Volume 56
Number 1

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Memoirs of the

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28 February

1997

ZOOLOGY OF VICTORIA

PL. 130

4 Insects )

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Cover: In Frederick McCoy's Natural history of Victoria, Prodromus of the zoology of Victoria, or,
figures and descriptions of all classes of the Victorian indigenous animals. Vol. 2, Decade 13, pub-
lished in 1886, he illustrated the mantis Archimantis latistyla. In this Memoir, Graham Milledge
redescribes the species and others related to it.

ISSN 0814-1827

MEMOIRS

of the
MUSEUM OF VICTORIA

MELBOURNE AUSTRALIA

Memoir 56
Number 1
28 February 1997

Director
Graham C. Morris

Director (Natural Sciences)
Ross P. Field

Editor
Gary C. B. Poore

Editorial Board
David J. Holloway Chung Cheng Lu Ken L. Walker

PUBLISHED BY ORDER OF THE COUNCIL

© Museum of Victoria Council 1997

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CONTENTS

Revision of the Tribe Archimantini (Mantodea: Mantidae: Mantinae)
G.A. Milledge

TOPA SN Oo memon AREA A eo oste e S DON rpo ERRORES TEE ETT E l
A new species of Progradungula Forster and Gray (Araneae: Gradungulidae) from Victoria

(aL AAT otro re p T LN RM A rit 65
Supplement to a revision of Australian members of the bee genus Homalictus (Cockerell)
(Hymenoptera: Halictidae)

AS. IO emos peo ds te ao p o a A Ia TR Taar UO 0254 69
Descriptions of new Leptophlebiidae (Insecta: Ephemeroptera) from Australia. I. Tillyardophlebia
gen. nov.

dA DEUS UE R repon EMT PRE nn ON REIP ELI EHE SUN Te h aea n THT 83
Phylogeny and biogeography of Australian genera of Chlorocystini (Insecta: Homoptera:

Tibicinidae)

dil METRI AI RIMA IO ORO 91
The type species of A/eurodicus Douglas, a whitefly genus of economic importance (Homoptera:
Aleyrodidae)

LIVRE Mk I cea eae EIS NUR UR CROAT sea EE E AA em ET He AR d ete 125
Additions to the taxonomy of the Limnoriidae (Crustacea: Isopoda)

[LL OOO SIT orea ERR ON POS A E E E LEN PTA, 129
A new genus of marine isopod (Crustacea: Flabellifera: Sphaeromatidae) from Australia and the
Indo-Pacific Ocean

INSEE RICO m EE E nen ko kn kan a yok menti kn, LEUR (ROTTI EM kask pop I CURRO 145

Spermatological evidence supports the taxonomic placement of the Australian endemic hairy stone
crab, Lomis hirta (Decapoda: Anomura: Lomidae)
(CE IO ET ed cde tefie Oo Uomo A a doj E E ae HT TTT SENEC e oe 235

A remarkable new pygmy seahorse (Syngnathidae: Hippocampus) from south-eastern Australia, with a
redescription of H. bargibanti Whitley from New Caledonia
Bald IGA ok um roei 5 te DES eno Mt D TE E Er EE Un Mr SSR E On iii 245

A new grenadier, genus Trachonurus, from New Zealand and Australia (Pisces: Gadiformes:
Macrouridae)
T. Iwamoto and P. McMillan

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Memoirs of the Museum of Victoria 56(1):1-63 (1997)

REVISION OF THE TRIBE ARCHIMANTINI
(MANTODEA: MANTIDAE: MANTINAE)

G. A. MILLEDGE

Museum of Victoria. 71 Victoria Crescent. Abbotsford. Vic. 3067. Australia

Abstract

Milledge. G.A.. 1997. Revision of the tribe Archimantini (Mantodea: Mantidae: Mantinae).

Memoirs of the Museum of Victoria 56:1-63.

The Australo-Papuan tribe Archimantini is redefined. The genera Pseudomantis Saussure
and Rhodomantis Giglio-Tos are excluded. The genus Austromantis Sjöstedt is recognised as
valid and included. One new genus. Cort/iylomantis. and four new species, Archimantis
gracilis, A.vittata. Austovates papua and Corthylomantis baldersoni, are described. Archi-
mentis minor Giglio-Tos is a new synonym of A.sobrina Saussure, Austromantis gracilis
Sjöstedt a new synonym of .L.albomarginata Sjöstedt and Coenomantis melanoptera (Tin-
dale) a new synonym of C.kraussiana (Saussure). Archimantis inermis Werner is transferred
to the neotropical genus Angela Serville. The subspecies Archimantis latistyla gigantea Beier
is rejected as invalid. Keys to genera and species are provided. Information on biology is
recorded. distributions given and relationships discussed.

Introduction

The Archimantini includes medium to large,
elongate mantids restricted to Australia (14
species) and Papua New Guinea (1 species). All
inhabit shrubs and/or tall grasses where they are
well camouflaged. They do not appear to be
abundant in nature but many species can be
encountered without much difficulty by search-
ing in appropriate habitats. Species diversity is
greatest in the tropical north of Australia, how-
ever several species occur in Victoria where the
southern limit of the group is reached. There
are no records of the Archimantini from
Tasmania.

Giglio-Tos (1917) proposed the Archimanti-
nae to include several Australian genera of the
Mantidae, characterised by the first and fourth
discoidal spines being not shorter than the sec-
ond, wings reduced in the female and the facial
shield wider than high. He divided the Archi-
mantinae into two groups: the Archimantes,
having the cerci compressed and the first dis-
coidal spine longer than the second, containing
the genera Archimantis, Coenomantis and Rheo-
mantis (=Archimantis), and the Pseudomantes,
having the cerci cylindrical and the first discoi-
dal spine of equal length to the secnd, containing
Pseudomantis and Rhodomantis. Beier (1935)
rejected this, including the Archimantinae
within the Mantinae. However Beier (1964)
later created the tribe Archimantini which was
essentially equivalent to Giglio-Tos’ Archiman-
tinae and included the genera Archimantis,

Austrovates, Coenomantis, Nullabora, Pseudo-
mantis and Rhodomantis. ln this work the
Archimantini is considered as equivalent to
Giglio-Tos’ Archimantes, characterised by the
first discoidal spine being longer than the second
(fig. 6) and including the genera Archimantis,
Austromantis, Austrovates, Coenomantis, Cor-
thylomantis, and Nullabora. The genus Austro-
mantis was synonymised with Pseudomantis by
Beier(1935) but is here considered a valid genus.
Pseudomantis and Rhodomantis, which have the
secnd discoidal spine as long or longer than the
first, are removed to Beier's Mantini.

The tribe Archimantini thus contains six gen-
era, one of which is new, and 15 species of which
four are new. The largest genus is Archimantis,
containing nine species of which two are new. Of
the nine species and one subspecies listed for
this genus by Balderson (1984) seven species are
recognised as valid. A.minor Giglio-Tos is syn-
onymised with A.sobrina and A.latistyla gigan-
tea Beier 1s considered a variant and accorded
no taxonomic status. The holotype male of
A.inermis Werner has been examined and deter-
mined to belong to the neotropical genus Angela
Serville. It was probably ascribed to Australia in
error. Five of the remaining six genera are con-
sidered monotypic, while the sixth, Austrovates,
contains two species of which one is new.

Methods and terminology follow those of Mil-
ledge (1990) except that the following abbrevi-
ations are used in the descriptions of male
genitalia: aa — anterior apodeme of right

d G. A. MILLEDGE

phallomere: apr — apical process of left phal-
lomere, dl — dorsal lamina of left phallomere;

dpr — distal process of ventral phallomere, Iph
— left phallomere: ml — membraneous lobe of
left phallomere: pa — phalloid apophysis; rph
— right phallomere; vl — ventral lamina of left

phallomere; vpl — ventral plate; vspr — ventral
sclerotized process (figs 7 — 8). Abbreviations
used for institutions where material is held are as
follows: AM — Australian Museum, Sydney:
ANIC — Australian National Insect Collection,
Canberra; BMNH — Natural History Museum,
London: NMV — Museum of Victoria, Mel-
bourne; NHRM — Naturhistoriska Riksmu-
seet, Stockholm; QM — Queensland Museum,
Brisbane; RNHL — Rijksmuseum van Natuur-
lijke Historie, Leiden: SAM — South Australian
Museum, Adelaide: UQ — University of
Queensland, Brisbane; WAM — Western Aus-
tralian Museum, Perth; ZMH — Zoologisches
Museum, Hamburg; ZSIC — Zoological Survey
of India, Calcutta.

Relationships

Two major groups can be recognized within the
Australian Mantodcan fauna. The first, smaller
and probably more recent element contains gen-
era which reach their greatest diversity outside
the Australian region, in most cases the East
Asian region. The Australian representatives of
the genus Acromantis (1 sp.) (Hymenopodidae)
and the genera Hierodula (4-5 spp.), Mantis (1
sp. Tenodera (2 spp.) and Statilia (1 sp.)
(Mantidae-Mantinae) fall into this category. It is
notable that all have macropterous females and
thus are probably capable of dispersal across
water barriers.

The second. largerand probably older element
consists of genera which are endemic to Aus-
tralia or to Australia and nearby islands (includ-
ing Papua New Guinea). Many of the species in
this group have brachypterous females. The
relationships of this older element are not
entirely clear. For example the nearest relatives
of the Australasian endemic subfamily Paraoxy-
pilinae are the Amorphoscclinae (both Amor-
phoscelidae) which appear to be essentially
African. An Australasian endemic subfamily of
the Mantidae, the Orthoderinae, is most closely
related to the Choreododinae, which occurs in
southern India and Sri Lanka, and in South and
Central America. This would imply a Gondwa-
nan origin for the older element.

However, further work is required on the
faunas of other regions before the relationships

of the Australian representatives of the large
subfamily Mantinac, including the Archiman-
tini, can be clarified. Within the Australian
fauna a possible candidate for sister group
status of the Archimantini 1s the genus Pseudo-
mantis, in which the 2nd discoidal spine
is of similar length to the Ist, possibly represent-
ing a stage in the progressive reduction of this
spine.

Relationships within the Archimantini are
also obscure but perhaps less so. Austromantis
appears closest in form to a presumed anscestor
of the tribe, displaying several pleisiomorphic
characters including only moderate elongation
of the body, macropterous females and cylindri-
cal cerci. Nullabora appears closely related to
Austromantis, sharing the aforesaid characters
as well as a bifurcate dpr in the male genitalia,
which is not found in other members of the
Archimantini. Corthylomantis also possesses
cylindrical cerci, however, the colour pattern of
the wing in the female may be a synapomorphy
linking this genus with Coenomantis. Austrov-
ates variegata displays a finely striped eye pat-
tern and median projections between the
abdominal tergites of juveniles, characters
shared by Coenomantis. Juvenile Archimantis
quinquelobata have abdominal projections and
also display subapical lobes on the mid and hind
femora similar to though less well developed,
than those found in Austrovates, possibly indi-
cating some relationship. The shallow U-shaped
arrangement of the egg cells in the ootheca of
Archimantis quinquelobata is shared with Coe-
nomantis, however the form of oothecae within
the Archimantini is not well enough known to
interpret the significance of this.

Within the genus Archimantis several species
groupings are apparent. Probable synapomor-
phies linking A./atistyla, A.armata, A.monstrosa
and A.sobrina are robust build, ventral colour
pattern of costal area of tegmen and broad based
uncinate dpr of the male genitalia. Within this
group the latter three species are also linked by
the compact, strongly shagreened pa of the male
genitalia. A.brunneriana and A.straminea share
probable synapomorphies of very compressed
head, angular eyes and mottled colour pattern
on ventral surface of forecoxa. Males of
A.gracilis and A.vittata are quite similar in
appearance but the form of their genitalia is dis-
similar. As the female of the latter species is
unknown, the rclationship between them
remains uncertain. Further work may lead to the
splitting of this genus, however, to do so now
would be premature.

