

[not] Lepralia subimmersa MacGillivray, 1879; 23, p!
35, fig. 5.

Calyptotheca thornelyae Dumont, 1981: 625

DESCRIPTION

Colony forming extensive, unilaminar or mul-
tilaminar, light reddish-brown sheets.
Autozooids rectangular, hexagonal or irregularly
polygonal. convex, separated by distinct sutures;
0.5-0.75 X0,.3-0,5mm. Primary orifice wider
than long, proximal border shallowly and evenly
concave between low, rounded, proximo-lateril
condyles. Frontal wall thick, finely nodular,
evenly perforated by numerous, small. round
pores: a low, crenulate, peristomial rim charac-
teristically present proximal to the orifice.
Ovicell prominent, domed, occupying most of the
fronts] wall of the distally succeeding autozonid;
calcification and perforation identical to that of
autozooid frontal wall; orifice of brooding zooid
shorterthan that of autozooid, with more shallow-
ly concave proximal edge. Adventitious
avicularia sporadic, infrequent; 0.1mm long, with
slender, acuminate rostrum; single or paired,
most usually situated proximo-lateral to orifice,
medially directed, occasionally budded along the
lateral margins of autozooids in later ontogeny,

REMARKS

Thornely’s (1905) two specimens of ‘Lepralia
subinonersa’ from Soi Lanka appear to be the only
known material of this species, prior to its collec-
tion from Heron Island; one of the specimens was
designated lectotype of Calypiotheca thornelyae
by Dumont (1981). Both specimens have been
compared with the Heron Island material, and
there is no doubt that the two sets are conspecific.
Calyptotheca subimmersa (MacGillivray) is
presently known only from the southem coasts of
Australia.

260

FIG. 18. a, Calyptotheca tharnelyae,
*70.

Emballotheca Levinsen, 1909

Colony encrusting. Autozooids with evenly
perforated frontal wall. Primary orifice shallowly
concave proximally, or distinctly sinuate; with
well-marked condyles; dimorphic, larger and
broader in brooding autozooids. No oral spines.
Avicularia adventitious; or absent. Ovicells large,
inflated, sutured; regularly porous.

Type species: Eschara quadrata MacGillivray,
1880.

MEMOIRS OF THE QUEENSLAND MUSEUM

90. b,c, Calyptotheca tenuata; b, * 70; ¢, x 60; d, Emballotheca pacifica,

Emballotheca pacifica Harmer
(Fig. 18d)

Emballotheca pacifica Harmer, 1957: 1088, pl. 67, figs
3-5.

DESCRIPTION

Colony an encrusting, multilaminar sheet.
Autozooids rectangular to irregularly polygonal,
convex, separated by distinct sutures. Primary
orifice wider than long, its proximal border with

BRYOZOA FROM HERON ISLAND

a rounded-triangular median process; with large,
downcurved lateral condyles. Frontal wall thick,
coarsely granular, perforated by large, irregular
pores, which tend to fuse in later ontogenetic
thickening. Ovicell prominent, obscuring much
of distally succeeding autozooid, as wide as long,
surface granular and coarsely perforated, and
typically crossed by sutures; the ofifice of
ovicelled autozooids 1s broader than that of ordi-
nary autozooids, with a shallowly concave
proximal edge. Avicularia situated proximo-
lateral to orifice, directed proximo-medially; of
variable length, up to 0.3mm Jong; the smallest
with a semielliptical mandible, the largest dis-
tinctly spoon-shaped,

DISTRIBUTION

Previously from isolated localities in the
western Pacific, from the South China Sea to
Torres Strait.

Robertsonidra Osbum, 1952

Colony encrusting. Autozooids with tubercu-
late frontal wall, bordered by conspicuous mar-
ginal pores. Primary orifice with proximal sinus,
frequently with a pronounced lateral suboral
usmbo; spines present or absent. Avicularia ad-
ventitious. Ovicell hyperstomial, closed by
autozooid operculum; surface tuberculate, with
minute perforations. Vertical walls with very
small uniporous septula; basal walls typically
raised above the substratum on stout, projecting
processes.

Type species:
Kobestsan, 1908,

Schizoporella oligopus

Robertsonidra argentea (Hincks)
(Fig. 19b)

Schizoperella argentea Hincks, 1881b: 158, pl. 9, figs
6, 6a.

Rhamphostomella argentea: Harmer, 1957: 1102 [par-
tim).

|not] Robertsonidra argentea; Powell, 1967; 169,

Colony a small, yellowish-white patch;
unilaminar. Autozooids oval to broadly
hexagonal, convex, separated by distinct sutures;
0.6-0.75 X0.4-0.5mm. Primary orifice about as
long as wide; proximal edge deeply, symmetri-
cally concave, lacking condyles; two or three,
thin, evanescent spines present in early ontogeny;
frequently a short, asymmetrically-placed sub-
oral umbo, Frontal wall regularly tuberculate,

with a single row of round marginal pores,
Avicularia dimorphic, one or both types present
on a single autozooid, some autozooids without
either; lateral-suboral, acute to frontal plane,
laterally directed, with short, hooked rostrum, or
normal to frontal plane, proximo-laterally
directed, with elongate triangular rostnum.
Ovicell spherical, regularly tuberculate, with
minute perforations.

OCCURRENCE

Asingle specimen collected from Heron Island,
Stn 23, encrusting coral rock, constitutes the first
Australian record.

REMARKS

Harmer’s (1957) account of R. argentea was
based on several different species, and there is no
indication that he examined either the type
specimen (BMNH reg.no. 1899.5.1.970, loc.
‘Africa’) or material of the genus type,
Schizoporella oligopus Robertson, which he
placed in the synonymy of R. argentea, Two of
the Siboga specimens listed by Harmer
(1957:1101) have been compared with the type
specimen of R. argenfea and found to represent
two different, undescribed, species of
Robertsonidra. Specimen 399D (Stn 164, BMNH
reg.no. 1975.7.24.22) has a pear- shaped orifice
with three distal oral spines and a disto- laterally
orientated, lateral suboral avicularium, with an
elongate triangular rostrum; the autozooids are
only loosely coherent. Specimen 108F (Stn 144,
BMNH reg.no. 1975.2.24.14) has a transversely
oval orifice with a distinct U-shaped sinus, lacks
spines, and has a distally-directed, lateral-suboral
avicularium with a short, broadly triangular
rostrum. Powell's (1967) figure of ‘R. argentea'
from the southern Red Sea depicts yet another
species of Robertsonidra, R. oligopus, which is
known from the Galapagos Islands and Southern
California is almost certainly distinct from &,
argentea, and from any of the other species dis-
cussed here. The type locality of R. argenteu is
unknown, although its habitat, ‘on coral’, sug-
pests il hes somewhere on the East African coast.

Robertsonidra novella sp, nov.
(Fig. 19a)

TYPE MATERIAL
HOLotyre: GH5231), Heron Island, Stn 22, 15 Ape
1972.

262 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 19, a, Robertsonidra novella, * 60, b, Robertsonidra argentea, * 60. c,d, Schizomavella inelusa; c, X70;
d, X 140. e, Phonicosia circinata, * 70,

BRYOZOA FROM HERON ISLAND

ETYMOLOGY
Latin novella, diminutive of novus, new.

DESCRIPTION

Colony forming small, silvery, unilaminar
patches, Autozooids oval to hexagonal, strongly
convex, separated by distinct grooves; 0.5-
0.6 0.35-0.4mm. Primary orifice transversely
oval, proximal border with a short, U-shaped
sinus flanked by small condyles; one or two short,
evanescent spines present distally, their bases
persisting especially in ovicelled autozooids; a
short umbo situated lateral to the sinus in many
autozooids. Frontal wall regularly tuberculate,
glistening under low magnification, with a single
series of round marginal pores. Avicularia
sporadic, lateral suboral; rostrum elongate trian-
gular, directed proximo-laterally. In some
autozooids a second, shorter avicularium occurs
lateral to the sinus, with the rostrum acnte to
frontal plane and laterally directed. Ovicell
slightly wider than Jong, globular, the surface
tuberculate, with minute perforations between
the tubercles.

REMARKS

This species is distinguished from &. argentea
by its oval orifice, which has a distinct proximal
sinus bounded by small condyles. It was present
in three of the Heron Island samples.

Family SCHIZOPORELLIDAE Jullien, 1883

Colony encrusting or erect. Autozooids with
cryptocystidean frontal wall; evenly perforated or
with marginal pores only, Primary orifice typical-
ly with distinct proximal sinus and conspicuous
condyles; oral spines present or absent.
Avicularia adventitious and/or vicarious or inter-
zooidal; or absent. Ovicells present. Vertical
walls with basal pore chambers or mural septula.

Schizomavella Canu & Bassler, 1917

Colony encrmusting. Frontal wall of autozooid
with regularly distributed perforations, Primary
orifice with distinct sinus and well-marked con-
dyles; oral spines present or absent. Avicularia
adventitious, typically suboral, but also lateral-
oral, or frontal: dimorphic in some species,
Ovicell hyperstomial, variably perforate, closed
by autozooidal operculum. Vertical walls with
small septula.

gore species: Lepralia auriculata Hassall,
1842.

263

Schizomavella inclusa (Thomely}
(Fig. 19c¢,d)

Sechizoporella inclusa Thomely, 1906; 450,

Schizomavella inclusa: Harmer, 1957: 1028, pl. 66,

figs 4, 6,7; Winston & Heimberg, 1986: 18, figs 43,
44,

DESCRIPTION

Colony forming thin, flat encrustations.
Autozooids irregularly rectangular, flat,
separated by distinct sutures; 0,35-0,5 x 0.25-
0.35mm. Primary orifice as wide as long,
broadest distally and appearing pear-shaped, with
a deep, U-shaped proximal sinus below short.
distinct condyles. No oral spines. Frontal wall
thick and vitreous, irregularly nodular, with
numerous, small, round pores; marginal pores
larger and more distinct. Avicularium suboral,
proximally directed, 0.05mm long, with oval or
bluntly triangular rostrum, A dimorphic
avicularium, With enlarged, spatulate mandible
was described by Harmer (1957). Ovicell wader
than long, partially immersed, with numerous
small pores, most densely distributed around
edges.

DISTRIBUTION

S. inclusa has been described from scattered
localities, from Madagascar, Mauritius and Sn
Lanka, to Indonesia, New Guinea and New
Britain, Harmer (1957) recorded a specimen from
the Red Sea, but it has not been recorded pre-
viously, with certainty, from Australian seas.

Arthropoma Leyvinsen, 1909

Colony encrusting. Autozooid with evenly per-
forated frontal wall. Primary orifice with narrow
medio-proximal sinus; operculum articulated, No
oral spines. Avicularia absent. Ovicell
prominent, imperforate, closed by autozooid
operculum, Vertical walls with large, multi-
porous septula, Ancestrula identical to later
autozooids.

Type species: Flustra cecilii Audowin, 1826.

Arthropoma cecilii (Audouin)
(Fig. 16f}

Flustra cecilii Audouin, 1826; 239; Savigny pl. 8, fig.
3.

Arthropoma eecilii: Harmer, 1957; 1001, pl. 72, fig,
23; text- fig. 108; Gordon, 1984; 85, pl. 30, fig, A.

264

DESCRIPTION

Colony forming small, glistening, white
patches; usually unilaminar. Autozooids lightly
calcified, strongly convex, with densely per-
forated frontal wall; commonly 0.5-0.7X
0.5mm. Primary onfice slightly wider than long,
D-shaped, with a very narrow, medio-proximal
sinus. No avicularia, although Harmer (1957)
reports 4 polymorph with an operculum twice the
size of that of ordinary autozooids. Ovicell
prominent, convex, smoothly calcified, opaque
white.

DISTRIBUTION

A. cecilii is widespread and common in all
shallow, tropical, subtropical and warm
temperate seas. It has been reported previously
from reef habitats in the Caribbean, the Indian
Ocean, the East Indies and the GBR.

Phonicosia Jullien, 1888

Colony encrusting. Autozooid frontal wall with
conspicuous marginal pores, and sometimes with
additional pores proximo-lateral to orifice, but
with central area usually imperforate. Primary
orifice with proximal sinus; operculum articu-
lated; distal oral spines present, Adventitious
avicularia occur sporadically. Ovicell prominent;
ectooecium membranous; entooecium calcified;
imperforate, except for small basal pores around
periphery; not closed by autozooid operculum.
Basal pore chambers present.

Type species: Phonicosia jausseaumet Jullien,
1888 [= Lepralia circinata MacGillivray, 1869}.

Phonicosia circinata (MacGillivray)
(Fig. 19¢)

Lepralia circinata MacGillivray, 1869: 134.
Arthropoma circinatum: Harmer, 1957: 1003, pl. 72,
figs 29-30; Hayward & Cook, 1983: 69, fig. 17c.
Phonicosia circinata; Gordon, 1984; 87, pl. 30, figs

E,F; 1989: 46,

DESCRIPTION

This distinctive species is readily recognized by
its D-shaped orifice, with slit-like sinus and
broad, denticulate condyles, surrounded by eight
or nine spines,

DISTRIBUTION

P. circinata has a wide geographical distribu-
tion, ranging throughout the western Pacific
Ocean, from Japan to New Zealand, eastwards to

MEMOIRS OF THE QUEENSLAND MUSEUM

South Africa, Tristan da Cunha, the Patagonian
Shelf and Tierra del Fuego; also at Marion Island
and Kerguelen.

REMARKS

In some populations a large adventitious
avicularium occurs proximal to the sinus, and the
density and distribution of frontal wall pores also
varies greatly. However, these variations may be
observed over a relatively small geographic dis-
tance, and perhaps even within single popula-
tions.

Escharina Milne-Edwards, 1836

Colony enerusting or erect. Frontal wall of
autozooids with distinct marginal pores, and
evenly scattered, minute perforations. Primary
orifice with well marked sinus; oral spines
present. Avicularia adventitious or interzooidal.
Ovicell hyperstomial, imperforate, closed by
autozooid operculum. Small basal pore chambers
present.

Type species: Eschara vulgaris Moll, 1803.

Escharina pesanseris (Smitt)

Hippethea pesanseris Smitt, 1873: 43, pl. 7, figs 159,
160.

Escharina pesanseris: Harmer, 1957: 998, pl, 67, figs
12-14,18,19; Gordon, 1984: 84, pi. 29, figs A, B.

DESCRIPTION

Colony encrusting, forming small rounded
patches; rather inconspicuous, Autozooids oval,
convex; about 0.8 X 0.5mm, frontal wall finely
granular, fine perforations revealed only by high
magnification. Primary orifice longer than wide,
with a narrow, slit-like sinus between broad con-
dyles; with eight or nine stout distal oral spines,
and a low proximal peristomial rim. Avicularia
paired, lateral to orifice, with distinctive, duck's
foot-shaped mandibles distally directed. Ovicell
relatively small, wider than long, with a
transverse frontal ridge; six spines only in
ovicelled autozooids.

DISTRIBUTION
E. pesanseris has a circumtropical distribution
in shallow water, and occurs commonly in reef
habitats.
Stylopoma Canu & Bassler, 1920

Colony encrusting, multilaminar, developing

BRYOZOA FROM HERON ISLAND

extensive nodular growths. Frontal wall of
autozooids evenly perforated by small pores, with
ii marginal series of larger pores. Primary orifice
with well developed condyles and deep proximal
sinus; no oral spines. Avicularia adventitious,
small, and vicarious, large. Ovicell globular,
prominent, perforated; overhanging and even-
tually occluding onfice of brooding autozooid.
Vertical walls with numerous, small, recessed
septula.

Type species: Eschara spongites Pallas, 1766.

The genus Stylepoma has a pantropical dis-
tribution in shallow, coastal waters, and most
species described seem to be commonly found in
reef habitats. Most are brightly pigmented.

Stylopoma duboisii (Audouin)
(Fig. 20a,b)

Flustraduboisii Audouin, 1826; 239; 1828: 66; Savig-
ny pl. 8, figs 41, 42.

Styvlopoma duboisit: Harmer, 1957; 1033, pl. 74, figs
1-7; Winston & Heimberg, 1986; 20. figs 45-57.

DESCRIPTION

Autozooids irregularly polygonal, rather flat,
separated by distinct sutures; 0.5-0.60,35-
0.45mm. Primary orifice wider than long, D-
shaped, the straight proximal border with a slit-
like median sinus; condyles low and broad, oc-
cupying whole of proximal border on each side
of sinus, distal edges finely toothed. Frontal wail
densely perforated by small round pores, and with
asingle series of larger marginal pores; becoming
coarsely nodular in later ontogeny. Typically, a
single, small avicularium proximo-lateral to
orifice, acute to frontal plane, disto-laterally
directed; 0.05mm long, with short, triangular
mandible. Additional avicularia may be budded
in later ontogeny, especially in brooding
autozooids, and most usually in proximity to the
orifice. Vicarious avicularia infrequent, as large
as autozooids, with broadly spatulate. mandible.
Ovicells globular, prominent, 0.5mm diameter,
occupying the frontal surface of several adjacent
autozooids; densely porous, with the aperture
overhanging and obscuring the orifice of the
maternal autozooid,

DISTRIBUTION

This species is commen in shallow waters
throughout Malaysia and Indonesia, and on the
coasts of New Guinea and Queensland. It ranges
westwards to Sri Lanka and the Red Sea, and was
reported from West Africa by Cook (1968b).

Seven of the Heron Island samples contained
large colonies of S. dubvisii.

Stylopoma bora Livingstone
(Fig

Schizoporella viridis var. thornelyi Livingstone, 1926:
84, pl. 8, figs 8, 9-

DESCRIPTION

Autozooids rectangular to polygonal, convex,
separated by distinct sutures; 0.6-0.8%0.35-
0.5mm. Primary orifice about as wide as long,
proximal border with 4 broad V-shaped sinus
occupying about half its width; condyles rectan-
gular, distinct. Frontal wall densely perforated by
stall, evenly distributed pores, each ina shallow
pit. A single, small adventitious aviculanum
present on many auitozooids, lateral to sinus;
rostrum 0.08mm long, acute to frontal plane,
distally directed, supporting a short, triangular
mandible, Vicarious avicularia sporadic, slightly
smaller than autozooids, with waisted, spatulate
rostrum.

DISTRIBUTION

This species is known with certainty only from
the Great Barrier Reef, although part of the In-
donesian material described by Harmer (1957) as
§. parviporasum Canu & Bassler may represent
8, thornelyae,

Stvlopoma viride (Thomely)
(Fig. 20d)

Schizoporella viridis Thomely, 1905: 116, fig. 3.
Stylopoma viride: Harmer, 1957: 1036, pl. 74, figs
8-13; Winston & Heimberg, 1986; 21, figs 48-50.

DESCRIPTION

Autozooids rectangular. diamond-shaped or
polygonal, convex, separated by distinct sutures;
0.8-1.2* 0.5-0.6mm_ Pnmary orifice orbicular,
withadeep V-shaped sinus occupying two-thirds
of the proximal width: condyles small, knob-like.
Frontal wall densely perforated by small, round
pores, the marginal series slightly larger and more
distinct; calcification thick, forming taised,
nodular ridges between the pores. Majority of
autozooids with characteristic disto-lateral ad-
ventitious avicularium, rostrum slender, pointed,
035mm long, extending disto-medially above
the orifice. A smaller suboral aviculanum present
in many aulozooids, adjacent ta sinlis, with yary-
ing orientation. Vicarious avicularia identical to

266 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 20, a,b, Stylopoma duboisii; a, X 80; b, X70. ¢, Stylopoma thornelyae, * 80, d, Stylapoma viride, 80.
cf, Cigclisula areolata, €, autozooids, * 50; f, two ovicelled zooids, * 50,

BRYOZOA FROM HERON ISLAND

disto-lateral adventitious avicularia; rare. Ovicell
globose, us large as an autozooid, with identical
calcification and perforation, and with an inde-
pendent, frontally situated aperture,

REMARKS

The massive, multilaminar colonies of §. viride
have a characteristic green coloration, muted in
living colonies by a thick, greyish-brown frontal
cuticle. The pigment is contained within the cal-
cification and persists after the death of the
colony.

DISTRIBUTION

This species 1s known from the Red Sea, Sri
Lanka and other localities in the Indian Ocean; it
has been widely reported from Malaysia, In-
donesia, New Guinea and the Philippines, and is
distributed on north and northeast coasts of
Australia, and along the Great Barrier Reef.

Family STOMACHETOSELLIDAE Canu &
Bassler, 1917

Colony encnisting, forming a thick, multi-
laminar sheet; orerect, as folded, bilaminar plates
attached by an encrusting base. Autozooids thick-
ly calcified; frontal wall with marginal and fron-
tal pores. Primary orifice sinuate or bell-shaped.
encircled by a deep, thick peristome. Ayicularia
adventitious and/or vicarious; or absent. Ovicells
present, typically with few, large, frontal lacunae.

Cigelisula Canu & Bassler, 1927

Colony encrusting, or erect, unilaminar or mul-
lilaminar, often nodular or massive. Frontal wall
of autozooid with marginal perforations (same-
times in double or triple series), with stout inter-
areolar ridges developing in later ontogeny;
central area of frontal wall imperforate. Primary
orifice more or less bell-shaped, the proximal
border shallowly concaye between distinct con-
dyles. No oral spines. A thickened peristomial
tim developing in later ontogeny, charac-
leristically incorporating a proximal suboral
avicularium, eccentrically placed and laterally
directed. Additional small, adventitious
avicularia may be present elsewhere on
autozooid; in some species large, vicarious
aviculatia also occur. Ovicell characteristic:
prominent, grossly inflated, sutured, with a few,
large, frontal lacunae,

Type species: Escharoides occlusa Busk, 1884.

Cigclisula areolata (Kirkpatrick)
{Fig. 20e,f)

Lepralia occlusa var. areolala Kirkpatrick, 189(a:
618, pl. 16, figs 7, 7a.

Porella areolata: Livingstone, 1926: 91, pl. 8, fig. 4.

Cigclisula areolata: Hastings, 1932; 434,

DESCRIPTION

Colony forming thick, extensive multilaminar
sheets. Autozooids oyal to hexagonal, strongly
convex, thickly calcified, separated by distinct
sutures; commonly 0.6-0.7 *0.4-0.5mm. Primary
orifice as wide as long, bell-shaped, with concave
proximal border; condyles small, rounded. Frontal
calcification coarsely granular, becoming dis-
tinctly nodular in later ontogeny, with a double or
triple series of marginal pores, but the central area
of the frontal wall imperforate, As the frontal wall
thickens, stout ridges develop peripherally, even-
tually forming a series of large, deep areolae, eacti
of which may enclose two or more of the original
pores. Orifice encircled by a thick, peristomiil
rim; a single, smal] avicularium proximo-medial-
ly; its rostrum perpendicular to frontal plane and
facing laterally, immersed in peristomial cal-
cification, but with tip of rostrum hooked and
distinct. Large vicarious avicularia occur sporadi-
cally. Ovicell wider than long, almost as large as
autozooid, globular and prominent, with the aper-
ture overhanging the autozooid orifice; bordered
by Jarge ureolae, and with four to eight large
lacunae frontally.

DISTRIBUTION

This species is known only from Torres Swait
and the Great Barrier Reef, At Heron Island large
colonies were present in two samples from reef
crest locations.

Family SMITTINIDAE Levinsen, 1909

Colony encrusting or efect. Autozooids with
cryptocystidean frontal wall, evenly perforated or
with marginal perforations only. Primary orifice
with or without a median proximal denticle
(lyrula) and proximo-lateral condyles; oral spines
present or absent. Avicularia adventitious, or ab-
sent. Ovicell prominent, perforate or imperforate,

Smittoidea Osburn, 1952
Frontal wall of autozovids with marginal pores

only. Avicularium situated medially on the fron-
tal wall, immediately proximal to the orifice,

268

often partly enveloped by the peristome; typically

proximally directed. Ovicell prominent, evenly

perforated. Vertical walls with mural septula.
Type species: Smittoidea prolifica Osburn,
952.

Smittoidea pacifica Soule & Soule
(Fig. 24e,f)

Smittoidea pacifica Soule & Soule, 1973: 380, figs I,
F-H.

DESCRIPTION

Colony a small, silvery-white, unilaminar
patch. Autozooids oval to hexagonal, flat or
slightly convex, separated by indistinct sutures;
0.4-0.5 X 0.2-0.3mm. Primary orifice wider than
long; lyrula anvil-shaped, broad and deep, oc-
cupying entire width of orifice; condyles short
and blunt, indistinct. No oral spines. Peristome
completely encircling orifice, its edge thickened
and slightly lobed in later ontogeny; with a con-
spicuous U-shaped notch medio-proximally, the
inner edges of which may be thickened to define
a descending groove. Frontal wall coarsely
nodular, bordered by large and conspicuous mar-
ginal pores. A single suboral avicularium on each
autozooid, directed proximally or proximo-
laterally; rostrum elongate, finely tapered,
rounded at tip, its edges sometimes finely den-
ticulate; crossbar thickened, the proximal opesia
circular. Oyicell slightly wider than long, frontal
surface with numerous small pores; developing a
coarsely nodular ooecial cover, which extends to
the distal edge of the peristome, leaving only the
central frontal area of the ovicell exposed.

REMARKS

Soule & Soule (1973) founded this species on
specimens collected from Hawaii. They included
Osburn’s (1952) record of S. reticulata (J. Mac-
gillivray) from the Galapagos Islands in the
synonymy of S. pacifica, while noting that one
zooid in Osburn’s specimen bore six oral spines.
The Heron Island specimens here described have
been compared with Hawaiian paratype material
of S. pacifica, which they match exactly.

S. reticulata is distributed in the eastern Atlan-
tic region from the Mediterranean to the Barents
Sea, and almost certainly does not occur in the
Pacific. Osburn’s Galapagos material does not
belong to S. reticulata, and is possibly not S.
pacifica either.

MEMOIRS OF THE QUEENSLAND MUSEUM

Parasmittina Osburn, 1952

Colony encrusting, unilaminar or multilaminar.
Frontal wall of autozooid with marginal pores
only. Primary orifice with medio-proximal lyrula
and paired lateral condyles; oral spines usually
present; typically with a well developed peris-
tome. Avicularia adventitious and interzooidal,
frequently polymorphic, often numerous; charac-
teristically, one or more occur lateral to the
orifice. Ovicell hyperstomial, prominent, with
frontal pores; not closed by autozooid operculum.
Small basal pore chambers present.

Type species: Lepralia jeffreysi Norman, 1876.

Species of Parasmittina are common, and often
abundant, in shallow reef habitats. Species diver-
sity is usually high (e.g., Soule & Soule, 1973;
Hayward, 1988) and it is probable that many
tropical species remain to be recognized.

Parasmittina hastingsae Soule & Soule
(Fig. 21a-c)

Parasmittina hastingsae Soule & Soule, 1973: 417,
figs 9C,D.

Parasmittina hastingsae: Winston & Heimberg, 1986:
26, figs 59-63.

DESCRIPTION

Colony forming broad, yellowish, multi-
laminar sheets. Autozooids oval, hexagonal or
irregular, convex, separated by distinct sutures;
commonly 0.35-0.5 X c.0.25mm. Primary orifice
slightly wider than long, rounded; lyrula occupy-
ing about half proximal orifice width, with con-
vex edge and bluntly pointed corners; condyles
acute, distinct. Two or three (rarely four) short,
distal oral spines present in early ontogeny; peris-
tome developed as paired lateral folds, with a
deep, medio-proximal sinus between; erect and
slightly flared, enclosing distal border of orifice
in later ontogeny. Frontal wall of autozooid finely
nodular, with a single series of round marginal
pores. As calcification thickens in later ontogeny,
the boundaries between individual autozooids be-
come increasingly obscure, and the marginal
pores much more irregular in outline. Avicularia
of three main types, all becoming more frequent
in later ontogeny: most characteristically a short,
oval type, proximo-lateral to orifice, proximally
directed; less frequently, a larger type with
rostrum elongate, tapered, rounded distally and
serrated along its edge, directed proximally;
finally, a large interzooidal type with rostrum
tapered, rounded and slightly flared distally, its

BRYOZOA FROM HERON ISLAND 269

FIG. 21. a-c, Parasmittina hastingsae; a, autozooids and a giant avicularium, 60; b, part of the Hawaiian
paratype material, X55; c, primary orifice, x 250. d-f, Parasmittina galerita; d, primary orifice, 220; e,
autozooids and a giant avicularium, x 60; f, ovicelled autozooids, x55.

edge serrated, distally directed. Ovicell more or ing onto ils frontal surface, forming a complete
less spherical, with 20-30 thickly rimmed, rela-~ tube, distal portion of which becoming immersed
tively large, frontal pores; the peristome extend- below an encroaching ovecial cover.

REMARKS

This species was especially abundant in the
Heron Island collections. [tis distinguished from
other species of the genus by the combination of
small, oval avicularia (proximally directed when
lateral to orifice), and large avicularia with
tapered, serrated rostra, which are proximally
directed when lateral to the autozooid orifice, but
distally directed when interzooidal. P_ hastingsae
was. described from Hawaii by Soule & Soule
(1973) and subsequently reported from Komodo,
Indonesia, by Winston & Heimberg (1986).

Parasmittina galerita sp. nov.

(Fig. 21d-f)

Smittina parseyalii: Harmer, 1957: 941 [part], pl. 65,
fip. 5 [only].

TYPE MATERIAL
Ho.otyPe: GH5231, Heron Island, Stn 16, 13 Apr
1972,

ETYMOLOGY
Latin galerum, a helmet.

DESCRIPTION

Colony a multilaminar sheet. Autozooids
small, rectangular to irregularly polygonal, con-
vex, separated by thickened sutures. Primary
orifice about as wide as long; lyrula short, oc-
cupying about half total width of orifice; its edge
slightly curyed; condyles small, rounded,
downcurved, with finely denticulate edges; a
single, inconspicuous, mid-~distal oral spine
present in early ontogeny. Penstame widely open
proximally, at first consisting simply of paired,
subtriangular lateral lappets, extending to enclose
the distal border of the onfice in later ontogeny.
Frontal wall coarsely nodular, bordered by large,
and conspicuous marginal pores. Avicularia fre-
quent, polymorphic: each autozooid typically
with two, lateral suboral avicularia, the first short,
0.1mm long. oval or shoe-shaped, proximally
directed, the second longer, 0,15mm, with acute
tnangular rostrum directed disto-medially onto
the peristome rim; giant avicularia sporadic, as
long as autozooids, the rostrum narrow, spatulate
distally, with smooth tim. Ovicel] relatively
small, wider than long, with about 20 smal] pores
frontally; developing a nodular, peripheral ooe-
cial cover, and a distinct frontal lip where the
peristome traverses it.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.51+0.06;

MEMOIRS OF THE QUEENSLAND MUSEUM

width 0,.3340.04; orifice length 0.10+0.007;
width 0.10+0.005,

REMARKS

The specimens assigned to Cellepora par-
sevalii Audouin by Harmer (1957) encompass at
least two species. The Red Sea specimen listed by
him (Ghardaga, 4D) is quite distinct from that
from Makassar Strait (381, Siboga Sin 77), which
is the same species as that described here from
Heron Island. P. parsevaliformis Soule & Soule
also has just a single mid-distal oral spine, and a
similar array of avicularia, with a short oval type
proximally directed, and an acute triangular type
directed disto-medially onto the peristome. How-
ever, P. parsevaliformis differs from P. galerita
in its longer orifice, narrower sinus, with distinct-
ly cusped corners, and in its ovicell, which is
smooth frontally, with a peripheral band of rela-
tively large pores.

Parasmittina latiavicularia (Kirkpatrick)
(Fig. 22a-c)

Smittia latiavicularia Kirkpatrick, 1888: 80, pl. 10, fig.
3.

Parasmittina latiavicularia; Hayward, 1988; 301, fig,
5a,

Parasminina leviavicularia Soule & Soule, 1973: 409,
figs 7E-G.

DESCRIPTION

Colony a small, irregular, multilaminar patch.
Autozooids oval to hexagonal, convex, separated
by distinct sutures; 0.45-0.7 x 0.35-0.45mm,
Primary orifice Jonger than wide, distinctly oval,
the distal border with a few indistinct denticula-
tions medially; lyrula occupying about half width
of proximal edge. slender, with curved edges and
cusped corners; condyles oval, prominent,
downcurved, with finely denticulate edges. One
ortwo slender, distinctoral spines present in early
ontogeny. Peristome consisting almost entirely of
paired lateral lappets; widely open proximally.
Frontal wall nodular, bordered by a single or
double series of large pores. Avicularia
numerous: characteristically, single or paired
lateral oral, with acute triangular rostrum disto-
medially directed, ascending the peristomial lap-
pet; either or both replaced by a short, oval or
shoe-shaped avicularium, with proximally
directed rostrum; in some autozooids the trian-
gular avicularium may be medio-proximally
situated, with the rostrum distally directed, along
the edge of the peristome; in older parts of the

BRYOZOA FROM HERON ISLAND 271

FIG, 22. a-c, Parasmittina latiavicularia; a, primary orifice, X 240; b, autozooids and a giant avicularium, * 90;
c, ovicelled autozooids, X80. d-f, Parasmittina onychorrhyncha; d, ovicelled autozooids, X 60; e, primary
orifice, x 400; f, autozooids and giant avicularia, x90.

272

colony the small oval avicularia may proliferate
along the margins of the autozooids. Giant
avicularia rare, occasionally replacing the lateral
suboral avicularium, rostrum shorter than
autozooid, broadened or rather angular distally,
with an irregularly cusped rim. Ovicell spherical,
prominent, with about 20 small frontal pores;
becoming coarsely nodular, developing a distinct
frontal lip above the secondary orifice, and often
with a well developed umbo.

DISTRIBUTION

This rather inconspicuous species has been pre-
viously reported only from Mauritius and Hawaii.
It was present in two of the Heron Island samples.

Parasmittina onychorrhyncha sp. nov.
(Fig. 22d-f)

TYPE MATERIAL
Ho.otyPe: GH5232, Heron Island, Stn 25, 18 Apr
1972.

ETYMOLOGY
Greek Onychos, a claw, and rhynchos, a snout.

DESCRIPTION

Colony an irregular, multilaminar sheet.
Autozooids rectangular, hexagonal or oval at
growing edges, but often irregularly polygonal in
frontally budded laminae; convex, separated by
distinct sutures. Primary orifice wider than long;
lyrula short and broad, with straight edge, oc-
cupying about half proximal border; condyles
thin, pointed, downcurved; typically with a
single, slender, mid-distal oral spine. Peristome
widely open proximally, consisting largely of
paired, subtriangular lateral lappets. Frontal wall
distinctly nodular, bordered by conspicuous mar-
ginal pores. Avicularia sporadic, frequent on
some parts of colony, rare elsewhere, polymor-
phic: most often short, shoe-shaped, about 0.1mm
long, proximo-lateral to orifice and proximally
directed; less often, about 0.2mm long, distal or
disto-lateral to orifice, narrowly spatulate or
bluntly triangular, directed proximo-laterally; oc-
casionally gigantic, occupying half frontal area of
autozooid, the rostrum broadly spatulate, deeply
cupped, with a coarsely toothed distal rim.
Ovicell globose, prominent, closely perforated
frontally by numerous small, irregular pores;
developing a granular ooecial cover, often slight-
ly raised and frilled where it borders the perforate
frontal region of the ovicell.

Measurements (means and standard deviations

MEMOIRS OF THE QUEENSLAND MUSEUM

of 20 values, mm): autozooid length 0.59+0.05;
width 0.33+0.05; orifice length 0.10+0.007;
width 0.12+0.006.

REMARKS

P. delicatula (Busk, 1884), described from
Hawaii, and P. fistulata (Harmer, 1957), from
Indonesia, are very similar species which can be
separated principally by the morphology of the
primary orifice and the occurrence of particular
types of avicularia. These features also serve to
distinguish both species from P. onychor-
rhyncha,

Parasmittina serrula Soule & Soule
(Fig. 23e,f, 24a)

Parasmittina serrula Soule & Soule, 1973: 386, figs
3D-F.

Parasmittina serrula: Gordon, 1984: 96, pl. 35, figs B,
OF

DESCRIPTION

Colony forming small, glistening white
patches, rarely more than 5mm in diameter.
Autozooids oval to hexagonal, small, convex,
separated by deep grooves; 0.3-0.5 Xc.0.25mm.
Primary orifice wider than long, with denticulate
distal border; lyrula occupying about half width
of proximal border, with slightly curved edge and
cusped corners; condyles downcurved, rounded,
with very finely denticulate edges. Two to four
long, stout distal oral spines; obliterated in brood-
ing autozooids but persisting into late ontogeny
in others. Peristome most prominent laterally and
proximally, with a well-marked medio-proximal
notch, often flanked on each side by a stout, erect
process. Frontal wall finely nodular, bordered by
large and distinct marginal pores. Lateral suboral
avicularia typically paired, equally-sized; the
rostrum slender, parallel-sided, rounded at tip,
with very fine serrations along its distal edges;
characteristically inclined so that the rostrum lies
on its side. Giant avicularia occasional, as long as
autozooid, with the rostrum slightly flared distal-
ly and very coarsely serrated. Ovicell spherical,
smooth, with 15-20 regularly dispersed pores;
peristome encroaches onto frontal surface of
ovicell in later ontogeny, but an ooecial cover
does not seem to develop.