REVISION OF THE ARCHIMANTINI (MANTIDAE)

Archimantini

Diagnosis. Head anterio-posteriorly com-
pressed, frontal shield transverse, first discoidal
spine longer than second, margin of fore femur
between outer spines smooth.

Notes. Members of the Archimantini can be col-
lected by searching in shrubs and tall grasses,
preferably at night with a spotlight. They tend to
prefer denser growth, probably because it pro-
vides more cover from predators. Males are
readily attracted to light traps at night. Speci-
mens can be successfuly reared to maturity in

flywire cages on a diet of live insects. The height
of the cages should be at least three times the
length of the mantis to provide adequate room
for moulting and temperatures should be kept
warm (25-30C). Individuals should be housed
seperately to avoid canabalism and not in visual
contact with one another to avoid stress.

Dry pinning is the preferred method of pres-
ervation for late instar nymphs and adults. Early
instar nymphs are better preserved in alcohol.
Specimens to be pinned should be gutted and
stuffed as per the method of Rentz (1985: 9) to
preserve body colour.

Key to genera of Archimantini (adults)

l Distal segments of cercus laterally compressed ............ 4
— Distal segments of cercus cylindrical .................... 2
2(1). Costal area of tegmen with white marginal band, females
MACPOPIGKOUSA e M. r TIENS I e ERI erga rieti 3

— Costal area of tegmen without white marginal band, females
TACH MES FOU SYON LT e a a Corthylomantis

(2). Eye tapering to blunt point, cerci stout ........... Nullabora
— Eye not tapering to blunt point, cerci slender ... Austromantis
4(1) Hindwings smokey black s rean hr oee sten Coenomantis
— Elindwinesainostlyihvaline; ss 4 X44 4 ebo. lane sk 5
5(4). Mid and hind femora with distinct subapical lobe on posterior
melO I dadre, dr, Mr e RE Y Austrovates

Mid and hind femora without distinct subapical lobe on posterior

matem ne e dotati

Austromantis Sjöstedt

Austromantis Sjòstedt, 1918: 28. Type species Aus-
tromantis albomarginata Sjòstedt, here designated.

Diagnosis. Head moderately anterio-posteriorly
compressed, eye margin rounded angular;
macropterous in both sexes; cerci cylindrical;
dpr of male genitalia bifurcate.

Description. Body of moderate size and build.
Head about 1.5 times as wide as high, moder-
ately compressed anterio-posteriorly, slightly
concave anteriorly (more so in male). apical
margin slightly curved, more strongly in para-
ocular regions; eyes forming broadly rounded
angle at dorsolateral margin; frontal shield with
dorsal margin broadly arched with indentations
below antennae and ventral ocellus, lateral mar-
gins gently curved, with distinct subantennal
ridge; antennae longer than prothorax in male.
shorter and finer in female.

Pronotum moderately elongate, more slender
in male, supracoxal expansion distinct; lateral
margins slightly lamellate, virtually smooth in

EE o Ho etone Archimantis

male, finely denticulate in female; prozona dor-
sally with very fine scattered tubercules or
smooth in some males, few more distinct
tubercles ventrally; metazona smooth with faint
mid-dorsal keel.

Fore coxa elongate, shorter than femur,
anterior margin distinctly toothed; fore femur
with 4 discoidal spines, the 2nd very short, the
3rd quite long, with 4 outer spines, the proximal
pair closer together, and 15 inner spines, claw
groove centrally located, ventral surface with
scattered tubercles and slight depression
anterior to the 4th discoidal spine; fore tibia
with 8-9 outer and 13-14 inner spines. Mid and
hind legs slender; femora without genicular
spine or subapical lobe.

Macropterous in both sexes, wings as long as
or longer than abdomen: tegmen with costal area
and costal margin of discoidal area opaque,
remainder hyaline; hind wing with costal area
semi opaque, remainder hyaline.

Abdomen elongate, slender in male, broader
in female; cerci of medium length, cylindrical,

4 G. A. MILLEDGE

distal segments slightly bead like. Male genitalia
with dpr a strongly sclerotized bifurcate hook
curved dextrad, medial lobe small but distinct;
apr curved sinstrad and tapering to rounded tip.
pa separated from vl bv membranous area, ml
elongate and finely hirsute; rph with dorsal arm
partially sclerotized, aa elongate becoming
squamiform sinstrally, vpl continuous with
main body of phallomere, vspr compact u
shaped.

Austromantis albomarginata Sjöstedt
Figures 1-8, 166

Vustronuntis albomarginata Sjöstedt 1918: 28.
lastromantis gracilis Sjöstedt 1918: 30. Syn. nov.

Material examined. Syntype male of A. albomargi-
nala Kimberley district, N.V.Austr., Mjöberg, Febr..
(221 89)", Syntype female of La/bomarginata, Kim-
berley district. N. V. Austr.. Mjöberg, Febr., (222 89)”.
Holotype male of 1.gracilis. “Broome, V.Austr., Mjó-
berg. (223 89)" (AIL NHRM ).

Otherspecimens examined (576, 89, 2 juv). Old. Le,
Barcaldine. 10 Feb 1981. 8. 24 km N of Einasleigh, 31
Dec 1989. d. o, Mareeba, 7 Dec 1952. (All AM). g,
Armstrong Ck Crossing. 13 km NNW of Guthalungra,
26 Jan 1982. 0.3 km SW of Barduthulla, 26 Dec 1961.
4, 200 km SSW of Normanton, 14 Oct 1965.0. 5 km W
of Maggieville HS, N of Normanton, 10 Apr 1962
(All ANIC). Jo. Burke and Wills Junction
19914'5. 140721'E. 16 Jan 1993. 26, Dajarra Rd., 19
km SW of Cloncurry. 20°50’S, 140°23’E, 2 Jan 1993,
g. Glenore Pumping Station. Norman River (17°51’S,
141708 E), 13 Jan 1993. g, 16 km SE of Hann River
Crossing. NW of Laura, 12 Jan 1990. s, Karumba,
7 May 1989 (all NMV). c, Normanton, 6 May 1963
(SAM). 20, Cloncurry. 8. 17 Apr 1947. g, Flinders
River. 42 km SW of Normanton. 26 May 1972
(AIL UQ).

NT. €. 15 km SW of Alroy Downs HS 197248.
135°58’E, 10 Apr 1976. 9, 28 km SE of Anthony
Lagoon 1809/5, 135°44’E, 11 Apr 19766, Baroalba Ck
Springs 1247'S, 132^51'E, 19 km ENE of Mt Cahill.
28 Oct 1972, 28, 36 km SW of Borroloola 16198.
136705/E. 4 Nov 1975. ljuv, Bukalara Range, 46 km
SSW of Borroloola 16°28’S, 136" I0'E, 22 Apr 1976. g,
| km N of Cahills Crossing. East Alligator River
L2725'8, [32758 E, 31 Oct 1972.6, Caranbirini Water-
hole, 33 km SW of Borroloola 16”16”S. 136905'E, 21
Apr 1976. p. L7729'S, 133°30"E, 8 km NNW of Elliot.

14 Oct 1972, 9, McArthur River HS. 80 km SW of

Borroloola. 13 May 1953. 20, 16°47’S. 135°45°E,
McArthur River. 14 km SW of Cape Crawford, 25 Oct
1975.g. 12950'5. 132'51'E, 16 km NE of Mt Cahill, 23
May 1973, 9, 123178. 132°54# E. 9 km NE of Mudgin-
berri HS, 30 Oct 1972, (All ANIC), c, Darwin, May
1977 (NMV). 9, Groote Eylandt, Feb 1922. 4. Lake
Woods. 15 km SW of Elliot, 15 Oct 1977 (Both SAM).
Je, Horn Islet, Sir Edward Pellew Group. 1-7 Feb
1968 (UO),

WA. c, Margaret River, 110 km WSW of Halls Creek,
8 Feb 1977 (AM). g, ljuv. 1547'S, 128°40’E, Bandi-
coot Range. 7 km W of Kununurra. 7 May 1983. 26,
17°19’S, 122*10'E, 8 km S of Cape Bertholet, West
Kimberley distr. 16, 17 Apr 1977. c. IFI7S,
122*10^E. 5 km SSW of Cape Bertholet, West Kimber-
ley dist, 21 Apr 1977, 38, Fitzroy River Crossing,
Derby-Broome Rd, 3 Nov 1978. d, 15746'S, 12844'E,
Kununurra, 11 Apr 1985. d, 16°34’S, 122'49'E, 4 km
WNW of Martins Well, West Kimberley dist., 20 Apr
1977. 9, 21°34’S, 117903'E, 3 km NW of Millstream
HS. 5 Apr 1971 g, 21°35’S, 11704'E. 1 km NNE of
Millstream HS, 16 Apr 1971. g. 21°36’S, 11 TOTE, 4
km ESE of Millstream HS. 31 Oct 1970. 20, 21737'S,
117°06’E. 5 km SE of Millstream HS, 12 Apr 1971. 4g,
21*35/S. 11 OVE, 2 km ENE of Millstream HS, 22, 30
Oct, 4 Nov 1970, 26. 21735'S. 11 704'E. | km N of
Millstream HS, 23, 28 Oct 1970. g, 21*35'S. 117704" E,
0.5 km WNW of Millstream HS, 7 Apr 1971. o,
14°49’S, 125*42'E, Mitchell River Falls, Kimberley
dist.. 12 May 1983. d, 40km ENE of Pardoo HS, Great
Northern Hwy, 23 Nov 1973. 9, 50 km SW of Sandfire
Flat, Broome-Port Hedland Rd, 29 Oct 1978. (AN
ANIC). c, Kununurra, 7 Jul 1969 (NMV). 0, Fortescue
River. Hammersley Range (SAM). 25, Derby, 7 May
1964 (WAM).

Description. Body green; metazona of prothorax
with margins and ventral surface purplish (red-
dish brown in dry specimens); inner face of fore
coxa mauve posteriorly, yellow anteriorly (not
apparent in dry specimens); femoral spines
blackish brown, tibial spines tipped in blackish
brown; proximal half of mid and hind femora
yellow; costal area of tegmen with marginal
white band and submarginal mauve and green
bands, costal margin of discoidal area greenish;
costal area of wing greenish; abdominal tergites
2-5 with central yellow patches, sternites 2-6
with black anterior margin.

Male genitalia (figs 6, 7) with posterior arm of
bifurcate dpr much shorter than anterior arm;
apr elongate gently curved; pa rather smooth
with elongate, narrow, pointed posterior pro-
cess, sharply curved dextrad and small knob like
anterior process, directed dorsad.

Measurements(mm). Body length, & 50-62, 9
66-68. Head width, d 6.5-7.7, 9 9.5. Head
depth, & 3.5-4.6, 9 6.8. Pronotum width, ¢ 3.0-
4.4, 9 5.5. Pronotum length, g 15.0-20.2, 9 23.1.
Fore coxa length, ¢ 9.2-12.7, Q 17.1. Fore femur
length, & 11.0-15.4, 9 18.7. Tegmen length, €
31.9-39.6, 9 40.7. Cercus length, d 4.0-4.6, 9
6.1.

Immature stages. Nymphs similar in appearance
to adults, ootheca unknown.

REVISION OF THE ARCHIMANTINI (MANTIDAE)

Austromantis albomarginata. 1. male: 2. female (pz = prozona, mz = metazona).

Scale = 10 mm.

Figures 1-2.

G. A. MILLEDGE

5

(l

Figures 3-8. Austromantis albomarginata. 3. male head; 4, female h 15, fi

BA i ; .3.n ; 4. ead; 5, female cercus; 6, female foreleg, insi
id1-3— discoidal spines 1-3): 7, male genitalia. dorsal (aa = anterior apodeme, apr = apical process I
process. Iph = left phallomere, pa = phalloid apophysis, rph = right phallomere); 8, male genitalia ventral
(ml = membraneous lobe, vph = ventral phallomere, vpl = ventral plate, vspr = ventral sclerotized process)

Scales = 2 mm.

Distribution and habits. Known from the north-
ern parts of Western Australia, Northern Terri-

REVISION OF THE ARCHIMANTINI (MANTIDAE) 7
hindwing with costal area semi opaque.
remaider hyaline.

Abdomen elongate; cercus of moderate

tory and Queensland (fig. 166). I have collected
this species from shrubs in northern Queens-
land. The large proportion of males in collec-
tions probably indicates that most specimens
were collected at light traps.

Remarks. This species shows some variation in
size. Sjostedt's type male of A.gracilis is at the
small end of the range, however its genitalia are
virtually identical to those of the male syntype of
A.albomarginata examined.

Nullabora Tindale

Nullabora Tindale, 1923: 442. Type species Nulla-
bora flavoguttata Tindale by monotypy.

Diagnosis. Head strongly anterio-posteriorly
compressed, eyes tapering to blunt point, both
sexes macropterous, cerci cylindrical and stout,
dpr of male genitalia bifurcate.

Description. Body of medium size, rather slen-
der, female more robust than male. Head at least
twice as wide as high, strongly anterio-poster-
iorly compressed, slightly concave anteriorly,
apical margin horizontal but curving downward
little toward eyes; eyes tapering to blunt point at
dorso-lateral margin; frontal shield transverse,
about 3.5 times as wide as high, surface flat, dor-
sal margin broadly curved with indentations
below antennae and ventral ocellus, lateral mar-
gins slightly curved; antennae of male slightly
longer than prothorax, those of female finer and
about half length of prothorax.

Pronotum elongate, slender, supracoxal
expansion moderate; margins slightly lamellate,
almost smooth in male, finely denticulate in
female; prozona slightly constricted in both
sexes, moderately tuberculate beneath; meta-
zona with very faint mid dorsal keel.

Fore coxa slender, shorter than femur,
anterior margin with 6—7 small blunt teeth inter-
spersed by smaller denticles; fore femur slender,
claw groove at or slightly distal of midpoint,
with 4 discoidal spines, the 2nd shortest, the 3rd
longest, with 4 outer spines and 16 inner spines,
ventral surface with scattered tubercles and shal-
low depression anterior to 4th discoidal spine;
fore tibia with 1 1 outer and 16 inner spines. Mid
and hind legs slender, femora without genicular
spine.

Macropterous in both sexes, wings as long as
abdomen in female, slightly shorter in male,
tegmen with costal area and costal margin of
discoidal area opaque, remainder hyaline;

length, cylindrical and rather stout, terminal seg-
ment pointed. Male genitalia with dpr a sharply
curved bifurcate hook; apr tapering to blunt
point, slightly sinuate; dl of Iph reduced; vsph
small, stoutly u-shaped.

Nullabora flavoguttata Tindale
Figures 9-17, 166
Nullabora flavoguttata Tindale. 1923: 443.

Material examined. Holotype female . Kingoonya.
S.Aust., R.Harvey, I.14070. Paratype nymph, North-
ern Territory. Capt.S.A. White (both SAM),

Other specimens examined (9c, 149, 11 juv.). Qld.
26, 9, Butcher Creek, 20 km W of Cloncurry, 21 Jan
1977 (AM). I juv., 48 km E of Camooweal. 24 Aug
1960. c, 21°41’S, 140°30’E, Selwyn Mine, 160 km SE
of Mt Isa, 28 Dec 1991. 9, 21741'S. 140°30"E, Selwyn
Mine, 160 km SE of Mt Isa. 17 Oct 1990. 1 juv.,
17°24’S, 138" 1 8/E, 9 km SSE of Westmoreland HS, 29
May 1975 (all ANIC). 29, g, 20'43'S, 140°21’E,
Butcher Ck., 16 km W of Cloncurry, 1 Jan 1993, e,
19*1 7S, 140729'E, 16 km ESE of Burke and Wills
Junction, 15 Jan 1993. 9, 19°17’S. 140727 E, Crocodile
Ck., 14 km ESE of Burke and Wills Junction, 14 Jan
1993. c, 29, | juv., 20°50’S, 14023'E, Dajarra Rd., 19
km SW of Cloncurry, 2 Jan 1993 (all NMV).

NT. 9, 16°16’S, 136'05'E, Cararbirini Waterhole, 33
km SW of Borroloola, 21 Apr 1976. 1 juv., 195178,
136°02’E, 14 km SSE of Dalmore Downs HS, 25 Apr
1976.9, | juv., 20 km S of Elliot, 19 Nov 1966. 1 juv..
20km NNW of Renner Springs, 1 5 Oct 1969. 1 juv., 30
km E of Vaughan Springs HS, 25 Jul 1968 (all
ANIC).

South Australia. d, 26°09’S, 130°35’E, 56 km W of
Amata, Musgrave Ranges. 20-21 Jan 1982 (ANIC). g,
Anna Creek HS, 6 Dec 1974 (SAM).

WA. 9, 18°49’S, 12371 7'E, 163 km ESE of Broome, 4
Aug 1976. 9. 11 km ESE of Dandaraga HS, nr Sand-
stone, 23 Jan 1974, 9, 9 km NW of Leonora, 19 Jan
1974. 1 juv., McSpeery Gap. Napier Range. SE of
Kimberley Downs HS, 27 Oct 1969. 9. 10 km E of Mt
Thorlan, NW of Christmas Creek HS. 22 Oct 1969, 1
juv.. 10 km E of Mt Thorlan, NW of Christmas Creek
HS, 28 Oct 1969, 1 juv., 28°30’S, 115*45^E, 25 km NE
of Mullewa, 17 Oct 1984 (all ANIC). | juv., Hogarth
Well. Jul 1956 (SAM). c, 24°39’S, 128°45’E, Bunga-
biddy Rockhole, Walter James Range, 15-16 Jan
1990. 9, 23^59'S, 117°32’E, Pingandy Station. 13 Mar.
1980 (all WAM).

Description. Dry specimens yellowish green.
Face with broad whitish horizontal band; meta-
zona of pronotum edged dorsally in reddish
brown, white along mid dorsal line, almost com-
pletely reddish brown to black ventrally: inner
face of fore coxa yellowish with anterior

G. A. MILLEDGE

10

Figures 9-10, Nullabora flavoguttata. 9, male; 10, female. Scale = 10 mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE) 9

11 12

Figures 11-17, Nullabora flavoguttata. 11, male head; 12, female head; 13, female cercus; 14, female abdomen;
15, female foreleg, inside; 16, male genitalia, dorsal; 17, male genitalia, ventral. Scales = 2 mm.

10 G. A, MILLEDGE

marginal teeth black; inner face of fore femur
yellowish in ventral half, femoral spines dark
brown: fore tibia with spines tipped dark brown;
meso and meta sternum reddish brown to black:
costal area of tegmen with white outer band and
green to purplish inner band, outer margin of
costal area greenish, remainder of tegmen hyal-
ine; costal area of hindwing yellow-green,
remainder hyaline; abdomen with broad mid
dorsal longitudinal band, purplish centrally and
white to either side, ventrally (fig. 14) with large
yellow patch bordered by reddish brown on mid
anterior margin of sternites 2-6 (d) or 2-5 (9).
Male genitalia (figs 16-17) with compact,
strongly shagreened pa, consisting of smaller
anterior and larger posterior knobs, the larger
one with transverse furrow.

Measurements (mm). Body length, d 60, 9 66-
70.Head width, g 8.6.9 10.5. Head depth, g 3.8, o
4.6-4.8. Pronotum width, ¢ 2.6, 9 3.5. Pronotum
length, & 19.2, o 25.2-25.9. Tegmen length, c
34.1. 9 41.8. Fore coxa length, d 12.1, 9 15.4.
Fore femur length, d 14,3, o 16.5-17.6. Cercus
length. g 3.3, 9 5.0-7.2.

Immature stages. Nymphs similar in appearance
to adults. Ootheca unknown for certain, how-
ever, | have collected oothecae from shrubs in
which adults and nymphs were also taken and
which probably belong to this species. The
oothecae are small, globular and whitish in
colour with an unusual flap-like structure over-
laying the dorsal exit region.

Distribution and habits. Known from the drier
parts of Queensland, Northern Territory, West-
ern Australia and South Australia (Fig.166). I
have collected this species from acacia shrubs in
the Gulf country of Queensland.

Remarks. Despite being widespread this
species is poorly represented in collections, per-
haps indicating scarcity in nature,

Corthylomantis gen. nov.
Type species. Corthylomantis baldersoni sp. nov.

Diagnosis. Body of moderate size, elongate.
Head about as wide as high, moderately com-
pressed anterio-posteriorly, slightly concave
anteriorly. apical margin curved toward parao-
cular regions: dorso-lateral margin of eyes
rounded; frontal shield with dorsal margin
broadly arched, identations below antennae and
ventral ocellus, lateral margins slightly curved,
with distinct subantennal ridge; antennae of
male about as long as prothorax, of female about
half the length of prothorax.

Pronotum elongate, more slender in male,
supracoxal expansion distinct; lateral margins
slightly lamellete, particularly around prozona
and supracoxal expansion, only lightly denti-
culed in anterior half; prozona smooth dorsally
in male, few scattered tubercles in female, lightly
tubercled ventrally; metazona with distinct mid
dorsal carina.

Fore coxa shorter than femur, with 6 or 7
small blunt teeth on anterior margin; Fore femur
with 4 discoidal spines, the 2nd shortest, the 3rd
longest, with 4 outer spines and 15 inner spines,
claw groove centrally situated; Fore tibia with
14-15 inner and 8-10 outer spines. Mid and
hind femora with small genicular spine.

Male macropterous, wings just surpassing 6th
abdominal segment, tegmen with proximal third
of costal area and costal margin of discoidal area
opaque to subopaque, remainder hyaline,
hindwing with costal area opaque, remainder
hyaline. Female brachypterous, wings covering
first 2 abdominal segments, tegmen completely
opaque, hindwing subopaque.

Abdomen elongate, more slender in male;
cerci short, distal segments only very slightly
compressed laterally, tip of apical segment a
rounded point. Male genitalia with dpr a
strongly sclerotized hook curved dextrad, some-
times with small secondary spine near tip;
medial lobe not prominent. Apr curved sinstrad
and tapering to rounded tip; pa separated from
vl of Iph by membranous area; ml elongate,
finely hirsute. Rph with dorsal arm partially
sclerotized; aa elongate, squamiform sinstrad;
vpl continuous with main body of rph; vspr
compact, u-shaped.

Etymology. The first component of the generic
epithet is derived from the Greek korthylos
meaning little king or chieftain and relates to the
derivation ofthe first component of Archimantis
i.e. from archon meaning chief, ruler.