DISTRIBUTION

Soule & Soule (1973) found this to be the
commonest species of Parasmittina on Hawaiian
reefs. It was subsequently described and figured

BRYOZOA FROM HERON ISLAND

from the Kermadec Islands by Gordon (1984) and
from Belize by Winston (1984). It occurred in
three of the Heron Island samples but has not
previously been reported from Australia.

Parasmittina turbula sp. nov.
(Fig. 23a-d)

TYPE MATERIAL
HoLotTyPe: GH5233, Heron Island, Sin 25, 18 Apr
1972.

ETYMOLOGY
Latin terba, disorder.

DESCRIPTION

Colony encrusting, multilaminar, developing
an irregular, mamillate form, the later generations
of autozooids lacking regular orientation.
Autozooids oval to irregular. convex, separated
by distinct sutures; frontal wall irregularly
nodular, bordered by large marginal pores, with
thickened struts between; 0.5 X 0.3mm. Primary
orifice about as wide as long (0,12mm); lyrula
occupying about one-third of proximal width,
short, rectangular, with a gently curved edge;
condyles oval, downcurved, with finely crenulate
edges. A single disto-medial spine present; peris-
tome complete, initially with a delicately flared
lip, a deep distal embayment accommodating the
spine, and a shallow mid-proximal notch flanked
by thickened internal ridges. With ontogenetic
thickening the peristome becomes deeper, the
uistal spine ts obliterated and the proximal notch
accentuated. Aviculana polymorphic, numerous;
one or two lateral to the peristome in newly
budded autozooids, usually narrow, parallel-
sided, from 0.2-0.3mm long, proximally directed,
occasionally shorter, triangular and disto-medial-
ly directed; additional avicularia develop in later
ontogeny, typically clustered around the peris-
tome, and directed towards the orifice: gigantic
avicularia present in a few autozooids, laterally
situated and proximally directed, with flared,
slightly spatlate mandible. Gvicell spherical,
with numerous fine perforations, rapidly en-
veloped by secondary calcification; with a num-
ber of smal] ayicularia developing on its frontal
surface, directed towards the autozooidal onfice,

REMARKS

The slender, parallel-sided ayicularia recall
those of P. areolata Soule & Soule. P. turbula is
distinguished from that species by its single, dis-
tal oral spine, the morphology of the lyrula and

273

condyles, and by the clusters of medially directed
avicularia developed around the penstome in
later ontogeny.

Plearocodonellina Soule & Soule, 1973

Colony encrusting. Autozooid frontal wall with
marginal] pores only. Primary orifice with strong-
ly developed condyles; proximal border concave,
developed as a broad or shallow sinus. Peristome
developing in early ontogeny, completely sur-
rounding orifice, its inner edge often thickened
medio-proximally and forming a secondary
sinus. Distal oral spines present. Avicularia a
venlilious, dimorphic, typically situated
proximo-lateral to orifice, Ovicell with small,
scattered pores, Vertical walls with small
uniporous septula.

Type species: Plewracodonellina lahainae
Soule & Soule, 1973.

Pleurocodonellina signata (Waters)
(Fig. 24b)

Smittia signata Waters, 1889: 17, pL 3, figs 4-6.

Smittina signata: Harmer, 1957: 928, pl. 63, figs 27-29.

Parasmittina signata: Winston & Heimberg, 1986: 21,
figs 51-54.

DESCRIPTION

Colony forming extensive, pmkish brown,
unilaminar sheets. Autozooids commonly
0.5 0.3mm, with a finely neduJar frontal wall,
bordered by a single series of large and con-
spicuous areolae, Primary orifice wider than
long, with broad, flat condyles delimiting a nar-
row, rounded, proximal sinus; one to three distal
oral spines present in early ontogeny. Peristome
thin, erect, completely surrounding onfice und
obliterating spine bases, Most sutozooids with a
single avicularium proximo-lateral to orifice;
rostrum slender, elongate triangular, proximally
directed. Ovicell recumbent on succeeding
autozooid, wider than long; frontal surface slight-
ly flattened, with small distal pores.

REMARKS

Cheetham & Sandberg (1964) tentatively as-
signed this species to Rimulosroma Vigneaux,
while noting that the published accounts of both
the genus { Vigneaux, 1949) and its Miocene type
species (Duvergier, 1921) were inadequate for
detailed morphological comparisons. Soule &
Soule (1973) suggested that Smittia signate
Waters might be placed in their genus

274 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 23. a-d, Parasmittina turbula; a, X70; b, *95; c, primary orifice, X 340; d, giant avicularia, x 60. e,f,
Parasmittina serrula; e, primary orifice, * 350; f, * 80.

FIG. 24. a, Parasmittina serrula, X130.b, Pleurocodonellina signata, * 90. c,d, Pleurocodonellina clavicula;
c, X60; d, * 60. e,f, Smittoidea pacifica; e, X80; f, * 155.

275

BRYOZOA FROM HERON ISLAND

Pleurecodonetlina, and that suggestion is fol-
lowed here. In all morphological characters S.
signata conforms to the diagnosis of
Pleurocodonellina. In particular, the flaring
peristomial rim which obliterates the distal spines
present in early ontogeny is closely similar to that
of the type species, and only a more pronounced
development of the paired lateral condyles: is
necessary to produce the narrow, rounded sinus
seen in P, signata,

DISTRIBUTION

This species is practically circumtropical in
distribution. It ranges throughout the Indo-West-
Pacific realm, and has been reported from
Mauritius, East Africa, West Africa, and the
Caribbean. It is quite common in shallow reef
habitats, and at Heron Island occurred in five of
the samples studied.

Pleurocodonellina clayicula sp. noy,
(Fig. 24¢,d)

TyPé MATERIAL
HoLotyre; GH5234, Heron Iskand, Stn 12, 12 Apr
1972,

ETYMOLOGY
Latin clavicula, diminutive of clavis, a key.

DESCRIPTION _

Colony a broad, encrusting, multilaminar sheet.
Autozooids rectangular, hexagonal or irregularly
polygonal, rather flat, separated by distinct, thick-
ened sutures. Frontal wall thickly calcified, coar-
sely nodular, bordered by a single series of large
marginal pores. Primary orifice longer than wide,
the proximal third comprising a deep, browd, U-
shaped sinus, delimited by flat, rounded trian-
gular condyles; encircled by a thin, low,
peristomial rim. No oral spines, Avicularia
present lateral to orifice in newly budded
autozooids, later developed elsewhere along mar-
gins of autozooids; slender, parallel-sided,
rounded distally and proximally; not infrequently
a giant avicularium present, extending down
whole length of autozooid, the rostrum 0.4mm
long, broadly spoon-shaped, deeply cupped dis-
tally, with a coarsely denticulate rim. Ovicell
prominent, globular, wider than long, witha cres-
centic series of irregular pores, developing a coar-
sely nodular ooecial cover.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.55+0.05;

MEMOIRS OF THE QUEENSLAND MUSEUM

width 0.37+0.04; orifice length 0.11+0.008;
width 0.1040,0).

REMARKS

The sinuate orifice, encircling peristomial rim,
lateral suboral avicularia and perforate ovicell all
indicate that this undescribed species is ap-
propriately placed in Pleurocodonellina. A single
large colony was present in sample 12, from the
reef flat west of the cay.

Family PETRALIELLIDAE Harmer, 1957

Colony encrusting; or forming erect, laminate
plates.. Autozooids with perforated frontal wall,
Primary orifice with or without oral spines and
condyles. Avicularia adventitious, often as-
sociated with orifice. Ovicell with smal! frontal
pores. Vertical and basal walls with multiporous
septula.

Sinupetraliefla Stach, 1936

Colony enctusting, or erect and bilaminar.
Aulozooids with regularly perforate, cryptocys-
tidean frontal wall. Primary orifice with asym-
metrically developed condyles, defining a small
sinus; bordered proximally by a low peristome
incorporating a small avicularium. No oral
spines. Frontal adventitious avicularia present.
Ovicell with entooecium finely perforated, ec-
tooecium largely membranous, developing
patches or basal struts of calcification in later
ontogeny. Vertical walls with multiporous sep-
tula.

Type species: Petralia litoralis Livingstone in
Hastings, 1932.

Sinupetraliella litoralis (Livingstone
in Hastings) (Fig. 25a)

Petralia litoralis Livingstone MS, Hastings, 19232:
438, pl. 1, fig. A, text-fig. 15.

Stnupetraliella litoralis: Harmer, 1957: 706, pl. 45, figs
11-15.

DESCRIPTION

Colony developing broad, loosely encrusting,
unilaminar sheets; deep brown in colour.
Autozooids rectangular to hexagonal, convex,
separated by thin sutures; 0.8-0.9 x 0.4-0.6mm.
Frontal wall thickly calcified, densely perforated
by large, round pores. Primary orifice about as
wide as long, with a pair of pointed condyles,
asymmetrically developed, delimiting an oval,

BRYOZOA FROM HERON ISLAND

proximal sinus. Peristome a low rim around
proximal and lateral border of orifice, incorporat-
ing a small avicularium, 0.1mm long, transverse-
ly orientated, its plane perpendicular to orifice,
situated immediately above the sinus; in later
ontogeny the avicularium partly hidden by a stout
mucro developed above it. Frontal adventitious
avicularia sparsely developed, usually along mar-
gins of autozooids; rostrum slender, elongate
oval, variably orientated. Ovicell slightly wider
than long, partly overhanging autozooid orifice:
entooecium finely perforate, appearing dimpled,
ectooecial calcification limited to a narrow,
peripheral band and a few radiating struts.

DISTRIBUTION

S. litoralis was described from the Low Isles
(Hastings, 1932) and reported by Harmer (1957)
from Celebes. Large colonies were present in
three of the Heron Island samples.

Family CREPIDACANTHIDAE Levinsen,
1909

Colony encrusting. Frontal wall of autozooids
with marginal pores and peripheral spines.
Orifice with convex or distinctly rectangular
proximal edge, appearing trilobed. Avicularia ad-
yventitious, with setiform mandibles. Ovicell hy-
perstomial, with frontal pores. Vertical walls with
small basal pore chambers. Ancestrula tatiform.

Crepidacantha Levinsen, 1909

Colony encrusting, unilaminar. Autozooid
frontal wall with marginal pores only; primary
urifice contracted laterally, with pronounced
lateral condyles, the proximal edge frequently
concave, and thus appearing tifoliate; no oral
spines. Avicularia adventitious, usually paired,
lateral to orifice; mandible setiform. Ovicell hy-
perstomial, globular, closed by autozooid oper-
culum; with an uncalcified area of ectooecium
frontally, bordered by small pores in the en-
tooecium. Small basal pore chambers present,
with intervening spinous processes visible in
autozooids at the growing margin. Ancestrula
tauform.

Type species: Flustra poissonii Audouin, 1826.

Crepidacantha crinispina Levinsen
(Fig. 25c)

Crepidacantha poissoni var. crinispina Levinsen,
1909: 266, text- figs.1-3, 5, 6.

Crepidacantha crinispina: Brown, 1954; 248, fig. LC:
Gordon, 1984: 100, pl. 37, fig. B.

DESCRIPTION

Colony forming broad, irregular sheets.
Autozooids oval to hexagonal, convex, separated
by deep grooves: 0.35-0.50 0.25-0.45mm-
Frontal wall finely granular, with indistinct mar-
ginal pores, Primary orifice longer than wide,
with conspicuous lateral processes, appearing
distinctly trifoliate. Avicularia paired, situated
level with proximal comers of orifice; rostra oval,
about 0,05mm long, directed proximally or
proximo-medially. Ovicell about as wide as long,
entooecium entirely exposed frontally, with only
a peripheral band of ectooecium.

DISTRIBUTION

C. crinispina ranges in the western Pacific from
New Zealand to the Philippines, and westwards
to Indonesia and Thailand. Gordon (1984)
reported it as a common intertidal species around
North Island, New Zealand. It has also been
reported from the southwest Atlantic (Lopez
Gappa, 1977) and from Gough Island and Ascen-
sion in the central Atlantic (Hayward, 1980).

Crepidacantha carsioseta Winston
& Heimberg (Fig. 25d)

Crepidacantha carsioseta Winsto & Heimberg,
1986; 27, figs 64-66.

DESCRIPTION

Autozooids oval to hexagonal, 0.35-
0.45 0.3mm, Frontal wall smooth, bordered by
inconspicuous marginal pores. Primary orifice
longer than wide, trifoliate, with a broadly
rounded proximal lip. Avicularia paired,
proximal to orifice, directed medially; mandible
setiform, Ovicell spherical, smooth, with 2
curved band of pores crossing the frontal surface
distally.

DISTRIBUTION

This species has recently been described
(Winston & Heimberg, 1986) fram two localities
in Indonesia and is here reported from Australia
for the first time. The material from Heron Island
consists of small colonies from sample 25.

Family INVERSIULIDAE Vigneaux, 1949

Colony encrusting. Aulozooids with tubercu-
late frontal wall evenly perforated by small pores:

278 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 25. a, Sinupetraliella litoralis, X 40. b, Inversiula inversa, 70. c, Crepidacantha crinispina, * 90. d,
Crepidacantha carsioseta, *75.e,f, Microporella orientalis, e, *% 220; f, X50.

BRYOZOA FROM HERON ISLAND

a prominent suboral ascopore present. Orifice
with straight distal rim and concave proximal rim,
the operculum hinged along its distal edge; no
condyles; no oral spines. Avicularia adventitious.
No ovicells.

Inversiula Jullien,1888

Colony encrusting, unilaminar, Autozooids
with tuberculate frontal wall perforated by
numerous simple or stellate pores; primary
orifice with reversed orientation, the semicircular
operculum hinged along its straight distal edge;
aScus openly via a well-defined ascopore
proximal to the orifice; no oral spines. Avicularia
adventitious; typically paired, lateral to orifice,
Embryos. brooded intemally. Small basal pore
chambers present,

Type species: /nversiula nutrix Jullien, 1883.

Inversiula inversa (Waters)
(Fig. 25b)

Porina? inversa Waters, 1887: 95, pl. 4, fig, 23.
inversiula inversa: Harmer, 1957: 956, pl 72. figs 24,
32, 33.

DESCRIPTION

Colony forming small, irregular patches.
Autozooids irregularly hexagonal, separated by
distinct sutures; 0.55-0,65%c.0.5mm. Frontal
wall with raised ridges and tubercles, which form
4 radiating series around the edges; with distinct
marginal pores immersed between thickened
ridges, and Jarge, stellate pores over the central
part, each on a raised tubercle, Ascopore slightly
larger than frontal pores, stellate, also on a thick-
ened tubercle, immediately proximal to orifice.
Avicularia proximo-lateral to orifice, paired,
shortly columnar; with oval rostrum, 0.06mm
long.

DISTRIBUTION

!. inversa has been reported sporadically along
the eastern seaboard of Australia, from New
South Wales to Torres Strait, and from the Philip-
pine Islands, It was present in two of the Heron
Island samples.

Family MICROPORELLIDAE Hincks, 1877

Colony encrusting; or forming erect, bilaminar
plites, attached by an encrusting base; or arbores-
cent, freely branching. Autozooid frontal wall
evenly perforated, or with marginal pores only; a

conspicuous ascopore present proximal to orifice.
Onifice typically D-shaped, lacking condyles; oral
spines present or absent. Avicularia adventitious,
or absent. Ovicell hyperstomial. Vertical walls
with basal pore chambers or mural septula.

Microporella Hincks, 1877

Colony encrusting. Autozooids with nodular
frontal wall perforated hy scattered pores; as-
copore situated in distal half of autozooid, con-
spicuous. Primary orifice D-shaped. Distal oral
spines present, but sometimes small, and imper-
manent. Avicularia adventitious, mandible
setiform, Ovicell prominent, imperforate or with
few, minute pores; closed by autozooid oper-
culum. Large basal pore chambers present.

Type species: Eschara ciliata Pallas, 1766,

Microporella orientalis Harmer
(Fig. 25e.f)

Microporella orientalis Harmer, 1957: 962, pl. 62, figs
25-28, 38,

[not] Microporella orientalis; Gordon, 1984, 103, pl.
39, fig. C.

DESCRIPTION

Colony developing broad, unilaminar sheets,
Autozooids oval to hexagonal, convex, separated
by distinct sutures; 0.5-0.7*0.35-0.5mm.
Primary orifice wider than long, 0.08 X 0.12mm,
the proximal edge finely denticulate. Three
slender, distal oral spines present in early on-
togeny, missing in most autozooids distant from
growing edge. Frontal wall appearing distinctly
nodular, densely perforated by small, round
pores. Ascopore separated from proximal edge of
orifice by a distance slightly less than one orifice.
length, with slightly thickened rm and coarsely
denticulate, crescentic lumen. A single
aviculanum on each autozooid, situated lateral to
ascopore, with broad, thick crossbar and semicir-
cular proximal opesia; the rostrum abruptly
tapered and narrowest distally; mandible heavily
sclerotized and appearing dark brown, 0.2-
0.25mm long, setiform, or variously broadened
on one or both sides of the median rachis, and
typically with a pair of short, hooked, lateral
processes close to the base, Ovicell more or less
spherical, nodular, with minute pores scattered
over its surface; brooding autozooids develop a
thickened peristomial rim, peaked medially,
which passes distal to the ascopore and fuses on
each side with the ovicell.

REMARKS

Harmer (1957) possibly included more than
one species in his account of M, orientalis, and
his synonymy needs to be carefully re-examined.
The Heron Island specimens have been compared
with the holotype {Siboga Stn 50, W. Flores}, and
found to be almost identical to it; the size and
shape of the primary onfice are characteristic for
the species, as are the three delicate, distal oral
spines. The avicularian mandible varies in form
throughout a single colony and there is no utility
in denoting those specimens with lanceolate man-
dibles as ‘orientalis var,” (Harmer, 1957:963). It
is probable that M. orientalis occurs generally
throughout the western Pacific, and perhaps ex-
tends into the Indian Ocean, but the wider
geographical distribution accorded it by Harmer
(1957) should not be accepted until all of the
specimens and records listed by him have been
scrutinized again. The M. orientalis described
and figured by Gordon (1984) has four to six oral
spines, a smooth proximal border to the orifice,
an incomplete penstome in brooding autozooids,
and a striking, saucer-like rim to the ascopore; it
clearly represents a distinctly different species.

Microporella fimbriata sp. nov-
(Fig. 26a-c)

TYPE MATERIAL

Ho.oryre: GH5235, Heron Island, Sut 25, 18 Apr
1972.

PARATYPE: GH5236, as for holotype.

ETYMOLOGY
Latin fimbria, a fringe,

DESCRIPTION

Colony forming an extensive unilaminar sheet.
Autozooids oval to hexagonal, broad and rather
flat, separated by distinct sutures; 0.5-0.65 0.5-
0.6mm. Primary orifice wider than long,
proximal edge with a slightly curved, coarsely
denticulate inner rim, the outer tooth on each side
especially prominent. Four slender distal oral
spines present in the youngest autozooids, lost in
later ontogeny. Frontal wall coarsely nodular,
densely perforated by regularly distributed,
round pores; ascopore close to orifice, separated
from its proximal edge by a distance equivalent
only to one-third of onfice length; rim oval, thick-
ened, smooth, lumen shallowly crescentic, with
fine denticulations. Each autozooid with a single
avicularium lateral to ascopore, without a
pronounced rostrum, but with its frontal surface

MEMOIRS OF THE QUEENSLAND MUSEUM

symmetrical about a pair of stout, median con-

les; mandible thin, setiform, up to 0.7mm long.
laterally directed when at rest. Ovicell compara-
tively small, slightly wider than long, coarsely
nodular, with scattered, minute pores; aperture
rather angular, with a smooth, thickened border.

REMARKS

This is possibly the species descnbed from
Sumbawa, Indonesia by Harmer (1957). who
referred it to the North Pacific species M.
vibraculifera Hincks.

Fenestrulina Jullien, |838

Colony encrusting, unilaniinar, Frontal wall of
autozooid with pores in marginal series, or scat-
tered over its entire surface; pores typically stel-
late or complexly reticulate; ascus opening via a
conspicuous ascopore proximal to primary
onfice, often denticulate or reticulate. Primary
onfice D-shaped, with straight proximal edge:
$pines present or absent. Avicularia absent.
Ovicell with membranous ectooecium, calcified
entooecium, and marginal pores, Large basal
pore chambers present.

Type species: Cellepora malusii Audouin,
1826,

Fenestrulina caseola Hayward
(Fig. 26d)

Fenestrulina caseola Hayward, 1983: 325, pL 10, fig. &

DESCRIPTION

Colony forming flat, unilaminar sheets.
Autozooids oval to irregularly hexagonal, broad,
gently convex; commonly 0.6 X 0.5mm. Frontal
wall densely perforated by large, round, stellate
pores; ascopore separated from proximal edge of
orifice by a distance equivalent to orifice length,
with crescentic, denticulate foramen. No oral
spines. Ovicell broader than long, depressed and
nol very prominent; frontal surface with a thick-
ened transverse ridge; marginal pores small, in-
distinet.

DisTRIBUTION

A single large colony was present in sample 25.
This species was recently described from the
Mascarene Islands, Indian Ocean (Hayward,
1988), and the Heron Island record thus marks
only its second reported occurrence.

BRYOZOA FROM HERON ISLAND 281

FIG, 26. a-c, Microporella fimbriata; a, * 80; b, X 170; c, uncleaned autozooids, to show avicularian mandibles,
* 80. d, Fenestrulina caseola, X90, e,f, Celleporina reginae; e, primary orifice, X330; fF, «110.

282

Family MARGARETTIDAE Harmer, 1957

Colony erect, branching, jointed, attached by
cuticular rhizoids, Autozooids with ridged frontal
wall, perforated by minute pores, with a median
ascopore, Primary orifice rounded, hidden by a
projecting, tubular peristome. No spines. No
avicularia. Ovicell submerged in peristome,
which is conspicuously inflated in brooding
autozooids. Ancestrula erect, resembling later
autozooids, with 4 chitinous basal attachment.

Margaretta Gray, 1843

Colony erect, branching, jointed, attached by
cuticular rhizoids; consisting of cylindrical,
straight or curving. internodes. linked by brown
cuticular nodes, each arising from a short, cal-
cified basis rami developed by the parent inter-
node. Each node and basis rami consists of single,
double or triple tubes. Autozooids ordered in
whorls of two to eight around the axis of the
internode; frontal wall granular, densely per-
forated and appearing reticulate; a disto-medial
ascopore present. Primary orifice orbicular,
visible only in earliest ontogeny, obscured by a
long, cylindrical peristome, typically perforated
and longitudinally ridged. No spines. No
avicularia. Ovicell peristomial, hidden; brooding
autozooids with dimorphic peristomes, typically
longer, curyed, or with a modified secondary
orifice.

Type species: Celluria cerevides Ellis &
Solander, 1786,

Margaretta tennis Harmer
(Fig. 27a)

Margaretta tenuis Harmer, 1957; 840, pl. 55, figs
13-18.
Margaretta tenuis: Hayward, 1988: 330, pl, 11, fig, a,

DESCRIPTION

Colony generally sparsely branched, inter-
nodes 2-3mm long, 0.5mm wide. Aulozooids in
triple whorls, 0.8-1.0* c,0.4mm, fusiform, with
finely and regularly reticulate frontal shields, the
boundaries between adjacent autozooids not dis-
tinct. Basis rami tripartite, nodes consisting of
short, tripartite tubes, Peristomes of fertile
autozooids projecting at right angles to internode
axis, but otherwise not markedly different from
others,

MEMOIRS OF THE QUEENSLAND MUSEUM

DISTRIBUTION

M. tenuis has been reported from Queensland,
Torres Strait, the Philippine Islands, and from
many Indonesian localities; it ranges westwards
to Mauritius and the Red Sea.

Margaretta triplex Harmer
(Fig, 27b)

Margaretta triplex Harmer, 1957: 841, pl. 55, figs
Margaretta triplex: Ryland, 1984: 72, fig. 39.24;
Winston & Heimberg, 1986: 30, figs 70-72

DESCRIPTION

Colony stout, richly branched, up to 100mm
high; internodes straight or curved up to 30mm
ong, Imm wide. Autozooids in whorls of four,
1.2-1.4Xc.0.6mm, elongate oval, the frontal
walls finely reticulate, boundaries between
autozooids generally distinct. Basis rami and
node tripartite. Peristomes of ferule antozooids
projecting at right angle to internode axis, short,
very broad based and appearing volcano-like in
profile.

DISTRIBUTION

M. triplex is known from Queensland, Torres
Strait and Indonesia, and hus also been reported
from Zanzibar.

Superfamily CELLEPOROIDEA Johnston, 1838
Family CELLEPORIDAE Johnston, 1838

Colony encrusting, multilaminar, pisiform,
nodular or mamillate; or erect, attached by an
encrusling base, irregularly lobed or branched.
Autozovids without regular orientation; frontal
wall smooth, with marginal pores. Orifice with
concave or conspicuously sinuate proximal bor-
der; condyles present; spines present or absent.
Aviculariaadyentitious, and/or yicarious or inter-
zooidal; frequently polymorphic. Ovicell present,
vanously perforated, not closed by autozooid
operculum,

Turbicellepora Ryland, 1963

Colony encrusting, multilaminar, often nodular
or massive, or erect and branching. Frontal wall
of autozooid with marginal pores only, Primary
orifice with sinus; oral spines present or absent,
Avicularia polymorphic: adventitious, lateral-
oral; vicarious, varying in size, Ovicell with scat-

BRYOZOA FROM HERON ISLAND 283

FIG, 27. a, Margaretta tenuis, branch tip, X 50. b, Margaretta triplex, part of a branch showing a triple branch
base, X50. c,d, Turbicellepora ampla; c, X50; d, *55. e, Schedocleidochasma porcellanum, * 90.

284

tered frontal pores, not closed by autozooid oper-
culum. Basal pore chambers present.

Type species; Cellepora coronopus Wood,
1844,

Turbicellepora ampla (Kirkpatrick)
(Fig. 27¢,d)

Schizoporella anipla Kirkpatrick, 1888: 76, pl. 7, fig.
4,

Stephanosella bernardi(; Harmer, 1957: 1051, pl. 74,
figs 21-23.

Turbicellepora ampla: Hayward, 1988: 341, figs 6d,e.

DESCRIPTION

Colony forming multilaminar, nodular encrus-
tations. Autozooids strongly convex, with dis-
tinct, lobed, boundaries; the frontal wall
smoothly calcified, with indistinct, radiating
folds; marginal pores small and mconspicuous.
Primary orifice 0.12mm long, with a broad, V-
shaped sinus comprising about one-third its
length; with a pronounced distal and lateral peris-
tomual rim; no oral spines. Lateral oral ayicularia
single or paired; rostrum oval, acute to, frontal
plane, disto-laterally directed. Vicarious
avicularia dimorphic: with oval rostrum similar
to, but larger than, the lateral oral type, or larger
with broadly spatulate rostrum. Ovicell with
numerous small, papillate frontal pores.

DISTRIBUTION

This species was originally described from
Mauritius (Kirkpatrick, 1888). It is known from
a number of localities in the Indo-Malayan
region, but its presence in the Red Sea has yet to
be substantiated (Hayward, 1988).

Celleporina Gray, 1848

Colony encrusting, pisiform or lenticular, mul-
ulaminar, Orifice with a sinus; flanked by paired
Jateral oral avicularia, with columnar chambers,
incorporated into the peristome, Ovicell
prominent, spherical, with a perforated area of
entooecium, the tabula, frontally; not closed by
the zooidal operculum. Small basal pore cham-
bers present.

Type species: Lepralia hassallii Johnston,
1847.

Celleporina reginae sj), oy,
(Fig. 26e.f)

TYPE MATERIAL

MEMOIRS OF THE QUEENSLAND MUSEUM

HoLoTyPe: GH5237, Heron Island, Stn 16, 13 Apr
1972.

ETYMOLOGY
Latin regina, queen.

DESCRIPTION

Colony encrusting, mamillate. Autozooids at
growing edge convex; 0.4-0.45 x 0.25-0.35mm;
frontal wall smooth, with few, relatively large
marginal pores. Primary onfice longer than wide,
with a broad, U-shaped sinus comprising more
than one-third its total length: condyles thickened
and rounded; no oral spines. Paired lateral oral
avicularia present, with slender, columnar cys-
tids; rostra elliptical, with finely dentate distal
edges, acute to plane of orifice and invariably
directed laterally. A low, thin peristome encloses
the proximal half of the orifice, between the two
aviculana, but does not obscure it. Vicarious
avicularia frequent, up to 0.25mm long, the
rostrum broadly spatulate and deeply cupped dis-
tally; crossbar stout, lacking a columella, Ovicell
slightly longer than wide; ectooecium thick and
smooth, with a thickened ridge bordering the
frontal area of entooecium; frontal marginal
pores large, elongate and slit-like, so that the
entooecium appears to consist of 4 radiating
series of fused struts.

Measurements (means and standard deviations
of 20 values, mm): onfice Jength 0,11:40.007;
width 0.10+0.007,

REMARKS

C. reginae is distinguished from other westem
Pacific species of Celleporina by its deeply
sinuate orifice, and laterally directed lateral oral
avicularia. A single large colony was present in
sample 16, from the reef crest northeast of the cay.

Family PHIDOLOPORIDAE Gabb & Horn,
1862

Colony encrusting, unilaminar or multilaminar,
often massive; or erect, attached by an encrusting
base, forming bilaminar plates, or dichotomously
branching arborescent forms, or complex, anas-
tomosed, reticulate structures. Autozooid frontal
wall with marginal pores only. Primary onfice
with concave or conspicuously sinuate proximal
border, the distal border finely denticulate; con-
dyles present; oral spines present or absent.
Aviculatia adventitious, and/or vicarious or jnter-
zooidal. Ovicells present, with uncalcified area of
ectooecitm frontally; often with a well developed

BRYOZOA FROM HERON ISLAND

labellum, median longitudinal fissures charac-
teristic of some genera: not closed by aulozvoid
operculain,

Reteporella Busk, 1884

Colony erect, attached by an encrusting base;
branching dichotomously, developing an open
arborescent structure, or with the branches anas-
tomosing to form regular reticulate structures;
fan-shaped, or cup-shaped. or complexly folded.
Autozooids in two to many longitudinal series, on
a defined frontal surface; abfrontal surface of
colony consisting of blind kenozooids, usually
bearing adventitious avicularia. Pimary orifice
with denticulate distal border, frequently
obscured by a sinuate peristome. Avicularia ad-
yentitious, polymorphic, Ovicell with a median
longitudinal fissure, variably developed-

Type species: Refeporella flabellata Busk,
84

re

Reteporella graeffei (Kirchenpauer)
(Figs 28c,d, 29a)

Retihornera graeffei Kirchenpaucr, 1869: 30.

Reteporella graeffei: Harmer, 1934: 573, pl. 35, figs
12-15; pl, 38, figs 13-15; text-figs TSE, 31, 32;
Ryland, 1984: 74, figs 38.11, 40.31-34.

DESCRIPTION

Colony forming an irregular, open or shallowly
cupped fan; branches narrow, rather flat, mregular-
ly curved or twisted, dividing dichotomously at
imegular intervals, remaining free or anastomos-
ing to Jeave long, narrow fenestrulae between.
Autozooids in four to eight alternating lon-
gitudinal rows, flat, or only slightly convex,
broadly hexagonal, separated by distinct raised
sutures in early ontogeny, but boundaries later
rather indistinct. Frontal wall smooth, with two to
four proximal marginal pores. Primary orifice
wider than long, the proximal border shallowly
concave between small, rounded condyles;
obscured by a tall proximal peristome, with a
longitudinal median fissure, broadened basally to
form a labial pore; penstomial rim with two or
three short processes proximally, and with one
long spine on its inner (i.e., axial) lateral margin,
and four on its outer lateral margin. In later on-
togeny only a single spine persists on each side.
All spines segmented, up 0.3mm long. Avicularia
present on frontal walls of autozooids, polymor-
phic: most frequently 0.1mm long, with semi-el-
liptical rostrum, and the beni part produced

a8 a projecting, linguiform process, occasionally
larger, with the rostrum acute, and hooked or
dentate distally: gigantic avicularia with abruptly
hooked rostrum, up to 04mm long, present on
some marginal autozooids. Ovicell slightly longer
than wide, with an elongate median slit. In lajer
ontogeny the proximal regions of the colony are
grossly thickened, peristomes become deeply itn-
mersed, and additional avicularia are budded.

DISTRIBUTION

R. graeffei is common in shallow reef environ-
ments throughout the tropical westem Pacific,
from Queensland to Japan; it ranges throughout
the Indo-Malayan region and westwards to Sri
Lanka and East Africa.

Triphyllozoon Canu & Bassler, 1917

Colony erect, fenestrate, attached by an
encrusting base. Fan- or cup-shaped, or with the
edges complexly folded to form a squat, three-
dimensional bun shape. Autozooids opening on
one face only of fenestrate sheet, in alternating
series along the branches (trabeculae), which
anastomose at regular intervals defining oval
fenestrulae. Basal surfaces of colony crossed by
thickened sutures. Primary orifice with denticu-
late Tim, obscured by a deep penstome with a
longitudinal slit (pseudosinus), often partly
closed except for a proximal pseudospiramen,
typically with a small peristomial avicularium,
Oral spines present. Frontal wall with few, large
marginal pores. Avicularia adventitious,
polymorphic. Ovicell prominent, with a con-
spicuous, trilobed frontal suture, and a well
developed labellum overhanging the aperture.

Type species: Retepora monilifera Muc-
Gillivray, 1860.

Triphyllozoon mucronatum (Busk)
(Fig. 28a,b)

Retepora mucronata Busk, 1854: 120, pl. 26, figs 6a-e.

Triphyllozoon mucronatum: Harmer, 1934: 620, pl. 35,
fig, 35; pl. 40, figs 13-20; text-fig. 42.

Triphyllozogn sp.: Ryland, 1984: 74, fig. 40.36.

DESCRIPTION

Colony developing a low bun shape, up to
30mm high with a horizontal spread of up to
60mm, consisting of finely reticulate, deeply
lobed folds, anastomosing frequently to form a
compact structure. Fenestrulae small, regularly
oval, 1,0 0.5mm; trabeculae comprise three to

286 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 28. a,b, Triphyllozeon mucronatum; a, X 80; b, X 120. c,d, Reteporella graeffei; c, X 60; d, X 130,

BRYOZOA FROM HERON ISLAND

five longitudinal autozooid series, Antozooids
commonly 0.4% 0.2mm, with smooth, convex
frontal wall, bordered by a few large. widely
spaced marginal pores; sutures beiweeno
aulozooids distinct at growing edge, later
progressively obscured. Primary orifice D-
shaped, hidden by a deep peristome enclosing its
proximal] and lateral borders; lip of peristome
reflected, with 4 series of coarse creoulations on
its inner edge: medio-proximally a short fissure
connects the penstomial rim with the small labial
pore, to one side of this is a small peristomial
avicularium, acute to frontal plane, laterally
directed, with rounded, finely denticulate
rostrum. Four short, distal oral spines present in
newly budded autozooids, only the proximal pair
persisting in later ontogeny. Frontal adventitious
aviculana frequent, resirum 0.1mm Long, eval or
slightly spatulate. A single large vicarious
avicularnium present distal or proximal to each
fenestrula, with a swollen cystid supporting a
narrowly crescentic rostrum: other, larger
avicularia occur sporadically, typically within
each fenestrula an avicularium with a narrow,
elongate rostrum, broadened and denticulate dis-
tally; elsewhere, elongate tnangular avicalaria
may be present. Ovicell slightly longer than wide,
with a short, square-edged labellum; median
frontal fissure longer than proximal fissures.

DISTRIBUTION

This species is known from a few localities in
the Philippine Islands, northern New Guinea and
eastern Indonesia. It has not been identified pre-
viously from Australia.

Schedocleidochasma Soule, Soule & Chaney,
1991

Colony encrusting, uni- or multilaminar.
Primary orifice of autozooid cleithridiate
(keyhole-shaped), with a deep, arrowhead-
shaped or transversely oval sinus below
prominent condyles; distal rim of orifice smooth.
Oral spines present or absent, Frontal wall with
few marginal pores. Avicularia adventitious.
Ovicell hyperstomial, partly immersed, not
closed by autozooid operculum; with a frontal
area of uncalcified ectooecium, and a well-
marked labellum. Vertical walls with basal or
mural pore chambers.

Type species: Schedocleidochasma porcel-
laniforme Soule, Soule & Chaney. 1991,

237

Schedocleidochasma porcellanum (Busk)
(Fig, 27e)

Lepralia porcellana Busk, 1860: 283, pl. 31, fig. 3.

Cleidechasina porceilanum: Harmer, 1957: 1042, pl.
71, figs 5, 16; Cook 1964b: 11, pl. 1, fig. 4; pl. 2,
figs 1, 2; text-figs 4A-E; Winston & Heimberg 1986;
34, figs 85-87.

Schedocleidechasma porcellanen; Soule, Soule &
Chaney, 1991: 481.