Corthylomantis baldersoni sp. nov.
Figures 18-28, 167

Material examined (29. o). Holotype: d. 154318.
128739'E, Valentine Rockhole, 12 km WNW of Kunu-
nurra, Western Australia, 8 May 1983. D.C.F.Rentz
and J.Balderson, Stop 4 (ANIC),

Paratypes: 9, 8.3 km SSW of Canobie HS, Queens-
land, 13 Apr 1962, K.H.L.Key and E.L.Corby (ANIC,
13565). 9. 17°23’S, 124°44’E, Lennard River Crossing,
Gibb River Road, Kimberley. Western Australia. Apr
1988, T.F.Houston 678-9 (WAM. 90/489).

Description. Body colour yellowish brown. Pros-
ternum with dark colouration between bases of

REVISION OF THE ARCHIMANTINI (MANTIDAE)

18

On

Tx mova

Figures 18-19, Corthylomantis baldersoni. 18, male; 19, female. Scale = 10 mm.

12 G. A. MILLEDGE

Figures 20-28. Corthylomantis baldersoni. 20. male head: 21, female head: 22. male foreleg, inside; 23, female
hindwing: 24, male cercus: 25, female cercus; 26, male genitalia, dorsal; 27. male genitalia, ventral: 28. male
genitalia, right lateral. Scale = 2 mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE) E

fore coxae; inner face of fore coxa (fig. 22) pale
distally, becoming darker with pale spots proxi-
mally; fore femur with inner face pale, Ist and
3rd discoidal and large inner spines entirely
blackish brown, 2nd and 3rd discoidal, outer
and small inner spines more or less tipped with
blackish brown; inner face of tarsal segments
some times blackish. Opaque areas of tegmen
similar to body colour above, intense reddish
brown beneath; hindwing of male with costal
area brownish, remainder hyaline, hindwing of
female (fig. 23) smokey black in colour, cross-
veins of discoidal area flushed white. Male geni-
talia (figs 126-128) with dpr rather narrow
basally, apr only slightly swollen medially,
anterior section of ventral lamina of Iph small in
area with short projection at junction with pa, pa
a single uncinate dextrad projection, surface
lightly ridged and shagreened.

Measurements (mm). Body length, ¢ 75-78, 9
70. Head width, & 6.2, 9 7.5. Head depth, & 5.4, 9
6.3, Pronotum length, 9 22.2-22.5. 9 24.5. Pron-
otum width, g 3.6-3.7, 9 5.0, Fore coxa length, 9
11.6-11.8, 9 13.5. Fore femur length, d 13,5-
13.9, 015.5. Tegmen length, à 40.5-43,5,9 15.5.
Cercus length, 9 5.7, o 5.8.

Immature
unknown.

stages. Nymphs and ootheca

Etymology. This species is named for John Bal-
derson for his contributions to the collection and
study of Australian praying mantids.

Distribution and habits. Known only from 3
localities, 2 in north Western Australia and 1
from northwestern Queensland (fig. 167). The
habits of this species are unknown, however, the
female paratype has a small label attached stat-
ing the following, ‘active in grass tussocks at
night’,

Remarks. This appears to be a widespread but
uncommon species.

Coenomantis Giglio-Tos

Coenomantis Giglio-Tos, 1917: 45. Type species
Pseudomantis kraussiana Saussure, by monotypy.

Thorodia Vindale, 1923: 452. Type species Thorodia
melanopiera Tindale, by monotypy.

Diagnosis. Male macropterous, female brachyp-
terous, prothorax covered with scattered
tubercles, discoidal area of hindwing pigmented,
cerci short and stout.

Description. Body of medium size, moderately
elongate. Head slightly wider than high, moder-
ately anterio-posteriorly compressed, apical

margin broadly arched, paraocular regions dis-
tinctly lower; eyes prominent, margins rounded:
upper margin of frontal shield curved, with
indentations below antennae and median
ocellus, with distinct subantennal ridge, ventral
margin broadly arched; antennae about the
same length as pronotum in male, distinctly
shorter in female.

Pronotum moderately elongate, with distinct
supra coxal expansion; lateral margins distictly
tuberculate in prozona, moderately tuberculate
in metazona of female, almost smooth in meta-
zona ef male, lamellate, especially around supra-
coxal expansion; dorsal surface with scattered
tubercles, more distinct in female, metazona
with distinct median keel; ventral surface with
scattered tubercles, more distinct in female.

Fore coxa with small scattered tubercles on all
surfaces, anterior margin with 4-5 teeth and
number of smaller tubercles; fore femur with few
small tubercles on all surfaces, 4 discoidal
spines, 2nd very short, 3rd very long, with 4
outer and 13-14 inner spines, ventral surface
with slight depression anterior to 4th discoidal
spine; fore tibia with 7-9 inner and 13-14 outer
spines; hind legs slender, with or without
genicular spine, coxae with small lobe at vento-
lateral corner.

Male macropterous, female brachypterous,
tegmen with costal area opaque, discoidal area
opaque in female but only partially so in male,
hindwing pigmented.

Abdomen moderately elongate, cerci short
and stout with distal segments laterally com-
pressed. Male genitalia with dpr of vph uncinate,
apr elongate and tapering to blunt point, vspr u-
shaped.

Coenomantis kraussiana (Saussure)

Figures 29-51, 167

Coenomantis kraussiana (Saussure), 1873: 25.

Thorodia melanoptera Tindale. 1923: 453, Syn.
nov.

Thorodia melanoptera major Tindale, 1923: 453.

Material examined. Syntype male of T.melanoptera.
Murray River, SA, H,S.Cope, L 14064. Syntype female
of T.melanoptera, Mindarie, SA, L14064. Syntype
male and syntype female of T.melanoptera major.
Kingoonya, SA, R.Harvey, 1.14065. (All SAM).

Other specimens examined (1636, 495. 27 juv.). Qld.
d, 4 km SSW of Birdsville. 4 Dec 1974. 28, 287138,
150717 E. 37 km N of Goondiwindi, 14 Apr 1982, 1
juvy.. 18 km N of Tara, 22 Mar 1962 (all ANIC), g, 10
km W of Etjabuka Stn. 3 Jun 1975 (NMV).

NSW. 9, Brewarrina, May 1962. g, 33 km WSW of
Coolabah, 2 Nov 1963. 2. Round Hill Nature Reserve,
28 Dec 1976. d, Stephens Creek, 24 km ENE of Broken

A. MILLEDGE

G.

14

P

SE
LEZ

I

LTT

Figures 29-30, Coenomantis kraussiana. 29, male; 30, female. Scale = 10 mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE) 15

wo ate ty
NIRE n
hi
T

ies,

Figures 31-39, Coenomantis kraussiana. 31, male head: 32, female head; 33, female eye, colour pattern; 34, male
cercus: 35. female cercus; 36, female wing; 37, female foreleg, inside; 38, mid instar nymph, lateral abdomen; 39,
ootheca, dorso-lateral. Scale = 2 mm.

16 G,

A. MILLEDGE

Figures 40-41,
Scale = 2 mm.

Coenomantis kraussiana. 40,

Hull. 28 Jan 1976 (all AM). &, 32°08’S, 149900'E, 4 km
WSW of Elong Elong. nr Mendooran, 2 Jan 1971. c,

319057S, 14°142’E, Fowlers Gap Research Stn, N of

Broken Hill, 1979-1981. 5, Gilgandra, 17 Nov 1984, 9,
Lightning Ridge, 8 Jul 1977, 9, 32°30°S, 140°20’E, Kin-
chega Nat Pk, Jan 1986. 20, Trangie Exp Stn, 6 km NW
of Trangie, 13-21 Sep 1978. 9, Wittabrenna Creek, 20
km N of Tibooburra, 2 Nov 1971. 49, Wittabrenna
Creek, 20 km N of Tibooburra, 25 Mar 1972 (all
ANIC). 26.9, l juv., 25 km S of Bourke, 21 Feb 1989. 9.

5 km NNW of Silverton, 30 Jan 1990. 9. 86 km N of

Wentworth, 29 Jan 1990 (all NMV), c, Broken Hill, 10
Nov 1976.6, 25 km NE of Broken Hill, 10 Jan 1963. c,
Broken Hill, | Dec 1906. 3, Broken Hill, 10 Mar 1900.
d, 16 km SW of Silverton, 16 Dec 1963 (all SAM).
NT. d Alice Springs, 20-25 Apr 1955. d, 23°42’S,
133°53’E. Alice Springs, 1 Nov 1980. 109, 23°41’S,
133°52’E, Alice Springs, 6-8 Nov 1988. 20, 23°41’S,
133°52’E, Alice Springs, 3-4 Nov 1988. 38, 24118,
133°31’E, Stuart Hwy, 65 km SSW of Alice Springs, 27
Oct 1988. g, Alice Springs, 17 Nov 1965. c, Alice

Springs. 29 Sep 1955. 60, 23°41°, 134715 E, 39 km E of

Alice Springs. 25-26 Sep 1978. d, 25°39’S, 134°38’E,
10 km SE of Finke, 30 Sep 1972. d, Gosses Bluff, 13
Apr 1969. c, 24*10'S. 135°24’E, Hale River Crossing,
7-8 Sep 1955. 9, 24°36’S, 133°12’E, 7 km SW of Hen-
bury HS. 2 Oct 1972, 39, 26°00’S, 131°25’E, 26 km
WSW of Mulga Park HS, 18 Jan 1982. 28, 22°47’S,
136918/E, Plenty Hwy, 268 km ENE of Alice Springs,
14 Oct 1978. 29, 60 km NW of Tanami, 13 Apr 1963. ¢,
2572]'S, 131*03'E. Uluru Motel, Ayers Rock, 4 Nov
1980. (all ANIC). d, Alice Springs, 27 Nov 1954. o,
Kings Canyon Rd. 31 km W of Stuart Hwy, | Oct 1987

male

ventral.

male genitalia,

genitalia, dorsal; 4l.

(all NMV). d, Anthony Well, 8 Aug 1930. 49, 1 juv.,
Double Punch Bowl, Henbury, 15 Oct 1985. 9, Heav-
itree Gap, MacDonnel Ranges. 9, Mulga Park HS, Oct
1960. c, Mount Liebig, Aug 1932. d, West tip
Peterman Range 3 Oct 1963(all SAM).

South Australia. g, Innamincka, Coopers Creek, 24
Jan 1976. d, 27 km S of Innamincka, 28 May 1967
(both AM). I juv., Blinman, 16 Oct 1966. l juv., 40 km
ESE of Burra, 28 Jul 1956. 1 juv., 20 km SE of Burra,
15 Aug 1961. c, Coopers Creek, 19 Oct 1949. 5, Corny
Point. 8, 78 km W of Ernabella Mission, 30 Sep 1960.
9, 33°30°S, 135°54’E, Hambidge Nat Pk, Eyre Peninn-
sula. 16-17 Dec 1970. 1 juv., 47 km NNE of Kingoo-
nya, 16 Jul 1956. c, Lowan Station, 7 km S of Sherlock,
27 Dec 1954. g, 26°20’S, 134956'E,2 km SW of Mount
Barr. SSE of Abminga, 25 Sep 1972. d, 32°17/S,
140°19’E, 25 km WNW of Olary, 20 Dec 1970. 1 juv.,
267078. 138°56’E, Old Alton Downs HS, WSW of
Birdsville, I9 Sep 1972. 1 juv., 1.5 km E of Pimba,
13Jul 1956. 29, Slippery Dip Camp, Brachina Creek,
4. 9 Nov 1987. | juv., Tintinara, 17 Oct 1977. 44,
31°20’S, 138737 E, Trezona Camp, Brachina Creek, 7
Nov 1987. d, 31°20 S, 13837'E, Trezona Camp, Bra-
china Creek. 8 Nov 1987. 1 juv., 60 km NNW of The
Twins HS, 17 Jul 1956. 8, Watson, 22 Oct 1960, 28, 50
km SW of Whyalla, 20 Nov 1985 (all ANIC). 1 juv.,
Abminga Railway Stn, 22 Sep 1987. 5 juv., 5.5 km
NNW of Alberrie Creek Railway Stn, 18 Sep 1987. 1
juv., 9 km SE of Bloods Creek Bore, 22 Sep 1987. 1
juv.. Eringa HS. 190 km N of Oodnadatta, 21 Sep
1987, d, Halidon. d. Hesso, 3 Nov 1987. 29, 22 km W
of Merna Morna HS, 16 Sep 1987. | juv., Moralana
HS. 16 Sep 1987. l juv., Neales Creek, 3 km N of

REVISION OF THE ARCHIMANTINI (MANTIDAE) 17

Figures 42-51, Coenomantis kraussiana, dorsal prothorax. 42, male, Kingoonya, S.A. (syntype, Thorodia mel-
anoptera major); 43, male, Alice Springs, N.T.; 44. male, 6 km NW of Trangie, N.S.W.; 45, male, Murray River
S.A. (syntype, T!melanoptera), 46, male, 2 km SW of Mt. Barr, S.A.; 47, female, Kingoonya, S.A. (syntype,
T.m.major); 48, female, 20 km NE of Carnarvon, W.A.; 49, female, Kings Canyon Rd., 31 km W of Stuart
Hwy., N.T.; 50, female, 13.6 km NW of Lascelles, Vic.; 51, female, 133 km NW of Oodnadatta, S.A.

Scale = 5 mm.

18 G. A. MILLEDGE

Oodnadatta, 20 Sep 1987. 9. 133 km NNW of Ood-
nadatta, 20 Sep 1987 (all NMV). g, Adelaide. o. Alford.
13Jan 1892. 9, Callimurra Waterhole, 1 1.3 km ENE of
Innaminka HS. 31 May 1976. 9, Cameron Corner, 16
May 1976. 7. Coopers Creek, May 1925. 7. Coopers
Creek Ferry Crossing, 30 Nov 1977. g, Dalhousie
Springs, 4 May 1976, c. Farina Creek, 30 May 1976.
28. Fowlers Bay. ¢. Henley Beach, Adelaide. 39. 29,
Kingoonya, 7. Koonalda Stn, Nullabor Plain, Jan
1955. 3, Koonalda, 5 Jan 1960, g. Lake Eyre, 13 May
1963. 29. Lucindale. 7c, Madigan Gulf, Lake Eyre,
Nov 1966. 58. Madigan Gulf, Lake Evre, 16 Feb 1956.
3, Lameroo. 5g, Marree Picnic Ground, 2 Nov 1955. 9,
Mereenie Bluff. 20, 32757'S, 137°24’E, Middleback
Stn, 1 Oct, 25 Nov 1983. 9, Mindarie. o, Mernmerna
HS, 1949, 28, Mudla Bore, 38 km NE of Billa Kalina
HS. 4 Dec 1974. d, 9, Murray River. £, New Kalamu-
rina Stn, Warburton Range, 10 Mar 1972. c, Nth
Prescott Point, Madigan Gulf, Lake Eyre, 2 Nov 1965.
| juv.. Old Billa Kalina Ruins, 4 Dec 1974. 3, Olympic
Dam Site. 2-5 Nov 1987. 9. Ooldea. g, Owieandana, N
Flinders Range, Nov 1924. 9, Pimba. d, Poole Creek.
42 km W of Marree, 29 Sep 1989. | juv., Port Arthur,
22 Apr 1968. d, Port Augusta. 9, Purple Downs, 267
km NW of Port Augusta. 28, 18 km S of Radium Hill,
30 Oct 1962. 2d, Salt Creck, Coorong, 16 Nov 1967.
2d. Simpson Desert, Jul-Aug 1972. c, Strezlecki Creek
Crossing, 15 km SW of Tinga Tingana, 8 May 1976. g,
Talaculanna Waterhole, 2 Dec 1964. c. West of Wel-
bourne Hill. 24 Oct 1953. c, White Wells Reserve, 12
Jan 1960. o, Wirreandah Creek Crossing, 30 km S of
Hawker. 26 Nov 1975. e, Woomera Village. 2 Dec
1961. 8.9, Yurga. Dec 1955 (all SAM). d, Ettaduna Stn,
Birdsville Track, 3 Oct 1972, 4, 5 km NE of Kitto Kit-
talooloo Foreshore, 5 Aug 1971. o. western mid shore
of Lake Palankarinna, Aug 1971 (all WAM).

Western Australia. o. 20 km SSW of Boologooro HS.
NE of Carnarvon, 16 Sep 1961. | juv., 29"49'S,
| 14759'E, 27 km W of Eneabba, 9 Sep 1981.6, LI km N
of Geraldton, 13 Dec 1972. g, 30°45’S. 121°28"E, Kal-
goorlie, 17 Jan 1978. 26, Learmonth, Exmouth Gulf,
17 Sep 1961. 1 juv.. 31°57’S, 126742 E, 32 km SW of
Madura, 8 Oct 1968. d, 57 km ENE of Pardoo HS,
ENE of Port Hedland, 18 Apr 1963. 1 juv., 33°24’S,
1207] "E, 27 km NNE of Ravensthorpe. 16 Nov 1969
(all ANIC). 9. Norseman. Oct 1906 (NMV). 25,
31°26'S, 123°33’E, Buningonia Springs. 18-25 Nov
1978. d, 1.5 km E of Lake Ngapakaldi, 2 Jul-2 Aug
1971 (all WAM).

Victoria. 4, Camel Pad Lake, Hattah Lakes Nat Pk,
24 Jan 1986. 9. 3556'S, 141°40’E. Chinamans Well,
Big Desert. 11-14 Dec 1984. di, 34°12’S, 141°35’E, 8.8
km N of Culluleraine. 17 Nov 1985. d, 34912'S,
141736'E. 8.2 km N of Culluleraine. 14 Nov 1985. 26,
9, Hattah Lakes Nat Pk, 21 Oct 1982, 29, 34"48'S,
142°07°E, 15.5 km WSW of Hattah. Jan 1986. 9.
34 50'S. 14270 T'E. 17 km SW of Hattah. 21 Jan 1987.
7. Irymple, 12 Dec 1955. d, Kewell. Western District.
April 1888. 9, 3 km ENE of Lake Wallawalla. 20 Feb
1986. 9, 34709'S. 141°08"E, 5 km NW of Lake Walla-
walla, 20 Nov 1985.9, 35335, 142°26°E, 15.2 km NW
of Lascelles. 21 Jan 1987. 9. 35°31’S, 142?28'E, 14.9

km NW of Lascelles, 23 Jan 1987. 9, 13.6 km NW of
Lascelles, 20 Jan 1987. d, 9, 34°48’S, 142722'E. Len-
drook Plain, 9 km SE of Hattah, 23 Oct 1985. 1 juv.,
347458, 141°04’E, 3.7 km N of Millewa South Bore,
Feb 1986. 26, 34°36’S, 14103 E, 19.4 km N of Millewa
South Bore, 13, 14 Nov 1985. 9, 34°50’S, 142°33’E, 7.8
km SW of junction of Murray Valley Hwy and
Annuello Rd, Jan 1987. 9, 35°25’S, 141*10'E, 16.8 km
SSW of Murrayville, 23 Feb 1987. 9, Ouyen, Nov
1914. 6, 35°13’S, 143°19’E, 21.3 km NE of Patchewol-
lock, 29 Jan 1987. 9, 5.7 km N of Round Swamp, Big
Desert, Mar 1985. 9, 21.3 km W of Sunset Tank, 19
Nov 1987 (all NMV).

Description. Body grey-brown with numerous
paler and darker blotches and spots giving the
appearance of woody material. Inner face of fore
coxa (fig. 37) and ventral surface of metazona of
pronotum with large pale spots, femoral spines
blackish, eyes finely striped (fig. 33). Tubercles
on ventral surface of metazona elongate in some
specimens, lamellate margins of pronotum of
variable width anteriorly. Tegmina of male
extend to just short of tip of abdomen, costal
area and anterior part of discoidal area opaque
brown above, black and reddish brown below,
remainder virtually hyaline. Tegmina of female
reaching posterior margin of 2nd abdominal seg-
ment, colour similar to male but discoidal area
completely opaque and anal area smokey grey
with pale cross veins. Wings (fig. 36) with costal,

discoidal and anal area smokey brown to black
with bluish sheen; discoidal area with extensive
to almost absent transparent patches between

cross veins; anal area with whitish cross veins.

Abdomen with distinct to not obvious bilobate
intersegmental projections along mid dorsal line
between abdominal segments 2-6; anterior mar-
gin of sternites 2-6 blackish. Male genitalia (figs
40-41) with pa a single caudally directed unci-

nate projection, curving ventro-laterally toward

distal end, margins rather uneven, occasionally

with small ventral projection about mid point.

Measurements (mm). Body length, ¢ 45-69, o
45-62. Head width, 9 4,2-6.5, 9 5.8-7.4. Head
depth, d 3.3-4.3, 9 3.8-5.4, Pronotum length, d
11-17, 9 12-19. Pronotum width, 3 2.4-4.8, o
3.4-6.4. Fore coxa length, 9 7,0-10.5, o 7.6-
11.0. Fore femur length, ¢ 7,5-12,7,99.0-14.0.
Tegmen length, ¢ 29-42, o 13-23. Cercus length,
d 2.5-3.9, o 3.0-4.0.

Immature stages. Nymphs similar in appearance
to adults, dorsal abdominal projections more
distinct (fig. 38); ootheca (fig. 39) of moderate
size, dirty cream in colour, outer spongy layer
very thin to moderate in depth, eggs arranged in
u-shape.

REVISION OF THE ARCHIMANTINI (MANTIDAE) 19

Distribution and habits. Found in the drier
regions of all mainland states of Australia (fig.
167). Individuals are most commonly found
close to the ground in small woody shrubs where
they are excellently camouflaged. Oothecae can
be found attached to the stems of these shrubs
and are often parasitised by wasps of the genus
Podagrion. Adults are usually collected from
spring to early summer. Occasional runt sized
adults are found, probably resulting from growth
under conditions of food shortage.

Remarks. The holotype of this species cannot be
found (Balderson, 1984) Tindale (1923)
described two subspecies of Thorodia melanop-
tera which Beier (1935) later elevated to species
level without giving reasons. After examination
of a large number of specimens I find that the
pronotal variation used by Tindale as the basis
for distinguishing the subspecies shows a grad-
ual range of variation (figs 42-51), also the male
genitalia of the type specimens of T.melanoptera
and T.m.major show no significant difference,
Therefore I consider these taxa synonomous.

Austrovates SjOstedt

Austrovates Sjòstedt, 1918: 35. Type species Aus-
trovates variegata Sjòstedt, by monotypy.

Heterarchimantis Werner, 1922: 121. Type species
Heterarchimantis lobata Werner, by monotypy.

Diagnosis. Body elongate, male macropterous,
female brachypterous, mid and hind legs with
small subapical lobe, cerci with distal segments
laterally compressed.

Description. Large, elongate and stick-like in
form; male macropterous, female brachypte-
rous. Head wider than high, anterio-posteriorly
compressed, concave anteriorly, apical margin
slightly arched; eyes prominent, margins
rounded; frontal shield transverse, apical mar-
gin slightly arched and strongly indented below
antennae and vental ocellus, lateral margins
sinuate, with distinct subantennal ridge; anten-
nae little shorter than pronotum in male, dis-
tinctly shorter in female.