REMARKS

This familiar species has a circumtropical/sub-
tropical distribution in shallow coastal waters,
and may be particularly common in reef flat
habitats, on coral rubble, or other hard substrata.
At Heron Island it forms flat, silvery, unilaminar
sheets, often exceeding 3cm*. It is readily recog-
nised by the shape of the primary orifice, the
lateral suboral avicularium, and its thick, poreel-
lanous secondary calcification. It was present in
abundance in seven of the Heron Island samples,

The major revision of Cleidochasma by Soule
et al. (1991) was published after this paper had
been submitted for publication. The specimen
illustrated here appears to conform to their con-
cept of §. porcellanum, especially in its shallow,
transversely oval sinus but, in view of the fact that
several previous West Pacific records of this
species were shown by Soule et al. to represent
theit new species, 5. parcellaniforme, it will be
necessary subsequently to review the Heron Is-
land matenal attnibuted to 5S. porcellanum.

Rhynchozoon Hincks, 1895

Colony encrusting, multilaminar; discoid,
domed, or more usually irregularly spreading,
extensive, nodular, Autozooids distinct only al
growing edges of colony, or in new, frontally
budded laminae: elsewhere immersed to a vary-
ing degree in thick secondary calcification.
Primary orifice with distal border (anter) beaded
or denticulate; proximal border (poster) broadly
concave to narrowly sinuate; condyles usually
distinct. Oral spines sometimes present in newly
budded antozooids, Onfice encircled, and Jater
partly obscured, by marked peristomial thicken-
ing, frequently produced into erect or cylindrical,
knob-like processes; typically with a distinct
notch (pseudosinus) in the proximal border, A
transversely or obliquely laterally orientated
avicularium located on the proximo-lateral bor-
der of the orifice, large and conspicuous, with
hooked rostrum and apical process (umbo), ot

288

small and concealed within peristome; often with
a stout process (uncinate process) projecting
towards the orifice from the inner, proximal edge
of the avicularium. Additional, large or small,
avicularia present on frontal wall of autozooid, or
around peristome rim, often numerous. Frontal
calcification of autozooids thick, frequently pear-
ly or distinctly nodular, with marginal pores only.
Ovicell typically elongate oval, with a distinct lip
(labellum) above the aperture; frontal surface
with the outer layer (ectooecium) uncalcified,
exposing a smooth or granular area of en-
tooecium (tabula); most often immersed in thick-
ening calcification in early ontogeny and
becoming indistinct. Vertical walls with small
basal pore chambers.

Type species: Lepralia bispinosa Johnston,
1847.

Species of Rhynchozoon are common on all
tropical reefs and can be particularly difficult to
identify. Ontogenetic thickening of the frontal
calcification, accompanied by progressive im-
mersion of the primary orifice, and the prolifera-
tion of frontally budded avicularia may
completely mask those features important for
identification. In practice, it is usually impossible
to identify any specimen of Rhynchozoon without
first bleaching a sample to reveal details of the
primary orifice. Size and shape of the primary
orifice (in particular the sinus), and the morphol-
ogy of the proximal suboral avicularium are the
most important taxonomic features in this genus.
In some species, the frontal avicularium and the
ovicell provide useful, confirmatory characters.

Rhynchozoon ardeolum sp. nov.
(Fig. 30d,e)

TYPE MATERIAL
HOLOTYPE: GH5238, Heron Island, Stn 25, 18 Apr
1972.

ETYMOLOGY
Latin, diminutive of ardea, a heron.

DESCRIPTION

Colony forming broad, multilaminar patches.
Autozooids irregularly oval to polygonal, strong-
ly convex, separated by well-marked sutures;
commonly 0.45-0.5 x 0.35-0.4mm. Frontal wall
thickly calcified, distinctly corrugated, with a
single series of round marginal pores. Primary
orifice slightly wider than long, with a short,
rounded sinus, bounded by thickened cusps,
below blunt, rounded condyles. No oral spines.

MEMOIRS OF THE QUEENSLAND MUSEUM

Many autozooids bearing a very large lateral oral
avicularium, with a swollen cystid occupying
much of the frontal wall of the autozooid; rostrum
broadest proximally, the distal portion, bearing
the mandible, elongate, narrow, curved and
rounded at the tip, acute to the frontal plane of the
autozooid and directed disto-laterally; total
length of rostrum 0.3-0.35mm. Additional
avicularia sparsely developed, shorter than the
lateral oral type, with a narrowly triangular
rostrum distal to the mandibular condyles. In later
ontogeny frontal calcification thickens, the mar-
ginal pores become much accentuated, and three
or four stout, conical processes develop around
the disto-lateral borders of the orifice, which,
however, remains distinct, though partially im-
mersed, and never completely hidden. Ovicells
not present.

Measurements (means and standard deviations
of 20 values, mm); orifice length (incl. sinus)
0.12+0.004; width 0.12+0.00S.

REMARKS

This species is readily distinguished from all
other presently known species of Rhynchozoon
by its unusually large lateral oral avicularium,
and by its very distinctive primary orifice. It was
present in two of the Heron Island samples.

Rhynchozoon bifurcum Harmer
(Fig. 29b-d)

Rhynchozoon bifurcum Harmer, 1957: 1070, pl. 70,
figs 6, 13-15.

DESCRIPTION

Colony a multilaminar, mamillate sheet.
Autozooids at growing edge oval, convex,
separated by shallow grooves; about 0.5 X
0.3mm. Primary orifice wider than long, about
0.16 X0.10mm; proximal border shallowly con-
cave between inconspicuous proximo- lateral
condyles; no oral spines. Frontal wall nodular,
with closely spaced, conspicuous marginal pores.
Lateral suboral avicularium present in all
autozooids, prominent, with swollen cystid, and
broadly triangular, distally hooked rostrum;
proximally, a slender subtriangular uncinate
process projects above the primary orifice. A
stout mucro developing on avicularium, con-
tributing to the peristomial rim, which develops
early in ontogeny, obscuring the avicularium and
orifice, and eventually consists of six or seven
stout, conical processes more or less evenly
spaced around the secondary orifice. Frontal

BRYOZOA FROM HERON ISLAND 289

FIG. 29. a, Reteporella graeffei, 90. b-d, Rhynchozoon bifurcum; b, X55; e, an avicularium with bifurcate
mandible, x 130; d, primary orifice, * 320. e,f, Rhynchozoon scopulorum, &, %* 50; F, primary orifice, x 270.

290)

avicularium clongate oval to narrowly spatulate,
0,25-0.5ram long, the mandible in the largest
always deeply forked. Ovicell elongate, the fron-
tal surface flat and very finely tuberculate.

DISTRIBUTION

This species is known from Torres Strait, Sin-
gapore, and the Chagos Archipelago, but has not
been recorded previously from Queensland. The
specimen from Japan discussed by Harmer
(1957) perhaps does not belong to &. bifurcurn.

Rhynchozoon compactum (Thornely)
(Fig. 30b,c)

Cellepara compacta Thomely, 1905: 126, pl. fig. 17.
Rhynchozoon compactunt: Harmer, 1957; 1076, pl. 70,
figs 21-25.

DESCRIPTION

Colony forming irregular, multilaminar
patches. Autozouids at growing edge oval to
hexagonal, strongly convex. smoothly calcified,
about 0,5-0.6 *0.35mm. Pnmary orifice almost
circular, about 0.09mm diameter, with a small,
U-shaped sinus. No oral spines. A single, lateral-
oral, avicularium present jn many, but not all,
autozooids, the hooked rostrum perpendicular to
the plane of the orifice and proximo-laterally
orientated. Ovicell as wide as long, the frontal
entooecium smooth, labellum indistinctly
curved. Jn later ontogeny the frontal calcification
of the autozooid thickens and becomes. faintly
corrugated; boundaries between autozooids
remain distinet, however, and the few smal] mar-
ginal, pores are accentuated. A broad, thick ridge
develops around the distal and lateral borders of
the primary orifice, characteristically developing
two or three knob-like processes, but the orifice
remains clearly visible in frontal view, With the
development of frontal buds, the primary orifices
of the first layer of autozooids become immersed
in deep, peristomial tubes, but are not completely
hidden. Additional frontal aviculana are sparsely
developed, most frequent in later ontogeny; these
are up to 0.15mm leng, with an elongate oval
rostrum.

REMARKS

Heron Island specimens of this distinctive
species have been compared with the type
material from Sri Lanka, and found to be identical
to it, Harmer (1957) listed a specimen from Tor-
res Strait, which it is probably safe to assume
belongs to R. compactum, but his records from

MEMOIRS OF THE QUEENSLAND MUSEUM

Japan need to be confirmed following re-ex-
amination of the specimens listed.

Rhynchozoon baha Hayward
(Figs 30f, 31a,b)

Rhynchozoon haha Hayward, 1988: 334, pl. 11, figs
Lg

DESCRIPTION

Colony developing broad_ irregular, spreading
sheets. Autozooids at growing edge oval to
hexagonal, or rectangular, convex, separated by
distinct sutures; 0.5-0.6 * 0.3-0.35mm. Primary
orifice wider than long, with a broad, U-shaped
sinus occupying about three-quarters its proximal
width; condyles knob-like, small, no oral spines.
Frontal wall smooth, with a single series of large,
distinct marginal pores. A large, lateral suboral
avicularium present on most autozooids; cystid
swollen, frequently with a small umbo on its
frontal surface; rostrum acute to frontal plane,
directed disto-laterally, acuminate but with only
a slight distal hook; proximally the avicularian
rostrum thickens to form a short, blunt, uncinate
process. Frontal adventitious ayicularia charac-
teristic: typically one per autozooid, 0.2mm long,
directed proximally or proximo-laterally,;
lozenge-shaped. with the narrowly triangular
rostrum about twice as long as the proximal
opesia. Ovicell short, smooth frontally, complete-
ly immersed by ooecial cover and not readily
visible in frontal view, With ontogenetic thicken-
ing the primary orifice becomes deeply im-
mersed, hiding the suboral avicularium: four ar
five stout, conical umbones develop on the peris-
tomial rim, with the proximalmost pair flanking
a pronounced proximo-medial notch. The frontal
avicularia remain distinct, however, and the
autozooid boundaries can be recognised by the
deep, conspicuous marginal, pores.

DISTRIBUTION

R. haha was founded on material collected
from reef habitats around Mauritrus (Hayward,
1988). This is only the second occurrence, but it
will probably prove to be generally distributed in
the Indo-West-Pacific region, in shallow reef
habitats.

Rhynchozoon limatulum sp. nov.
(Fig, 31c-e}

TYPE MATERIAL

BRYOZOA FROM HERON ISLAND 291

FIG. 30. a, Rhynchozoon scopulorum, X70. b,c, Rhynchozoon compactum;, b, * 350; c, * 80, dye, Rhynchozoon
ardealum; d, *350; e, x60. f, Raynchezoon haha, * 70.

292

HOoLotyPe: GH5239, Heron Island, Stn 14, 12 Apr
1972.

ETYMOLOGY
Latin, diminutive of limatus, polished.

DESCRIPTION

Colony forming irregular, multilaminar sheets.
Autozooids at growing edge broadly oval, con-
vex, separated by distinct sutures. Primary orifice
distinctly wider than long, with a deep, U-shaped
sinus occupying most of proximal width; con-
dyles thickened, conspicuous. No oral spines.
Frontal wall smoothly calcified, bordered by
small, widely-spaced marginal pores; typically
developing two or three prominent, conical
processes proximally, forming part of the peris-
tome of the preceding autozooid. Peristome
spinous, typically with a single medio-proximal
process, paired lateral processes, and two or three
distal processes, all prominent, tapered, and oc-
casionally bifurcate apically. Most autozooids
with a lateral oral avicularium enclosed within
the peristome, the rostrum slender, triangular,
hooked distally, and laterally directed. One to
three smaller avicularia present on frontal wall;
rostrum slender, triangular, laterally directed.
Ovicell with a very short labellum and a con-
spicuous frontal fenestra; rapidly immersed in a
smooth ooecial cover.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.41+ 0.04;
width 0.3040.04; orifice length 0.08+0.004;
width 0.10+0,003.

REMARKS

This species is distinguished from others in the
Heron Island reef flat fauna by its short, wide
primary orifice, and comparatively deep sinus.
The smooth frontal calcification, spinous peris-
tomes and small frontal avicularia are also char-
acteristic.

Rhynchozoon scopulorum sp. nov.
(Figs 29e,f,30a)

TYPE MATERIAL
HoLotyre: GH5240, Heron Island, Stn 16, 13 Apr
1972.

ETYMOLOGY
Latin scopulosus, rocky.

DESCRIPTION
Colony forming broad, spreading sheets.

MEMOIRS OF THE QUEENSLAND MUSEUM

Autozooids at growing edge oval to hexagonal,
convex, separated by distinct sutures. Primary
orifice about as wide as long, with shallow, U-
shaped sinus occupying about half proximal
width; condyles rounded, distinct; distal and
disto-lateral denticulations fine, closely spaced.
No oral spines. Frontal wall distinctly nodular,
with few, widely spaced, marginal pores. Lateral
suboral avicularium large and prominent;
rostrum acute to frontal plane, laterally directed,
hooked distally; proximally, the rostral rim is
produced as a stout, conspicuous uncinate
process, almost reaching the opposite side of the
peristome, delimiting a rounded pseudosinus. A
small adventitious avicularium present on the
frontal wall of most autozooids, the rostrum tri-
angular, acute to frontal plane, usually directed
proximally. Large frontal avicularia sporadic:
0.4mm long, occupying most of autozooid frontal
wall, proximally directed; rostrum lanceolate,
often asymmetrical, with upturned tip. Ovicell
longer than broad, projecting well above peris-
tomial rim and remaining distinct in later on-
togeny; frontal surface slightly flattened,
minutely tuberculate. In later ontogeny the sub-
oral avicularium becomes submerged within the
peristome, and the proximal peristomial rim bears
a pair of short, conical umbones, with a shallow,
U-shaped notch between.

Measurements (means and standard deviations,
mm)): autozooid length (n= 20) 0.56+0.05; width
0.33+ 0.03; orifice length (n = 10) 0.11+ 0.005;
width 0.14+ 0.000.

REMARKS

This species is similar to the Indonesian R.
detectum Harmer, 1957, recently described and
figured by Winston & Heimberg (1986). How-
ever, R. detectum is distinguished from R.
scopulorum by its primary orifice, which is only
very shallowly concave proximally, and by its
large frontal avicularia, which are narrower and
more distinctly pointed than in R. scopulorum.
The large avicularia of the latter are in fact almost
identical to those of R. triangulare Harmer, 1957,
also described from Indonesia; the primary
orifice of R. triangulare, however, has a slit-like
proximal sinus. Rhynchozoon scopulorum seems
to be common at Heron Island; large colonies
were present in six of the samples studied.

BRYOZOA FROM HERON ISLAND 293

— i a
FIG. 31, a,b, Rhynchozoon haha; a, X 450; b, X55. c-e, Rhynchozoon limatulum; c, * 70; d, X 300; e, < 140.
f, Rhynchozeon splendens, * 80.

244

Rhynchozoon splendens Hayward
(Figs 31, 32a.b)

Rhynchozoon splendens Hayward, 1983. 335, pl. 13,
figs fg.

Rhynchozoon rostratum: Winston & Heimberg, 1986:
38, figs 95-98.

DESCRIPTION

Colony forming spreading, multilaminar
sheets, Autozooids at growing edge aval, convex,
separated by distinct grooves; commonly 0.45-
0.50% 0.3mm. Primary orifice 0.12mm_ wide,
about as long as wide; sinus short, U-shaped,
occupying about half proximal width of orifice;
condyles indistinct. No oral spines. Frontal wall
at first smooth, but becoming coarsely and ir-
regularly nodular, bordered by large and con-
spicuous marginal pores. Peristomial rim
markedly thickened, produced into a series. of
projecting, knob-like, conical process, the most
constant being a medio-proximal pair which may
be especially prominent; most autozooids with a
single, lateral, suboral avicularium enclosed
within the peristome, with triangular, hooked
rostrum directed laterally or disto-laterally; the
inner proximal corner of the rostral rim is
produced as a short, blunt, uncinate process.
Small frontal avicularia present, typically one
proximo-lateral to peristome, and sometimes a
second, more proximally situated; rostrum
diamond-shaped, with variable orientation.
Ovicell with a short, broad labellum; developing
a finely nodular ooecial cover, generally rather
iNCONSpIcuoUs,

REMARKS

R. splendens was recently described from
Maunitius (Hayward, 1988), where it was found to
be common in shallow reef habitats, Comparison
of Heron Island specimens with Mauritian
maternal confirms that they are all one species; the
shape and size of the primary onfice, the confor-
mation of the peristome, and the characteristic
diamond-shaped aviculana are all remarkably
similar in the two series of specimens, The species
reported from Sanur, Bali, by Winston & Heim-
berg (1986) as R. rostratun (Busk) seems to be R.
splendens. R. restratum is similar to R. splendens
in its diamond-shaped frontal ayiculana and lobed
penstome, Which encloses a large lateral-suboral
avicularium, but differs in its primary orifice,
which has a wide, shallow sinus, and propor-
tionately larger condyles. R. splendens was the
Mast commen species of Rhynchozoon collected

MEMOIRS OF THE QUEENSLAND MUSEUM

at Heron Island; small colonies, and Jarger sheets,
were frequent in eight of the samples described
here.

Rhynchozoon tubulosum (Hincks)
(Fig. 32c¢-e)

Mucronella (7) tubulosa Hincks, 1880d: 383, pl. 17,
lig. 7.

Mucronella tubulesa; Thornely, 1905: 124.

Rhynchozoon tubulosum: Harmer, 1957; L064, pl. 65,
figs 16-19; Gordon, 1984: 125, pl. 52, figs C-B;
Hayward, 1988; 336 fin part; not pl. 14, figs a, b).

DESCRIPTION

Colony forming small, unilaminar patches.
Autozooids at growing edge oval to hexagonal,
steeply convex, separated by distinct sutures (in-
distinct im later ontogeny); 0,45-0.5 *0.25-
0.35mm. Primary orifice as wide as long;
proximal edge broadly and shallowly concave
between small, rounded, proximo-lateral con-
dyles;, distal/disto-lateral denticulations small
and widely spaced. Frontal wall irregularly
nodular, with few, large, widely-spaced marginal
pores: proximal peristomial lip prominent, raised
medially asa stout, knobbly mucro, occasionally
spiky, or distinctly bifid apically; additional,
shorter mucronate processes sometimes
developed lateral to the median. Suboral
avicularium transversely orientated, on the inner
surface of the mucro, the rostrum sharply hooked
distally, but without an associated uncinate
process. Large frontal avicularia sporadically
present, most frequent in older parts of colony;
rostrum linguiform, about 0.35mm long, normal
to frontal plane and proximally directed. Tiny
adventitious avicularia, with short triangular
mandible, often developed towards tip of mucro,
and rarely elsewhere on the frontal wall. Ovicell
about as wide as long, slightly flattened frontally,
with a finely tuberculate surtace, projecting well
above rim of peristome.

REMARKS

With ontogenetic frontal wall thickening the
primary onfice becomes deeply immersed and
the mucronate proximal peristome especially
prominent, so that autozooids appear to be verti-
cally orientated, Hinck’s (1880d) unfortunately
inadequate figure shows a group of autozooids in
late ontogeny, with prominent, mucronate peris-
tomes, tuberculate ovicells clearly visible above
the peristome rims, and large, linguiform frontal
avicularia. No type specimen is known for R.

BRYOZOA FROM HERON ISLAND 295

FIG. 32. a,b, Rhynchozoon splendens; a, x 60; b, X 350, c-e, Rhynchozoon tubulosum; c, X 45; d, * 260; e,
x70.

tubuloswn, Hineks' original description is incom-
plete, and the identity of the Heron Island species
described here depends entirely on the
resemblance of older autozooids to Hincks' fig-
ure. Subsequent authors (see Harmer, 1957;
1064) considered that Riynchozoon longirostre
(Hincks), described from southern Australia, was
the same as RK. tubulosum, an assertion which was
denied by the author himself (Hincks, 1891). In
Hincks’ original specimens of R longirosire
(e.g, BMNH 1899.5.1.1161, 1163), the suboral
avicularium is associated with a massive uncinale
process, which partly obscures the primary
orifice, a feature not seen in any of the material
of R, tubulosum examined here. Consequently,
the synonymy of the two taxa should be rejected.
Thornely’s (1905) material of R. tubulosum from
Sri Lanka (BMNH 1936,12,.30,48A) and
Harmer's (1957) Indonesian specimens are ihe
same species as that occurring at Heron Island;
Harmer’s synonymy, however, cannot be ac-
cepted until all listed records of R. tubulosum, R.
longirostre and R. profundum (MacGillivray)
have been re-examined. The material described
from Mauritius by Hayward (1988) unfortunately
comprised two species, and the specimen figured
(Hayward, 1988, pl. 14, figs a,b) is aot R.
fubviosum, but an undescribed species of
Rhynchozoon.

CONCLUSION

The selection of Bryozoa described in this
preliminury account comprises a single cyclos-
tome, and 80 species of cheilostome, encompass-
ing 46 genera in 27 families. Of these 80 species,
14 are new (o science and 24 are reported for the
first time from Australian seas. The remaining 43
species include at least 25 which are known to be
widely distributed in the Indo-West Pacific
realm, some of which {v.g., Chorizopora
brongniartii and Arthrepoma cecilii) appear to
have almost world-wide distributions in shallow
tropical and temperate seas. This group of 81
species includes the most abundant, most familiar
and taxonomically least problematic members of
the bryozoan fauna of the Heron Island reefs.
While the range of genera represented gives some
indication of the taxonomic diversity of the Fauna,
the total number of species is likely to be greatly
increased is the rest of the collection is studied,
[tis worth noting that few of the species listed by
Livingstone (1927) or described by Hastings
(1932) from GBR localities have so far been
found in these collections, Unfortunately,

MEMOIRS OF THE QUEENSLAND MUSEUM

progress in laxonomic research on Wopical
Australian Bryozoa is hampered by the necessity
to review all previous published sources critical-
ly, and whenever possible to refer to type and
original specimens. Harmer’s (1957) fourth
Siboga Keport, presently the most important of
these sources, has serious shortcomings in that
the synonymies of many of the species it
describes can be shown to be erroneous, while a
substantial number of others are founded on
material which clearly consists of more than a
single species.

LITERATURE CITED

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AUDOUIN, J.V. 1826. Explication sommiaire dos
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225-244 (Planches J-14 par J.C, Savigny).

182%. bid. Edition 2.

BASSLER, R.S. 1935. Bryozoa, Generum et
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BENNETT, |. 1971. ‘The Great Barrier Recf*.
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British and adjacent waters. Bulletin of the
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{1): 1-63.

BORG, F. 1926. Studies on Recent Cyclostomutous
Bryozoa. Zoologiska Bidrag frin Uppsala 10;
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BROOD, K, 1976, Cyclostomatous Bryozoa from the
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BROWN, L.A, 1952, "The Tertiary Cheilostomatous
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1954. On the polyzoan genus Crepidacantha Levin
sen. Bulletin of the British Museum (Natural
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BUSK, G, 18523. An account of the Polyzoa and
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the Louisade Archipelago, ete. Appendix no. IV.
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the Voyage of H.M.S, Rattlesnake.” (T.&W.
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1852b. 'Calalogne of marine Polyzoa in the collec-

BRYOZOA FROM HERON ISLAND

tion of the British Museum,’ |; 1-54, (Trustees of
the British Museum; London),

1854. ‘Catalogue of marine polyzoa inthe collection
of the British Museums.” 2: 55-120. (Trustees of
the British Museum: London).

1860. Catalogue of the Polyzoa collected by J.Y.
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1860, with descriptions of the new specics.
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1859. ‘A monograph of the fossil Polyzoa of the
Crag’, (Palaeontographical Society; London),
1861. Zoophytology. Descriptions of new or imper-
fecily known Polyzoa. No. 1. Quarterly Journal
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2

FOOT-FLAGGING DISPLAYS OF SOME
AUSTRALIAN FROGS, Memuirs of the Queensland
Museum 32(0): 2, WYZs- Visual displays involving
movements of the feet have been recorded for four anuran
species: Aylodes ayperas (Leptoelactylidae), Smmnpis paryus
(wuberiingwis) and § latepalmains (Ranidac), and
Tuudactylus eungellensix (Myobatrachidae) (Davison, 1984;
Harding, 1982; Heyer et al. 1990; Winter & McDonald,
1986), Al of these species are diurnal and live along rainforest
sireams. This led Heyer et af, (1990) to suggest that visual
displitys might be at substitu for vocal communication dn
fesponse to naisy streamside environments where vocal
angoals are difficult to hear, and may be Jess risky for frogs
that have easy access to a safe refuge. ‘They also nored
morphological similarities between Fylodes asperus and the
Australian hylid Literia nannotis, and questioned whether /.
nannotis alse might exhibit Visual displays.

In this note We confirm that 4. wannoris does exhibit foot-
Nogging displays, and describe visua) displays by (hree olher
Australian hylid frogs. These include the first records of
nocturmal fool-flugging and fout-apging displays exhibited
ouide a rainforest environment. Meld observations were
qnade in [he wet lropics of northern Queensland and in nenh-
ern New South Wales, between March 1989 wad Marely 1992,

Liioria genimaculata: Three inales were observed dis-
playing on mossy rocks nex! to Birthday Ck, an upland
rainlorest stream in Mt Spee National Park. Observations
Wert wade over a 15 minute period beginning at 1LO0h on
16.2,90, One mate slowly raised the ght rear leg, swerched
i) out perpendicular io the body and rapidly vibrated the foot
ona vertical plane with the toes outstretched. It then Jeapt onto
a second mite about Imi away. This second male retreated a
further metre and exhibited the same display, once wilh cach
rear FOOL, A thied male perched Within 50cm then displayed
vigorously more than a dozen tines, with go apparent se-
quence of use of the two feet. There followed 4 series of leaps
anto each other, several wrestling byuts, and sporadic displays
ball Uneee mules unnl they dispersed approximately 15
ihututes after the displays were initiated. All of the males
called continously during these bouts,

Litoria nannotis: Observations were made. over a two tiour
period in November, 1990, on capiive specimens maintained
in a large aquarium. Visual displays were initiated by a
resident Male when a second male was placed into the same
aquarium. Immediately following the introduction, the resi-
dent male called vigorously as he approached the intruder, and
then extended the nghe roar leg our and back in an are,
spreading the toes tg reveal the webbing, belore relurning the
Jeg to the side of the body. This displiy was repeater’ coo.
Hintiously with both back legs vatil the intruder retreated into
a crevice. The display was reinitiaied gach nme the intruder
appeared, Male L. sannesis have been observed in the field to
wave their front legs in an arc above the head in similar
iyter-nale displays (A, Dennis, pers, comn.).

Litoris rheseola: Three males were observed displaying,
along rainforest streams inthe vicinity of Mt Finnigan, north-
orn Queensland, All observations were made at might berween
2200 and 233007 17.12.91 All Involvedinteractons between
calling males. The first male was perched on a leaf ata height
of 38cm and facing another calling male 40cm away. The first
male waved its Troot right foor twice in an acc over its head,

MEMOIRS OF THE QUEENSLAND MUSEUM

fy then made two rupid Hicks with (he rear leg so that the leg
was extended fully, 1 4 lateral plane perpendicular to the lowg.
axia of the body, and returned immediately to the side of the
body. A second male displayed only with the rear legs, ex-
hibiting seven simular leg flicks in Tour minutes. Six Hicks
were with the left leg and one with the right, A third calling
male was observed for five minuies and did Hot display until
another male called one metre away. It immediately began
displaying with both rear legs in the manner described above,
These displays were observed by moonlight.

Litoria fallax: Observations were made between 2150and
22100 08 01.01.91 in the vicinity of Brays Ck, northern NSW,
under well-moonlit conditions. The teraction was between
aculling male perched 50cm above the water and a suicllite,
non- calling nule, initially perched 35cm away. The calling
male began a rapid clicking encounter call, when approached
fron behind by the second male, A wrestling bout ensued for
approximately one minute during which the first male culled
continuously and ended with ihe second male sitting behind
and facing the same direction as the first, about 1Oem lower
on tte same branch, The first male then foot-flagged by
aliermmately extending rear legs, starting with a stretch side-
wands wad slowly moving each leg through a backward are to
end in the lateral resting position. This was repeated seven
times, ending when the second male leaped away, The reniain-
ing male resumed calling four minutes later, No species
reported here haa brightly coloured webbing between the toes.

Vhe hylid species reported here to exhibit foot flagging
behaviour are all nocturnally active and all use voealisations
(o allract conspecitic females. Displays were only observed in
the presence of rival males. Isolated males, or males in the
presence of females, have not been observed displaying.

The functional significance of fool-flagging appears to be
télated more (o male-male aggression than to courtship, Pre-
viously reported displays may also prove to have an aggres-
sive raiher than a courtship function. More investigations of
frog behaviour may reveal a greater diversity of visual dis-
plays than previously suspected. Also, observations are re-
quired te detwemine the effect of light intensity (moonlight)
on visual displays in noctumally active frogs.

K. McDonald and A. Richards assisted in the field. Pield
work was tunded by ARC grants A18715284.and AI8931875
toR. Alford, anda grant from the Wet Tropics World Heritage
Agency. C, James was supported by an Australian Post-
gredunce Research Award,

Literature Cited
Davisen, G. 1984. Foor-fagzing displays in Bornean Frogs,
The Sarawak Museum Journal 33; 177-178.
Airding, K.A. 1942, Courtship display in a Bornean Prog.
Proc. Biological Society of Washington 95; 621-624
Heyer, WR. Rand, AVS,, Wa Cruz, C.A.G., Peixoto, O,L, &
Nelson, C.B, 1990. Progs of Boraceia. Arquivos de
Zoologia 3); 241-410.

Winter, J. & McDonald, K.R. 1986. Bungella: The land of the
cloud, Australian Natural History 22; 39-43,

Stephen J. Richards, Zaalogy Department, James Cook
University, Townsville, Queensland 4811, Australia;
Christina James, Deparunent ef Zoology, University of
Queensland, Queensland 4072, Australia; 21 April, 1992,

A NEW SPECIES OF LIZARD (TILIQUA)
FROM THE MIOCENE OF RIVERSLEIGH, QUEENSLAND

GLENN M. SHEA AND MARK N, HUTCHINSON

Shea, G.M. & Hutchinson, M.N, 1992 06 29: A new species of lazard (Tiligua) Irom the
Miocene of Riversleigh, Queensland, Memoirs of the Queensland Museum 32(1); 303-310.

Brisbane. ISSN 0079-8835.

A new species of Tiliqua, T, pusilla, is described fromm fossil dentaries from the Gag locality,
Dwornamor local fauna, Middle Miocene of Riversleigh, Queensland. The new species is
characterised by its distinctive rounded check teeth and very small size. (J7iliqua, Scincidae.

dentition, Miocene, Riversleigh.

Glenn M. Shea, Department of Veterinary Anatomy, University of Sydney, New South Wales
2006, Australia; Mark N. Hutchinson, South Australian Museum, North Terrace, Adelaide,
South Australia 5000, Australia; 24 June, 19917.

The limestone deposits of the Riversleigh area
of northwestern Queensland (recently reviewed
by Archer et al., 1989) are becoming widely
known as a source of excellently preserved ver-
tebrate fossils, ranging in age from the late
Oligocene to the Pleistocene, The limestone
mutrix has enabled acid preparation of delicate
bones (e.g. bat skulls) in perfect condition and
many thousands of specimens have already been
recovered. The detailed preservation and the evi-
dent terrestrial nature of some of the faunas are
factors suggesting that small reptiles such as
lizards, which normally fossilise poorly, would be
preserved, and such is the case. Members of the
families Agamidae, Varanidae, Gekkonidae and
Scincidae have been identified in the collections
from Riversleigh, An initial survey of the agamids
has just been completed (Covacevich et al., 1990)
and we report here on one of the scincid lizards.
The Scincidae is the most diverse family in the
moder Australian fauna. Ttincludes many highly
characteristic forms, suggesting a long period of
evolution in Australia about which almost nothing
is known. The Riversleigh deposits are the first
from the Australian Tertiary known to be rich in
fossil skinks (Hutchinson, 1992),

To date, most of the remains identified as lizards
are partial or fragmentary lower jaws. These are
difficult to idenufy because there are too few data
concerning taxonomic variation in skeletal char-
acters of the very diverse modern Australopapuan
fauna. However, one of the most distinctive
Australian genera, Tiligua (s.1.), has long been
recognised as having unusual teeth (e.g. Smith,
1937), with several of the posterior cheek teeth
markedly enlarged and having conical or rounded
crowns, Shea (1990) described the Tiliqua
lineage: a tevised Tiligua (including

Trachydosaurus), pilus Cyclodomorphus, the lat-
ter including the slender species gerrardii,
casiiarinage and the branehialis complex. Both
genera have characteristic dental morphology
which allows iselated dental arcades to be iden-
tified to species (Archer and Brayshaw, 1978;
Bartholomai, 1977; Shea, pers, obs.),

Amongst the material from Riversleigh are twu
partial dentaries with the enlarged, round-
crowned teeth of the Tiligua lineage, These
specimens are here described as a new species of
Tiligua, the first extinct member of the genus to
be described.

MATERIALS AND METHODS

Prepared fossil specimens of lizard jaws were
provided by Di M, Archer, University of N.S, W-
These had been sorted from concentrate obtained
by the acid processing of limestone blocks from
Riversleigh Station, northwest Queensland. Ref-
erence material of modern taxa was obtained from
the the following collections: American Museum
of Natural History, New York; Australian
Museum, Sydney; Queensland Museum, Brisbane
(QM); Queen Victoria Museum, Launceston;
South Australian Museum, Adelaide (SAMA):
Western Australian Museum, Perth; Zoologisches
Museum, Humboldt Universitit, Berlin.
Specimens were examined, measured and drawn
using a Wild M3¥Z stereomicroscope with an
eyepiece micrometer and drawing tube.

SYSTEMATICS
Family SCINCIDAE
Subfamily LYGOSOMINAE Mittleman, 1952

Tiliqua Gray, 1825

Mer
Tiliqua pusilla sp. nov.
(Figs 1-2)
MATERIAL EXAMINED

Howorree. QM F16876 (formerly M. Archer collec-
tion neo. ARS337#)-

PARATYPE. QM FIG877 (formerly M, Archer collec-
tion no. AR4183).

Tyee Locatiry: Gag Jocality, one of the lowest of the
Tertiary System C group of sites on the Gay Plateau,
Riversleigh Staion, Queensland (precise detuils of site
available from University of NSW Vertebrate Palacon-
tology Laboratory). Matrix of freshwater limestone,
The Gag site has yielded the Dwornamor local fauna
and is of probably early Middle Miocene age (c.15 Myr
BP; Ascher ct al,, 1989),

DIAGNOSIS

A small species (estimated dentary tooth row
length of the holotype, Ymm), cheek teeth en-
larged, with crowns blunt and slightly rounded in
lingual profile, more conical in mesial profile,
apices bearing a shallow central pit from which
radiate striae.

DESCRIPTION

Hovoryee; (Fig, | a-b) A partial left dentary,
missing its rostral third and the caudal, non-tooth
bearing extremily, including all of the coronoid
process and the apex and ventral margin of the
angular process, Meckelian groove closed, no
trace of suture. Rostral end of ventral margin with
a low ridge, presumably the caudoventral ex-
tremity of the symphysial crest. Apex of splenial
notch (rostral margin of inferior alveolar foramen)
lies at midpoint of interval between coronoid bor-
der of dorsal medial lamina and caudal margin of
syphysial crest, Coronoid margin of dorsal
medial lamina obtusely angulate, Lateral surface
with a deep, well-defined arcuate muscle scar
caudal to level of last tooth, presumably for the M.
adductor mandibulae externus superficialis
(AMES) (part IV, Wineski and Gans, 1984).

Internal septum fused ventrally along entire
length to ventral medial lamina, completely
separating infenor alveolar canal from Meckelian
canal. Caudal margin of internal septum with two
notches (caudedorsal and sostroventral) for ar-
ticulation with surangular,

Labial wall of dental sulcus tall and robust,
‘ingual wall almost absent, but lingual margin
well-defined and sharply angulate. Dentition
pleurodont. Seven closely-packed teeth of even
occlusal height and two unfilled tooth loci present,
in order from mesial to distal: a tooth locus for a

MEMOIRS OF THE QUEENSLAND MUSEUM

similar-sized tooth to those immediately caudal,
at the Jevel of the caudal extent to the ventral
symphysial crest; a tooth, missing crown, with a
small resorption pit at base; an intact, unresorbed
tooth; a tooth largely resorbed at the base, but with
intact crown; a tooth with a small resorption pit
basally, but with mesial margin of crown missing,
a very large empty tooth locus for a tooth ap-
proximately twice the diameter of the preceding
teeth) a large tooth (only slightly Jess in diameter
than the preceding locus) with intact crowa, but
with @ large resorption pit basally; two intact teeth.
the second with a small resorption pit, both abrupt-
ly narrower and shorter (occlusal surface lower)
than the preceding tooth. Tooth crowns bluntly
rounded in lingual profile, slightly more conical
in mesial profile, the largest extant tooth even
more flattened; mesial teeth with a shallow apical
notch, from which striae radiate towards the
periphery of the crown; similar, shallower striac
present un more calidal teeth.

PARATYPE: (Fig. 1 c-d) The entire rostral half of
aleftdentary, broken at the level of the 11(h tooth
and apex of splenial notch, but with most teeth
either broken or missing. The portion overlapping
with the holotype (tooth loct 5-11) is similar to it
in all respects, but slightly smaller. In dorsal view,
dentary nearly straight. Symphysis fattened,
angle between symphysial surface and lingual sur-
face obtuse, slightly rounded. Angle between
mandibles (as estimated by apposition of a mirror
image) 41° internally, 50° externally.

Lateral surface with six irregularly spaced men-
tal foramina. Teeth and loci are present as follows,
from mesial to distal: Five teeth, obliquely frac-
tured at base to leave only the implanted portion,
the fifth Jevel with the end of the ventral sym-
physial crest; a tooth missing lateral surface of
crown; an empty tooth locus of simular size to the
preceding teeth; a tooth broken transyersely al
level of margin of mandible, missing entire crown;
an entire, unworn tooth with a fracture line below
the crown; a very large emply tooth locus for a
tooth approximately twice the diameter of the
preceding teeth, and a large tooth stump, similar
in diumeter to the preceding empty tooth locus,
missing crown, Resorption pits are present in the
bases of the second, fifth, eighth and eleventh
teeth. The single intact tooth (ninth) slanting
labiully about 30° trom vertical, Im mesial profile,
lingual wall nearly straight, labial wall of crown
slightly bulbous, apex moderately narrow, coni-
cal, but with a prominen\ mesiocistal apical notch,
In lingual view, crown more rounded, slightly

NEW MIOCENE SKINK

305

FIG. 1. Tiliqua pusilla sp. nov. Holotype (QM F16876, A-B) and paratype (QM F16877, C-D) partial left dentaries
in lingual (A, C) and labial (B, D) views. Scale bar equals 5mm. Abbreviations: ds, dental sulcus; iaf, inferior
alveolar foramen; is, internal septum; mames, scar for mandibular adductor musclulature; mf, mental foramina,
sc, symphysial crest; sn, splenial notch; numbers indicate tooth positions.

higher distally. Crown with radiating striae in
occlusal view.

ETYMOLOGY
From pusilla, Latin adjective meaning very
small.

COMPARISON WITH OTHER TAXA

The closed Meckelian groove of T. pusilla is a
synapomorphy of both the Egernia and Eugen-
gylus groups within the subfamily Lygosominae
(Greer, 1979). Within these two groups, presence
of enlarged teeth in the caudal part of the dental
arcade, with the tenth or subsequent dentary tooth

largest, is a synapomorphy of the Tiliqua lineage
(Shea, 1990).

Both specimens are nearly identical in size and
morphology and presumably come from different
individuals of the same species. The difference in
shape of the tooth crowns in mesial profile is
probably due to wear in the holotype. The paratype
is complete enough to determine that the tenth and
(especially) the eleventh teeth are enlarged and, by
matching the positions of the symphysial crest and
inferior alveolar foramen, it is possible to overlay
the two specimens to show that the large tooth
preserved in the holotype is the twelfth. Figure 2
shows a composite restoration of the mandible.

Tiliqua pusilla differs from all but one of the

FIG. 2, Tilique pusilla sp. nov. Diagrammatic restora-
tion of left dentary (A, labial view. B, lingual view)
based on holotype and paratype specimens. Scale bar
Smm.

modem species of Tiligua in its lesser size and also
in tooth shape. With an estimated dentary tooth
row length of 9mm, the holotype is smaller than
even neonates of all Tiliqua except T, adelaidensis
(neonate dentary tooth row length of other Tiligua
>10mm,; neonate dentary tooth row length of T.
ddelaidensis not known, but adult dentary tooth
row length of T. adelaidensis (8.2-9. 3mm, n =3).
Based on the proportions of living members of the
Tiligua lineage, this tooth row length indicates a
midline skull length of no more than 20mm. Of
the seven other species of Tiligua, T. nigrolutea,
T. occipitalis and T. rugosa (Fig, 3 a-c) have
conical tooth crowns in both lingual and mesial
profiles, with a cutting edge directed obliquely
across the crown, T. multifasciata (Pig. 3d), while
haying lower, broader teeth, also has. a conical
apex to the crown, but with a pronounced apical
notch. Tiliqua adelaidensis (Pig. 3¢), while more
similar in size to 7. pusilfa has teeth with parallel
sides, capped by u low, angulate crown jn lingual
profile, and slightly compressed in mesiodistal
view. Tilique scincaides and 7) gigas (Fig. 3f-e)
five rounded crowns to the largest teeth and
slightly more conical crowns to the more mesial
tceth, but in both species a low crest obliquely
crosses the occlusal surface. In T pusilla, a dis-
hinct notch 1s present, both on the single unwom
tooth of the paratype, and the slighty more wom
tecth of the holotype-

Tiliqua pusillaalso differs mm dental morphology
from Cyclodomerphus. The teeth of the C.
branchialis complex (Pig, 36-1) and C. casvarinae

MEMOIRS OF THE QUEENSLAND MUSEUM

(Fig, 3}) have more acutely conical crowns, with
the apex towards the distal margin in lingual
profile, or even slightly distally recurved. The
other two Cyclodomorphus species, C. gerrardii
(Fig. 3k) and C. maximus, have very rounded
crowns to the largest teeth, In C. gerrardii, the
teeth are more widely spaced, and the single en-
larged tooth is both grossly larger than adjacent
teeth, and much lower than in 7. pusilla, In C.
maximus, the teeth lack the apical notch and are
relatively narrower relative to their height. Fur-
ther, the mandible of C. maximus is relatively
more gracile, which is reflected by the much nar-
rower splenial notch in the caudal part of the
dentary, and the dentary is larger overall,

Our placement of the species in Tiliqua rather
than Cyclodomorphus is based on one sy napomor-
phy and one character of uncertain polarity. Tili-
qua pusilla, like modern Tiliqua, has a long, low,
ventral symphysial crest, extending caudally to
the level of the fifth tooth. Cyelodomorphus and
the closest outgroup to the Tiliqua lineage, Eger-
nia (Shea, 1990), have amuch shorter crest, which
may project medially giving a hooked outline.
Secondly, the angulate nature of the coronoid
margin of the dorsal medial lamina is similar to
that seen in modern Til/iqua, where it is due to
enhancement of a medial buttress on the rostral
process of the coronoid. In most Cyclodamarphus
and Egernia the buttress is less developed and
more oblique, leaving a more evenly sloping
coronoid margin to the dentary.

Tiliqua pusilla ig unusual compared with living
Tiliqua in \wo features: its small size and the low
number of dentary teeth, Both small adult size and
reduced number of dentary teeth are common in
Cyclodomorphus. Thus an explanation for the
similarity between 7. pusilla and Cyclodemar-
phus is needed. The first possibility is that the
similarity in size and reduced tooth number is
because the specimens of T. pusjlia are juveniles,
However, the specimens are clearly not juvenile
as they have a well-defined dental sulevs with a
sharply angulate lingual margin, two large teeth
ahd the more mesial teeth subequal in size and
showing several cycles of tooth replacement. In
juveniles of all Tiligua and Cyclodomorphus
species, the dental sulcus is poorly-defined, its
Jingual margin blending with the adjacent dorsal
medial lamina. There is only a single large tooth,
and the teeth mesial to this one show z distinct
alternation in size (Shea, MS).

Since both specimens are adult, the number of
dentary teeth is lower than for adults of other
Tiligua species, which have more than 14 dentary

NEW MIOCENE SKINK 307

FIG. 3. Lingual views of dentaries of living species of the Tiliqua lineage showing variation in proportions, tooth
number and shape. A and H are isolated dentaries showing the internal structure and E lacks the coronoid; the
remainder are intact mandibles. A, T. rugosa (SAMA R27584). B, T. eccipitalis (SAMA R35758). C, T.
nigrolutea (SAMA R2725). D, T. multifasciata (SAMA R35757). E, T. adelaidensis (SAMA R8589). F, T.
scincoides (SAMA R27037).G, T. gigas (SAMA R11419). H, Cyclodomorphus cf branchialis (SAMA R38021).
I, C. melanops (SAMA R37565). J, C. casuarinae (SAMA R35682). K, C. gerrardii (SAMA R35761). Scale

bars 5mm.

teeth, and show a progressive distal shift im the
number of (he largest Looth with age, as estimated
by increasing size (Shea, MS). In the Tiliqua
lineage the two attnbutes (small size, reduced
teoth count) are probably linked since in order to
have teeth of sufficient absolute size, small
species of the Tiliqua lineage may have to make
do with fewer teeth overall in cach dental arcade.
Compared with the similarly-sized T° adelaiden-
sis, T. pusilla has larger but fewer tecth (14, 11-12
largest, versus 16, 12-14 largest) and a more
heavily-bailt jaw, and was evidently more strictly
adapted for durophagy than T. adelaidensis,
which has thinner, sharper-crowned teeth, This
comparison supports the idea that the low number
of teeth in T. pusillais aconsequence of allometric
sealing, with a limit to reduction in tooth size of
crushing teeth imposed by the necessity of
preserving tooth strength.

DISCUSSION

Tiligua is one of the most charactonstically
Australian scincid genera. Morphologically
(Greer, 1970, 1979; Shea, 1990), biochemically
(Hutchinson, 1981; Baverstock and Donnellan,
1990) and karyologically (Donnellan, 1985), the
only living relatives of Tiligva are Cyclodomer-
piues, Egernica and Corucia, collectively compris-
ing the Egernia group (Greer, 1979), Little is
known of the history of Tifigue (or of any other
skink genus) in Australia,

Tiliqua has been recorded numerous times from
Pleistocene ta Recent deposits (Krelft. 1867,
1870, 1871; Stirling, 1889; Hale and Tindale,
1930; Tindale, 1933; Cook, 1960, 1963; Mul-
vaney, 1960; Mulvaney et al., 1964; Trezise,
1970; Lampert, 1971; Thome, 1971; Marshall,
1973: Bowdler, 1974; Ryder, 1974; M. J. Smith,
1976, 1982; Bartholomai, 1977; Hope et al,, 1977;
Archer and Brayshaw, 1978; Molnar, 1978, 1982;
Williams, 1980; Hope, 1981; M.A, Smith, 1982:
Wells ctal., 1984) butis almostunknown from the
Teruary. There is one record from the Pliocene
Wellington Caves (Hand et al, 1988) and one
from the Miocene, based on a statement by Estes
(1984) that a specimen of Tiligua had been iden-
lified by J. Gauthier among the Kutjamarpu local
fauna (Middle Miocene of northern South
Australia; Stirton et al., 1967). Both records
remain undescribed, The Kutjamarpu local fauna
is considered {Archer ct al., 1989) to be just older
than the Dwornamor local fauna, making the
former the the oldest record of the Tilique lincage.
The existence of this specimen in South Australia
and T. pusilla in Queensland shows that the Ti/i-

MEMOIRS OF THE QUEENSLAND MUSEUM

gua lineage was Widespread in Australia by the
Middle Miotene.

Identification of the Riversscigh species as Tili-
gua rather than Cyclodomorphus suggests that
these sister taxa had diverged prior to the Middle
Miocene, pushing the diversification and ultimate
origin of the Egernia group even further back,
potentially to pre-Miocene times. This scenano 1s
incompatible with a previous hypothesis for Eger-
nia of rapid Pleistocene speciation following a
Pliocene origin (Horton, 1972), A preliminary
dating via the ‘albumin clock” of around 12 Myr
BP for the Egernia- Tiliqua split (Baverstock and
Donnellan, 1990) ts thus also too recent.

The mammal fauna of the Dwornamor local
fauna is dominated by arboreal browsers and has
avery high diversity (Hund, 1985; Archer & Flan-
nery, 1985; Flannery & Archer, 19874; 1987b;
Archer et al., 1988) with minimally 58 species
present (Archer et al,, 1989, Table 1). Such civer-
sity would today be found only in a rainforest
environment, which has supported the idea of the
Dwornamor local fauna os # tropical rainforest
fauna, With the exception of T. gigas, living
species of Tiliqua do not inhabit rainforests, so
that 7. pusilla, combining small size and rain-
forest-dwelling hubits, represents a radiation of
the genus with no modern parallel, Possibly Tii-
gia Wis a more diverse laxon in the past, and/or
modem bluctongues represent a recent open-
country radiation descended from dwarfish rain-
forest ancestors.

ACKNOWLEDGEMENTS

MNH thanks M. Archer for the opportunity of
describing the Riversicigh skinks. We thank R-G_
Zweite), ALE. Greer, R. Sadlier, ). Covacevich, R-
Green, G. Stor and R. Giinther who provided
access to material in their care, and gave permis-
sion to extract skulls where necessary. A. E. Greer,
H, Godthelp and R, Sadlier improved the
manuscript With their coniments. Fig, | was
prepared by K. Bowshall-Hill. 8. Donnellan
provided useful information on New Guinean
rainforest skinks. Fieldwork by Archer and
coworkers at Riversleigh has been funded from
the following sources: Australian Research Grants
Scheme; Australian Depatiment of Arts, Sport,
the Environment, Tourism and Territories; Na-
tional Estate Programme Grants Scheme; Univer-
sity of New South Wales: Wang Computer Pty
Ltd; Australian Geographic Pty Ltd; Mount Isa
Mines Pty Lid; The Queensland Museum) the
Australian Museum: The Royal Zoological

NEW MIOCENE SKINK

Society of NSW; the Linnean Society of NSW;
Ansett/Wridgways Pty Ltd; MtIsa Shire Council;
The Riversleigh Society and Friends of Rivers-

leigh.
LITERATURE CITED

ARCHER, M. AND BRAYSHAW, H. 1978. Recent
local faunas from excavations al Herveys Range,
Kennedy, Jourama and Mount Roundback, north-
eastern Queensland. Memoirs of the Queensland
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ARCHER, M, AND FLANNERY, T. F. 1985. Revision
of the extinct gigantic rat Kangaroos (Potoroidae:
Marsupialia), with a description of anew Miocene
genus and species and a new Pleistocene species
of Propleopus, Iournal of Palaeontology 59:
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ARCHER, M., HAND, 8. J. AND GODTHELP, H.
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ARCHER, M. GODTHELP, H., HAND, S. J. AND
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29-65.

BARTHOLOMAI, A, 1977, The fossil vertebrate fauna
from Pleistocene deposits at Cement Mills, Gare,
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BAVERSTOCK, P. R. AND DONNELLAN, S. C.
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BOWDLER, S. 1974. An account of an archaeological
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COOK, D. L, 1960, Some animal remains Jound in
caves near Margaret River. Western Ausiralian
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1963. The fossil vertebrate fauna of Strong's Cave,
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COVACEVICH, J., COUPER, P., MOLNAR, R. E.,
WITTEN, G. AND YOUNG, W, 1990, Miocene
dragons from Riversicigh: new data on the history
of the family Agamidae (Reptilia: Squamata) in
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29; 339-360.

DONNELLAN, S. G. 1985. The evolution of sex
chromosomes in scincid lizards, (Unpublished
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ESTES, R. 1984, Fish amphibians and reptiles from the

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FLANNERY, T.F. AND ARCHER, M. 1987a.
Sirigocuscus reidi and Trichosurus dicksani, Wo
new fossil phalangerids (Marsupiatia:
Phalangeroidea) from the Miocene of
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cher, M. (ed.), ‘Possums and opossums: studics it
evolution’. (Surrey Beatty and Sons and the Kayal
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FLANNERY, T. F. AND ARCHER, M. 1987b. Hyp-
siprymnodon barthelomaji (Potoroidac: Mar-
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PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

GARY D. WEBSTER AND PETER A, JELL

Webster, G.D. & Jell, P.A, 1992 06 29: Permian echinoderms from Western Australia.
Memioirs of the Queensland Museum 32(1): 311-373. Brisbane, ISSN 0079-8835.

A new study of the Permian echinoderms of Western Australia included field collection and
restudy of several earlier collections, Significant morphologic and stratigraphic details are
provided about previously described taxa. New taxa are the crinoids: Neocamptecrinus
barrabiddyensis,N. millyitensis, Stomiocrinus merlinieighensis, Neoplaiycrinuscallyiharaere:
sis, N, miamiaensis, Wrightocrinus woeramelensis, Litocrinus pansus, L. projuberans,
Lampodosverinus variabilis, Cosmetocrinus? middalyaensis, Spheniscocrinus australis, Min-
ilyacrinas williamburyensis, Texacrinus geachensis, T. hardmanensix, T. gascoynensis,
Galateacrinus australis, Cymbiocrinus cherrabunensis, Jimbacrinus donnellyensis, and
Baryschiyr? carnarvonensis and the blastoids: Neosthisma extensum, and Rkopaloblastis
cuspidus and the stellaroid Monaster coleyi. Echinoid remains are recorded for the first time
in the Permian of Western Australia, Ranges of several Carboniferous crinoids are extended
into the Permian, including a coelocrinid and pentameric steni-bearing forms. The youngest
PaJacozoic crinoids known are described from the Tatarian Hardman Formation of the Canning
Basin.

Echinoderms in the Callythara Formation include 16 camerales, 37 inadunates, and 1 flexible
among the crinoids, 5 blastoids, and | echinoid. Younger Permian strata yielded 3 camerates,
12 inadunates, | blastoid, and 1 starfish. Lineages of Neocamprocrinus, Neaplarycrinus,
Texacrinus, Jimbacrinus, Tapinocrinus, and Stomiocrinus are recognised and offer potential
for additional correlation of Permian strata throughout Western Australia. [7] palacontelogy,
Crinaids, Permian, Western Australia.

Gary D. Webster, Depariment af Geology, Washington State University, Pullman, Washingion
99364-2812. USA; Peter A. Jell, Queensland Museum, PO Box 3300, Souilt Brisbane,

Qiieensland 4101; 20 December, 1991.

Echinoderms from the Permian of Wester
Australia were first recorded from crinoid stems
by Foord (1890) whe described them from the
Gascoyne River bed and by Etheridge (1903) who
reported an endophytic fungal infestation of some
crinoid columnals from the northern Carmmarvon
Basin. Teichert (1949, 1954) and Webster (1987,
1990) on crinoids, Breimer & Macurda (1972) on
blastoids and Kesling (1969) on starfish provide
the only detailed works on the fauna to date;
Teichert (1951) gave a useful summary of the
fauna as he knew itthen. This paper describes new
crinoids and blastoids as well as providing much
new information on known species and recording
echinoid remains for the first time. Fossil
echinoderms are known from the Permian of the
Perth, Carnarvon (previously referred to as the
Northwest Basin and Canning Basins with the
oldest occurring in the Sakmarian to Artinskian of
the Carnarvon Basin and the youngest in the
Tatarian of the Canning Basin, Echinoderm fossils
dealt with herein include a crinoid cup from the
Fossil Cliffs Formation of the Perth Basin, and a
few crinoid and blastoid specimens from the

Noonkanbah and Hardman Formations of the
Canning Basin but principally derive from the
Camarvon Basin where the Callytharra Formation
has yielded the most prolific echinoderm fauna.
Permian geology and stratigraphy of Western
Australia has been detailed through the regional
mapping efforts, since the late 1940s, of the
Bureau of Mineral Resources and the Geological
Survey of Wester Australia in the 1:250,000
geological map series of the BMR. These results
owe some of their success to earlier reconnais-
sance work of several people but in the context of
Permian palaeontology that of Teichert was most
significant, He contributed numerous papers and
reporis but none more significant than his
landmark study (Teichert, 1949) on the crinoid
Calceolispongia and biostratigraphic utility of its
distinctive species. This work provided an in-
trabasinal biostratigraphic scheme for the Carnar-
von Basin which remains virtually unchanged
today. Studies of the Ammonoidea (several papers
listed in Quilty, 1975) have been used to provide
an international correlation. Summaries of the
geology and stratigraphy were provided by the

312

Geological Survey of Western Australia (Playford
et al., 1975), for the Carnarvon Basin by Condon
(1954, 1967), and for the Canning Basin by
Veevers & Wells (1961). We have confined our
study to palaeontology and relied heavily upon
these earlier works as a framework within which
to place our findings. A generalised locality map
and stratigraphic columns of the Carnarvon and
Canning Basins were given in Webster (1990).

PRESERVATION

At Callytharra Springs most echinoderm
material is disarticulated except for microcrinoids,
microblastoids, and pleuricolumnal sections of
stems. A few exceptions of articulated cups of
crinoids and blastoids are known but disarticu-
lated plates of these species are also found as-
sociated. Associated but disarticulated cup plates,
brachials and columnals of particular species are
present on the outcrop indicating minor displace-
ment but definite disaggregation by currents or
bioturbation. Distortion and fracturing, resulting
from compaction, is most common in large rela-
tively thin ossicles or thinner parts of more robust
faunal elements. This is most noticeable on large
camerates such as Neoplatycrinus and the stem of
Neocamp-tocrinus, a result of the ease of seeing
such features on the larger faunal elements. Care-
ful collecting and microscopic investigation show
that this also occurs in smaller elements of the
fauna,

To the north, exposures of the Callytharra For-
mation from the vicinity of Mt Sandiman through
Williambury Station and into the Gooch Range
show carbonate grainstones with minor marls and
shales forming hogback ridges with the formation
tilted, usually to the west and eroded. Large
pluricolumnals, dominated by Baryschyr? car-
narvonensis sp. nov. up to 10cm long, are common
in the lower part of the Callytharra Formation.
Articulated crinoid cups and crowns, although
rather infrequently found, occur on bedding sur-
faces. Specimens are exceedingly difficult to ex-
tract because they are the same hardness as the
enclosing grainstones. Most specimens tend to
weather at the same rate as the matrix and solution
weathering of ossicles often precludes identifica-
tion. The articulated crowns, often with proximal
stem attached, deny postmortem transportation.
Most are considered to have died in situ; the cause
of death is not apparent. The matrix shows no
graded bedding that could imply debris flows or
tempestites, but bioturbation is obvious in some
grainstones.

MEMOIRS OF THE QUEENSLAND MUSEUM

Considerable time walking bedding planes and
looking at weathered scree, 1s required to find the
rare good specimen. Rarely, sets of arms and
crowns are found weathering from thin interbeds
of shale of marl. We suggest that numerous crinoid
species are to be found in these northern exposures
of the Callytharra Formation with careful and
patient collecting.

FAUNAL ANALYSIS

The faunal list of our collection from the Cal-
lytharra Formation (Table 1) shows that we found
most of the species recorded by Webster (1987)
except for Tapinocrinus macurdai, T. ingrami, T.
sp., Actinocrinites sp. cf. A. brouweri and a few of
the unidentified camerate and inadunate radials.

The Callytharra fauna contains 16 camerate
species, 37 inadunates and 1 flexible. As briefly
summarised by Webster (1987), most Permian
crinoid faunas worldwide are dominated by in-
adunates. Only the Timor and possibly Bolivian
faunas contain significant camerates and flexibles.
The Callytharra fauna is the second most diverse
Permian fauna in the world after Timor but the
virtual lack of flexibles leads us to suggest that the
flexibles could not adapt to the cooler water en-
vironment of the Australian Permian.

In this paper 2 camerates and 12 inadunates are
added to the faunal list as outlined by Webster
(1987). These include three holdover genera, a
coelocrinid (Dorycrinus), barycrinid indet., and
Cosmetocrinus, not previously recorded from the
Permian (Moore & Teichert, 1978). The youngest
possible ancestors are to be found in the Early
Carboniferous of North America. The new in-
adunates include Minilyacrinus gen. nov. and new
species of Lampadosocrinus, Texacrinus,
Galateacrinus and Baryschyr? not previously
recorded in the region. Only Texacrinus was pre-
viously reported from the Permian. The
microcrinoids described are the first described
from Australia. Wrightocrinus, Litocrinus and
Timorechinidae indet. are known from Timor thus
strengthening faunal links between the two areas.

The Basleo fauna of Timor is the most diverse
Permian crinoid fauna known in the world. It
contains 232 species as opposed to the 56 species
so far recorded in the Callytharra Formation. Al-
though study of the Timorese collections housed
in European museums leads one of us (GDW) to
believe that restudy of the inadunates may reduce
the number of species recognised it will remain an
incredibly diverse fauna. Teichert (1951) con-

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA 313
QML757 | QML758 ime QML760 ML761 | 757-761

indet. platycrinitid or dichocrinid radial 1 1
Actinocrinites cf. A. brouweri Sa ee a
Platycrinites wrighti =e
Neoplatycrinus callytharaensis 6 8
Pleurocrinus? sp. -radials (basals) 25(16 2 4 = ie
Neocamptocrinus sp. 8 17 32 =
indet. dichocrinid radials 1-3 17 5 5 ioe ss af
indet. camerate radials 1-5 10 6 : 4
Notocatillocrinus callytharaensis (partials) |__ 2(36 2(1 10 2) 111)
Synbathocrinus campanulatus (partials) 9 1(2
Synbathocrinus constrictus (partials) 2(6) 1
Cydonocrinus cf. C. tuberculatus partial 1 = =m
Tapinocrinus macurdai
Tapinocrinus ingrami
Tapinocrinus sp.
Parabursacrinus magnificus (radials) (150) 67 77 23) 1 86)
| Parabursacrinus granulatus radials 15
Parabursacrinus nefotassiensis radials 16 7
Calceolispongia spinosa basals 10
Calceolispongia cf. C. spinosa
Calceolispongia digitata basals 10 _| 4 | 6
Calceolispongiidae sp. indet.
indet inadunate radials 1-10 45 9 24 5 3 13
indet. flexible radial 1
| Apographiocrinus pumilus brachials 1 2 2

Phosphannulus pluricolumnals
Radials (unidentified)

| Tegmen spines
Radials with ribs

Neocamptocrinus radials

Infrabasal or basal circlet

Columnals (unidentified)

Neoschisma extensum plates

Notoblastus stellaris plates

Barycrinidae? indet. pentameric 15
pluricolumnals
Neoschisma verrucosum plates 4 1 | |

TABLE 1. Fauna of type section of Callythara Formation west of Callythara Springs on Wooramel River giving
specimen numbers from field collections by the authors. Numbers in parentheses refer to partial cups or specific
cup plates where both complete and partial cups were found.

sidered the Western Australian fauna closely al- Australian fauna as an impoverished Timor fauna
lied to that of Timor but, with the exception of | without indicating any reason for the impoverish-
Calceolispongia he regarded the Western ment. We suggest that the Timor fauna lived in a

314

Bulgadoo
Cundlego
Wandagee

oo
r=
a
=
Gj
O

Quinnanie Sh.

Neocampiocrinus millyitensis
Neocamptocrinus occidentalis

Neocampto. barrabiddyensis

MEMOIRS OF THE QUEENSLAND MUSEUM

The post Callytharra crinoids of
Western Australia, following the
work of Teichert (1949, 1954),
Webster (1990) and this paper, num-
ber 3 camerate and 12 inadunate
genera. These forms occur mainly in
sandstone or shale with rare excep-
tions such as Stomiocrinus merlin-
leighensis and Jimbacrinus

Noonkanbah
Cherrabun Mbr

| Fossil Cliffs

Stomiocrinus ferruginus

donnellyensis. Many occur in
clusters with multiple specimens of

Stomiocrinus merlinleighensis

Texacrinus hardmanensis

Texacrinus gascoynensis

few species along bedding planes.
Many specimens were disaggregated
in part indicating postmortem current

Texacrinus goochensis

or scavenger activity. Cross-bedding,
scour and fill and ripple marks were

Skaiocrinus granulosus
Jimbacrinus minilyaensis
Jimbacrinus bostocki
Jimbacrinus donnellyensis
Cymbiocrinus cherrabunensis

Trautscholdicrinus sp.

Occiducrinus australis

Rhenocrinidae gen. nov.

Eoindocrinus praecontignatus

xX
| | | tt tx] |

observed in these sandstones record-
ing current activity. Bioturbation is
evident but not extensively.
Crinoids of the Cherrabun Mem-
ber, Hardman Formation in the Can-
x |ning Basin (Table 2) are among the
youngest Palaeozoic crinoids known.
The age of this horizon, which also
yielded the ammonoid Cyclolobus
persulcatus Rothpletz, 1892, is dis-
cussed by Glenister et al. (1990) as

Tapinocrinus spinosus

Dzhulfian (i.e. within the Tatarian of

Family and genus indet.

Anechocrinus nalbiaensis

the European scale), All three species

represent lineages common in the
Carboniferous of North America.
Cymbiocrinus is known from the Per-

TABLE 2. Lineage chart and stratigraphic distribution of the mian of Thailand (Webster & Jell, in
echinoderm fauna of the post-Callytharra Fmn Permian of WA. Taxa prep.); Neocamptocrinus is known

from Teichert (1949) are not included as they were

well placed only from Australia (Willink, 1980;

stratigraphically in the original work. Callytharra through Wandagee Webster, 1990; herein).
Fmns are in stratigraphic sequence for the Carnarvon Basin. The Fossil
Cliffs Formation of the Perth Basin is temporally equivalent to the

Callytharra Fmn. The Noonkanbah Fmn (equivalent to the Coyrie
through Wandagee Fmns) and Tatarian Cherrabun Mbr, Hardman Fmn

are in the Canning Basin,

warm water environment favourable to a higher
carbonate productivity. In contrast the abundant
clastic sedimentation of the Western Australian
Permian and the type of sediment suggest not only
nearer shore, more turbid environments but also
cooler water environments with less carbonate
production, The cooler water environment of the
Permian deposits of Western Australia were
recognized by Condon (1967) and Dickins (1978)
among others. We consider the echinoderms of the
Western Australian Permian to be those that could
adapt better to the greater diversity of environ-
ments and note that a few are unknown in Timor
(Texacrinus, Cosmetocrinus and the coelocrinid).

LINEAGES

Teichert (1951) considered that
only one crinoid showed a lineage of stratigraphic
utility in Western Australia. Based on knowledge
at the time his view was correct. However, the
added information from the Callytharra and
younger fauna reveals several parallel lineages;
some may have biostratigraphic utility through
Western Australia, the rest of Australia and even
to Timor and India. The Calceolispongia lineage
detailed by Teichert (1949) remains the most use-
ful for Western Australia but it also needs revision
in the light of extensive new collections and its
potential for correlation in the Canning Basin has
yet to be addressed.

Other lineages that we recognise in Western

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

Australia (Table 2) are species within each of
Neocamptocrinus, Neoplatycrinus, Texacrinus,
Jimbacrinus, Tapinocrinus and Stomiacrinus. Al-
though these are considered independently it
should be noted that Neecamptocrinus,
Neoplatycrinus and Calceolispungia occur
together at most horizons m the Permian of
Western Australia suggesting that they needed
similar living conditions.

Stems of Neacamptocrinus are found in most
honzons with marine fossils from the Callytharra
to the Hardman Formations. They have not been
studied in detail. Specimens that we have ob-
served show greatest size in the Wandagee For-
mation (Artinskian) (Fig. 3M.N). Cups, thecae
and crowns of Neocamptocrinus are uncommon.
They are known from the Bulgadoo Shale N-
barrabiddyensis sp. nov., from the Cundlego For-
mation, N. occidentalis, and from the Hardman
Formation, W. millyitensis. These three species are
considered an evolutionary lineage with N. oc-
cidentalis evolving from N. barrabiddyensis by
elongation of the cup with nearly vertical walls
distally. Jn turn, N. millyitensis evolved from NV.
occidentalis with a more flattened basal cirelet,
reduction in the number of plates in the anal series
above the primanal and modification of the urna-
ment on tegmen plates. Webster (1990) discussed
the relationship of WV. eccidentalis and N. n. sp.
(=N_ millyitensis) with eastem Australian species
(Willink, 1980).

Stomiocrinus merlinieighensis m the Bulgadoo
Formation could be the ancestor of 8. ferraginus
in the Wandagee Formation by enlargement,
modification of the anms by shortening and thick-
ening the brachials and reduction of the relative
size oi the orals. Timor species of Stomiecrinus
are probably derived from S. merlinleighensis by
modification of oral plates and flattening of the
tepmen. Sfomiecrinus is unknown in eastern
Australia.

Evolution of Neeplatycrinus miamiaensis from
WN. callytharraensis by modification of the orna-
ment from one large prominent node to several
large nodes anc probably Natlening of the basal
circlet forming a low bowl-shaped cup. The
“twist” nodals and intemodals currently assigned
to Platyerinites/Neoplarycrinus spp. and com-
monly found associated with the disarticulated
plates of Necplatyerinus probably belong 10 the
genus, Unfortmnately they were probably static in
evolution during the Early Permian and show no
significant differences or evolutionary trends. Al-
though notreported, “twist” columnals are present
in the Oxtrack Formation of the Cracow urea,

315

Queensland, Additional study of the platycrinid
columnals of Australia is needed to fully evaluate
their stratigraphic potential.

The tapineerinids are known from Western
Australia and Timor. In Western Australia,
Tapinocrinas macurdai and T, ingrami ate known
from the Callytherra Formation with the former
the likely ancestor of T. spinosus from the
Wandagee Formation and Basleo fauna of Timor.
Development of T, spinosus was by flattening of
the cup into a disc shape. enlargement of the
radials into a blunt spine and loss of granular
ornament. Although the spine did not develop, 7°
timericus also lost the ornament and developed a
discoid cup. There i$ no record of Tapinocrinns in
easter Australia.

The texacrinids are represented in Western
Australia by four species starting with T. goocken-
sis in the Caliytharra Formation followed by the
more robust 7. gascoynensis in the Cundlego or
Wandagee Formation. No further form of this
heavier stock is known, Skaiocrinus granulosis is
a more elongate form with or without slightly
peneplenary radial facets and pinnules with lon-
gitudinal ridges. The youngest form 7!
hardmanensis is a slender elongate form which
may have evolved through T. goochensis from the
same stock as 8. granulosus. Texacrinids are un-
known in eastern Australia,

Teichert (1949) considered the origin of Cal-
cealispongia to be uncertain but perhaps derived
from the Cromyocrinidae, Moore et al. (in Moore
& Teichert, 1978) suggested that the Cal
ceohspongidae were derived through the Cym-
biocrinidse or Ampelocrinidae. We consider the
Cymbiocrinidae line to be most likely based on the
bowl-shaped cup, large single anal and cuneate
arms. The lineage of Jimbacrinus is not as clear as
that of Calceelispongia but may include a sc-
quence from Calecolispongidae? sp. (Webster,
1987) from the Callytharra Forrnation to J, deve
neltvensis in the Bulgadoo Shale to J. bostacks
Teichert, 1954 in the Cundlega Formation tw J
minilyaensis Webster, 1990 in the Wandagee For-
mation and possibly ta Cymbiecrinas cher-
rabunensis in the Hardman Formation.

Webster (1987) noted the resemblance of Cal-
ceohspongidae? sp. 10 the calceolispongids bur
considered differences in the lack of bulbous
basals and presence of an isotomous branching on
the third brachial in one ray. If the branching had
been on the secand brachial the specimen would
have beeo assigned to Cymbiocrinus. Cal-
ceolispongidae? sp. probably denived from the
sume ancestor as Calcealisnongia. J. dannellyen-

316

si may have evolved from Catceolispungidae? sp.
by thickening of the basals and loss of arm branch-
ing. In turn J, bostocki evolved by development of
the spines on the bulbous basals whereas J, nuni-
lyaensis has a reduction in the bulbous basals,
possibly representing a different lineage from Cal-
ceolispongidac? sp. Cymbiocrinus cherrabunen-
sis is related to the calceolispongids mostly by the
bulbous basals; the arm branching pattern, how-
ever, is not typical of that family.

The second largest Permian crinoid fauna of the
Camaryon Basin occurs in the Wandagee Forma-
tion in the east limb of a syncline on the north bank
of the Minilya River, approximately 1Okm
westnorthwest of Wandagee Homestead. This
locality is the type locality for several species of
Calceolispongia {Tetchert,1949) and other
crinoids {Webster.1990). Some confusion about
the stratigraphy has resulted from nomenclatural
changes, Teichert (1949) considered the
Wandagee Formation to include the unit later
named the Quinnanie Shale, Thus some of his
material is reported as from the upper Wandagee
on locality information at the University of
Western Australia when jt is actually from the
lower Wandagee Formation as defined by Condon
(1967) (see Material section under Occiducrinuy
australis and Skaiocrinus granulosus (herein;
Webster, 1990),

There has also been some confusion about the
locality, Webster (1990), in citing locality data
from the University of Westem Australia, listed
the locality as cast of Conikilya Pool. Teichert
(pers. comm. 1990 to GDW) corrected this to west
of Coolkilya Pool as shown on the Winning Pool-
Minilya 1:250 000 Geological map (sheet SGS0-
13, 49-16, 1984), When we stopped at Wandagee
Homestead the directions we received lu
Coolkilya Pool took us further west than the loca-
tion shown on maps. It is obvious, When on the
sile, that the size and location of the pool or pools
along the river bed shift with scour and fill during
major minolf and evaporation and infiltration
during subsequent dry seasons, Coolkilya Poo! as
designated on the map and as referred to by the
locals are not the same. Teichert apparently used
the local designation in his original freld locations
and recognized they differed when trying to
reconcile the Bureau of Mineral Resources map
Jogatinns.