Pronotum elongate and slender, supracoxal
expansion small but distinct; lateral margins
slightly lamellate, finely tuberculate in prozona,
same to almost smooth in metazona; prozona
slightly constricted laterally in centre, few scat-

tered tubercles dorsally, more distinctly
tuberculate below; metazona with distinct mid
dorsal keel and few scattered tubercles dor-
sally.

Fore coxa elongate, slender, anterior margin
with 4-5 small teeth, inner face finely tubercu-
late; fore femur elongate, slender, slightly sinu-
ous, 2nd discoidal spine shortest, 3rd longest, 4
outer and 14-16 inner spines, ventral face with
slight depression anterior to 4th discoidal spine;
tibia with 10 outer and 14-15 inner spines; mid
and hind legs slender with small subapical lobe
on inner margin of femur, genicular spine pres-
ent or absent, genicular lobes produced into
point; mid and hind coxae with small subapical
lobe at ventro-lateral corner.

Abdomen elongate, slender; cerci slightly to
markedly elongate, distal segments laterally
compressed. Male genitalia with 2 semicircular
membranous areas on right lateral margin of
vph, dpr a sharply curved hook; dl of Iph exten-
sive, apr elongate and tapering to blunt point;
vspr u-shaped.

Austrovates variegata Sjöstedt
Figures 52-53, 56-58, 60-66, 168

Austrovates variegata Sjöstedt, 1918: 36.
Heterarchimantis lobata Werner, 1922: 121.

Material examined. Syntype female of A.variegata,
“Noonkanbah, N.V.Austr., Mjóberg, Dec., 224 89"
(NHRM ). Holotype female of H./obata, Port Darwin
(N. Territory), N. Holl. (Australia), V.Bemm (RL).

Other specimens examined (269, 169, 23 juv.). Qld.
g, Almaden, Chillagoe dist., Jan 1932, 3, Bee Creek, 25
km SW of Nebo, 6 Feb 1981. c, Capella, 8 Feb 1981 (all
AM). I juv., 8 km E of Dunbar HS, 10 Oct 1975. g,
Longreach, Jan-Mar 1972. c, ¢, Longreach, 20-31 Jan
1972, I juv., Normanton, 13 Aug 1984. c, 21941'S,
140*30'E, Selwyn Mine, 160 km SE of Mt Isa, 23 Jan
1990 (all ANIC). c, 29, 19*14'S, 140°21"E, Burke and
Wills Junction, 16 Jan 1993. l juv., 20°43’S, 140°21’E,
Butcher Ck., 16 km W of Cloncurry, l Jan 1993. g,
20°51’S, 140°23’E, Dajarra Rd., 19 km SW of Clon-
curry, 2 Jan 1993, 2 juv.. 12 km E of Georgetown, 11
Apr 1991. 9, 4.5 km N of Georgetown, 12 Apr 1991. I
juv., 6 km N of Georgetown, 11 Apr 1991. 9, 17^51'S,
141°08’E, Glenore Pumping Station, Norman River,
13 Jan 1993. 29, 17744'S, 141°05’E, 6 km S of Nor-
manton, 10 Jan 1993 (all NMV). g, Eidsvold (QM). d,
Barcaldine, 6 Jan 1974. 3, Cunnumulla, 26 Dec 1973
(all SAM).

Key to species of Austrovates (males)
l. Tegmen with distinct white marginal band, cerci about same
length as subgenital plate ..................eereoou A.papua
— Tegmen without white marginal band, cerci about twice as long as

subgenital plate

TATOO aite o» A.variegata

20 G. A, MILLEDGE

NT. g, Nhulunbuy. Feb 1973. 9, Oenpelli. East Alli-
gator River, 16 Oct 1948 (both AM). c. 11709'S,
L32QYVE. Black Point, Cobourg Peninsula, 30 Jan
1977.8, 127258. 132758 E. 1 km N of Cahills Crossing,
East Alligator River, 29 May 1973. 1 juv., 19551'8.
136*02"E. 14 km SSE of Dalmore Downs HS, 25 Apr
1976. 9. Daly River Mission, 13-23 Jan 1974. 1 juv..
1471 37S, 130755 E. 57 km WNW of Dorisvale HS, 11
Sep 1968. | juv.. Katherine River Gorge, 18 Aug 1960.
3, Koongarra. 15 km E of Mount Cahill, 26 Apr 1974. 2
juv,. 1639/8, 135951'E, McArthur River HS, 80 km
SW of Borroloola, 13 May 1973. 1 juv. 12°40’S,
13254'E, Magela Creek. 9 km SSE of Mudginberri
HS. 25 May 1973. g. 117078. 132°08"E, Smith Point,
Cobourg Penninsula, 14 Nov 1977. d, 1431”8.
132°22’E, Tindal, 13 km ESE of Katherine, 6 Dec
1967. è, Victoria River Roadhouse, 21 Apr 1982 (all
ANIC). 1 juv., nr Adelaide River, Jul 1963. c, 2 juv.,
Darwin, 1 juv.. 44. Roper River. l juv., Stapleton. 29,
Tennant Creek (all SAM),

WA, 29, Broome, 5 and 9 Nov 1978. d, Broome, 21
Sept 1973. 20, 4? km ESE of Broome. 16 Apr 1969. |
juv.. 14749'S, 126749'E, Carson Escarpment, Kimber-
lev dist.. 9-15 Aug 1975. l juv.. 15°02’S, 126°55°E,
Drysdale River, Kimberley dist., 3-8 Aug 1975. l juv:,
14°53’S, 126°09’E. 5 km NW of King Edward River
Crossing. Kimberley dist.. 19 May 1983. 1 juv.
14°45’S. 12547'E. 10 km NNW of Mining Plateau 17
May 1983. 1 juv., 14"49'S, 125°42’E, Mitchell River
Falls. Kimberley dist. 12 May 1983. &. Mount Tom
Price. Jan-Feb, 1967, 9. Wyndham. 21 Jan 1930 (all
ANIC). 4, Crab Creck. E of Broome (SAM). 1 juv.,
Burma Rd. Reserve, 30 km E of Walkaway, Sep 1986.
ljuv., Hancock Gorge, 15 km S of Wittenoom, 15 May
1980. 7. Town Point. Barrow Island, 9 Sep 1973. I
juv.. Wotjulum. Sep 1955 (all WAM).

Diagnosis. Cerci much longer than subgenital
plate; male without white marginal band on teg-
men, with wings covering first 512 abdominal
segments and dpr narrow and strongly
recurved.

Description, Body brown with darker spots and
blotches: eye with fine striped pattern; inner face
of fore coxa rather darker proximally with pale
tubercles and darkish apical band; tegmen of
male with ventral portion of costal area blackish
proximally and whitish distally with black veins.
costal margin of discoidal area flushed brown
except for transparent stigma bounded at either
end by a dark patch, remainder hyaline: tegmen
of female similar but opaque portion of discoi-
dal area more extensive; hindwing with costal
area dark brown, more so apically, remainder
hyaline. Wings reaching or just surpassing pos-
terior margin of abdominal segment 5 in male,
segment 4 in female: cerci elongate (fig. 60-61),
distal segments broadened and laterally com-
pressed, apical segment with margin rounded

and sometimes with slight notch below apex;
anterior margin of abdominal sternites 2-5
blackish; antennae of male about two thirds the
length of pronotum. Male genitalia (fig. 65-66)
with dpr of vph a strongly curved elongate hook,
slightly sinuous distally; pa smooth with single
elongate pointed projection curving latero-
ventrally.

Measurements (mm). Body length, ¢ 92-94, 9
96-100, Head width, d 6.5-8.3, 9 8.5-9.2. Head
depth, d 3.7-4.2, 9 5.0-5.5. Pronotum width, d
3,1-3.6, 9 4.4-4.6. Pronotum length, 9 25.5-
31.5.9 34.3-36.9. Fore coxa length, 9 10.8-14.9,
9 17.2-17.8. Fore femur length, 9 13.0-16.7, 9
20.0-20.7. Tegmen length, d, 38.5-44.4, 9, 38.1-
44.0. Cercus length, ¢ 4.6-7.0, 9 6.2-7.9.

Immature stages. Nymphs similar in appearence
to adults but possessing mid dorsal intersegmen-
tal abdominal nodes (fig. 63). Ootheca (fig. 64)
elongate, brown and whitish, with insubstantial
outer foamy layer and sometimes thin projec-
tion at posterior tip of dorsal exit line. The
ootheca is dissimilar to those presently known
for other Archimantini.

Distribution and habits. Found in the northern
parts of Queensland, Northern Territory and
Western Australia (fig. 168). I have collected this
species from shrubs and small trees in open
woodland country in central north Queensland.
Although females have reduced wings they are
capable of at least short flights.

Austrovates papua sp. nov.
Figures 54-55, 59, 67-68, 168

Material examined, Holotype male, Doveta, Amazon
Bay area, New Guinea, 2400 ft, 24 Jul 1962.
W.W.Brandt (ANIC).

Diagnosis (male only). Cerci about same length
as subgenital plate, tegmen with white marginal
band, wings covering first 6⁄2 abdominal seg-
ments, genitalia with dpr broad and weakly
recurved.

Description (male only). Body of dry specimen
dark brown with darker and paler blotches and
spots, inner face of fore coxa dark with numer-
ous pale tubercles; costal area of tegmen with
outer white band, which diminishes distally and
inner blackish band, costal margin of discoidal
area cream in proximal quarter with dark spot at
proximal end of stigma and dark brown distally,
remainder of tegmen hyaline; hindwing with
costal margin blackish brown, remainder hyal-
ine: wings surpassing caudai margin of sixth

REVISION OF THE ARCHIMANTINI (MANTIDAE)

a

52

Austrovates variegata, male. Scale = 10 mm.

Figure 52,

22 G. A. MILLEDGE

TL

TI

=;

Figure 53, Austrovates variegata, female. Scale = 10 mm

REVISION OF THE ARCHIMANTINI (MANTIDAE)

Figure 54, Austovates papua, male. Scale = 10 mm.

24 G. A. MILLEDGE

Figure 55. Austrovates papua, male head.

Figures 56-58. A.variegata. 56. male head; 57. female head: 58, female head. eye pattern.

Figure 59, A.papua, male cercus.

Figures 60-64. A. variegata. 60, male cercus: 61, female cercus; 62, female foreleg, inside: 63, mid instar nymph,
lateral abdomen; 64. ootheca, dorso-lateral. Scale = 2 mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE)

Figures 65-66, Austrovates variegata. male genitalia. 65, dorsal; 66. ventral.
Figures 67-68. A.papua, male genitalia. 67. dorsal; 68. ventral. Scale = 2 mm.

ro
[97

26 G. A. MILLEDGE

abdominal segment: cerci (fig. 59) quite short,
distal segments broadened and laterally com-
pressed, whitish in colour, apical segment with
rounded margin. Genitalia (fig. 67-68) with dpr
a rather broad short hook, pa smooth with single
pointed projection directed posterio-laterally
and slightly curved ventrally.

Measurements (mm). Body length, 95. Head
width, 7.7. Head depth, 4.3. Pronotum length,
35.4. Pronotum width, 3.7. Fore coxa length,
15.2. Fore femur length, 17.4. Tegmen length,
49.5. Cercus length, 3.9.

Immature stages. Unknown.

Etymology. Specific epithet refers to the country
in which the holotype was collected. It is a noun
in apposition with feminine gender.

Distribution and habits. Known only from the
holotype locality in southeastern Papua New
Guinea (fig. 168). Habits are unknown but is
likely to bea shrub dweller. The vegetation ofthe
type locality is a mixture of grassland and wood-
land (Paijmans, 1975).