Tn the base of the Wandagee Formation as
defined by Condon (1967) Ovciduecrimes
australis, Eoindocrinus praecontignalus, Cal-
cealispongia ghundans, and Neocamptocrinus sp.
wecur along the sume bedding surface,

MEMOIRS OF THE QUEENSLAND MUSEUM

Tupinecrinus spinosus, Skaiocrinus granulosus
and Jimbacrinus minilyaensis occur at or close to
the same horizon. Stomiocrinus ferruginus may
also be from the equivalent horizon along the
Lyndon River to the north. Several species of
Calceolispongia occur at higher levels in the
aaa Formation as described by Teichert
(1949).

AGE AND CORRELATION

Discussion of the age and correlation of the
Callythara Formation at its type section was given
by Webster (1987), We accept the late Sakmarian
age for the base of the Callytharra Formation
based on ammaonoids (Glenister & Furnish, 1961),
New taxa reported herein do not alter Webster's
(1987) interpretation that the crinoids suggest a
slightly younger age of early Artinskian for the
Callytharra Formation. The Sakmarian-Ar-
tinskian boundary may be within the lower part. of
the Callytharra Formation,

Notiocatillocrinus callytharraensis, Synbath-
vcritus constrictus, Neoplatycrinus callytar
raensis ahd Neocamptocrinus sp, in the
Callytharra Formation throughout the Camarvon
Basin suggests a similar age for the unit over this
extent, Correlation of the Callytharra Formation
with the Fossil Cliffs Formation of the Perth
Basin, and Nura Nura Member of the Poole
Sandstone in the Canning Basin were discussed by
Teichert (1951) and Condon (1967), Webster
(1987) discussed the crinoid correlation of the
Callytharra Formation with that of eastern
Australian faunas reported by McKellar (1969)
and Willink (1978). The late Sakanarian-early Ar-
tinskian limestone interval in the basins of
Westen Australia, in the Oxtrack Formation in
Queensland, and the Berriedale Limestone and
limestone in the Crinoidal Zone of Tasmania rep-
resent a major sca level rise and change to warmer
conditions in the Australian part of the Tethyan
equivalent to the Sterlitamak and possibly Bir-
shevian cycles (Ross & Ross, 1985),

Correlation with (he Permian crinoids of Timor
remains somewhat of a problem, The crinoid bear-
ing Maubisse: Formation limestones of northern
Timor were imlerpreted as reefs and thought to
have been thrust onto the autochthonons non-
erinoida! Permian strata of southern Timor (Aud-
ley-Charles,1965,1968). He considered the
Timor Permian deposits to have formed in a
tmugh to the northwest of Australia. Charlton
(1989) suggested that the northem part of Timor
was allochthonous but thought much of the struc-

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA 317

FIG. 1. Coelocrinid indet. QMF21428. A, lateral view showing set of arms with distal part of spinose anal tube at
upper right, 1. B, incurved distal parts of flattened arms and spinose anal tube, X 1.6.

318

tural complexity was best explained by wrench
faulting , not by vertical movements as postulated
by Chamalaun and Grady (1978), All of these
interpretations agree that the Permian deposits of
Timor were formed in and along the distal side of
a trough that was par of the Australian block as
proposed by Hamilton (1979).

The crinoid bearing Maubisse Formation would
have been the most distal deposits, an unknown
distance, but perhaps as much as 1000km
northwest of the Canning and Camarvon Basins.
Plate reconstructions of the Permian (Zonenshain,
Kuzim and Kononoy, 1985, among others) pasi-
tion Australia along the eastern part of
Gondwanaland. The Camaryon and Canning
Basins would have been between 50° S and 40°S.
Timor would have been at approximately 25°—
30°S, The Timor faunas were considered Late
Permian on reports of fusulinids at Amarassi
(Schubert,1915) and cephalopods at Basleo
(Wanner.1931; Gerth. 1950). Haniel (1915) inter-
preted the cephalopods as middle Permian, on the
Early to Late Permian boundary. Brachiopods
(Broili,1916) and corals (Gerth, 1921) indicated a
Permian age but did not specify Early or Late,
whereas C. Wanner (1922) placed an Early Per-
mian age on the gastropods and bivalves. The
echinoderms were interpreted as Carboniferous
(Bather, 1914; Springer,1920, 1924), in contrast
io middle Permian (Wanner. 1924a) and Late
Permian (Wanner, 1931),

The lack of stratigvaphie control of the
numerous crinoid and blasteid localities from
Timor has been recognized for many years (Aud-
ley Charles, 1965; Glenister & Fumish.1987;
among others). Some cephalopod reports withand
without stratigraphic control at a few localities
include Sakmanan from Somohole, Artinskian
from Bitauni, and Late Pennjan from Amarassi
and Basleo (Glenister & Furnish,1987), Grant
(1976) considered the Bitauni brachiopods to be
of Artinskian age. Sections with stratigraphic con-
trol have yielded brachiopods from the Chiddruan
Maubisse Formation near Kasliu Village, West
Timor (Archbold & Bird,1989) and from the late
Sakmarian Khali and Bisnain Members of the
Maubisse Formation near Bisnain Village, West
Timor (Archbold & Barkham, 1989). The Bisnain
brachiopods were correlated with the Callythara
Formation by Archbold & Barkham (1989). Ar-
tinskian conodonts were recovered from matrix
materials and in situ limestone blocks collected on
early expeditions to Timor (Boogaard, 1987).
Thus Timor Permian strala is thought to range
from Sakmarian to Amarassian age. Wanner

MEMOIRS OF THE QUEENSLAND MUSEUM

(1931) reported that 90% of Timor crinoids and
75% of all other Permian fossils of Timor are from
the Basleo area. It is the age of the Basleo fossils
which we consider to be the most problematic
because a Late Permian cephalopod has been
reported by Glenister and Furnish (1987) from the
area. The stratigraphic relationship of the
cephalopod and crinoids is unknown,

Intemational correlation based on the crinoids
and blastoids suggests an Early Permian age,
probably Artinskian or Roadian ter the Baslea
echinoderms. The Basleo crincids and blastoids
cormlate best with Arinskian faunas from the
Callythara Formation (12 of the same genera, 3
species; 2 species with affinities and 3 genera with
close affinities to Timor genera), the Wandagee
Formation (4 of the same genera, 1] species) and
the Sarginsk Horizon of the Urals (15 penera),
There are few genera and no species in Common
with the Early Permian of Texas (3 genera),
Nevada (7 genera), and Bolivia (2 genus). Few
genera and no species are common to the Upper
Permian of Russia (1 genus), Sicily (5 genera),
Western Australia (1 genus), and eastern Australia
(2 genera). These data have been compiled from
crinoid occurrences indexed in Bussler & Moody
(1943), Webster (1973, 1977. 1986, 1988), and
unpublished data.

Until the structure and stratigraphy of the Bas-
len area are fully understood we consider the
echinoderm fauna to be Early Permian recogniz-
ing that part of the fauna could be Late Permian
based on the cephalopod reported by Glenister &
Furnish (1987),

SYSTEMATIC PALAEONTOLOGY

Crinoid terminology follows Ubaghs et al. (in
Moore & Teichert, 1978), with columnal patterns
after Webster (1974), Blastoid terminology is
from Beaver et al. (in Moore, 1967).

Material collected by us during July 1991 came
from localities entered in the Queensland Museum
Locality Register (QML), and is curated in the
Queensland Museum Palacontological Collection
(QMF). Other collections referred to are indicated
by the following prefixes: Commonwealth
Palaeontological Collection, Bureau of Mineral
Resources, Canberra (CPC), Museum of Victoria,
Melbourne (NMVP); Western Australian
Museum, Perth (OWAFP) and Department of Geol-
ogy, University of Western Australia, Perth
(UWAF).

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

319

Subclass CAMERATA Wachsmuth & Springer,
1885

Order age a Moore & Laudon,

Suborder COMPSOCRININA Ubaghs, 1978
Superfamily CARPOCRINOIDEA de Koninck
& Le Hon, 1854
Family COELOCRINIDAE Bather, 1899

Coelocrinid? indet
(Fig.1)

MATERIAL

QMF21428, a large, weathered, partial set of arms
with anal tube, found in three pieces, two of which fitted
together, from the middle Callytharra Formation in the
eastern line of the Gooch Range south of Carnarvon-
Williambury Station road, GR922549 Gooch Range 1:
100,000 topographical sheet (no. 1850), 1974 edition.

FIG. 2. A-C, Neocamptocrinus barrabiddyensis sp.
nov., A ray, posterior and basal views of holotype
CPC30668, X 1.2. D-F, Stomiocrinus merlinleighensis
sp. nov., X 10. D, holotype, CPC30669, lateral view of
crown. E, paratype, CPC30673, lateral view of crown
showing raised diamond-shaped oral level with the
arms. F, paratypes CPC30671, (lower right) and
CPC30672, lateral views; note distal arm branching on
CPC30672.

DESCRIPTION

Arms spatulate, at least 63.8mm long, biserial,
convex in section proximally (1.7mm wide), be-
coming externally flat distally (4.4mm wide); dis-
tally bearing short, laterally directed spines on
every 7th to 9th brachial on each side of the arm,
with the spines in alternating positions; distal tips
incurved towards a slender elongate anal tube
extending above the arms and bearing short spines
along the tube walls.

REMARKS

This specimen bears remarkable similarity to
the illustration (Wachsmuth & Springer, 1897, pl.
42, fig. 5) of Dorycrinus cornigerus (Hall, 1858).
Anal tube spines of D. cornigerus are longer,
extending beyond the arms when enclosed. Long
spatulate arms are known in several genera of the
Batocrinidae and Coelocrinidae (Wachsmuth &
Springer, 1897); however, only those of D. cor-
nigerus have short laterally directed spines along
the sides of the arms. Thus we questionably refer
the specimen to the Coelocrinidae.

This ts the first Permian report of the otherwise
Middle Devonian to Early Mississippian
European and North American family (Moore &
Teichert, 1978). Other camerates, such as Ac-
tinocrinites, Pleurocrinus and Camptocrinus, ini-
tially restricted to Early Carboniferous strata of
Europe and North America were discovered in the
Permian of Timor (Wanner, 1916, 1924a). Thus,

320

a possible coelocrinid in the Permian of Western
Australia adds to the list of camerate holdovers
lacking a Late Carboniferous record.

Superfamily HEXACRINITOIDEA
Wachsmuth & Springer, 1885

Family DICHOCRINIDAE Miller, 1889
Subfamily aa atte Broadhead,

Neocamptocrinus Willink, 1980

TYPE SPECIES

Neocamptocrinus bundanoonensis Wiil-
link,1980, from the Kazanian Berry Formation,
New South Wales; by original designation.

REMARKS

Willink (1980) recognised Neocamprocrinus on
the inflated tegmen and a column that js elliptical
throughout its length instead of round adjacent to
the cup as is Camptocrinus. Carboniferous and
Permian camptocrinid columns show that the el-
liptical shape is similar in both, that is nearly
round, but definitely slightly elliptical, adjacent to
the cup becoming progressively elliptical in a few
centimetres distally and remaining elliptical
throughout the remainder of the known column,
The long axis of the columnal attains its greatest
size in the Permian of Western Australia (Fig.
3M,.N). Cups assigned by Willink (1980),
Webster, (1990) and herein show an evolutionary
and stratigraphic record of potential value for the
Permian of Australia and Timor, We question {he
degree of inflation of the tegmen as a generic
feature; however, for convenience we apply the
name to these Permian forms including
Camptocrinus cf, C. indoaust-ralicus Wanner
(Webster, 1987; 102, fig. SJ-M).

Columnals of Neocamptocrinus are common in
many parts of the Permian of Westem Australia
(Webster, 1987, 1990; herein). Even proximal,
nearly circular columnals are easily recognised by
the distinctive, off centre, axial canal and unequal
length of culmina on the opposing long sides of
the articulum, Additional study of
Neocamptocrinus columnals in Western Australia
is necessary to determine their strat-igraphic
value. Columnals of Nevcamptocrinus sp. are
present in the Callytharra Formation at all
localities we collected. No cups were found, but
the radials listed as indeterminate camerate radial
4 and 5 by Webster (1987) probably belong to a
new species of Neacamptocrinus-

MEMOIRS OF THE QUEENSLAND MUSEUM

Neocamptocrinus barrabiddyensis sp, nov.
(Fig. 2A-C)

ETYMOLOGY
For Barrabiddy Creek.

MATERIAL

Holotype CPC30668, Bulgadoo Shale, east limb of
Barrabiddy Anticline, Barrabiddy Paddock, 1/4 mile
south of mill.

DIAGNOSIS
Radial notches relatively narrow; radials more
expansive,

DESCRIPTION

Cup bowl-shaped, wider (29.lmm) than high
(21.8mm), base truncated, widest at base of radial
facets, sutures flush, no ormmamentation. BB 2,
suture faint, 5.6mm wide, 5.5mm high, upflared,
slightly concave walls. RR 5, longer (194mm)
than wide (16.0mm), gently convex transversely,
convexo-concave longitudinally. Radial facets
angustary, c. 2/3 radial width; transverse ridge
elevated, with coarse ridges normal to ridge
length; brachial canal circular in section; muscle
fields elliptical, long axis subparallel to transverse
ridge, moderately impressed; ambulacral groove
narrow, sloping inward. Anal longer (19.6mm)
than wide (11.3mm), narrower than and in line
with radials. Columnal facet elliptical, 8.7mm
long, 63mm wide, impressed; articulum narrow;
culmen coarse, cog-like, areola wide, flat. Stem
and arms unknown,

REMARKS

This robust cup was weathered out of red clays-
tone containing abundant disarticulated plates of
Calceolispongia cf. barrabiddyensis. It has
narrower radial notches, radials that distally ex-
pand transversely ata greater rate and lower radjal
facets than N. occidentalis Webster, 1987 or N.
millyitensis sp. noy. These two species probably
evolved from N. barrabiddyensts.

Neocamptocrinus millyitensis sp, nov,
(Fig. 3A-L)

Neocainptocrinas sp. nov. Webster, 1990:57, pl |,
figs 7-11.

ETYMOLOGY
For the Millyit Range,

MATERIAL
Holotype CPC27054, paratypes CPC27455-27457
from the Millyit Range. Other material, QMF21420-

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

FIG. 3. A-K, Neocamptocrinus millyitensis sp. nov. A,
paratype radial QMF21421, showing radial facet, 3.
B, paratype partial theca QMF21420, showing anal
plates, anal opening, inverted U-shaped ridge above
anal opening, and axillary primibrachial of C ray,
X 2.8. C.D to K,L, articular facet and lateral views of
pluricolumals, paratypes QMF21422-21426, respec-
tively; showing proximal to distal changes in
transverse section and modifications in articular facet,
X 2.2. M,N, Neecamptocrinus sp. M, lateral view of
medial to distal pluricolumnal, WAM88.1070, show-
ing attached cirri distally, ¥ 0.4. N, lateral view of
medial pluricolumnal, WAM88.1069, x 0.4.

321

21426 from QML772,Millyit Range, Canning Basin
from the Cherrabun Member, Hardman Formation,

DESCRIPTION
See Webster, 1990:57.

REMARKS

Webster (1990) left material available at that
time in open nomenclature because its precise
locality and horizon were unknown. We have now
collected further material from a well-defined
locality and horizon, which may well be the type
locality.

The partial theca shows 3 anal plates, with a
large primanal in the radial circlet, a small anal
opening above the central anal plate and an in-
verted V-shaped ridge on the overlying oral. Both
orals bear a coarse central node. Radial facet has
a transverse ridge with a parallel shallow outer
ligament pit, two shallow muscle fields and a
T-shaped ambulacral track aborally.

Pluricolumnals are mostly short segments but
show the change in sectional shape from nearly
circular proximally to distally elliptical. The ar-
ticular facet of the proximal columnals has long
coarse culmina on the upper side of the columnal
and very short culmina on the under side. Culmina
become coarser laterally on the facet. The areola
is higher on the upper side with faint traces of
culmina across the surface. On the underside, the
areola is smooth and half the width of that on the
upper side, Axial canal is small and round.

Neocamptocrinus sp.
(Fig. 3M,N)

MATERIAL

Pluricolumnals WAMB88.1069-88.1070 from nonh
bank, Minilya River in the fourth massive sandstone al
the Wandagee Formation, Coolkilya Poo) area,
Wandagee Station. Collected K.J. McNamara.

REMARKS

These are the longest articulated pluricolumnals
of Neocamptocrinus known from Western
Australia. They are 28cm and 28.5em long with
columnals 30mm wide, 9mm ihick and 2.2mm
high. In a proximal direction noditaxes consisted
of 22, 23, 24. and 26 columnals each on
WAM88,1069. Citti are initiated at the junction
between two columnals and may grow to extend
over the height of four columnals; the axial canal
is also shared by the two calumnals, Columnal
growth after initial growth of the cirrus extends
beyond the distal edge of the proximal cirral to
partly cover the proximal end of the second cirral,

MEMOIRS OF THE QUEENSLAND MUSEUM

Cimi are round in section. Crenularium 1.8mm
wide, slightly less than half cirral radius (4.1mm);
culmina fine. Areola planar. Axial canal small and
round. Cirri taper distally with semipointed ends.
These large columnals lack the more rounded
proximal and distal sections of the stem, They
show that considerable lengths of the stem acted
as a runner with the cirri extended out and down
into the substrate. It is unknown if the flexures in
the two stem sections reflect flexibility around
substrate irregularities or diagenetic distortion,

Stomiocrinus Wanner, 1937

TYPE SPECIES
Stomiocrinus subglobosus Wanner, 1937 from
the Permian of Timor, by original designation.

Stomiocrinus merlinleighensis sp. nov.
(Fig, 2D-F)

ETYMOLOGY
Por the Merlinleigh Subbasin, Carnarvon Basin.

MATERIAL

Holotype CPC30669, paratypes CPC30670-30676
froma micaceous limestone in the top of the Bulgadoo
Shale, at type locality, near Donnelly’s Well.

DIAGNOSIS ;
Orals elongate, steeply upflared, visible in
lateral view.

DESCRIPTION

Crown small, cylindrical, Cup bullet-shaped,
widest at radial summit, fine shagreen ornamenta-
tion. BB 2, equidimensional, forming lower 1/3 of
cup, upflared steeply; interbasal suture in plane of
symmetry. RR 5, longer than wide, outflaring
gently longitudinally, moderately convex
Iransversely, straight to slightly convex lon-
gitudinally. Radial facets angustary, approx-
imately 1/3 greatest radial width, subhorizontal.
Brachials rectilinear, longer than wide, strongly
convex transversely, straight longitudinally; 1Br2
or 3 axillary; one additional branching in some
rays, all branchings isotomous. No pinnules. Anal
in radial circlet, shghtly narrower than radials. OO
5, elongate, steeply upflared, raised medial section
flush to slightly inset to proximal brachials,
mutual lateral edges underlay proximal brachials
and form base of ambulacral groove from the base
of the arms to the mouth, Posterior oral largest,

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

323

FIG, 4. Neoplatycrinus callytharraensis sp. nov. all from QML757, * 1, A-C, basal circlets QMF21594, 21593,
and 21592, respectively. D-H, radials with primibrachs and in some cases secundibrachs attached, QMF21595,

21599, 21597, 21598, and 21596, respectively.

MEASUREMENTS (mm)

3,6 (incomplete:
LS
16
13
0.7 0.7
0.6 0.6
1.0 0.8

Radial width =| 08 | 7

REMARKS

Stomiocrinus 1s reported from the Permian of
Timor, Russia (Broadhead, 1981) and Western
Australia (Webster, 1990). Broadhead (1981) as-
signed Stomiocrinus to the Camptocrininae based
on Yakovley’s (in Yakoyley & Ivanov, 1956)
combination of disassociated thecae and bilateral-
ly symmetrical columnals. The crowns of S. mer-
linleighensis occur on three small blocks
associated with Jimbacrinus donnellyensis and
Calceolispongia acuminata. None of the crowns
of S. merlinleighensis has the stem attached

whereas the crowns of J. donnellyensis have the
proximal stem attached. It is unknown if the
proximal stem of C. acuminata is attached.

In the matrix there are numerous thin round
pluricolumnals which contain columnals of 4mm
diameter and 7mm length that might have been the
stem of 5. merlinleighensis. They differ from the
short cirral segments (7mm diameter, 2mm long)
which are articulated with the attached stems on
J. donnellyensis. There are no bilaterally sym-
metrical columnals in the matrix. This leaves the
subfamily affinity of Sromiocrinus in doubt until
articulated thecae and columnals are found.

Superfamily PLATYCRINITOIDEA Austin &
Austin, 1842
Family PLATYCRINITIDAE Austin &
Austin, 1842
Neoplatycrinus Wanner, 1916

TYPE SPECIES
Neoplatycrinus dilatatus Wanner, 1916, from
the Permian of Timor; by original designation.

324

MEMOIRS OF THE QUEENSLAND MUSEUM

Neoplatycrinus callytharraensis sp. nov.
(Fig.4)

Neoplatycrinus aff. N. somoholensis Wanner, 1916;
Webster, 1987:100, figs 4A-O.
Pleurocrinus? sp. Webster, 1987;101, fig. 4P,Q.

ETYMOLOGY
For Callytharra Springs.

MATERIAL
Holotype UMMP61862. Paralypes UMMP61863-

FIG. 5. Neoplatycrinus miamiaensis sp. nov. A,B, oral
and oblique views of nodose oral, holotype CPC30677,
2. C.D, interior ( X 1.7) and exterior ( X 2.3) views of
weathered nodose ambulacral plate, paratype
CPC30678. E, external view of paratype radial
CPC30679, X2. F, exterior of paratype basal circlet
CPC30680, showing azygous basal at upper right and
elliptical stem facet, x2.

61867; WAM84.638, 84.639; UWAFS5638,
UWAF7860 and QMF2 1592-21599 all from QML757.

DIAGNOSIS
Oral and ambulacral plates with large node or

coarse ridges.

DESCRIPTION
As given by Webster, 1987, p. 100-101.

REMARKS

This species is based on disarticulated oral and
ambulacral plates bearing one very large node and
secondary smaller nodes or very coarse discon-
tinuous ridges. Basal and radial plates assigned
lack these coarse nodes as is the case in N.

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

miamiaensis, described below. We tound the or-
namented orals and ambulacral plates disas-
sociated in proximity to the unornamented basal
circlets and radials.

Webster, 1987, referred the Callytharra Forma-
tion specimens to N. aff. N. somoholensis, a
species based on internal molds (Wanner, 1916,
1924a). The ambulacral plates of V, somoholensis
are divided distally by adeep V-shaped separation
(Wanner, 1924a, pl. 3, fig, 2). Division on the
umbulacral plates of N. callytharrensis is not as
deep (Webster, 1987, fig. 4A), The upper distal
part of oral plates of NV. semoholensis would over-
hang the proximal vertical section whereas they
do not on NV, callytharracnsis. All other described
species of Neoplaryerinus lack Omamentation on
the oral and ambulacral plates.

Neoplatycrinus miamisensis sp. nov.
(Fig. 5)

ETYMOLOGY
For Mia Mia Station,

MATERIAL

Holotype CPC30677, paratypes CPC30678-3068 1 all
from the lowermost fossiliferous horizon in the Ar-
tinskian Coyric Formation, cast of Mia Mia Homestead,
23°21'°S, 114°34'B, northern Carnarvon Basin, Other
material CPC30711-307153, a colummal, 2 racials and
2 tegen plates from the lower part of the Noonkanbah
Formation at the type section (BMR loc, KNF'73), Cane
ning Basin.

DIAGNOSIS

Basal circlet subhorizontal; orals and am-
hulacrals bearing coarse nodes in radiating to ir-
regular patterns.

DESCRIPTION

Basal circlet large, estimated 32mm diameter,
subhorizontal; bears elliptical stem facet, 2.4mm
long, 9.3mm wide. BB 3, 2 large, 1 small (16mm
long, 15.5mm wide) surface coarsely granulate,
with coarse node (2.8mm diameter) on one plate
RR 5, wider (28,4mm) than long (20,7mm), con-
vex longitudinally and transversely, surface with
coarse granules, no nodes. Radial facet wide, 2/3
plate width, narrow, crescent shaped. OO 5, large,
estimated 35mm long, 28.2mm wide, polygonal.
proximally vertical, distally subhorizontal or
upflared, with 8 to 18 coarse nodes (4.4m
diameter, 2.8mm. high) in radiating or irregular
pattern and coarse granular ornament. Proximal
aimbulacral quadrangular, 18.9mm long, 19.3mm
wide, sharply pointed proximally and distally,

325

side vertical, exterior with up to 9 coarse nodes
and coarse granular omamentalion,

REMARKS

The descnpton is based on disarticulated os-
sicles from one horizon in the Coyrie Formation.
The theca is estimated ta have been up to 75mm
wide and 45mm high, with a low bow] shape
having a flat base, low out-flared radials, and a
bread flat to upflared tegmen. Interpretation of the
theca is based on: 1, a flat wide basal circlet (Fig.
SF); 2, radials attached to the vertical facets on the
basal circlet such that the proximal end of the
radial was horizontal, expanding the low bowl
laterally, 3, radials are convex longitudinally
resulting in the distal parts of the radial being
essentially vertical; the radial facet would have
sloped down and aut at an estimated 80°; and 4,
the proximal end of the orals was probably nearly
vertical and the distal part subhorizontal to
upflared perhaps as much as. 40°. Mutual facets on
the radials and oruls are not well preserved
precluding firm estimate of reconstruction.

Secundibrachials associated with the thecal
plates bear the same coarse nodes and granulate
ornamentation. They are U-shaped with two deep
V-shaped ambulacral grooves, one large and one
smaller and lateral, Most plates show growth
bands on the aboral side. Soluuion weathering has
modified the surtaces of most plates, removing or
obliterating the granular ornament is common,
and in extreme examples nodes are reduced ta
small stellate elevations. Diagenetic distortion and
fracturing followed by calcite cementation sealing
the fractures is present on some plates.

Numerous internodals and nodals of the “twist”
nodal colummnals were found in association with
the thecal plates. They are assigned to the species
because al! other crinoids in the fauna are non
platycrinitids. No difference could be discerned
between these colunmals and ones found in the
Callytharra Formation, This suggests that mosaic
evolution was occurring in the theca while the
columns were stable.

Neoplatyerinus 1s restricled to the Permian of
Timor (Lane in Moore & Teichert, 1978) and
Wester Australia (Webster, 1987). The genus is
recognized by the large oral plates. N. miamiaen-
sis is distinguished from N. callytharraensis by ws
oral and ambulacral plate ornamentation, Irdiffers
from WN. somoholensis by having a flat base instead
of being upflared at 30° to 40° Evolution of NV,
micinigensix From N, callytharraensis occurred by
modificauion of oral and ambulacral ornament,

The original field Iabel on specimens of NV.

3

6

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 6. Platyerinites/Neoplatycrinus spp. A,B, articular
facet({ * 2.7) and lateral ( X 2.5) views of pluricolumnal
QMF21429. C, lateral view of QMF21430 showing
cirral facets, X 1.5. D, articular facet of “twist” nodal,
QMF21431, X 2.5. E, microcrinoid (?) attachment disc
followed by 5 columnals overgrown by proximal cirral
on pluricolumnal QMF21432, X12. F, fenestrate
bryozoan attachment overgrown by columnals in
pluricolumal QMF21433, x5,

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

miamiaensis gave the stratigraphic unit as the
Bulgadoo Shale. However the latitudinal coor-
dinates place the locality in the Coyrie Formation
as mapped on the Winning Pool-Minilya
1:250,000 Geological Sheet SF50-13, 1984.

The columnal, radials and tegmen plates from
the Canning Basin bear the same ornamentation
on the oral and brachial as described above.

Platycrinites/Neoplatycrinus spp.
(Fig. 6)

MATERIAL

Figured material QMF21429,21431 from QML760,
QMEF21430, 21432, 21433 from QML757 and other
material 181 internodals, 102 nodals and 39 pluricolum-
nals QMF21578 (203) from QML757, QMF21579 (26)
from QML758, QMF21580 (14) from QML759,
QMF21581 (13) from QML760, QMF21582 (9) from
QML761, QMF21583 (19) from QML758-761,
QMF21584 (18) from QML769 and QMF21585 (6)
from QML770.,

DESCRIPTION

Internodals elliptical, opposing fulcral ridges
parallel; nodals rectangular to subquadrate, op-
posing fulcral ridges at 15-80° with most 67° or
80°. All columnals wider than high, elliptical axial
canal, very fine shagreen ornamentation.
Noditaxis variable, 1-5 internodals. Articulum
divided in half by prominent fulcral ridge. Fulcral
ridge depressed below articular rim adjacent to
axial canal, bears transverse vermiform to short
medium coarse ridges proximally, 3 to 7 medium
coarse slightly radiating culmina distally.
Crenularium narrow; culmina short, medium
coarse, lengthening near fulcal ridge, producing a
fan shaped pattern, unbranched. Areola divided by
fulcral ridges, each half crescent in outline,
smooth, sloping into shallow trough paralleling
fulcral ridge, deepest abaxial of axial canal. Axial
canal small, elongate parallel to fulcral ridge.
Latus gently to moderately convex on lateral
sides, moderately convex to vertical to gently
concave on ends of internodals, rarely bears small
nodes. May bear cirri in distal most parts of stem.

Cirri round transversely; articulum round.
Crenularium 1/3 radius; culmina coarse, cog-like,
may branch isotomously once. Areola smooth,
flat. Axial canal round.

REMARKS

The elliptical columnals vary from elongate
(18.6 x 9.4mm) to subcircular (21.8 x 19.5mm) in
adults. The more subcircular specimens are most-
ly cirral bearing and judged to be more distal

327

columnals, whereas the more elongate elliptical
forms lack cirri and are considered to be more
proximal. Thickness of individual nodals and in-
ternodals is uniform except in the cirral bearing
distal stem parts where they show pinching and
swelling in lateral view.

Cirral facets occur on nodals or internodals and
slant distally 5-10°. They are most commonly
developed on one or both ends of the long axis but
also occur along the short axis of the facet. There
are up to 4 cirri per columnal, with one the most
common. First cirrus is wedge shaped with distal
facet sloping distally thereby increasing the distal
slope of the cirrus. Cirri short, tapering distally.

Most specimens are weathered with loss of sur-
face articulum and axial canal details and enhan-
cement of growth bands. Up to 11-14 growth
bands weathered as ridges and grooves. Similar
growth bands on large cup plates of
Neoplatycrinus aff. N. somoholensis, Platy-
crinites wrighti and Pleurocrinus? sp. were il-
lustrated by Webster (1987). These columnals
probably belong to those species and are therefore
left in open nomenclature.

Organisms attached to the living column of
Platycrinites/Neoplatycrinus spp. include
bryozoans (Fig.6F) and other crinoids (Fig.6E) as
both are found overgrown by columnals or cirri.
Phosphannulus infestations are rare on
Platycrinites/Neoplatycrinus spp. Columnals, as
postmortem detrital grains were used as attach-
ment sites by various sessile organisms but most
commonly bryozoans. Diagenetic compaction
distorts the areola of numerous specimens.

Subclass INADUNATA Wachsmuth &
Springer, 1885
Order DISPARIDA Moore & Laudon, 1943
Superfamily ALLAGECRINOIDEA Carpenter
&Etheridge, 1881
Family ALLAGECRINIDAE Carpenter &
Etheridge, 1881

Wrightocrinus Moore, 1940

TYPE SPECIES
Allagecrinus jakovlevi Wanner, 1929 from the
Permian of Timor; by original designation.

Wrightocrinus wooramelensis sp. nov.
(Fig. 7A,B, I-K)

ETYMOLOGY
For the Wooramel River.

MATERIAL
Holotype QMF21438 from QML757, paratypes

328

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 7. A,B, I-K, Wrightocrinus wooramelensis sp, nov. oral and C ray views of partly crushed paratype
QMF21439, A * 30;B x40. I-K, oral, posterior and A ray views, of holotype QMF21438, x 30. C-H, Litecrinus
pansus sp. noy. C-E, AB interay, oral and D ray views of holotype QMF21448, x 40. F-H, A ray, oral and posterior

views of paratype QMF21449 x 40.

QMF21439 (QML760), CPC30692 (QML757-761)
and three crushed thecae QMF21554 (QML759),
QMF21555(QML761), all from west of Callytharra
Springs.

DIAGNOSIS
Radials protuberent; radial facets narrow; basals
short; orals with an elongate central depression.

DESCRIPTION

Theca 1.1mm high, 1.0mm wide, globose, con-
vex walls in lateral view, rounded pentagonal
outline in oral view, walls smooth. BB 2, suture

slightly rotated off A-CD plane, low, proximally
horizontal, bearing round articular facet for stem,
distally upturned, flaring outward, forming lower
1/5 of cup. RR 5, 3/4 as wide as long, widest at
midlength, outflared proximally, incurved distal-
ly, moderately convex longitudinally, transverse-
ly gently convex proximally, strongly convex
distally, with one facet each; radial facets
peneplenary, sloping inward, pit-like. Slight anal
notch on mutual shoulders of C and D radials. OO
5, elliptical, slightly longer than wide, central
linear depression bordered by rounded low ridge,

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

329

FIG. 8. Litocrinus protuberans sp. nov. A-C, BC interray, oral and posterior views of paratype QMF21441, x40.
D-F, A ray, oral and posterior views of paratype QMF21442, D x 40; E x 50; F x 40, G-l, A say, oral and E ray

views of holotype QMF21440, x 40.

form flat-topped elevated circlet slightly inset
from radial summit. Posterior basal largest, as
wide as long, separating BC and DE orals.

REMARKS

The holotype has a small calcite deposit on the
EA oral extending onto the mutual shoulders of
the Eand A radials. Atacursory glance this falsely
appears to be the posterior oral. The posterior oral
is recognised by the suture with the other four
orals and the small anal notch below it. All

specimens of W. wooramelensis may be imma-
ture, lacking the development of additional arms
in the A, Cand D rays.

The globose appearance of W. wooramelensis is
similar to immature specimens of W. jakovlevi
(Wanner, 1929) but distinguished from it by
having more protuberant radials, narrower radial
facets, shorter basals and orals bearing an elongate
central depression. Webster & Jell, in press, dis-
cussed the reasons, essentially that one is a

MEMOIRS OF THE QUEENSLAND MUSEUM

330

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA 331

FIG. 10, A-H, Synbathocrinus constrictus (Wanner, 1916), initial brachials, A-C, aboral, lateral and brachial facet
views, QMF21478. x 25. D-F, brachial facet, lateral and aboral views of QMF21479, A 25; B,C * 30 .G,H,
oral and C ray views of abnormal specimen with only 4 rays and 4 oral plates, QMF21462, < 30.1-M, Microcrinoid
holdfasts gen. et sp. indet. I,J, oral and lateral views of rectangular shaped base, QMF21477.1 *25; J 30. K,L,
oral and lateral views of hexagonal base QMF21475, K X 25; L X 30. M, oral view of rectangular base QMF21476,
* 30.

FIG. 9. Synbathocrinus constrictus (Wanner, 1916). A-C, oral, A ray and E ray views of QMF21461, A x50; B,C
x40. DE, oral and B ray views of QMF21464, 30. FG, oral and C ray views of abnormal specimen, A and B
rays semifused, QMF21467, X 25. H,I, oral and D ray views of pre arm stage QMF21460, x 50. J,K, oral and B
ray views of one armed stage, QMF21465, * 30. L,M, oral and A ray views of 2- armed stage, QMF21466, * 20.
N, oral view of 4-armed stage, QMF21463, x 20.

juvenile of the other, for placing Allagecrinus topulo.1981, from the Lower Carboniferous of
quinguebrachiatus Wanner, 1929 in synonomy Tennessee; by original designation,
with W. jakovlevi.
Litocrinus pansus sp. noy.
Litocrinus Lane & Sevastopulo, 1982 (Fig. 7C-H)

TYPE SPECIES ETYMOLOGY
Kallimorphocrinus punctatus Lane & Sevas- Latin pansus, spread out, extend.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 11. A, Spheniscocrinus australis sp. noy., basal view of holotype, showing base of anal tube in 6 o'clock
position on cup, CPC30682, x 3. B-D, Occiducrinus australis Webster, 1990, A ray, posterior and basal views
of QMF21427, 1. E, Tapinocrinus spinosus (Wanner, 1924a), posterior view showing distal tips of arms,

QMF21427, x 1.6.

MATERIAL

Holotype QMF21448 from QML760, paratype
QMF21447, 21449 from QML757, QMF21446 from
QML759, and other material QMF21556 (1) from
QML757, QMF21557 (3) from QML759, QMF21558
(1) from QML760 and QMF21559 from QML761.

DIAGNOSIS
Theca expanded markedly above narrow basal
circlet; oral circlet dome-shaped.