Archimantis Saussure
Archimantis Saussure, 1869: 56. Type species Man-
tis latistylus Serville, by monotypy.
Rheomantis Giglio- Tos. 1917: 44. Type species Fis-
vheria quinquelobata Tepper. by monotypy.

Diagnosis. Body elongate, eyes with rounded to
distinctly angular lateral margins, males
macropterous, females brachypterous, mid and
hind femora of adults without preapical lobes,
cerci with distal segments broadened and
laterally compressed.

Description. Body large, elongate, slender to
rather robust. Head wider than high, anterio-
posteriorly compressed, apical margin straight
to slightly arched; frontal shield strongly trans-
verse, usually with subantennal ridge; eyes with
rounded to distinctly angular lateral margins;
antennae as long or longer than pronotum in
male, less than half the length of pronotum in
female.

Prothorax long and slender with slight to pro-
nounced supracoxal expansion; prozona smooth
tolightly tuberculate above, strongly tuberculate
beneath; metazona with more or less distinct
median carina; pronotal margins smooth to
strongly toothed.

Male macropterous but wings not entirely
covering abdomen, female brachypterous; teg-
men of male with costal area and costal margin
of discoidal area opaque, remainder hyaline,
tegmen of female similar to male or completely
opaque, opaque areas strongly coloured beneath
in both sexes; hindwing hyaline except for costal
margin. Fore coxa shorter than metazone of

Key to the species of Archimantis
Fore margin of anterior coxa with 3 to 5 mostly large teeth, black
beneath v. e gen Ron T DO n T A.quinquelobata
Fore margin of anterior coxa without such teeth .......... 2
Lateral margins of pronotum strongly toothed ............ 3
Lateral margins of pronotum finely dentate or smooth ..... 4
Supracoxal expansion of pronotum broad (figs 102, 103)

T e eT RERO Pe Oo ta e Mir Ce de, ten pe A. monstrosa
Supracoxal expansion of pronotum narrow (figs 93, 94)

e SA AUI SEU UE, Lie e Arti NS PU Cees KA EISE A. armata
Lateral margin of eye distinctly angular, terminal segment of
cereus, distinetlyspormited n. at imc TRE EE 5
Terminal segment of cercus rounded or if pointed then eye
inasgin/moutdedaes. ina m cet his noes ER 5 HE 6
Cerci extremely elongate, pattern on inner face of fore coxa with
large pale areas distally ....................... A. straminea
Cerci moderately elongate, pattern on inner face of fore coxa dark
OSE, H O IENE A. brunneriana
Robust species, inner face of fore coxa pale proximally ..... 7
Slender species, inner face of fore coxa dark proximally .... 8
Tegmina of female short, covering only the first two abdominal
terga: parat maleconmpadi Ee ee inno . A. sobrina
Tegmina of female medium, covering almost the first four
abdominal terga, pa of male elongate spiniform ... A. latistyla
Tegmina of male distinctly banded ................. A. vittata
Tegmina of male not banded „...................; A. gracilis

REVISION OF THE ARCHIMANTINI (MANTIDAE) 27

pronotum, anterior margin lightly to strongly
armed; fore femur slender with 15 inner and
four outer spines, Ist discoidal spine much
longer than second, claw groove centrally situ-
ated; fore tibia with 8-11 outerspines and 13-17
inner spines; mid and hind legs long and slender,
femora of adults without preapical lobes.
Abdomen elongate, slender and dorso-ven-
trally compressed, more so in male; anterior
margins of abdominal sternites distinctly
coloured; supraanal plate transverse, cerci
slightly to distinctly elongate with distal seg-
ments strongly laterally compressed. Vph of
male genitalia with medial lobe prominent and
more or less curved dorsally; dpr more or less
produced into single, strongly sclerotised hook
curving dextrad; Iph with apr curved sinstrad
with rounded tip; pa separated from vl of Iph by
membraneous area; ml elongate and finely hir-
sute; rph with medial arm partially sclerotised,
aa elongate but squamiform sinstrad; vpl con-
tinuous with main body of rph; vspr small,
heavily sclerotised and more or less u-shaped.

Archimantis latistyla (Serville)
Figures 69-80, 170

Mantis latistylus Serville, 1838: 179

Archimantis latistylus (Serville).-Saussure,
1869: 65

Archimantis latistyla (Serville).-Giglio-Tos,
1912: 164

Mantis fuscielytris McCoy, 1886: 118.

Archimantis latistylus gigantea Beier, 1963: 9.
Syn. nov.

Material examined. Two syntype females of Mantis
fuscielytris, Victoria (both NM V). Holotype female of
Archimantis latistylus gigantea, Rockhampton (Qld),
ex Museum Godeffroy, 2677 0.273 (ZMH).

Other specimens examined (1043, 1599, 110 juv.).
ACT. 1 juv., Black Mt, 30 Dec 1929. o, Black Mt, 15
Dec 1964. c, Black Mt, 13 Jan 1965. 9, Black Mt, 22
Dec 1964. c, Black Mt, 10 Jan 1966. c, Black Mt, 23
Dec 1969. 1m, Black Mt, 22 Jan 1967. c, Black Mt, 24
Dec 1965. c, Black Mt, 23 Jan 1964. d, Black Mt, 16
Dec 1982. 26, Black Mt, Jan 1969. 63, 9, Black Mt, Dec
1961.¢, Black Mt, Oct 1961.¢, Black Mt, Jan 1969. 3c,
Black Mt, Jan 1963. 2, Black Mt, Dec 1963. 9, Can-
berra, 15 Jan 1978. 9, Canberra, 21 Jan 1962. 9, Can-
berra, 19 Jan 1987. 9, 35°18’S, 148*58'E, 1 km E of Mt
McDonald, 11 Dec 1980. d, 35°22’S, 148 5T'E,
Murrays Corner, Paddys River, nr Canberra, 29 Dec
1985. o, Uriarra, 28 Jan 1984 (all ANIC). o, Black Mt
Reserve, Canberra, 26 Jan 1982 (NMV).

Qld. d. Cathu State Forest, N of Eungella Nat Pk, 21
Jan 1990. 9, Clermont, 7 Feb 1981. 9, Clermont, Jan
1929. 28, Coominglah Range, 24 km N of Monto, 6
Jan 1975. 49, 8 km E of Emuford, 30 Dec 1989. 28,
Forty Mile Scrub, 65 km W of Mt Garnet, 19 Dec

1974. 9, Millstream Falls, nr Ravenshoe, 5 Jan 1967. 9,
Mount Tamborine, Dec 1925 (all AM). d, 9, 2 juv., 20
km W of Arcadia HS, 24 Oct 1967. d, 2o, | juv.,
2526'S, 15123'E, Binjour Plateau, 31 km N of Gayn-
dah, 19 Apr 1982. 9, Bluff Range, nr Biggenden, 2-12
May 1971. d, Bodumba Creek, nr Kenilworth, 12 Jan
1986. 28, 3o, 11 juv., 22°31’S, 148°43’E, 9 km ESE of
Bombandy HS, 24 Apr 1981. d, Bribie L. 18-26 Dec
1972. 9, 28°02’S, 153°26’E, Broadbeach, 28 Dec 1970.
l juv., 20 km NE of Bundaberg, Apr 1971. 2 juv..,
Bundaberg, Apr 1971. 1 juv., Bundaberg, Mar 1971.
29, 1 juv., Bundaberg, May 1971. 1 juv., 3 km N of
Bundaberg, 19-21 Apr 1971. d, 9, 4 juv., 26°00’S,
153*05'E, Camp Milo, Cooloola Nat Pk, 16 Apr 1982.
9, Castle Hill, Townsville, 3 Apr 1962. g, Ceratodus, 31
Dec 1955, 8, 8 km S of Clermont, 18 Mar 1982. 1 juv.,
13°56’S, 143°12’E, Coen, 18 Jul 1986. d, 18°35’S,
144°44’E, 12 km N of Conjuboy HS, 12 Nov 1981. d,
Cunninghams Gap, MacPherson Range, 3-4 Dec
1982. 9, 25°21’S, 151" 13'E, 9 km NE of Eidsvold, 20
Apr 1982. 9, 18°03’S, 144°52’E, Forty Mile Scrub Nat
Pk, 52 km SSW of Mt Garnet, 22 Jul 1986. 9, Gordon-
vale, 19 Jan 1962. 9, 9, 11°41’S, 142°28’E, 3 km NE of
Gunshot Creek, 17 Mar 1992. d, 11*49'S, 142°30’E, 6-
10 km SSE of Heathlands HS, 14 Mar 1992. 29,
11*49'S, 142*30'E, 6-10 km SSE of Heathlands HS, 28
Mar 1992. d, 11°45’S, 142°35’E, Heathlands HS, 10
Mar 1992.6, 11°46’S, 142°41’E, 11 km E of Heathlands
HS, 11 Mar 1992. d, 17 km NNW of Hivesville, 3 Jan
1965. 1 juv., 1514'S, 14507'E, 7 km N of Hope Vale
Mission, 4 Oct 1980. c, o, 4 juv., 23°46’S, 149°06’E, 2
km S of Horshoe Lookout, Blackdown Tableland, SW
of Dingo, 1-2 Feb 1981. 1 juv., 18°27’S, 146°08’E, 22
km N of Ingham, 28 Sep 1979. d, Innot Hot Springs, 11
Jan 1962. d, 17°25’S, 145°04’E, 15 km W of Irvine-
bank, 27-28 Nov 1981. 3juv., 18km SW of Lake Nuga
Nuga, Carnarvon Range, 25 Oct 1967. d, 150 km N of
Marlborough, 9 May 1955. o, 28°04’S, 153°27’E,
Miami, 31 Dec 1970. 29, 2 juv., 27°51’S, 150721'E, 15
km S of Moonie, 22 Apr 1982. 26, 39, 7 juv., 15910'S,
145907'E, 3.5 km SSW of Mt Baird, nr Cooktown, 3-5
May 1982. 9, Mt Cook Nat Pk, 11 Oct 1980. 9, 26°54’S,
152°54’E, Mt Coonowrin, Glasshouse Mts, 9 Aug
1985. 3 juv., 26°54’S, 152°38"E, Mt Ngungun, Glas-
shouse Mts, 9 Aug 1985. d, 29, 4 juv., 25926'S,
152°56’E, Mt Tibrogargan, Glasshouse Mts, 15 Aug
1985. 9, 6 juv., 12°44’S, 143*13'E, 2 km NNE of Mt
Tozer, Iron Range Nat Pk, 3 Jul 1986. 7 juv., 12°42’S,
143°20’E, 13 km ENE of Mt Tozer, nr Iron Range Nat
Pk, 10 Jul 1986. 29, 9, 15°03’S, 145°09’E, 3 km NE of
Mt Webb, nr Cooktown, 30 Apr-3 May 1981. 3 juv.,
1503'S, 145°09’E, 3 km NE of Mt Webb, 2 Oct 1980. 9,
12 km W of Moura, nr Banana, 30 Dec 1955. d,
23°41°S, 149°51°E, Mourangee Stn, NW of Edungalba,
nr Duaringa, 2 Jul 1980. 2 juv., 3 km SE of Planet
Downs HS, E of Rolleston, 25 Oct 1967. 1 juv.,
17*38'S, 145/26 E, 7 km WSW of Ravenshoe, 12 Nov
1981. d, 23°43’S, 150'42'E, 50 km S of Rockhampton,
13 Dec 1968. 3 juv., 18km N of Roma, 24 Oct 1967. d,
o, 15°17’S, 145*13'E, 1 km N of Rounded Hill, nr
Cooktown, 5-7 May 1981. 28, 9, 15°17’S, 145°14’E, 3
km ENE of Rounded Hill, nr Cooktown, 5-7 May

G. A. MILLEDGE

70

GESTUS

Figures 69-70, Archimantis latistyla. 69, male; 70, female. Scale = 10 mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE) 29

istyla. 71, male head; 72, female head; 73, male cercus; 74, female cercus; 75,

Figures 71-80, Archimantis lat
8, male genitalia, dorsal; 79, male genitalia, ven-

ootheca, lateral; 76, ootheca, dorsal; 77, ootheca, transverse; 7
tral: 80, male genitalia, right lateral. Scale — 2 mm.