DESCRIPTION

Theca small, equidimensional, 0.4-0.6mm,
pyriform, walls concave. expanding rapidly above
basal circlet, unornamented. BB 5 or fused, smail,
forming lower 40% of cup, gently outflared. RR
5S, longer than wide, moderately convex transver-
sely, concave Jongitudinally with straight to con-

vex distal ends, may be weakly protruding distal-
ly. OO 5, rounded proximal edge in oral view,
proximally steeply upflared, distally subhorizon-
tal, may bear weakly elevated protrusion at upflar-
ing to subhorizontal flexures, forming domed arch
with diameter less than thecal max. just below
radial summit; posterior oral larger, separating BC
and DE orals.

REMARKS

L. pansus is yariable in development of pro-
trusions on orals and may develop a weak node
distally on the radials. Without careful observa-
tion LZ, pansus could be considered a juvenile or
variant of L. protuberans. However, the species
differs from L. protuberans by the greater lateral
expansion, lack of strongly protruded radials and

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

more rounded dome shape of the oral circlet. It is
the smallest form in the Callytharra fauna.

Litocrinus protuberans sp. nov.
(Fig. 8)

ETYMOLOGY
Latin protuberosus, protuberant; for the radials.

MATERIAL

Holotype QMF21440 from QML759, paratypes
QMF21441 from QML758, QMF21442, 21443 from
QML759, QMF21444 from QML760, QMF21445
from QML757, and other material QMF21560 (8) from
QML757, QMF21561 (3) from QML758, QMF21562
(15) from QML759, QMF21563 (19) from QML760,
QMF21564 (3) from QML761, QMF21565 (5) from
QML757-761.

DIAGNOSIS
Radials with evenly rounded large protuberances.

DESCRIPTION

Thecae small, largest specimen 1.1mm high,
0.9mm wide, turbinate to pyriform, walls expand
rapidly from base of radials to large protuberances
in distal part of radials, capped by oral circlet,
strongly pentalobate in oral view, unornamented.
Basals fused, bear circular stem facet proximally,
gently outflaring distally, forming lower 1/3 of cup.
RR 5, slightly longer than wide, expanding rapidly
upward, longitudinally concave proximally to
strongly convex distally, transversely gently convex
proximally strongly convex distally; large smooth
growth protuberance on distal part of radial. No anal
notch on plates. OO 5, slightly longer than wide to
subequal to wider than long, slightly irregular outer
surface, vertical to slightly inflared proximally, sub-
horizontal to gently arched distally. Posterior oral
largest, separating BC and DE orals, bearing small
hydropore or madreporite. Oral circlet slightly inset
on radial summits.

REMARKS

Growth sequence from 0.4-1.1mm present in the
67 specimens studied. Initially the theca is equi-
dimensional and the radial protuberances are
recognisable but not domineering as the cup wall
flares moderately and evenly from the stem facet
to the radial protuberances. As growth continues
radials elongate and the protuberance expands to
become the most noticeable morphologic feature
of the theca. The base of the cup is relatively
narrow and most of the wall flaring is with the
radials.

333

No other species of Litocrinus has as strongly
protruded distal parts of the radials as L.
protuberans. There is a large node in the centre of
each radial on the Late Permian L. palermoensis
(Strimple & Sevastopulo, 1982) from Sicily that
is a later evolved form.

Family CATILLOCRINIDAE Wachsmuth and
Springer, 1886.

Notiocatillocrinus Willink, 1978

TYPE SPECIES

Notiocatillocrinus oakiensis Willink, 1978,
from the Lower Permian of NSW; by original
designation.

Notiocatillocrinus callytharraensis
Webster, 1987 (Fig. 14E)

Notiocatillocrinus callytharraensis Webster, 1987:
108, figs 7, 8.

MATERIAL

One weathered crown NMVP 127202 from Fossil Hill,
Bidgermia Station, GR351884 on Glenburgh 1:250,000
Geological Sheet SG 50. Collected by G.A. Thomas.

REMARKS

This is the largest specimen known. It is twice
the length of the holotype and displays an excel-
lent cross-section of the cup and tegmen.

Easily recognisable radials, showing the mullti-
ple arm facets, cups and the horseshoe-shaped
tegmen plates of N. callytharraensis were found
at every locality visited. Often they were em-
bedded in the grainstones and not retrieveable
without excessive labour. These plates make ex-
cellent index fossils in the Callytharra Formation
throughout the Carnarvon Basin.

Superfamily SE One Miller,
Family SYNBATHOCRINIDAE Miller, 1889
Synbathocrinus Phillips, 1836

TYPE SPECIES
Synbathocrinus conicus Phillips, 1836 from the
Lower Carboniferous of England; by monotypy.

Synbathocrinus constrictus (Wanner, 1916)
(Figs 9, 1OA-H)

MATERIAL
Figured specimens QMF21460, 21461, 21463, 21465,

334

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 12. Barycrinid? gen. indet. A, articular facet of pentameric columnal QMF21434, X 1.4. B, articular facet of
pentameric pleuricolumnal QMF21435, » 1.7. C-E, root holdfast with branchlets or citi, QMF21436, * 1.4. C,
enlargement of articular facet of non-pentameric rootlet cirri. D,B, lateral views of holdfast in possible growth
position, F,G, articular facet and inner lateral views of weathered pleuripentamere QMF21437, * 1.8.

21466, 21478, 21479 from QML759, QMF2 1462 from
QML758, QMF21464, 21467 from QMI757 and other
material QMF21571 (23) from QML757, QMF21572
(3) from QML758, QMF21573 (37) from QML759,
QMF21574 (7) from QML760, QMF21575 (1) from
QML761, and QMF21576 (6) from QML758-761.

REMARKS

Articulated thecae of juvenile and adult
specimens of Synbathocrinus consirictus are
moderately abundant throughout the interbedded
arenaceous shales and limestones of the lower
fossiliferous part of the Callytharra Formation al
Callytharra Springs. Juvenile spec-imens are 0).4-
2.5mm high and include growth stages from the
armless embryonic through 5-armed forms.

Early growth stages are dominated by inflated

orals and protuberant radials with a low narrow
basal circlet. Radials and orals increase in length
rapidly, are approximately equal in height and are
the most obyious plates in the theca during early
development. Radials show an early development
of the medial longitudinal ridge, whereas orals are
initially inflated then gradually develop a shallow
central depression. The CD oral is largest, adjoin-
ing all other orals, separating the BC and DEorals,
and bearing a small central protuberance, the
hydropore or madreporite. At 1.3mm thecal height
radials are approximately twice the height of the
oral circlet and the first radial facet develops on
the E radial. Growth continues in a linear increase
greater than transverse until approximately
2.5mm height when lateral expansion is rapid.
Arms develop in sequence in the C, B, D and A

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

335

FIG. 13, Lampadosocrinus variabilis sp. nov. A-C, A ray, oral and C ray views of paratype QMF21451, x40.
D-F, A ray, oral and DE interray views of paratype QMF21452, x 40. G-I, Eray, oral and C ray views of holotype

QMF21450, x 40.

rays. Only single specimens of most of the dif-
ferent arm stages are known. It is not known if the
same sequential arm development is regularly fol-
lowed. Orals are retained with the development of
the fifth arm and the anal plate is not developed
although a faint depression may occur in the radial
circlet at the mutual shoulders of the C and D
radials below the posterior oral. Macro specimens
lack the oral circlet and were illustrated by
Webster (1987).

Two abnormal specimens lack part or all of the
fifth ray. One (Fig. 8G,H) has 4 radials and 4 orals.
Except by careful examination it appears to be a
normal 5 rayed theca. The other (Fig. 7F,G) has a

partial fusing of the A and B radials and orals are
distorted.

Small brachials up to 2.5mm long in the disar-
ticulated washings are judged to be the beginning
growth stages of arms of S. constrictus on the basis
of the articular facet and the medial longitudinal
ridge. Two of these brachials are illustrated to show
the variation in longitudinal curvature, shape, am-
bulacral groove and distal tips.

These thecae are assigned to S. constrictus be-
cause they show the prominent radial ridge.
Webster (1987) noted the gradation between S.
constrictus and S. campanulatus so some of these

336

may be immature specimens of §. canpenalatus
but (hat form is not recognised until its adult stage.

Family UNCERTAIN

Microcrinoid holdfasts gen. ct sp. indet.
(Fig. 10I-M)

MATERIAL
QMPF21475-21477 from QML759 and QMF21577
from QML757-761 west of Callytharra Springs.

DESCRIPTION

Base hexagonal, rectangular or irregularly
polygonal; sides scalloped to irregular; thickened
centrally or towards one end extending as base of
round columnal. Base concave.

REMARKS

Holdfasts 1,0-2.0mm long; 0.5-1.5mm wide
and 0.6-1.0mm high. They range from Early Car-
boniferous in the United States (Webster, unpubl.
data) to the Permian, They are illustrated for
comparison in future studies and to demonstrate
the scope of the Callytharra echinoderm fauni,

Order CLADIDA Moore &Laudon, 1943
Suborder CY ATHOCRININA Bather, 1899
Superfamily Cen ‘OIDEA Bassler,

Family CY ATHOCRINITIDAE Bassler, 1938
Occiducrinus Webster, 1990

TYPE SPECIES

Occiducrinus australis Webster, 1990 from the
Aftinskian Wandagee Formation. Minilya River,
W.A.; by original designation.

Occiducrinus australis Webster, 1990
(Fig. 11B-D)

Occiaucrinus australis Webstet, 1990:60, pl. 2, figs T-
3,
MATERIAL
QMF21427 from the third thick sandstone bed above
the base of the Artinskian Wandagee Formation, cast
limb of syncline, approximately 150m north of the
Minilya River, Coolkilya Pool area, Wandagee Station,
Western Australia.

REMARKS

This cup was weathering out as associated dis-
articulated plates on a bedding surface which also
yielded Eoindocrinus praecontignatus and Cal-
ceolispongia abundans. The reconstrmicted cup

MEMOIRS OF THE QUEENSLAND MUSEUM

jacked the anal, the D radial and the ornls; 1t is
nearly identical in size with the holotype.

The proximal columnal showing the articulum
provides information not preserved on the
holotype. Crenularium narrow; culmina short,
widely spaced; crenellae wide, flat base, bearing
rudimentary short ridges and V-shaped
protrusions along irregularly walled outer side.
Arcola three times width of crenularium, Disar-
ticulated columnals should be easily recognised.

Family BARYCRINIDAE Jaekel, 1918

Barycrinid? indet.
(Figs 12, 22D)

MATERIAL

Figured specimens QMF21434, 21436 trom
QML760, QME21435 fram QML761, QMF21437
from QML757, Other material QMF21586 from
QML757 and QMF21587 from QML760,

DESCRIPTION

Columnals circular, pentameres; latus gently
convex; heteromorphic type I pattern; symplexy
articulation. Crenularium 1/3 radius on small
columnals, may be more on large columnals,
projects into areola at pentamere boundaries on
some specimens. Culmina irregular, medium
width, wider distally, may divide isotomously
once; new culmina may be inserted distally.
Areola 1/3 radius on small columnals, may be less
on large columnals, sloping inward, smooth. Axial
canal strongly pentastellate; claustrum narrow,
thin, wedge-shaped, forms base of areola, Cirr
circular in section, short, tapering rapidly distally,
not divided into pentameres; latus straight; ar-
ticulum same as pentamere articulum; axial canal
narrow, slightly dumb bell-shaped transversely.

REMARKS

All pluripentameres (up to 4,.95em length) and
pluricolumnals of articulated pentameres were
recovered as loose specimens. Weathering
destroyed many morphologic features but espe-
cially the claustrum on pluripentameres. Differen-
tiated weathering resulted in irregular wavy
thickness on external surfaces of some columnals
(Fig. 22D). Most of the larger specimens are part
of the holdfast or distal parts of the stem. The
robust size suggests they supported a sizeable
crown. It is unknown if the pentameres extended
to the cup.

Pentameric columnals are present on several
inadunates and camerates of Ordovician through
Devonian age (Moore & Jeffords, 1968). Middle

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

Mississippian Barycrinus from lowa are the pre-
viously recognized youngest pentamenc stems
(Moore et al. in Moore & Teichert, 1978). We
refer these specimens questionubly yo the
Barycnnidae and illustrate them because they are
easily recognizable, have potential as Permian
index fossils in the Carnarvon Basin and extend
the geographic and stratigraphic range of pen-
tameric columnals.,

ag ey y CODIACRINOIDBA Bather, 1890
amily $ TREBLOCRINIDAE Lane, 1967

Lampadosocrinus Strimple & Koenig, 1956

TYPE SPECIES
Dichostreblocrinus minulius Peck, 1936 from
the Lower Carboniferous St Joe Formation, USA.

Lampadosocrinus variabilis sp, nov.
(Fig. 13)

RTVYMOLOGY

Latin variabilis variable; refers to the variation
in shape of the oral circlet, and number of in-
frabusal plates.

MATERIAL

Holotype QMF21450 from QML757-761 undifferen-
tiated, paratypes QMF21451 from QML759,
QMF21452 from QML757-761, QMF21453 from
QOML757, QMF2i454 from QML758, QMF21455-
21459 from QML759 and other material QMF21566 (5)
from QML757, QMF21567 (2) trom QML758,
QMF21568 (24) from QML759, QMF21569 (2) from
OML760, QMF21570 (5) from QML757-761.

DIAGNOSIS
Oral plates of variable shupe; theca inflated;
busals tumid.

DESCRIPTION

Theca small, conical, subcylindrical to widely
inflated, widest at or slightly below oral summit,
height/width ratio 1.0-1.4, surface ornament of
minute pits.

IBB 1-5, most commonly fused; circlet low,
narrowly to moderately outflared, pentalobate to
circular transversely, forming lower 1/2 of cup,
1/4 of theca, bearing small impressed circular
stem facet. BB 5, with variable length to width
ratio, wider than long, subequal to longer than
wide, convex transversely and longitudinally,
tumidity gentle to moderately strong, rarely
projecting into a short spine. OO 5, roundly 10
sharply protruded at or barely below thecal sum-
mit, oral surface inflated to depressed centrally,

337

sutures flush to eleyated along mutual narrow
ridges. CD oral largest, protuberance more
rounded than others. separates BC and DE orals.
Oral circlet forming 2/5 thecal height, with slight-
ly concave summit, horizontal or slightly inflated,
sutures align with basal suture, rotated slightly or
in normal allernate position. BC and DE oval
protuberances commonly slightly down and out-
flared, EA and AB oral protuberances normally
subhorizontal. No anal plates, BC basal left
shoulder summit slightly below CD right shoulder
summit; all other basal summits align.

REMARKS

Variation among the 47 specimens studied
reflects growth changes and intraspecific varia-
tion. Two changes with growth are height/width
ratios of the oral plates. In general elongation of
the theea progresses from subequal height and
width in immature forms to height greater than
width in adult forms. Ratios in a growth serics of
4 specimens calculated as 1.0, 1.4, 1.06, 1.11,
smallest to largest. Oral plates of immature
specimens are commonly roundly protuberant,
rarcly pointedly protuberant, and Jack the nmmed
depression on the summit typical of larger
specimens. In immature forms the five
protuberances project above the summit slightly,
forming a slightly depressed summit. With growth
the rimmed depression develops on cach of Lhe
orals while the central oral summit is normally
gently inflated forming the highest point on the
theca. On the largest individuals the summit is
commonly horzontal.

Intraspecific variation is noted in the following
features. Most specimens are inflated moderately
with only a few specimens having mildly inflated
subcylindrical walls above the infrabasals, The
angle of outflaring to the protuberances on the
orals is variable. Tumidity of the basals ranges
from slight to moderate and rounded to elongate
in outline; rarely the tumidity is a short sharpened
projection or blunt spine. Sharpness of the oral
tumidily varies from rounded to angular and
directed subhorizontal to gently upflared.

Suborder POTERIOCRININA Jaekel, 1918
Superfamily SCYTALOCRINOIDEA Moore &
Laudon, 1943
Family APHELECRINIDAE Stnmple, 1967

Cosmetocrinus Kirk, 1941
Type Srecres

C. gracilis Kirk, 1941 from the Mississippian of
Indiana; by original designation.

338

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 14, A, Cosmetocrinus? middalyaensis sp. nov., posterior view of holotype UWAF113385, x0.7. B-D,
Trautscholdicrinus sp. A ray, basal and posterior views of WAM67.529, <3. E, Notiocatillocrinus callythar-
raensis Webster, 1987, weathered crown showing cross-section of cup and tegmen, NMVP 127202, X 1.1.

Cosmetocrinus(?) middalyaensis sp. nov,

(Fig. 14A)
ETYMOLOGY
For Middalya Station.
MATERIAL

Holotype UWAF113385 from QML770, Collected by
John Williamson.

DIAGNOSIS
Right tube plate not in contact with the radianal.

DESCRIPTION

Crown tall (119mm), cylindrical (23.3mm wide
maximum), plates smooth. Dorsal cup conical,
flattened, approximately equidimensional,
22.6mm long, 22mm wide at summit, base flat,
walls slightly convex proximally, slightly out-
flared distally. IBB 5?, large, 7mm long, 7mm
wide, steeply upflared. BB 5, large, CD longer

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

(13.1lmm) than wide (7.5mm), mildly convex
transversely and longitudinally. RR 5, large,
7.8mm long, 7.8mm wide, mildly convex trans-
versely and longitudinally. Radial facets plenary,
sloping outward, with well developed transverse
midge and outer ligament pit; slight gape between
radials and primibrachials, Anals 3, radianal nar-
row, in oblique position, adjoins C radial, BC and
CD basal and anal X; anal X largest, 6.3mm long,
4.5mm wide, adjoining right tube plate, C radial,
radianal, CD basal, D radial, and 2 tube plates,
distal up above radial summit; right tube plate
nearly excluded from cup, only proximal tip
below radial summit, not in contact with radianal.
1Br one per ray: in C and D rays, longer (8mm)
than wide (6.6mm), constricted distally, strongly
convex transversely, axillary. HBrr, W1Brr and
IVBrr cuheste, strongly convex transversely. All
branching isotomous. Second branching on II Brg-
19, one additional branching in some rays. Total
number of arms exceeds 20 per ray if branching
uniform in all rays. Pinnules small, slender, Anal
six: tall, slender, mostly covered. Stem round
transversely, heteromorphic, thin columnals, type
i patterns for proximal 40mm, homeomorphic
thick columnals distally; total of 130mm of stem
preserved, Columnal latus convex, symplexy ar-
liculation. Non-cirriferous.

REMARKS

Radials of this species were described as. in-
udunate radial #7 (Webster, 1987} based on loose
radials found at Callytharra Springs approximate-
ly 250km Lo the south. C.? middalyaensis is
widespread in the Camarvon Basin and the radials
are sufficiently distinct to be identified when
disurticulated -

The crown is crushed in situ as itretains the arms
and considerable length of the stem, Erosion has
removed, but left the impression of, the distal

30mm of the arms, Generic assignment remains.

tentative because the A ray is not exposed to
determine if branching occurs on the first
pomibrachial or a higher primibrachial as in
Paracosmelocrinus. Some species of Paracos-
mefocrinus have a strongly cicriferous proximal
Sem, which is not present in C.(?) middalyaensis.
The number of infrabasals is also uncertain as only
one complete und one partial infrabasal are ex-
posed. Their size and position suggests there are
only three infrabasals on this form.

Crinoids associated on the slab are a weathered
crown of Texacrinus granulosus sp. nov., disat-
ticulated columnals of a camptocrinid and ar least
five unidentified crinoids, In addition several

339

specimens of the spiriferid brachiopod

usispirifer carnarvonensis Archbold & Thomas,
1987 and fenestrate bryozoans are present.

Elevation of the nght tube plate until itis nearly
eliminated from the cup is an advanced condition,
suggesting a younger age than the Early Carbonif-
erous age of previously described species (Moore
& Teichert, 1978). Aphele-crinids were pre-
viously known only from the Lower Car-
boniferous of North America and Europe (Moore:
&Teichest, 1978),

Superfamily AGASSIZOCRINOIDEA Miller,
Family AMPELOCRINIDAE Kirk, 1942
Spheniscocrinus Wanner, 1937

TYPE SPECIES
Spheniscecrinus spinosus Wanner, 1937 from
the Permian of Timor: by original designation.

Spheniseocrinous australis sp. noy-
(Fig. 11A)

ETYMOLOGY
Latin australis, southern.

MATERIAL

Holotype crown CPC30682 from the Artinskian upper
Bulgadoo Shale or lower Cundlego Formation, 7.5km
on bearing 287° from Wandagee Homestead, Minilya 4
mile map sheet, Carnarvon Basin.

DIAGNOSIS
Ornament shagreen with small pits; sutures
flush; spines on brachials without nodes.

DESCRIPTION

Crown small, low, outspreading laterally, ¢s-
timated 60mm diameter; ornament of smal)
depressions on very fine shagreen surface. Cup
low bow! to discoid-shaped, 5.7mm wide, 2.2mm
high (estimated), shallow apical pits. IBB 5, im-
pressions small, probably subhorizontal. BB 3,
small, convex transversely and longitudinally, un-
equal size; EA estimated 1.3mm long, 1.2mm
wide; CD 2.1mm long (estimated), 2mm wide. RR
5, wider (1.8mm) than long (1.3mm), gently con-
vex longitudinally, strongly convex transversely,
thick, spreading laterally proximally resulting in
wide gap between radials at summit. Radial facet
plenary, slope steeply outward, thick, small gaps.
Anals 3, in intermediate position. Radianal in oblj-
que position intermediate in size. Anal X largest.
strongly convex transversely, adjains posterior
basal. Righttube plate smallest, notin contact with

340

FIG. 15. Minilyacrinus williamburyensis sp. nov.,
posterior and B ray views of weathered holotype
crown, WAM91,697, x 1. C, fragmentary set of arms,
UWAF1 13504, * 1.5.

MEMOIRS OF THE QUEENSLAND MUSEUM

radianal, proximal end barely below radial sum-
mit, admedial to anal X, barely visible in lateral
view. IBrr 2 per ray, rectilinear, wider than long,
thick, straight longitudinally, strongly convex
transversely; axillary IBrr2 constricted slightly
proximally, 11Brr and JI1Brr cuneate, wider than
long, thick, strongly convex transversely. All
branches isotomous on IBr2 and Brg, minimum
4 arms per ray. Pinnules moderately large but not
ramules, one per brachial on alternate sides of arm.
Stem and tegmen not known.

REMARKS

This crown is inverted with the arms splayed
outward on the substrate. The stem, infrabasal
circlet and parts of most basals are not preserved
but impressions of the inner side of the infrabasals
and basals are moderately clear. The anal series
extending into the base of the anal tube or sac is
large, nearly the size of the base of the arms.

Spheniscocrinus was defined by Wanner, 1937,
as having only one anal in the cup, however, the
holotype and only specimen appears to be slightly
distorted in the base of the C ray and adjacent anal
series. It may have an intermediate arrangement
of the anals as described above.

This is only the second member of the genus and
is distinguished from the type species by the fine
shagreen ornament, flush sutures and lack of
nodes or spines on the brachials. Generic distribu-
lion is extended to Western Australia strengthen-
ing the link between Timorese and Western
Australian Permian faunas.

Supertamily DECADOCRINOIDEA Bather, 1890F
amily DECADOCRINIDAE Bather, 1890

Trautscholdicrinus Yakovlev & Ivanov in
Yakovlev, 1939

TYPE SPECIES

Trautschaldicrinus miloradowitschi Yakovlev
& Ivanov in Yakovlev, 1939, Upper Carbon-
iferous Moscow Basin; by original designation.

Trautscholdicrinus sp.
(Fig. 14B-D)

MATERIAL

Partial crown WAMG67,529, from the Artinskian Fos-
sil Cliffs Formation, north branch Irwin River, Perth
Basin. Found by Mrs J, While.

DESCRIPTION

Cup medium bowl-shaped, shallow basal in-
vagination, walls subvertical, fine granular orna-
ment. IBB 5, small, equidimensional (2.0mm)

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

kite-shaped, slightly upflared, not visible in lateral
view. BB 5, hexagonal, wider (5.8mm) than long
(5.0mm), moderately convex transversely, strong-
ly convex longitudinally, proximally form base of
cup, distally subvertical forming nearly half cup
height, slightly tumid. RR 5, wider (5.7mm) than
long (3.4mm) slightly convex longitudinally,
moderately convex transversely. Radial facet
plenary, sloping gently outward, bearing
transverse ridge, outer ligament pit and furrow,
and wide shallow muscle areas adorally. Anals 3,
in intermediate position. Radianal lurgest, pen-
tagonal, adjoins C radial, BC and CD basals, anal
X and right tube plates. Anal X pentagonal, trun-
cated CD basal, adjoining one tube plate distally,
Right tube plate not preserved. proximal end bare-
ly below radial summit. A cay 1Bry axillary,
strongly convex transversely. Distal brachials
slightly cuneate, strongly convex transversely,
bearing one pinnule each distally. Stem impres-
sion round.

REMARKS

The cup is crushed from compaction but plates
are well preserved. The infrabasal circlet is not
visible in lateral view and sutures appear slightly
impressed because the tumidity of the basals
projects above them. This is the second crinaid
reported from the Fossil Cliff Formation after C.
divitara (Teichert, 1949), The specimen is of in-
sufficient quality to serve as a type. Radiuls like
these occur in the disarticulated ossicles at Cal-
lytharra Springs.

Superfamily canis Salata hl Bather,
Family CROM YOCRINIDAE Bather, 1890
Minilyacrinus gen. nov,

TYPE SPECIES
Minilyacrinus williamburyensis sp. nov.

ETYMOLOGY
For the Minilya River.

DIAGNOSIS
Cup bawl-shaped, with single anal; arms
uniserial, cuncate, numbering 20.

Minilyacrinus williamburyensis sp. nov.
(Fig, 15)

ETYMOLOGY
For Williambury Station.

341

MATERIAL

Holotype WAM91.697 from an undesignated locality
on Williambury Station. The specimen was enclosed in
a quartz and mica bearing grainstone of the Callytharra
Formation, It probably came from the vicinity of
Donnelly’s Well or other exposures of the Callytharra
Formation in the eastern line of the Gooch Range in the
southwestem part of Williambury Station. One partial
set of arms (UWA ) from the eastern line of the Gooch
Range along the Minilya-Williambury Station road.

DESCRIPTION

Crown moderately large, 42.8mm long incom-
plete, average 13.1mm wide, cylindrical. Anal sac
small cylindrical, formed of eight rows of plates.
Cup moderately large, 13.9mm high, 23.lmm
Wide wt summit, bowl-shaped; walls gently oul-
flared distally; moderately deep basal inyagina-
tion (7.1mm diameter); unormnamented; sutures
flush. IBB small, covered by proximal columnals,
confined te basal invagination, probably
downflaring. BB 5, large, slightly wider (8.8mm)
than long (7-8mm), strongly convex longitudinal-
ly, moderately convex transversely, proximal
ends within basal invagination, forming basal
plane of cup; CD basal symmetrical, truncated
distally for adjoined radianal. RR 5, large, wider
(14.7mm) than long (9.7mm), moderately convex
transversely, mildly convex longitudinally.
Radianal longer (8.6mum) than wide (6.5mm),
Slightly over half above radial summit, adjoined
distally by one lube plate. Radial facets plenary,
sloping gently outward, moderate gape between
radials and primibrachials, {Brr one per ray, wider
(14.5mm) than long (& 1mm), moderately convex
transversely, mildly convex longitudinally, axil-
lary. By 2 per ray, wider (8.4mm) than long
(4.1mm), strongly convex tranversely. 1] Brr
wider than long, slightly cuneate, strongly convex
transversely, thick, pinnulate. Ambulacral groove
narrow, deep. Branching isotomous, 20 arms.
Stem heteromorphic; columnals circular transver-
sely, lumen pentagonal. symplexy articulation,

REMARKS

The partial crown has been deeply weathered on
exposed surfaces, the distal part of the arms and
anal sac ure lost, slight crushing occurred oblique
to the A-CD symmetry plane, and arms adjacent
to the anal series have been disarticulated in part.
Weathering along the flush sutures of the cup
plates and into cleavage planes in the calcite plates
appear as deep impressions. A small portion of the
Band Crays, including radials and adjacent basals
were cnclosed in matrix and unweathered.

342 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 16, Texacrinus goochensis sp. nov. A-E, paratype UWAF115099. A, weathered crown showing attached
stem, X 1. B-D, oral, posterior and basal views of dorsal cup with attached primibrachial, < 3. E, enlargement of
part of A showing parts of arms to upper left and directly above cup and change in heteromorphic patterns,
proximally type II to distally Type I, X 2. F,G, lateral and posterior views of holotype crown UWAF 113387, F
xX2;G X3.5.

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

Removal of the matrix exposed the lack of or-
namentation and flush sutures.

The partial set of arms show the tapering distal
arm tips of slight cuneate shapes typical of the
cromyocrinids, Thickness of the plates, narrow
arobulacral grooves, strong external convexity
and slight cuneate shape of the brachials is similar
to that of uniserial brachials in the
Cromyocrinidae. However, cup plates lack the
impressed sutures typical of smooth plated
cromyocrinids with unisenal arms (Webster,
1981).

The single anal and multiple arms reflect an
advanced evolutionary condition. This is the first
cromyocrinid bearing uniserial arms to be recog-
nised with more than 10 arms, whereas many
biserial armed cromyocriniods have 12 to 19 arms
(Webster, 1981).

Evolution of Minilyacrinus is uncertain, per-
haps through Cromyocrinus, Goleocrinus or
Mooreocrinus by increase in arm number, redue-
tion of the number of anals in the cup and increase
in depth of the basal invagination. Evolution from
Meapacrinus would only require increase in arm
number.

The Cromyocrinidae is an Early Carboniferous
to Early Permian family with greatest generic and
specific diversity in the Late Carboniferous
(Webster, 1981), They have been reported from
North and South America and Eurasia. Webster
(1981) considered all Permian forms from Timor,
Russia and India previously referred to the
cromyocrinids to belong to an unnamed genus or
genera of non-cromyocrinid stock. Only two
cromyocrinids, Moapacrinus and an unnamed
genus, are of Permian age. Both are from the
southwestern United States, have only one anal,
and the 10 arms are uniserial on Moapacrinus but
unknown on the unnamed genus (Webster, 1981),

Minilyaerinus is the first cromyocrinid from
Western Australia. The partial set of 10 arms
identified as Poteriocrinidae gen. ind. et. spec.
nov. ind. (Wanner, 1949, pl, 3, fig. 31) from Timor
probably belongs to a cromyocrinid.

Superfamily ERISOCRINOIDEA Wachsmuth
& Springer, 1886

Family GRAPHIOCRINIDAE Wachsmuth &
Springer, 1886

Tapinocrinus Webster, 1987

TYPE SPECIES
Tapinocrinus macurdai Webster, 1987 from the

343

Artinskian Callytharra Formation, W.A.; by
original designation.

Tapinocrinus spinosus Webster, 1990
(Fig. 11E)

MATERIAL

WAM91.696 from the Wandagee Formation, aarth
side Minilya River, Coolkilya Pool area, Wandagee
Station, Western Australia: collected by H. Coley.

REMARKS

A second specimen of Tapinocrinus spinosus,
42.5mm long, 20.l1mm wide, shows the delicate
shape and pomted apex of the fully enclosed
crown. A very fine shagreen ornamentation is
preserved along the arm sutures on brachials of the
D ray, This is not preserved on the specimen
teported by Webster (1990).

SuperfamilyTEXACRINOIDEAStrimple, 1961
Family TEXACRINIDAE Strimple, 1961
Texacrinus Moore & Plummer, 1940

TYPE SPECIES

Texacrinus gracilis Moore & Plummer, 1940)
from the Middle Pennsylvanian, Millsap Lake
Formation, Texas; by original designation.

Texacrinus goochensis sp. nov.
(Fig. 16}

ETYMOLOGY
For the Gooch Range.

MATERIAL

Holotype UWAF | 13387, a crown collected by Chris
Hearne; paratype UWAL15099 collected by John Wil-
liamson from QML?770.

DIAGNOSIS
Ornament granulose.

DESCRIPTION

Crown small, weakly outflared, anal sac un-
known, granulose ornament on cup plates and
proximal brachials. Dorsal cup small, low bow!l-
shape, nearly three times wider than high, widest
at radial summit, moderately deep basal invagina-
tion, sulures flush to gently impressed. moderate
pits at apices of basals and radials. IBB small,
confined to basal invagination, covered by
proximal columnals, probably downflared, BB 5,
wider than long, tumid, forming walls of invagina-
tion proximally, basal plane of cup medially,
lower part of gently outflaring walls of cup distal-
ly, distal lip incurved; posterior basal may be

344

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 17. A-D, Texacrinus gascoynensis sp. nov., posterior, basal, BC interray and B ray views of holotype
WAM91.698, * 1. E-G, Texacrinus hardmanensis sp. nov., basal, BC interray and posterior views of holotype

CPC30683, 1.5.

truncated distally for anal X. RR 5, approximately
twice as wide as long, mildly convex transversely
and longitudinally, tumid, outflared. Radial facets
plenary, sloping outward gently, with well de-
veloped transverse ridge, outer ligament pit and
muscle fields. Anals 3 in primitive or modified
primitive arrangement, variable in relative sizes.
Radianal pentagonal or hexagonal, may or may
not adjoin D radial, longer than wide. Anal X

longer than wide, 0.5-0.67 below radial summit,
adjoining one tube plate distally. Right tube plate
small, proximal 1/3 below radial summit, adjoin-
ing one tube plate distally. [Brr one per ray, wider
than long, constricted distally, strongly convex
transversely, unequal in length (A ray longest),
axillary. UBrr to VIBrr slightly cuneate, equi-
dimensional to wider than long, branching on 4th
or Sth brachial in each series. First branch

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

JAS

aseeae

FIG. 18. A, Galateacriuus australis sp, noy., anal view of holotype UWAF1)3388, 1. B, Family and genus
indet., lateral view of set of arms QMF21469, x 1.6. C,D, Timorechinid indet., lateral views of set of arms,

QMF21468, X 1.6.

isotomous, thereafter heterotomous bi-ex-
otomous, 30 to 50 arms, Stem round, columnals
thin heteromorphic type II pattern proximally,
thicker heteromorphic type I pattern distally, latus
convex, non-cirriferous, Jumen pentagonal.

MEASUREMENTS (mm)

Paralype
UWA113385

Basal ae
Radial length

IBrl length
IBrl width

Diam. proximal
columnal

REMARKS

The holotype is a crushed crown lacking the
distal parts of the arms with the anals in primitive
arrangement; the radianal is in oblique position
not in contact with the D radial. Weathering of the
paratype has removed all except one primibrachial
and distal parts of the arms of two rays, The
paratype has anals in a slightly advanced arrange-
ment; the radianal adjoins the D radial separating
anal X from the CD basal. The slightly advanced
arrangement of the anals has been reported in all
species previously assigned to the genus suggest-
ing that ithas been a fairly stable genetic character
since the middle Pennsylvanian.

Texacrinus has been reported from middle Pen-
nsylvanian to Early Permian strata of the United
States (Webster, 1973). Variables within the fami-
ly are degree of suture impression (slight to
moderate), depth of basal invagination (slight to
deep), development of apical pits (not present to
moderately deep), cup shape (slightly globose to
weakly outflared Jow to medium bowl), and
brachial form (nearly flat to strongly convex
transversely; rectilinear to slightly cuneate, very

346 MEMOIRS OF THE QUEENSLAND MUSEUM

|
|
|
a
a
|
|

FIG. 19. Skaiocrinus granulosus Webster, 1990, A, C

ray view of WAM91.669 showing bi-exotomous
branching pattern, X1.6. B, lateral view of
WAM91.668 showing recurved anal tube and interior
of some arms, X 1.6. C,D, lateral and posterior views
of WAM91.694, % 1.6; note anal tube on interior of
lateral view. E, lateral view of WAM91.671 showing
recurved anal tube and proximal pinnules, X 1.3.

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

short to short), Where known, the arms branch
bi-exotomous after an initial isotomous division.
We consider T° interruptus, T. irradiatus and T.
compactus named by Strimple (1952) and ull from
the Francis Shale of Oklahoma to be one species
showing slight to moderate variation in many of
the cup features listed above, We select T. ir-
vadiatus to be the senior synonym because the
holotype is a more complete individual and inter-
mediate in the range of variation.

Texacrinus goochensis extends the geographic
range of the genus to Western Australia and is the
first species named with granulose ornament.

Texacrinus hardmanensis sp, noy.
(Fig. 17E-G)

ETYMOLOGY
For Mount Hardman,

MATERIAL

Holotype CPC30683, a partial crown, from the
Tatarian Cherrabun Member, Hardman Formation, at
Mt. Hardman, Canning Basin. BMR localities KLB 11.

DIAGNOSIS

Anals in intermediate position with proximal
lips of anal X and radianal below summit; arms
elongate.

DESCRIPTION

Crown elongate (68.5mm, slightly incomplete),
eylindrical, widest at radial summit, no arm girdle,
fine vermiform to granular ornamentation on all
plates, crushed slightly normal to plane of sym-
metry. Cup bowl-shaped, wider (16,3mm
average) than high (9.7mm), base invaginated,
sutures shallowly impressed, shallow apical pits.
IBB 57, covered, small, probably downflared, not
visible in lateral view. BB 5, wider than long (C
basal 6.4mm wide, 5.9mm long), gently convex
transversely, moderately convex Jongitudinally,
forming base of cup proximally; CD basal trun-
cated distally for radianal. RR 5, wider (8.9mm)
than long (6.3mm), gently convex. Radial facets
plenary, subhorizontal, narrow gape. Anals 3, in
inlermediate position. Radianal large, 4.4mm
long, 3.8mm wide, in line of radials. Anal X and
right tube plate subequal, 2.5mm wide, 2.4min
lang, proximal ends barely below radial summit,
each adjoining one tube plate distally. [Brr one per
ray, wider (9.7mm) than long (5.2mm), slightly
convex longitudinally, moderately convex
transversely, axillary. [DAer, 1iBer, VB and
VBrr rectilinear, wider than long, strongly convex
transversely, branch on Sth to Sth brachial in each

7

series, First branching isotomous, thereafter
heterotomous bi-exotomous, minimum of 10 arms
per ray. Pinnules slender, borne by alternating
branchials. Stem pentagonal.

REMARKS

The crown was enclosed in a quartz wackestone.
It had been crushed slightly by compaction. I-
regular calcite cementation of the matrix and
hydrous iron oxide cementation and replacement
of plate edges along some sutures made prepara-
tion difficult.

The specimen is an advanced member of
Texacrinus with the most elongate slender arms
known within the genus. It probably evolved from
T. goochensis by elongation of the brachials and
modification of the anals. ft is one of the three
youngest known Palaeozoic crinoids,

Texacrinus gascoynensis sp. nov,
(Fig. i7A-D)

ETYMOLOGY
For the Gascoyne River.

MATERIAL

Holotype WAM91.698, from the Artinskian
Cundiego or Wandagee Formation on Jimba Jimba
Station, Carnarvon Basin. Exact horizon and locality
unknown.

DIAGNOSIS

Radial facets peneplenary: anals in a near ad-
vanced position with only proximal most tips of
anal X.and right tube plate below radial sumimul.

DESCRIPTION

Crown, robust, moderately short, 56.9mm long,
average 37. 4mm wide, cylindrical, Cup moderate
bowl-shaped, large, 10,0mm high, average
33.7mm wide; basal invagination 12.4mm
diameter, 4.0mm deep} sutures flush; shagreen
ornamentation. IBB 5?, covered, small, confined
to basal invagination, probably downflared. BB 5,
large, wider (12.9mm) than long (10,0mm),
strongly convex Jongitudinally, mildly convex
transversely, forming basal plane of cup; CD basal
truncated distally for radianal. RR ‘5, large, nearly
twice as wide (17,.6mm) as long (9.0mm), gently
convex longitudinally and transversely, mildly
outflaring. Radial facets peneplenary with narrow
notch, sloping gently outward, Radianal large,
slightly wider (8.1mm) than long (7.8mm), ad-
joined disially by anal x and night tube plate
proximal most tips of which are below radia)
summit. [Brr one per ray, pentagonal, wider

348

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 20. Cymbiocrinus cherrahunensis sp. nov. A, D ray arm showing cuneate brachials, * 1. B-D, posterior, C
ray and basal views of holotype CPC30684, X 0.75. E,F, internal and external views of paratype basal QMF21480,

xK2,

(14.5mm) than long (6.7mm), constricted distally,
strongly convex transversely, gently convex lon-
gitudinally, axillary. IBrr to VUBrr rectilinear,
wider than long, moderately convex transversely,
straight to gently convex longitudinally; brachials
number 2 I]Br; 3 IIBr, 41VBr, 4or5 VBr,5 VIBr,
3 or 4 VII Br, unknown number of VIII Br. First
branching isotomous, all others heterotomous bi-
exotomous; minimum 14 arm per ray. Anal sac
unknown, probably short. Stem round transverse-
ly, heteromorphic. Latus strongly convex; axial
canal round.

REMARKS

This nearly complete crown is replaced with
hydrous iron oxides and weathered a deep fer-
tuginus red. It was embedded in a medium-
grained quartz wackestone and slightly
compressed normal to the A-CD axis. The radial
facets are peneplenary and distinguish it within the
genus. If the base of the anal X and right tube plate

were any higher they would be out of the cup and
the form would be considered to have a single
anal.

Consideration was given to designating a new
genus based on the combination of the peneplen-
ary facets and near single anal condition. Until the
single anal condition is established we prefer to
retain this form in Texacrinus.

Galateacrinus Moore, 1940

TYPE SPECIES

G. stevensi Moore, 1940 from the middle Pen-
nsylvanian Oologah Limestone of Oklahoma; by
original designation.

Galateacrinus australis sp. nov.
(Fig. 18A)

ETYMOLOGY
Latin australis, southern.

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA 349

FIG, 21. Jimbacrinus donnellyensis sp. nov. A, posterior view of holotype CPC30685, *4. B, lateral view of
weathered paratype crown with arms splayed CPC30690, % 3, C, oblique view of infrabasal circlet and basal
invagination of paratype CPC30686 on same slab as A, % 3.5, D, posterior views of paratypes CPC30687 (right)
and CPC30688 (left) with Stomiocrinus merlinleighensis sp, nov, paratypes CPC30674 (to left of stems, OSem
helow base of CPC30688) and CPC30675 (extreme bottom right corer), * 3.

350

MATERIAL
Holotype UWAF1 13388 from QML770.

DIAGNOSIS

Cup low bowl-shaped, lacking radial notches;
brachials very short, locally becoming biserial
within the uniserial arms.

DESCRIPTION

Crown tall (85.9mm incomplete) moderately
outflaring (55.3mm wide at IBr1), anal sac stout.
Dorsal cup low bowl-shape, nearly discoid, twice
as wide (33.3mm maximum) as high (16mm
diameter); sides widely outflaring; wide deep
basal invagination (9.9mm); coarse vermiform(?)
ornamentation; sutures weakly impressed. IBB
restricted to invagination, probably downflared,
not exposed if preserved. BB 5, large, CD 8.8mm
long, 10.2mm wide, moderately convex lon-
gitudinally, gently convex transversely, proximal
ends in basal invagination, widely outflared dis-
tally, forming basal plane of cup; CD basal trun-
cated distally for adjoining anal X. RR 5, large,
wider (15.5mm) than long (9.6mm), outflared
widely; radial facets plenary, sloping strongly out-
ward, Anals 3, primitive arrangement, distal 1/3
of anal X and right tube plate above radial summit.
Radianal pentagonal, in oblique position, slightly
longer (7.9mm) than wide (6.7mm); adjoining C
radial, BC and CD basals, anal X and right tube
plate. Anal X hexagonal, longer than wide
(8.5x6.2mm), adjoins one tube plate distally.
Right tube plate hexagonal(5.5x5.5mm). [Brr
l/ray, wider than long (15.5x6.2mm), moderately
convex transversely and longitudinally, axillary.
I[Brr to VIBrr strongly convex transversely, wider
than long, cuneate with local biserial brachials.
First branch isotomous, thereafter heterotomous
bi-exotomous. Second branching on IIBrr4 or 5,
third through sixth branch on 6th to 8th brachial
in the series; 12 arms per ray. Pinnules, anal sac
and column not preserved.

REMARKS

Only the C and D rays and adjoining parts of the
dorsal cup are exposed and perhaps the only parts
preserved. Its lack of radial notches and posses-
sion of cuneate brachials leave it transitional be-
tween the Texacrinidae (lacking radial notches
with rectilinear to cuneate brachials), and the
Galateacrinidae (with radial notches and cuneate
brachials). The radial notches of the
Galateacrinidae are formed by the very bulbous
nature of the radials as the facets are actually
plenary but adoral of the aboreal tumidity. Thus
the Galateacrinidae are separated from the

MEMOIRS OF THE QUEENSLAND MUSEUM

Texacrinidae on the shape of the cup, discoid vs
low bowl respectively, not the length of the radial
facet as the presence of radial notches tends to
imply. We question if this is of sufficient dif-
ference to justify family separation.

This is the first record of a Galateacrinus in
Australia. The genus has previously been reported
only from the middle Pennsylvanian to Early Per-
mian of the United States (Webster, 1973, 1977,
1986). The low bow] shaped cup, lack of radial
notches and short brachials which locally become
biserial readily distinguish G. australis from all
other species of the genus.

Skaiocrinus Webster, 1990

TYPE SPECIES

Skaiocrinus granulosus Webster, 1990 from the
Artinskian Wandagee Formation, Minilya River;
by original designation.

Skaiocrinus granulosus Webster, 1990
(Fig. 19)

Skaiocrinus granulosus Webster, 1990:66, pl.2, figs
15-17.

MATERIAL

Cup WAM91.684; 6 partial crowns, WAM91.668,
91.669, 91.671, 91.692, 91.694, 91.696 and 3 sets of
arms WAM91.670, 91.691, 91.693 from the Artinskian
Wandagee Fmn, Minilya River, collected by H. Coley.

REMARKS

Several morphological features not visible or
showing variation not recognisable on the original
study material (Webster, 1990) are shown on the
new specimens. Cup morphology is quite consis-
tent with the exception of the width of the radial
facets. On the holotype they are nearly plenary but
must be considered peneplenary, whereas they are
plenary or very nearly plenary with slight varia-
tion on all new specimens. Arms may branch up
to 8 times (WAM91.692), resulting in 9 arms in
one half ray and 90 arms for the crown if branch-
ing is uniform and maximum in all rays. Pinnules
are very slender, taper slightly until the distal
termination, have a medial longitudinal ridge,
begin on the first secundibrach, and contain 9
plates in proximal pinnules. The slender tegmen
is formed of small polygonal plates with flush
sutures, nO appearant transverse or longitudinal
alignment of plates and is recurved. The column
is heteromoprhic Type II pattern.

Based on the peneplenary radial facets, Webster
(1990) assigned Skaiocrinus to the

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

Pachylocrinidae noting exotomous arm braching
and similarity to the Texacrinidae. The genus is
here transferred to the Texacrinidae because most
of the new specimens show plenary facets.

No formation was known for the original study
material. New specimens are from the Wand/agee
Fonnation, in a calcareous siltstone to medium-
grained quartz wackestone. Cementation is
generally weak but may include hydrous iron
oxides which are difficult to remove when prepar-
ing specimens, Natural weathering exposed most
of the features described above.

Family piblmatsa ine Strimple & Wat-
Ins

Cymbiocrinus Kirk, 1944

TYPE SPECIES

Cymbiocrinus romingeri Kirk, 1944 from the
early Chesterian St. Genevieve Limestone of
Alabama; by original designation,

REMARKS

The Cymbiocnnidae may be divided into 2
groups with differing arm structures Cymn-
biocrinus, Aenigmocrinus, Oklahomacrinus and
Allosocrinus, have cuneate brachials,
Leecobasicrinus, Aesiocrinus, Paracymbiocrinas
and Proallosocrinus have rectilinear brachials.
Both groups appear in the Chesterian and range
into the Virgilian in the United States. They
probably had differing ancestral lineages and may
represent different families, Additional i investiga-
tion of the Cymbiocrinidae is required to properly
resolve these problems.

Cymbiocrinus cherrabunensis sp, nov,
(Fig. 20)

ETYMOLOGY
For the Cherrabun Member,

MATERIAL

Holotype, CPC30684, a crown from the Tatar-ian
Cherrabun Member, Hardman Formation, east side of
Mt Hardman, Canning Basin. Coll. J,M.Dickins,
Paratype QMF21480 fram QML 772.

DIAGNOSIS
Primanal large; 1] arms incurled distally.

DESCRIPTION

Crown 72mm long (arms curled distally), 7imm
Wide, inflated pear-shaped with incurled distal
arms, no surface ornament. Cup medium bow!l-
shaped, wider (273mm) than high (11-8mm),

351

walls outflared, base shallowly invaginated, su-
tures impressed, moderately deep apical pits. IBB
5, small, kite-shaped, 4.8mm long, 4.7mm wide,
downflared, not visible in lateral view of cup, BB
5, large, 10,0mm long. 10.8mm wide, inflated,
moderately convex transversely and longitudinal-
ly. RR 5, wider (14.3mm) than long (8.8mm),
outflared, gently convex transversely and lon-
gitudinally. Radial facet plenary, bearing
transverse ridge and outer ligament pit, slightly
guped, IBrr two per ray, moderately convex
transversely, constricted distally. 1Bry,
trapezoidal, wider (14.3mm) than long (3.7mm).
IBry triangular, 6.6mm wide 5.3mm long, axillary.
All other brachials, unless axillary, strongly
cuneate, bearing one slender pinnule each on al-
ternate sides of arm. Arms 11? all branching
isolomous, Proximal columnal pentagonal,
4.5mm diameter; latus convex. Crenularium nar-
vow, culmina short, coarse. Areola wide, flut.
Axial canal subpentagonal.

REMARKS

Assignment to Cymbiocrinus is based on cup
shape, single anal, cuneate brachials, and branehi-
ing on the second primibrachial. A third arm on
the C ray (branching on the first secundibrachial
abanally) is not typical of the genus. Itis uncertain
if this represents an evolutionary development or
a variant. The bulbous basals show affinity to the
calceolispongiid, Jimibacrinus, however the cal-
ceolispongiids have inflated basals, 5 atomous
arms and most commonly a vestigal stem.

Hydrous iron oxides have replaced all plates and
the crown is embedded in a coarse-grained quartz
wackestone. The delicate arms are incurl-ed dis-
tally but visible in the natural weathered
specimen, Itis interpreted to have lived in or neur
the bunal site on a sandy substrate.

Family TIMORECHINIDAE Jackel, 1918
Timorechinid gen, indet, (Pig. 18C,D)

MATERIAL
QMF21468 a partial set of arms, QML770,

REMARKS

A small incomplete set of arms (25.0mm long)
with 3 series of brachials shows 2 isotorious
branchings, Brachials are wider than long
proximally, equidimensional medially, longer
than widedistully, strongly convex externally, and
slightly cuneate. Axillary brachials (5-7 in middle
series) have a central upward and outward
directed short spine. Most spines are broken off

352

=

and the weathered fracture is sometimes difficult
to detect. Isolated spinose axillary brachials
probably belonging to this specimen are
moderately common at Callytharra Springs.
Proximal series of brachials twice width (3.9 mm)
of middle series (1.9mm); distal series (1.1 mm
wide) >1/2 width of middle series. Except for
spines on axils and smaller size, this set of arms is
similar to Notiocrinus timoricus Wanner (1924a,
pl.12, figs 18,19) from Basleo, Timor.

Because the branching pattern cannot be deter-
mined and the cup is lacking, the arms are ques-
tionably referred to Timorechinidae, with iso-
tomous and heterotomous biendotomous arms.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 22. A-F, Baryschyr? carnarvonensis sp. nov. A,

weathered mediam stem pluricolumnal, paratype
QMF21470, X0.8. B, weathered distal stem
pluricolumnal with multiple cirral facets, paratype
QMF21471, X0.8. C, deeply weathered medial stem
pluricolumnal, paratype QMF21472, x 1.5. D,E, ar-
ticular facet and lateral view of distal stem of
pluricolumnal holotype CPC30693, x 1. F, weathered
medial stem pluricolumnal paratype CPC30694, x 1.
G, Barycrinid? indet., weathered exterior surface of
pluripentamere, QMF21437, X2 (see Fig. 12G for
internal view). This specimen placed here for com-
parison of weathering characters.

Superfamily CALCEOLISPONGIOIDEA
Teichert, 1954
FamilyCALCEOLISPONGIIDAEF Teichert, 1954
Jimbacrinus Teichert, 1954

TYPE SPECIES

Jimbacrinus bostocki Teichert, 1954 from the
Artinskian Cundlego Formation, Western
Australia; by original designation.

Jimbacrinus donnellyensis sp. nov.
(Fig. 21)

ETYMOLOGY
For Donnelly’s Well.

MATERIAL
Holotype CPC30685, paratypes CPC30686-30691

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

from the upper part of the Artinskian Bulgadoo Forma-
tion in the type section near Donnelly’s Well.

DESCRIPTION

Crown small, pear-shaped, arms incurled distal-
ly. Cup bowl-shaped, walls subvertical, base
slightly invaginated, moderately deep apical pits.
IBB 5, kite-shaped, longer than wide, proximally
downflared, distally upflared, visible in lateral
view of cup. BB 5, hexagonal, subequal to wider
than long, proximally subhorizontal, distally
upflared becoming subvertical at distal tip; CD
basal heptagonal, truncated distally for primanal.
RR 5, pentagonal, wider than long, subvertical,
bear shallow medial! longitudinal depression.
Radial facet plenary, bearing transverse ridge,
outer ligament pit and marginal shelf. Primanal
large, longer than wide, proximal 2/3 in radial
citclet, adjoins 2 tegmen plates distally. Arms 5,
atomous. Brachials rectilinear proximally,
cuneate distally, strongly convex transversely,
concave longitudinally, bearing faint longitudinal
ridges after 3rd or 4th brachial; IBr2 bearing two
coarse slightly diverging laterally projecting blunt
nodes. Pinnules large, elongate, one per brachial
on alternate sides of arm. Stem pentagonal in
section proximally; latus convex; heteromorphic
type 1 pattern; distally round, homeomorphic,
bearing few short cirri. Axial canal round.

MEASUREMENTS (mm)
| ond oi
5S | 58
Crown height
Crown width
Cup height
Cup width
Diameter IBB ‘2
IBB length 1:90* | 3.4
IBB width 3,
BB length 25 | 22
BB width
RR length 23 | 20[ |
RR width 31 | 29 | |
Primanal length 12 |
Primanal width 16 | 16] |
Columnal diameter

*estimated

353

FIG, 23. Neoschisma verrucosum Wanner, 1924b. A,B,
Cradials QMF21481] and 21482, * 10 and X 6, respec-
tively. C,D, oral and Jateral views of D radial
QMF21483, %5 and %6, respectively. E,F, external
and internal views of epideltoid QMF21484, * 7, G,H,
paberadl and internal views of hypodeltoid QMF21485,
x

354

REMARKS

This description is a composite based on the 7
specimens. The holotype has a well preserved cup,
proximal most column and C ray arm but exposed
hasals have been rounded by solution. One
paratype (Fig, 21C) shows the infrabasal circlet
and prominent protrusion on the basals. Two
specimens (Fig. 21A,B) have well preserved
brachials one and two, and two other specimens
(Fig.21D) have well preserved columnals show-
ing the gradanen from the pentagonal to round
sections within approximately one centimeter of
the cup. Weathenng has etched the surface of most
specimens producing nodes and grooves which
could be mistaken for ornamentation unless all
specimens ate considered.

Unlike most species of Calceolospengia that
rested on the substrate with extended basals
(Teichert, 1949), Jimbacrinus lived above the
substrate supported by the stem (Teichert, 1954).
All plates except the basals of J. donnellyensts are
relatively thin and the crown is small relative to
the stem; these features suggest an elevated life
habit.

This 1s the third species of Jimbacrinus recog-
nised and it is the oldest, It is easily distinguished
from J. bostocki and J. minilyaensis by the wider
basal plane and subhorizontally directed basal
protuberances. J. boslocki arobably evolved from
J, dannellyensis by development of the central
spine on the basals.

Family and genus indet.
(Fig. 18B)

MATERIAL
QMF21469 4 partial set of arms from OML770,

REMARKS

This set of arms includes parts of 4 half(?) rays
which could be a part or parts of one to three rays.
The brachials arc: moderately (proximally) to
slightly (distally) wider than long; externally
strongly convex transversely; bearing a medial
longitudinal ridge terminating distally in a very
short spine directed upward and outward; and
moderately (proximally) to strongly (distally)
cuneate. Brachials are slightly offset giving arms
a slightly zigzag appearance, which is enhanced
by the alternating direction of the medial
nidge/spine. Isotomous endotomous or exolomous
branching occurs twice on two of the partial rays
and possibly a third ime on onc. A slender pinnule
articulates with the branchial at the distal end of
the medial ridge aboreal to the spine,

Classification of this specimen is uncertain. The

MEMOIRS OF THE QUEENSLAND MUSEUM

zigzag brachials resemble some genera of the
Decadocrinoidea, but they only have 10 arms,
isotomously branching on the first or second
primibrachial. Four superfamilies of the
poteriocrininids include genera with endotomous
or exotomous branching. The Hydreionocrinoidea
have biserial arms and given no further considera-
tion, The Pirasocrinoidea typically have spine
rings capping the anal sac, brachials are rectilinear
to faintly cuncate, and few genera are actually
known to have endotomous or exotomous branch-
ing. Linocrinus, Tholocrinus and Exocrinus, of the
Zeacrinitoidea have ray ridges or cuncate
brachials but all lack the zigzag offset of the
brachials. Only Lingerinus, has both ray ridges
and cuneate brachials, Cuneate brachials are char-
acteristic to several genera of the Texacrinoidea.
Galateacrinus ernatus Moore, 1940 has medial
ridges terminated in short spines on cuneate
brachjals, but the brachials are not offset.

Lacking proximal parts of the arms and the cup
the specimen is left in open nomenclature, Tt is
illustrated to show recognised diversity in the
advanced poteriocrinid faunas from the Callythar-
ra Formation, The branchials and branching puat-
tern are sufficiently distinct to offer correlation
potential if similar specimens are found else-
where.

Family CYCLOMISCHIDAE Moore &
Jeffords, 1968

Barysehyr Moore & Jeffords, 196%

Crinoid stem Foord, 1890, p,104, pl.4, fig.11,
Enlarged crinoid stem-portions Etheridge, 1903, p.25,
pl.4, figs 1-9.

TYPE SPECIES

Baryschyr anosus Moore & Jeffords, 1968 from
the Pumpkin Creek Limestone Member, Domick
Hills Formation, Pennsylvanian of Oklahoma; by
original designation,

Barysehyr? carnarvoneusis sp- nov.
(Fig. 22A-F)

ErMoLocy
For the Camarvon Basin.

MATERIAL

Holotype CPC30693, paratypes CPC30694,
QMF21470, 21471,21474 from QML770, QMF21472,
21473 from QML757 and 368 specimens as follows
QMF21588 (11 specimens) from QML757, OMF21589
(25) from QML768, QMF21590 (11) from QML769,
QMF21591 (201) from QML770. BMRF1643 (28),

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

355

FIG. 24. Neoschisma extensum sp. nov. A, fragment of a C radial QMF21486, x 2. B, small radial QMF21487,
™“5.C.D, a fragmentary and a complete D ridial QMF21488 and 21489, *3. E, fragmentary articulated theca
NMVP127198, 2.5. F, large basal plate QMF21490, x 2.5.G,H, oblique lateral and lateral yiews of large radial
QMF21491, *2. I,J, oblique lateral and Jateral views of radial QMF21492, * 2.5 and & 2, respectively.

BMRF1644 (5), BMRF17226 (55), BMRF17601 (3),
BMRF19385 (4), BMRF19386 (6), BMRCC139 (12),
BMRG259 (3)and BMRGW$4 (4).

DESCRIPTION

Proximal columnals unknown; medial colum-
nals homeomorphic to heteromorphic type I; cir-
riferous distal columnals heteromorphic type I,
Columnals circular in transverse section, axial
canal pentagonal. Articulum wide, full facet width
with no areola to nearly full facet width with
narrow areola sloping into axial canal. Culmen
fine, branching isotomously at midwidth to distal
1/4 of articulum, 40 per em at mid width of ar-
ticulum. Symplexy articulation. Latus smooth,
straight. Axial canal narrows at moderately con-
vex claustrum on each columnal. Cirti circular in
transverse section, articulum like columnal ar-

ticulum. Cirral scars aligned in part, may give
columnal a rough pentagonal section, overlap |
then 2 adjacent columnals as growth increases;
cirri scars increase in size distally.

REMARKS

Large pluricolumnals, 5.0-9,4cm long and 1.7-
2.4cm diameter are especially common in ex-
posures of the basal part of the Callytharra
Formation throughout the Carnarvon Basin. They
have been collected for many years and are present
in most major museum collections throughout
Australia. The first illustrations of these colum-
nals was by Foord (1890) followed by Etheridge
(1903) who reported infestations of endophytic
fungus on some stem segments. More recently
Welsh (1976) has demonstrated that these infesta-
tions and others referred to myzostomids by Warn

356 MEMOIRS OF THE QUEENSLAND MUSEUM

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

(1974), among others, are caused by Phasyphan-
rtals.

Considerable morphologic and diagenetic
variation occurs in the pluricolumnals of B.? car-
narvonensis. Many pluricolumnals lack cirri
whereas others have a few and others many. Those
with cirri show a progradation of increasing size
of the facet distally and a corresponding alignment
of the facet and more pentagonal transverse sec-
tion of the columnals with the larger facet sizes.
feteromorphic noditaxial patterns change distal-
ly, Proximal patterns are 2 or 3 intemodals. be-
tween nodals tn fairly regular repetitions. Distally
internodals are fewer and occur infrequently, one
every 3 to 7 noduls. The distal imegular noditaxis
patterns correspond to the most cirriferous
pluncolumnals.

Weathering of the pluricolumnals is greatest
along articular facets resulting in pluricolumnals
with a V-shaped or cog-shaped latus and what
appears to be umpressed sutures, Weathering on
some specimens has been decp and results in
irregularly shaped transverse cross-sections. The
large axial canal collapsed under compaction after
burial and many pluricolumnals appear elliptical
in transverse section,

Class BLASTOIDEA

The only descriptions of Western Australian
Permian blastoids have been those of Breimer &
Macurda (1972) who named the fissiculates Weos-
chisma australe and Notoblastus stellarts and
identified the Timorese species Thawmatoblastus
lonigramus Wanner, 1924b in Westem Australia
and Breimer (1983) where the detailed specics
descriptions were provided, Apart fram providing
further information on each of these a new
spiraculale species is described and the Timorese
species Neoschisma yverruscosum is recorded in
W.A, Apart from this specific match all have
generic links to Timer suggesting u close
biogeographical link between the two arcas.

Order FISSICULATA Jackel, 1921
Family NEOSCHISMATIDAE Wanner, 1940

Breimer & Macurda (1972) discussed fissicu-

late phylogeny in great detail having established
their classification apparently phenetically. All
the fissiculates described herein are assigned ta
the Neoschismatiduc following Breimer &
Macurda. The only comment regarding inter-
generic relationships concerns the placement of
Pterotoblastus. Although placed in the
Codasteridas by Wanner (1940) and by Breimer
& Macurda (1972) Pterefeblastus appears to have
quite a deal in common with various neoschis-
matids, in particular Nofoblastus, The major dis-
tinguishing feature appears to be the reduction in
the number of slits per hydrospire field and loss of
slits on the anal deltoid in Prerereblasius. Breimer
& Macurda (1972) made an emphatic point that
“Hydrospire structures of neoschismatids are es-
sentially unchanged; ... There is no trend towards
reduction in the number of hydrospire slits, neither
in regular, nor in anal hydrospire groups”. How-
ever, their further discussion of hydrospires in the
Family reveals an abundance of slits in the earliest
representative, Hadroblastus, and several of the
Permian genera but in several species, effective
reductions are recorded, According to Breimer &
Macurada (1972, p. 346) these reductions are ef-
fected through filling of the slits by secondary
calcile deposits, They regard Tonoreblastus as a
form with most uf its slits completely closed off
by secondary calette, Prerotoblastus has more slits
than Tinoreblastis and the small number in each
genus could be due to the same process, calcite
infilling of most slits. In the anal interray groups
of slits, Neoschisma contains one species with
approximately the sianme number of slits as non:
anal groups because the hypodeltoid bears
numerous slits; a second species has a greally
reduced number where the hypedeltoid docs not
bear slits. In Nofoblastus there are 1-5 slits in cach
anal hydrospire field. This is significantly fewer
than in anal fields of certain species in other
genera where slits are present on both epi- arid
hypodeltoids. Even in Noloblastus, No cornuiuy
(McKellar, 1969) has 5 slits per group in the D
interray, N. ovens’ (Wanner, 1940) and NV. sreliieriy
haye 3 and NV. brevispinus hus 2. Therefore, we
consider that the difference between 2 and 0 slits
in the anal group no more significant than that
between 5 und 3. We suggest that Prerotoblastus

FIG. 25, Neoschisma extensum sp, nov. A,B, external and internal views of delund QMF21493, <4. C.D, external
and internal views of holotype hypodeltoid QMF21494, 4. E\F, external and internal views of small deltoid
QMF21495, * 7 and. *6, respectively, G, external view of small deltoid QMF21496, %6. HI, internal and
extemal views of hypodeltoid QMF21497, * 4, J-L, internal, internal oblique and external views of large deltoid
with laneet still attached OMF2)498, 4, MN, internal and externa) views of large deltoid QMP21499, «4.
OP external and internal views of deltoid QME21S00_ x4 and 5. QL, intemal and external views of deltoid

QMP2ZISUT, * 4,

MEMOIRS OF THE QUEENSLAND MUSEUM

358

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

may be more correctly classified with the essen-
tially Permian Southern Hemisphere group, the
Neoschismatidae.

Neoschisma Wanner, 1924b

TYPE SPECIES
N. verrucosien Wanner, 1924b from the Per-
mian of Timor; by oripinal designation.

Neoschisma verrucosum Wanner, 1924b
(Fig. 23)

Neoschisma verrucosum Wanner, 1924b-7, pl. 799,
figs 9,10,

Neoschisma verrucosum Wanner, Breimer & Macur-
da, 1972:31, pl.19, fig.5, pl.20, fig.7.

Neaschisma australe Breimer & Macurda, 1972;31,
pL, figs 6,7, [non figs 1-4, 8].

Neoschisma verrucosum Wanner;Breimer, 1983: 126,
pl.26, figs 5,7,9.

Neoschisma australe Breimer & Macurda; Macurda,
1983: pl.27, fig. 2 (only); pl.28, figs 7-9, 12.

MATERIAL

Western Australia - UMMP58682 (holotype of NV.
australe) from the type section of the Callytharra For-
mation, 0.88 km west of Callytharra Springs, WA,
QMF21485 an hypodeltoid, QMF21484 an epidelwid
and QMF21481-21483 radials from QML757.

DISCUSSION

Breimer & Macurda (1972) diagnosed N.
australe as having a flatrather than scalloped vault
and as having hydrospire slits developed on both
the epi- and hypodeltoids, The surface of the vault
was a variable feature in the Callytharra collec-
tions with the depth of the VY in the top of the
radials and the convexity of the deltoids varying
with growth and among individuals of similar
size. With the relatively few specimens available
this feature Would seem unsuitable for species
separation, However, development of hydrospire
slits on both epi- and hypodeltoids should be clear
cut. In the type species, the hypodeltoid does not
bear any hydrospire slits (Wanner, 1924b, pl.
199, fig, 10; Breimer & Macurda, 1972, pl, 19,

359

fig. 5). In the holotype of N. australe (Breamer &
Macurda, 1972, pl. 19, figs 6, 7) the hypodecltoid
is a flat, fairly upright plate, tapering udorally but
not pointed, with the anal aperture an indistinct
invagination into the adoral side and with a dis-
tinct double tubercle medially on the aboral RD
margin.

Another such plate (Fig.22G,H) is now avail-
able with the same shape, distinctive aboral
tubercle and ragged adoral margin. In this
specimen the protruding ragged adoral edge con-
ceals the anal aperture which is obvious on the
internal side. The anal opening cuts through a
prominent curving ridge that mins across the plate.
A low but distinct ndge runs from near the centre
of this gap to the RD margin towards the C side.
Moreover this specimen bears no hydtospire slits.
On this basis we assign the holotype of N. australe
and the isolated hypodeltoid mentioned above to
N. verrucasum based on the lack of hydraspire
slits on the hypodeltoid. Radials (2 C and | D
radials; Fig,23A-D) with truncated upper corners
are referable t this Species also; the truncation
corresponds to the RD margin along the hypodel-
toid.,

On the other hand, the hypodeltoid figured by
Breimer & Macurda (1972, pl. 19, fig. 1) and two
others (Pig. 25C,D,H,1) do bear numerous
hydrospire slits. These plates represent a separate
species anu ure described below. To that species
are ulso assigned several radials which may be
identified as having the “broad convex humped
area” described by Breimer & Macurda (1972,
p.32) in the hydrospire field adjacent to the
hypodeltoid near its widest point. However, many
radials, basals and delioids, from other than the
anal past of the theca are not able to be assigned
with certainty. They are here assigned oN, exten-
sum for convenience only,

The epideltoid (Fig.23E,F) is assigned to this
species because the anal opening is irregular, anal
collar very Jow and there are hydrospire slits un
both its aboral extremities, Despite the consider-
able size difference between this epideltoid and
available hypodeltoids it seems to fit with the
hypodeltoid described above in that the low collar

FIG, 26. Notoblastus stellariy Breimer and Macurda, 1972. A-D, external views of deltoids QMF21502-21505.
A,B, *7; C.D x6. E, oral view of small radial QMF21506, #4. F,G, lateral and oral views of small radial
QMF21507, %4. H,1, oral views of radials CPC30967 and QMF21508, X3 and X 5, respectively. J,K, lateral
views of small radials QMF21509 and 21510, %4, L-N, oral view of epideltoids QMF21511-21513, L,M 3;
N 7, OP, internal and external views of lancetQMF21514, *7and x 10, Q, external view of lancet QMF21515,
* 10. R, internal view of lancet QMF21516, *9.8,T, oral and lateral oblique views of fragment of a radinl with
lancet still in place CPC30695, x4. U, fragmentary radial and lancet QMF21517, *6, V, fragmentary radial
CPC30966, * 3. W.X, oral and lateral views of articulated but badly weathered theca NMVPI27 199, 2,5. ¥,Z,

Jateral and oral views of large radial QMF2Z1518, 3,5,

360

MEMOIRS OF THE QUEENSLAND MUSEUM

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

and irregular opening would fit better with the
overhanging anal opening described above.

Neoschisma extensum sp, noy,
(Figs 24,25)

Neoschisma australe Breimer & Macurda, 1972:31,
pl.19, figs 1-4, 8,

Neoschisma australe Breimer & Macurda; Breimer,
19832122, pl.27, figs 1, 3-12; pl.28, figs 1-6, 710,
11, 13-16, pl.30, fig.3.

ETYMOLOGY
Latin extensus, stretched out; referring ta the
anal collar,

MATERIAL

Holotype QMP21494, an hypodeltoid from QML760,
Puratypes hypodeltoid and D radial (Breimer & Macur-
da, 1972, pl. 19, fig. 1), C radial (Breimer & Macurda,
1972, p. 19, fig. 3, 4; BMR GW 137), D radial (Breimer
& Macruda, 1972, pl. 19, fig. 8) NMVP127198 an
articulated but crushed and incomplete theca from
20.3m in the type section of the Callytharra Formation,
O.88km west of Callytharra Springs. QMF21490 frag-
mentary basal, QMP21486-21489, 21491, 21492 al-
most complete radials, QMF21493, 21495.
21498-21501 deltoids, hypodeltoid QMF21497 from
ML757. One deltoid, QMF21496 from QML760.

DISCUSSION

The problem of assigning non-anal plates to WV.
verrucosumor this new species has been discussed
above as has the hypodeltoid with hydrospire slits
belonging to the new species, One hypodeltoid
(Fig.25C,D) is of the same maximum width as the
isolated hypodeltoid of N. verrucosum
(Fig.23G,H) so the different hypodeltoids may not
be attributed to different growth stages of the same
species. Any isolated plates of Neaschisma that
cannot be definitely referred to one or other of the
species is tentatively referred to this new species.

The description of N. australe as given by
Breimer & Macurda (1972, p. 32) and Macurda
(1983, p.122) should be applied to this new
species except as varied in the following. The
fragmentary basals suggest thai the BB sides of
the plates diverged strongly adorally to give the

31

FIG: 28, Deltoid indei. A, CPC30710, x4. B,
QMF2i553, x3,

theca a strongly flared base. In large ratials, lateral
cdges are not always straight; often having a gent-
ly sigmoidal curve over adoral half (Fig.24H). In
C and D radials hydrospire slits are absent in a
zone near to and on the anal interarea side of the
central and ambulacral notch, This zone was
described by Breimer & Macurda (1972) as that
where a “broad convex humped area formed”. In
Specimens available to us (Fig.24A,C,D) the con-
vexily is at right angles to the hydrospire slits and
parallel to the RD margin with sharp furrows
crossing the humped area also parallel ta RD
margin (Breimer & Macurda, 1972, pl.19, fig.4;
Fig.24A). The upper margin of this convex zone
is In contact with a convex face on the hypodeltoid

just adoral to the hydrospire slits. On the interior

side of this convex area on the radial is a shallow
concave channel that leads across the RD suture
along the deltoid to the anal opening. Lateral to
this conyex zone on the radial, the projection of
the radial limb is greatly reduced and in one
specimen the hydrospire field is declined aborally
(Fig.24D). In this same CD interarea position
there appears to be a greater than normal develop-
ment of secondary calcite in front of the slits and
one specimen (Fig.24D) a shield has grown almost
completely hiding the slits from lateral view.
One available deltoid (Fig-250,P) is much
shorter, in interradial direction, than other del-
toids, This specimen has the aboral tip slightly
downtumed forming a shallow aboral pocket on
the internal surface. It has 8 hydrospire slits on one

FIG. 27. A.F,.G, Thaumaioblaxtus sp. nov. fragments of radial projection from KNF73, A, CPC30698, 3. FG,
CPC30702, x3 and * 7,5, respectively, B-E, J-O, Thaumarablastus indet. B-B from KNF73; J-O from ML89,
B, fragment of radial projection showing branched ambulacral tract, CPC30699, *3. C,D, oral and lateral views
of radial with incomplete projection, CPC30700, X 3,5, E, radial, CPC30701, * 4, J-M, radials with incomplete
projections, CPC30705, 30706, 30707, 30708, respectively, * 2.5, *3, ¥3, *4, respectively, N,O, lateral and
ural views of radial with incomplete projection and upon which a javenile molluse shell (probably an oyster) has
attached, CPC30709, * 2,5, H,l, Thaumatoblastus longiramus Wanner, 1924b, radials with incomplete projec-
lions but retaining the lancet and side plates, CPC30703, * 4 and CPC30704, & 3,

362 MEMOIRS OF THE QUEENSLAND MUSEUM

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

side and only 3 or 4 on the other, whereas elongate
deltoids of comparable width may have 15 or 16
slits. The side with only 3 or 4 slits has a long
aboral zone without slits. Whether this is from an
aberrant specimen or there is some other explana-
tion is not known at present. We have no epidel-
toid to assign to this species. The two hypodeltoids
appear dissimilar principally because one has a
very high anal collar, subdued comarginal orna-
ment and prominent distal blade-like projection
whereas the other has no raised anal collar,
prominent sharp comarginal ridges and no distal
projection. On the latter specimen the anal collar
and distal projection may have bene removed by
postmortem abrasion. Internally (Fig.25D,H) the
two plates are identical with a ridge across the
plate at the adoral end of the hydrospire slits. On
the surface of the anal groove, adoral to the
transverse ridge, are low irregularly placed
tubercles. Adorally on the plate are a pair of flat
facets presumably for junction with the epideltoid.

One deltoid retains a lancet on one side (Fig.25J-
L) and one radial retains the short aboral tip of the
lancet in the ambulacral notch (Fig.24G,H). The
DD margin of the deltoid is straight adorally, then
slightly concave and convex dipping down aboral-
ly. This shape is well illustrated in the assembly
of Breimer & Macurda (1972, pl.19, fig.1) but the
dorsal half of deltoids should be in contact. In oral
view the lancet corresponds to this shape being
narrow and depressed adorally beneath the abut-
ting deltoids, but wider and flush, with the sides
of the deltoids in the main body area where del-
toids do not touch. In Breimer & Macurda’s (1972,
p.33) description it is not clear which part of the
lancet is being referred to as adoral and which is
aboral; in this work the adoral section is where the
ambulacral groove is supported entirely by the
deltoids, the main body is where the ambulacral
groove is supported by the lancet and adoral sec-
tion is on the radial. The lancet is widest at aboral
end of main body, tapering very slowly adorally it
disappears internally at adoral end of the main
body. Small transverse grooves are evident within
the ambulacral tract but the arrangement of side
plates or even their presence is not confirmed by
available material.

363

Notoblatus Brown, 1942

TYPE SPECIES

Notoblastus brevispinus Brown, 1942 from the
Early Permian of New South Wales; by original
designation.

Notoblastus stellaris Breimer &Macurda, 1972
(Fig. 26)

Notoblastus stellaris Breimer & Macurda, 1972:34,
pl.20, fig. 1; pl.21, figs 1,2,7,8.

Notoblastus stellaris Breimer & Macurda; Breimer,
1983:132, pl.29, figs 5-7,11,12; pl.30, figs 1,2,4-
10,15-17.

MATERIAL

Holotype UMMP58683 from the type section of the
Callytharra Limestone on the Wooramel River, WA.
NMVP127199 an articulated but weathered specimen
from locality BB929 which is 20.2m above the base of
Section 7 on Mt. Sandiman Station, 1.9km at 15° from
Mt Sandiman Woolshed, from the collection of Dr G.A.
Thomas, now housed in the Museum of Victoria.
Radials QMF21506-21510, 21517, 21518, CPC30965,
30967, deltoids QMF21502-21505, epideltoids
QMF21511-21513, and lancets QMF21514-21516
from QML757.

DISCUSSION

This species was described in detail by Breimer
and Macurda (1972:34) and Macurda (1983:132);
only where new information is available are com-
ments added here. However, an attempt is made
to illustrate several features and plates not obvious
in earlier papers.

The theca described here is taller than that of
Breimer & Macurda (1972) with the basals ac-
counting for some 25-30% of total height; the
height is 17mm and maximum width 16mm but
this theca is compressed laterally so the width is
overstated. In life the theca must have been con-
siderably higher than wide.

On the radials, the ambulacral sinus extends
along the radial projection to the point where the
projection turns upwards; in the smaller
specimens this means the sinus does not extend
onto the projection at all but in the largest radial
(Fig.26Y,Z) the sinus extends well out onto the
projection to the point where the upper surface of
the projection is about to turn upwards. There

FIG. 29. Rhopaloblastus cuspidus sp. noy. A-C, lateral, oral oblique and Jateral oblique views of holotype theca
QMF21519, A X3; B,C X7. D-F, lateral, oral and posterior oblique views of paratype theca NMVP94182. D
<3; E,F <4.5. G,H, lateral and oral views of fragment of a lancet with 6 side plates attached QMF21520, x 10.
I, basal circlet of small theca QMF21521, X 3. J,K, internal and external views of a lancet still in place on a radial

QMF21522, X4 and X15,

364

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 30. Rhopaloblastus cuspidus sp. nov. A,C, large
basals QMF21523, 21525, *4. D, internal of small
basal QMF21526, <4. E,F, internal and external of
basal QMF21527, X4. B,L,N,O,R-T, radials
QMF21524, 21530, 21532, 21533, 21535-21537, all
X4exceptR X3.G-I, oral, internal and external lateral
views of radial QMF21528, <3. J,K, oral and lateral
views of fragmentary radial QMF21529, 5. M, inter-
nal view of fragment of lancet QMF21531, <6. P,Q,
internal and external views of damaged radial
QMF21534, x4.

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

365

FIG. 31, Cidaroid indet. A-C, spine bases QMF21541-21543, * 4. D-L, fragmentary interambulacrals QMF21544-

21550, all ¥ 7 except E andl * 6.

appears to be some variation in the shape of radials
through growth but the small sample size and
degree of postmortem deformation make it impos-
sible to identify how much, In the CD interray,
hydrospire slits are reduced to one ortwo pet field
und these situated close to the ambulacrum. In the
smaller radials described here, both in the articu-
lated specimen and as disassociated plates, the
hydrospire fields are not evident in lateral view;
although the individual plates all appear to have
undergone a degree of distortion this is unlikely to
have accounted for the attitude of the hydrospire
fields in every case. In large radials
(Fig.268,U,Y,Z) the hydrospire fields are more
upright and evident in lateral view.

Epideltoid same as a deJtoid except that
hydrospire fields are reduced to | or 2 slits on cach
margin projecuion at lateral extremes; these
projections were sutured to radials. On inner sides
of these projections are elongate facets that
presumably sutured to the hypodeltoid. In the
centre of the plate was a large circular anal open-
ing. The adorally chevron-shaped ridge of the
deltoid is greatly accentuated. No plate in our
collections may be certainly identified as the
hypodeltoid but its shape is inferred from the
epideltoid and radial that would have bounded it.
The hypodeltoid must have been wide and rhom-
boidal with the adoral corner excavated by the anal

opening so that a shape approaching broadly ar-
cuate is possible.

Since isolated lancets and those still in the am-
bulacra of weathered radials (Fig.268,U) show the
lancet to be elongate rhomboidal in plan view. The
adoral part (i.e. from widest point to adoral tip) is
shorter than aboral part which tapers very gently
and extends to tip of ambulacrum. Laterally on
adoral part are the slightly protruding faces. that
abut the excavation into the sides of the deltoids.
On internal surface is an adoral diamond shaped
area bounded by a distinct ridge and with concave
surface. The aboral tip of the diamond fills the
V-shaped gap in the radial and the aboral exten-
sion with rounded inner surface lies in the am-
bulacral groove on the radial,

Thaumatoblastus Wanner, 1924b

TYPE SPECIES

Thawnatoblastus longiramus Warner,1924b
from the Permian of Timor; by original designa-
tion.

Thaumatoblastus longiramus Wanner, 1924b
(Fig.30H,1)
Thaumatoblasrus longiranius Wanner:Breimer &
Macurda, 1972:36, pl.24, fig.2; pl.25, figs 1-8:
pL28, fig.5.

MEMOIRS OF THE QUEENSLAND MUSEUM

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

Thaumatoblastus longiranus Wanner: Macurda,
1983:135, pl. 31, figs 7-10; pl.32, figs 1-10.

MATERIAL.
Radial plates CPC30703, 30704 from ML87.

REMARKS

One of these radials (Fig.27H) was figured by
Breimer & Macurda (1972, pl.25, fig.5) and
agrees with the other in size of the side plates
relative to the width of the ambulacrum, These
side plates appear to be the same size and shape as
figured by Wanner (1924b) and their assignment
to the type species is accepted.

Thaumatoblastus sp. noy.
(Fig.27A,G)

MATERIAL
Radial plates CPC30698,30702 from KNF73 and
QMF30705-30709 fram ML89,

REMARKS

The two specimens with lancet and side plates
preserved (Fig.27A,F,G) show 4 greater number
of side plates per centimetre of ambulacrum when
compared with the 2 radials described above as
Thaumatablastus longiramus. These two
specimens are-considered specifically separate but
because they are so incomplete no new specific
name is introduced,

Thanmatoblastus sp. indet.
(Fig.27B-F,J-O)

MATERIAL
Radial plates CPC30699-30701 from KNF73 and
QMEF30705-30709 from ML&9.

REMARKS

Assignment of these radials, that have lost their
lancet and side plates, to either of the species
mentioned above is not possible with the present
state of knowledge. However, it is likely that they
belong to one or other of the species occurring in
Wester Australia. All those radials figured by
Breimer & Macurda (1972, pl.24, fig.2; pl.25, figs
1-4, 6,8; pl.28, fig.5) which do not retaim a lancet
or side plates are in the same situation and definite
specific assignment will depend on more detailed
study of this and better material from Western

367

Australia and Timor if and when it becomes avail-
able.

Family UNCERTAIN
Deltoid indct
(Fig. 28)

MATERIAL

Two plates QMF21553 from QML757 and
CPC30710 from 20m above base of the type section of
the Callytharra Formation.

DESCRIPTION

These two plates are hexagonal with the six
sides virtually of equal length. Two adjacent sides
have 7 or 8 slits normal to the margin; interpreted
as hydrospire slits. Comarginal growth lines are
evident parallel to the other sides; a group of 6-9
low rounded tubercles are situated centrally with
the central tubercle highest and most prominent.

REMARKS

Although these plates are abraded and only one
is preserved so as to be confident of its features.
they are interpreted as the deltoid plates of some
blastoid because of the hydrospire slits and
general resemblance to the deltoids of Notoblastus
described above. However, the six even sides to
these plates and apparently unspecialised edges
(ie. other than those with slits) are features un-
known among the blastoids. Their identity even as
blastoid plaies, remains a mystery at present but
they are recorded to show the diversity.

Order SPIRACULATA Jackel, 1921 |
Family PENTREMITIDAE d’Orbigny, 1851

Khopaloblastus Wanner, 1924b

TYPE SPECIES
R. umericus Wanner, 1924b from the Permian
of Timor; by orginal designation.

DIAGNOSIS
See Breimer & Macurda,1965:216.

REMARKS

Breimer & Macurda (1965) considered
Rhepaloblastus to differ from other genera in the
Peniremilidae by its petaloid ambulacra and ex-
posed cryptodeltoids. The species described
below has exposed cryptodeltoids but ambulacra,

FIG, 32, Monaster coleyi sp. nov. AE, adoral end and oral view of arm fragment (could be part of holotype but
not certain) NMVP127201, X7 and «8, respectively, B-D, F-H, holotype NMVP127200. B, lateral view of A
ray, *5.C,D, oral view, *4 and * 7, F\E, aboral view, x4 and * 6, G, enlargement of madreporite area, * 10,

368

in the largest available specimen, are not petaloid
but rather elongate, much more in the style of
Calycoblastus. We have assigned the species with
the emphasis on the anal deltoid arrangement be-
cause that feature seems generically more consis-
tent through the blastoids. The smaller theca
(Fig.29D,E) exhibits more petaloid ambulacra but
in all other respects this species coincides better
with the type of Calycoblastus than Rhopaloblas-
tus. Calycoblastus was originally protrayed (Wan-
ner,1924b) as having eight spiracles and an
anispiracle. Brown (1942) credited C. casei from
New South Wales with 10 spiracles. However,
Breimer & Macurda (1965) re-examined the
holotype of the type species arguing from the
weathered surface that it possessed four spiracles
and an anispiracle. This is in accord with the
diagnosis of Rhopaloblastus and with the species
described below but C. casei Brown, 1942 has yet
to be re-examined to verify this feature and deter-
mine whether it should remain in that genus.
Radials of this new species closely resemble
radials ascribed to Rhopaloblastus (?) belfordi
(Crockford & Brown) (Brown, 1942:102) from the
Permian of the Hunter Valley and to Rhopaloblas-
tus? cf. belfordi (McKellar,1969:25) from the
Berserker Beds near Rockhampton. Without
knowledge of the anal deltoids it is impossible to
assign these isolated radials generically; their
elongate ambulacra indicate Calycoblastus but the
similarity to those assigned to Rhopaloblastus
below suggest the earlier assignments may be
correct. It is surprising that no reference was made
to Calycoblastus in discussions of these eastern
Australian radials by either author.

Rhopaloblastus cuspidus sp. nov.
(Figs 29,30)

ETYMOLOGY
Latin cuspis, point; refers to adoral projection of
radial at centre of ambulacra.

MATERIAL

Holotype QMF21519; paratypes articulated
specimens QMF21519, 21521, and NMVP94182,
basals QMF21523, 21525-21527, radials QMF21524,
21528-21530, 21532-21537, and lancets QMF21520,
21522, 21531 from QML757.

DESCRIPTION

Calyx tall, with height to width ratio 2, maxi-
mum width at aboral extremities of ambulacra
which are 0.3 of height from oral pole; vault
dome-shaped. Pelvis conical, with relatively

MEMOIRS OF THE QUEENSLAND MUSEUM

broad base (2mm), at least at calical height of
about 20mm. Stem unknown.

Basals three, approximately half pelvic height,
each occupying 120° of circumference, slightly
waisted just above stem attachment, with straight
flaring sides; upper margin concave to chevron-
shaped centrally, descending on both sides. Stem
attachment area terminal, flat to gently concave,
with narrow marginal crenullarium. Radials tall,
occupying about half calical height, with well
developed comarginal growth ridges in each
growth sector, externally convex in both
transverse and vertical section, with broad low
tubercle at origin a short distance below the aboral
end of ambulacral sinus, with a low ridge extend-
ing over that short distance and the ridge waisted
at its midlength. As the ridge widens towards the
ambulacrum it merges with a narrow smooth area
of secondary calcite deposition bordering the am-
bulacral sinus under which growth lines of the RR
sector simply disappear. At the aboral end of the
ambulacral sinus a short broad cusp projects orally
(often broken off on available material but if not
evident, the broken base is evident). Ambulacral
sinus of variable length but usually occupying
about half length of radial (almost all dissociated
radials are broken in the RD sector, making it
impossible to be certain of this proportion but in
some radials the length of the RB sector relative
to its width suggests that the ambulacral sinus may
occupy less than 20% of radial length), Ambulacra
narrow, widening only gently adorally, depressed
beneath outer surface of radial; lateral walls of
ambulacral sinus becoming higher towards RD
front, gently convex, crossed at low angle to
lateral margins by growth ridges that are coarser
than on the exterior, sigmoidal in section with
variable attitude from almost vertical and then the
sigmoidal section only weakly evident to much
lower angle exposing a stronger sigmoidal section
and producing a much wider ambulacral sinus; in
the latter case the groove formed by the sigmoidal
wall of the sinus is closer to the external surface
than the convex roll and the groove does not reach
the aboral end of the sinus. On the floor of the
ridges is a further groove to accommodate the
lancet plate; lancet extremely long, subquadrate in
section, covered by side plates. Side plates each
with single large prominent brachiolar facet and
hydrospire pore abradially between successive
side plates; side plates in contact along radial line
throughout ambulacrum. RD margin variably
transverse to chevron-shaped orally, and either
straight or slightly sigmoidal, overlying part of
deltoid; some isolated radials appear to have

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA

pointed limbs suggesting the RD margin could, in
that case, be rather short but articulated specimens
of that type are unknown. On the radial interior
secondary calcite near the aboral end of the am-
bulacrum has weathered away in some specimens
exposing the hydrospires in groups of 5,

Deltoid with tall triangular aboral part beanng a
median radial crest and prominent transverse
coarse growth ridges that run aborally down the
wall of the ambulacral sinus; oral end expanded
into large deltoid lip around the spiracle opening
which descends at a low angle beneath the trian-
gular part of the deltoid. Around the spiracle and
extending down the deltoid-ambulacrum bound-
ary are Small upnght spiracle cover plates, Anal
deltoid area destroyed by compaction in holotype
buta superdeltoid is evident with low ridged radial
omament on it in a dislocated situation, On
paratype theca (Fig.29E,F) the exposed cryptodel-
toids are evident with weathered remnants of the
hypodeltoid also apparent. Oral opening pen-
tagonal,

REMARKS

The complete holotype and paratype (Fig.29D-
F) provide most of the critical information on
arrangement of plates in the theca. In the holotype,
features may not be directly interpreted because
of compaction after death which has apparently
shortened the polar axis and probably increased its
diameter. Most isolated plates available are in-
complete or crushed butall in all a fairly good idea
of the species is available.

This species is distinguished by its thecal and
ambulacral shapes and has the unique feature of
the cusp in the aboral end of the ambulacral sinus,
The prominent growth lines across the deltoid also
appear to be distinctive.

Class ECHINOIDEA
Subclass REGULARIA Latreille, 1825
Order CIDAROIDA Claus, 1880

Cidaroid indet.
(Fig.31)

MATERIAL

Spine bases QMF21541-21543 and fragmentary inter-
ambulacral plates QMF21544-21552 all from the type
section of the Callythara Formation (QML761).

REMARKS

Teichert (1951:80) reported that echinoids are
quite unknown in the Permian of Western
Australia and none have been described since
then, In washed samples from the type locality of

369

the Callytharra Formation fragments of echinoids
have been found and are reported here to make the
faunal list more comprehensive.

The primary tubercles on these interambulacral
plates are typically cidaroid with their prominent
boss, platform, and tubercle,The noncrenulate
tubercles along with the long, solid spines with
longitudinal ridges suggest the Archaeocidandac
but the Miocidanidae cannot be excluded so fami-
ly assignment is aotattempted. Although different
stages of weathering could be inducing different
appearances of the plate ornament away from the
tubercles it seems highly likely that at least 2
different types are present. Therefore, the pos-
sibility should be kept in mind that more than one
species is present.

Archaeocidaris selwyni Etheridge,1892 from
the Permian of NSW so the group is known from
Australia but that species had much larger inter-
ambulacral plates than are available from the
Western Australian sample. Although these
species are probably closely related useful com-
parison is not possible at present.

Class STELLAROIDEA Lamarck, 1801
Family MONASTERIDAE Schuchert,

Monaster Etheridge, 1892

TYPE SPECIES

Palaeaster clarki deKoninck, 1877 from the
Permian of New South Wales; designated
Schuchert,1914.

Monaster coleyi sp. nov,
(Fig-32)

ETYMOLOGY
For the collector Mr J. Coley wha was book-
keeper at Wandagee Station tor many years.

MATERIAL

Holotype NMVP127200 and paratype NMVP127201
from the Wandagee Beds (horizon g-r), on the north
bank of the Minilya River in the eastern limb of the
syncline near Coolkilya Pool, Collected by J. Coley,
From the collections of C, Teichert deposited in the
University of Melbourne and transferred to the Museum
of Victoria m the early 1980s.

DESCRIPTION
This species is described by comparison with
Monaster wandageensis Kesling, 1969.
Stellate;atms tapering gradually to nearly paral-
lel-sided; widened adorally to produce smooth
curve on the interbrachial margin. Disc with
prominent coronet on aboral surface; RI subrec-

370

tangular, smooth, lying over Sm2 plates and abut-
ting Sm1 plates on adpolar corners. Plating inside
Sm1 circlet unknown. Interradially and distally
between Sm2 plates is a deep cleft. M large, over-
lapping two Sm2 and abutting two R1 plates, with
subradial striae.

Oral surface flat, mouth small; Adm] and Ax
not clear.

Arms strongly plated. Amb plates small, set
deep in ambulacral groove. Admm wider than
long, with transversely oblique row of 3 or 4 short
stout spines, with obtusely pointed margin on
ambulacral groove, with abradial margin in zone
of tiny plates apparently part of a shagreenate skin
covering the aboral surface and terminating at the
junction between Imm and Admm.

Imm longer than Admm, approximately sub-
quadrate, with obtusely pointed adradial margin,
with series of transverse flutings along abradial
edge. Radials small, subquadrate proximally, be-
coming more rounded distally, not in contact with
each other, in contact with Imm at certain points
only leaving gaps at sutures between Admm. All
covered by finely shagreenate skin with small
plates embedded in it.

REMARKS

This species is separated from the closely re-
lated M. wandageensis Kesling,1969 from the
same locality by its nontuberculate R1, smaller
Sm1, interradial clefts between Sm2 plates, and
finely shagreenate aboral skin. It differs from M.
carnarvonensis Kesling, 1969 also from the same
locality, by the subrectangular R1, larger M, less
tapering arms and finely shagreenate skin. It dif-
fers from the type species in the narrower, and
straighter arms and more numerous arm plates.

ACKNOWLEDGEMENTS

We thank Dr J. Mac Dickins, Bureau of Mineral
Resources, Canberra and Dr Ken J. McNamara,
Western Australian Museum, Perth for their
hospitality and for arranging for the loan of
specimens from their respective institutions. Dr
Des Strusz, Bureau of Mineral Resources, Can-
berra, Dr David Haig, Department of Geology,
University of Western Australia, and Dr David
Hollaway, Museum of Victoria are also thanked
for organising loan of material in their care. We
are grateful to the managers of Williambury,
Wandagee, and Cherrabun Stations for access to
collecting localities on their properties. We are
grateful to Margaret Eva, Librarian at the Geology
Department, University of Queensland for consid-

MEMOIRS OF THE QUEENSLAND MUSEUM

erable help with provision of necessary refer-
ences. GDW extends his appreciation to the Direc-
tor of the Queensland Museum for facilities and
office space during prosecution of this project.

LITERATURE CITED

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BATHER, F.A. 1914. British fossil crinoids.-10,
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BOOGAARD, M. 1987. Lower Permian conodonts
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LOCALITY REGISTER

QML localities are in the Callytharra Forma-
tion. Map sheets referred to are in the 1:250 000
Geological Series.

QML757 — North side of bladed track from Callytharra
Homestead to Byro Homestead in western most
exposures 1.5km west of type section south of
Wooramel River. 25°52’30"S, 115°29°E
Wooramel Sheet SG50-5, 1966.

QML758 — North side of bladed track from Callytharra
Homestead to Byro Homestead as track enters dry
wash tributary to Wooramel River; lowermost
fossiliferous shale and marl capped by limestone
in lower fossiliferous unit in Callytharra Forma-
tion. type section. 25° 52’30"S, 115° 30°05"E,
Glen burgh Sheet SG50-6, 1963.

QML759 — Second fossiliferous shale and marl capped
by limestone in type section. South side of track
and stratigraphically higher than QML758.

QML760 — Third fossiliferous shale and marl capped
by limestone in type section. Immediately south
of and stratigraphically higher than QML759.

QML761 — Fourth and fifth fossiliferous shale and marl

capped by limestone in type section. Immediately
south and stratigraphically higher than QML760.

QML767 — Flats south of low hill approximately 100-
150m northeast of turnoff to Merlinleigh Homes-
tead, 6.5km west northwest of Mt Sandiman
Homestead. 24° 23’S, 115° 20’E, Kennedy
Range Sheet SG50-1, 1963.

QML768 = Flats with low limestone exposures 50-
100m east of new road to Moogoorie on both sides
of northeast-southwest fenceline, 24° 18°57"S,
115° 18°40"E, Kennedy Range Sheet SG50-1,
1963.

QML770—2-3km east of Mundarrie Well in limestones
of west dipping hogbacks north and south of Car-
narvon-Lyndon road; GR292356, Winning Pool -
Minilya Sheet SF50-13, 49-16, 1984.

QML771 — Third massive sandstone above base of
Wandagee Formation, 50-150m north of Minilya
River in east limb of syncline; GR257375, Win-
ning Pool-Minilya Sheet SF50-13, 49-16, 1984.

QML772 - Bench in lower slope below cliff 0.5-1.0km
NNW of type section in Cherrabun Member of

PERMIAN ECHINODERMS FROM WESTERN AUSTRALIA 373

Hardman Formation, Millyit Range; GR767877 BMR loc. ML87 and ML89 — Bulgadoo Shale, lower-
Sueno ee (type sec- most fossiliferous horizon; campsite east of Mia
tion) at Brutens Yard. Mia Homestead. 23° 21’S, 114°34’E.

374

ASESTING COMMUNITY OF FALSE WATER RATS
XEROMYS MYOIDES ON THE MYORA
SEDGELANDS, NORTH STRADBROKE ISLAND.
Memeity of the Queensland Museum 22/1): 374, 1992:- The
False Water Rat Xeromys mycoides Thomas, 1889 was
described from Mackay. There are 14 in museums: 6 Northern
Territory (1. from Sth Alligator R. [Parker, 1973) 2, Daly R-
(Redhead & McKean, 1975]; 2, Tomkinsan R. [Magnusson et
al, 1976); 3 Melville Is. [Magnusson et al. 1976): 6
Queensland (3,, from Mackay [Thomas, 1889 fholotype),
MeDougall, 1944 (5, 3 have been lost)}; 1, Beerwah [Dwyer
et al, 1979]; I, Bribie Ts. [Qld Museurs JM2381); 1, Nth
Stradbroke Is. (Van Dyck, 1979]). In addition, specimens
have been sighted or trapped-and-released; N,T, - Tomkinson
R., 4, and Melville Is, where 3 were captured from an
extraordinary mud nest (Magnusson et al, 1976); Qld -
Proserpine, 2 (P. Campbell, pers. comm, 1YKL)}, Kinaba Is., t,
Covloolu, 1, und Beerwah, 1 (Dwyer etal, 1979),

This small species has been seen and collected in or near
Shallow Coastal wetlands auch as mangrove forests, freshwater
lagoons, swamps and sedged lakes close to foredunes. Little
is known of its ecology or social behaviour, Curiowsly, i113
not ever known (f the False Water Ral is aquatic.

This AGte reports the recent discovery of a number of active
nests of ¥ tyoldes in the Myuru sedgelands, North
Stradbroke Is., (see Van Dyck, 1992) On 2 November, 199),
a farge, reed-covered maund similar to that described by
Magnussoneéal. (1976) was Jound 13minland from the upper
Aiangrove limiton the Myore sedgclands, between Capenphah
and Cooroonpah Cks (27°27'58"S,153'25'47"E), Overnight
trapping at the mound was unsuccessful. The following day a
two hoursurvey along this narrow sedge strip revealed 3 more
nests at intervals of 80-410m, Ovemight trapping ut one of
these nests yielded an adult female (HB:120mm, TV:87mm,
MF(su):24, E(n):14.4mm, W:49g) which was photographed,
filted with a radio transmitter and released al the capture sile,
This andividual had small white spots over the dorsum,
posterior of the shoulder blades. Similar spots were evident an
JM238 1 callected at Myora Springs in 1978, On 15 Novem-
ber, 199], a ble survey of coastal sedgeland between Myora
Springs and Chigail Chiggil found another & possible nests
that vaned from relatively simple tunnels built into the supra-
hittoral bank ta more complex mounds, A cocently consiructed
mound was found near Myora Springs oq 4 Febroary, 1992,

All mounds were located within the narrow ribbon of reed
and sedgeland (1,2m logh) where salinity is reduced by fresh-
Water drainage. This zone, comprised almost exclusively of
Juncus kraussti, Baunea juncea, B. rubiginesa and Finvbris-
tilix ferruginea was up Lo 30m wide and was sharply
delineated from the upper mangrove woodland iract (A vicen-
nia marina, Rhizophora stylosa and Bruguieva gymnorrhiza)
and the lowest extent of the wallum (dominated by Melaleuca
quinguenervia, Bucalypies interrnedia, E, robusta, Leptosper-
mur liversidgei, Gahnia sieberiana and Coustis blokei),
Mounds were built on slightly elevated ‘islands', some being
found among smal) patches of stunted Meluleuca quinguerer-
vicand Cwsuaring glowca, ylhers on raised sedee hummocks,
Oye West Was ncarporMad andurd the steey Of a Livia Vitex
tefolla war, beolor,

Mounds were from 200m 10 600m high wilh basal circum-
ferences from |. fon te 4.4m, From | to 3 enieance holes were

MEMOIRS OF THE QUEENSLAND MUSEUM

found areach mound hut others may have been hidden under
the labyrinch of exposed tree moots into which mounds were
ofien incerporatedl These holes were oval in shape [ap-
proximately 601m by 40mm) and were positioned eather at
the top or the bottom of mounds; they Were from 30cm to 2m
apart. Although lower holes were Nooded ar high tide, the
mounds themselves were never entirely inundated.

Evidence of extemal mound ‘workings’ consisted of fresh
roof plasterings it whieli a thin pasi¢ of black mud nixed with
peat, dried leaves, sedges, or crab shell had been daubed over
the lop of the mound and worked between sedges (usually
Juncus kraussii). Areas ‘plastered’ in this way measured as
much aa 12cm" and fresh ‘plastering’ seemed to indicate
that the nest was occupied,

X, myoides has been variously described as ‘one of the rarest
of our native mammals’, ‘Vulnerable to changes caused by
man’, “rare, scattered (probably at risk in some habitats and
extinct in others)’ und 1 designated it as ‘vulneruble’ (Wan
Oyek, 1991), Itis not known whether the few reconds of the
species reflect penuine vary or a lack of searching effort in
appropriate habitats, However, an occupied band of mud nests
of X. myvides in the Myora sedgeland provides an excellent
opportunity for future conservation research,

The commitment of the Endangered Species Program
CANP WS) to funding such research is gratefully acknow-
Jedged. Dr Peier Bostock (Queensland Herbarium) provided
plant identifications.

literature Cited

Deyer, P., Hockings, M. & Willmer, J. 1979, Mamovals of
Cooloola and Beerwah. Proceedings of the Rayal Society
of Queensland 90; 65-84,

Magnusson, W.E., Webb, G.,W, & Taylor, JA, (976, Two
oew locality records, anew habitat and d nest description
for Xeramys myoides Thomas (Rodeniie Mundae),
Australia Wildlife Research 35 153-157,

McDougull, W.A. 1944, An investigahon of the rat problem
in Queensland cane fields, 2. Species and general habits.
Queensland Journal of Agricultural Science [= 48-78.

Parker, SA. 1973. An annotated checklist of the native land
mammials of the Northern Terettory. Records of the South
Australian Museum 16(11): 1-57.

Redhead, T,D, & McKean, J.L 1975, A new record of the
false water rat, Xeromys myvudes Viramas, T&R Pram tre
Northern Territory of Australia. Australian Joumal of
Mammalogy |: 347-354.

Thomas, O, 1889, Description of 4 new. genus of Muridas
allied to Hydromys, Proceedings of the Royal Zookugival
Soclety of London 1849; 247-250, pl, 29,

Van Dyck, 8. 1979. The false walter rat Xeromys myvides an
Stradbroke Island, w new locality in sowlhesstern
Queenshind, Proc. Royal Society Queensland 90, 84,

1991. The status af mammals Pp. 349-353. In Ingram, G.J.
& Raven, RJ. (eds), ‘An Atlas of Queensland's frogs,
reptiles, birds and mammals’.(Qld Museum; Brisbane),

1992. Parting the reeds on Myora's Xeromys kibbuiz,
Wilillife Australia 29( 1): 8-10

Steve Van Dyck, Queensland Museum, PO Dox 3300,
South Brisbane 210!, Queensland, Australi: Ellie
Burbidge, Point Lookout, North Stradbroke Island
4183, Queensland, Australia; 17 February, 1992,

CONTENTS
Bruce, N.L.
Anopsilana barnardi, a new species of estuarine cirolanid crustacean isopod from tropical
(oanbere Alea tralia: vps). st. hes) ote ah alg pat es anand et Sn ae seh Sale she pe Se es a ed eee SE 1
CHURCHILL, T.B. AND RAVEN, R.J.
Systematics of the intertidal trapdoor spider genus /dioctis (Mygalomorphae: Barychelidae)
in the Westem-Pacific with a new genus fromthe Northeast ...........2.0eee cece eee e cence 9
COLEMAN, R.A.
Shipwreck and historical survey: Chilcott Islet, Coral Sea ............02ec2eeeeesees 5,4: sitios 5g FNS
Couper, P. J. AND INGRAM, G.J.
A new species of skink of Lerista from Queensland and a re-appraisal of L. allanae

CE STAI ee rersts eee ote ee esoteetrd naabscta sia tines grace teen attonls aSierede. eo-ekioniy aces ao
DAVID, B., CHANT, D, AND FLOOD, J.
Jalijbang 2 and the distribution of pecked faces in Australia ........6.20c0ecceerseveveeeneseenen 61
DAVE, P.J.F
Revision of Sarmatium Dana (Crustacea: Brachyura: Sesarminae) with descriptions of
Wireb ie ieniese roe eects miele cherie (ak Soha a eis BS ae a tars Cana ct os oe brie cine CER Taek 79
Hooper, J.N.A. AND BERGQUIST, P.R.
Cymbastela, a new genus of lamellate coral reef sponges ........0.000 cece cee eee eevee rersenaen 99
HUGHES, N.C. AND DROSER, M.L.
Trace fossils from the Phe Formation (Lower Cambrian), Zanskar Valley, northwestern India ....... 139
JONES, D.S.
Scalpellid barnacles (Cirripedia: Thoracica) from the northeastern and central eastern
Australian continental shelf and slope ............2020ecee cece eee e net ee eee nneeenees 145

KENNETT, R.M., GEORGES, A., THOMAS, K. AND GEORGES, T.C.
Distribution of the long-necked freshwater turtle Chelodina novaeguineae and new

information on its ecology ............000 een eeeeeteneeee SF 2th; Si3.e: Pee ees tee reat LID

LAMBEI, K.J.

Re-examination of the venation of Osmylopsychops spillerae Tillyard from the Triassic of Queensland 183
LAMPRELL, K,

Notes on Spondylus Linnaeus with descriptions of two new species from Western

Australia (Mollusca: Bivalvia: Spondylidae) ........- 6.0006. e cece eee cece nee ee naan 189

ROZEFELDS, A.C.

The subtribe Hicksbeachiinae (Proteaceae) in the Australian Tertiary ............6. 0000 cceee eee 195
ROZEFELDs, A.C., CHRISTOPHEL, D.C. AND ALLEY, NLF.

Tertiary occurrence of the fern Lygodium (Schizaeaceae) in Australia and New Zealand ............ 203
RYLAND, J.S. AND HAYWARD, P.J.

Bryozoa from Heron Island, Great Barrier Reef ..........200 02 e cece eee ce sev cee ee eenveeeees 223
SHEA, G.M. AND HUTCHINSON, M.N,

A new species of lizard (Tiligua) from the Miocene of Riversleigh, Queensland ..............00+. 303
WEBSTER, G.D. AND JELL, P.A.

Permian echinoderms from Western Australia 2... 0.00 sce cecee renee see sereerenresrouseesaes 311
NOTES
COUPER, P.J.

Anomalopus pluto Ingram, a poorly known skink from Cape York Peninsula ...........--0++0eee0% 54
Couper, P.J., WHITTIER, J., MASON, R.T. AND INGRAM, G.J.

A nesting record for Coeranoscincus reticulatus (Giinther) ..........60c0 cree cece eee eee n een ees 60
DAVE, P.J.F.

The rare shrimp, Phyllognathia ceratophthalma (Balss), discovered in Moreton Bay ................ 98
RICHARDS, S.J.

The tadpole of the Australopapuan frog Rana daemeli ......6.0060 ccc eee e nee n eens eanen pete we
RICHARDS, S.J. AND JAMES, C. .

Foot-flagging displays of some Australian frogs .........0.0sccss ee eeeceesene nee neneercseers 302

VAN Dyck, S. AND DURBIDGE, E. b
A nesting community of False Water Rats Xeromys myoides on the Myora Sedgelands,
: Nort Sttadbroke Ishagidl oan) bg 5 Bis www aodle ome Wea spate b aheeosis piel ace 4 tox sortie eerie 374