30 G. A. MILLEDGE

1981. l juv., 15 km NE of St.George, 8 Nov 1974. o,
Stanthorpe, 11 Jan 1982. 9, Taroom, 3 Jul 1974. d,
28'56'S, 151°08"E, 9 km SSW of Texas, 24 Nov 1983.
2, l juv., 25°46’S, 152°38’E, 8 km SE of Tiaro, 14 Aug
1985. 2 juv., Watalgan Range, nr Bundaberg, 6 Nov
1971, 1 juv., 130°6’S, 142"56/E, Wenlock River Cross-
ing, Portland Roads Rd, 30 June 1986. d, o, 26"07'S,
152'47'E, Wilsons Pocket, 14 km ENE of Gympie, 18
Apr 1982 (all ANIC). 9, 15*19'S, 145°O1’E, 2.5 km S of
Bald Hills Stn, 4 Apr 1991. g, Brisbane, 3 Dec 1965. 9,
Bowen. | juv., Bruce Hwy, 4 km S of Bowen, | Apr
1991.3, 140 km S of Charters Towers, 19 Feb 1989. d,
Chillagoe, 27 Mar 1989. 9, 17°13’S, 145947'E, 5.5 km
N of Collins Weir, W of Atherton, 10 Apr 1991. 9,
1771 5'S, 145°17’E, 1 km NNE of Collins Weir, W of
Atherton, 10 Feb 1989. o, Cooktown, Jul 1970. g,
Cooktown, 7 Apr 1991. d, Cooktown, 8 Apr 1991. 2,
16°59’S, 145°33’E, Davies Creek Rd, 2 km E of Ken-
nedy Hwy, 8 Feb 1989, g, 16°43’S, 145°38’E, Ellis
Beach, 28 Nov, 1983. o, Forty Mile Scrub Nat Pk, 22
Feb 1989. o, Georgetown, 15 Apr 1991. 1 juv., 15°32’S,
145"| 2E, Hardwicke Creek, 0.5 km S of Annan River,
6 Apr 1991. 9, Hodzoic Rd, 0.7 km E of Peninsular
Development Rd, N of Mareeba, 15 Feb 1989. 26,
Julago, 13 km SE of Townsville, 3 Feb 1989. 9, 1843'S,
14438'E, Kennedy Development Rd, Spring Creek HS
turnoff, 17 Apr 1991. 29, Mt Molloy, 10 Mar 19*90. 29,
10 km S of Mt Ossa, 3 Feb 1989. d, 15°31’S, 145°08’E,
Oaky Creek Rd, 15 km SW of Cooktown, 9 Apr 1991
(all NMV). g, Brisbane, 25 Jan 1916. c, Brisbane, 22
Dec 1917. 7. Brisbane, 13 Oct 1914. 9, Brisbane, 14
Mar 1918, 9, Brisbane, 22 Oct 1917. 9, Emerald. 9,
Moreton Island, Apr 1916. 9, Stanthorpe, 30 Jan 1941.
29, Stradbroke Island, 5 Dec 1913. (all QM). c, Cairns.
9, Endeavour River, Cooktown. 39, Maryborough. g,
Somerset. d, Stanthorpe (all SAM). 3, Biloela, 11 Feb
1947. 9, Brisbane, 3 Nov 1952, d, Brisbane, | Apr
1961. 9, Brisbane, 16 Sep 1962. 4, Brymaroo, 30 Jan
1948. o, Mackay, 3 Feb 1934, 7. Murarrie, 4 Jun 1964.
9. Nambour, Jan 1962. o, Yuleba, Jan 1964 (all
UQ).

New South Wales. 29, Beacon Hill, nr Sydney, 7 Dec
1978. 9, Clark Island, 22 Jan 1930. 1 juv., Dobroyd
Head, Sydney Harbour, 14 Feb 1978. g, Eccleston, 15
Dec 1922. d, French's Forest, Sydney, 7 Nov 1922. g,
Hornsby, Mar 1911, 9, Jannali, 26 Nov 1933. 1 juv.,
Kurnell, 3 May 1931. c, Matraville, Nov 1925, 1 juv.,
North Head, Sydney, 14 Feb 1978. 9, St.Ives, nr Syd-
ney, Dec 1967. 28, Sydney. e, Vaucluse, Sydney, Apr
1927. 9, Wallaby Creek, Tooloorn, 5-13 Dec 1962. o,
Willoughby, Dec 1922. 9, Woronora River, Engadine,
16 Dec 1976 (all AM). 7. Bankstown, Sydney, Oct
1964. 9, 33°03’S, 151°40’E, Belmont Golf Club, nr
Newcastle, 19 Nov 1982. 2 juv., 3444/5, 147°54’E, 5
km NE of Bethungra, 13 Sep 1968. 9, Bomaderry, 25
Dec 1967. 1 juv., Bonville, 25 Aug 1977. 2, 6 km W of
Bonville, 12 Dec 1969. 3, Broulee, 9 Jan 1962. 9,
33°08’S, 151°38’E, 3km NNE of Catherine Hill Bay, nr
Newcastle, 19 Nov 1982. 9, Cheltenham, 12 Mar 1967.
d, 3558'S, 150°09’E, Congo, 8 km ESE of Moruya, 17
Dec 1983. I juv., 3501'S, 15049’E, Currarong, 17 Aug
1977. 2 juv., Enmore Falls, 19 Mar 1961. 1 juv.,

29°06’S, 153724'E, 3.7 km NW of Evans Head, 14 Dec
1971. o, 29°07’S, 153°25°E, 2.3 km WNW of Evans
Head, 14 Dec 1971. 9, 5 km N of Evans Head, 13 Dec
1955. d, 2o, 1 juv., 28°24’S, 152°53’E, Findon Creek,
27 km E of Woodenbong. 20 Nov 1983. o, Forster, 2
Feb 1967. 1 juv., Forster, 11 Dec 1947. d, 33°12’S,
151°36’E, Geebung Camp, Lake Munmorah, 15 Nov
1983. 1 juv., 32°53’S, 151°32’E, Great Sugarloaf
Mountain, nr West Wallsend, 16 Nov 1983. 9, 3502'S,
15035’E, Huskisson, 26 Sep 1984. 9, Killarney
Heights, 11 Oct 1969. o, Kincumber, Dec 1949, 29,
Kogarah, May 1925. 1 juv., Korora Bay, nr Coffs Har-
bour, 22 May 1966. d, 28°35’S, 153°08’E, 13 km ENE
of Kyogle, 20 Nov 1983. o, Lake George, 29 Jan 1972.
9, Lake George, 2 Feb 1972. o, 6 juv.., Lake George,
1974. 3 juv., Lake George, 1971. 1 juv., Lawrence, 22
Jun 1985, | juv., Lindfield, 23 Apr 1927. 9, 4 km S of
Mogriguy, | Jan 1971. d, 34°46’S, 150°37’E, Mt Tapi-
tales, 7 km NW of Berry, 30 Dec 1975. 9, Mt Wilson,
11 Jan 1929. 9, 8 km S of Murwillumbah, 31 Jan 1963.

9, Narrabeen, 28 Sep 1951. 29, Nelligen, 4 Dec 1950. o,
35°36’S, 150°04’E, 8 km NW of Nelligen, 15 Feb 1984.

] juv., 1.5 km SSW of Nerriga, 14 Nov 1962. 1 juv.,

Northbridge, 26 Oct 1927. 3, Northbridge, Nov 1928.

9, Point Perpendicular, Jervis Bay, 27 Dec 1966. 9,

31°26’S, 152°56’E, Port Macquarie, 31 Dec 1968. 9,

Queanbeyan, Feb 1985. d, Sans Souci, Georges River,

19 Jan 1955. 29, South Turramurra, 26 Mar 1967. 8, 9,

1.5 km SE of Southwest Rocks, NE of Kempsey, 3 Jan

1970. 1 juv., Sydney, 28 May 1951. d, Teachers Col-

lege, Wagga, 1968. 9, 2 km NNW of Teagardens, Port

Stephens, 15 Jan 1971. 9, Tea Tree Creek, Armidale, 7

Jan 1960. 9, 28°58’S, 151°43’E, 31 km WNW of Ten-

terfield, 23 Nov 1983, d, 3 km Sof Tooraweenah, 6 Jan

1956. 1 juv., 3 km S of Ulladulla, 15 Oct 1966 (all

ANIC). d, 25 km ENE of Goulburn, 30 Dec 1990

(NMY). 9, Sydney (SAM).

Vic. 29, Rosebud, Mornington Penninsula, 16 Jan
1970. 9, 37°59’S, 147°43’E, Rotamah Island, 20 km
SSE of Bairnsdale, 30 Nov-1 Dec 1984. 9, 20 km ENE
of Stratford, 2 Dec 1956. 1 juv., 39°01’S, 146°18’E,
Tidal River, Wilsons Promintory, 5-6 Nov 1985 (all
ANIO). l juv., Balwyn, 27 Aug 1908. 9, Bullbeef Creek,
2 km W of Mt Tanjil, 19 Apr 1982. d, 34°43’S,
142"26'E, Chalka Creek, Hattah Lakes Nat Pk, 21 Jan
1987. o, Croydon, 4 Dec 1904. 1 juv., Fernshaw. 29, 2
juv., Frankston. 9, Gippsland. 9, Kow Plains, Dec
1910. 9, Launching Place, 9 Nov 1983. d, 9, Moorool- |
bark, Mar 1977. o, Murrayville Track, 13 km S of
Murrayville, 26 Feb 1987. 39, 35258, 141?10'E, 16.8
km SSW of Murrayville, 18 Feb 1987. o, Rhyll, Phillip
Island, 8 Mar 1948. 2 juv., Rotamah Island, Gippsland
Lakes, 9 Mar 1986. o, Trafalgar, Gippsland, 9 Dec
1892. 9, 35°32’S, 142°32’E, Waithe Fauna Reserve, 24
Jan 1986. d, o, 35°32’S, 142°25’E, Waithe Fauna
Reserve, 21 Jan 1987. d, 35°32’S, 142°29’E, Waithe
Fauna Reserve, Feb 1987 (all NMV). 29, Healesville
district. | juv., Healesville, 20 Oct 1910. 9, Mt Yule,
Healesville, 14 Nov 1911 (all QM).

SA. I juv., 33°53’S, 136°36’E, 5 km NE of Arno Bay,
21 Oct 1982. d, 34°08’S, 135°22’E, 3 km SSE of Mt
Hope, 20 Oct 1982. 1 juv., 8 km SSE of Mt Hope, 29

REVISION OF THE ARCHIMANTINI (MANTIDAE) 31

Oct 1969. 3. 9, 1 juv., 35*59'S, 137°11’E, Vivonne Bay,
Kangaroo Island, 8 Dec 1977 (all ANIC). 9, Adelaide, 3
Jan 1887. d, Kangaroo Island. 9, Lucindale, 29 Jan
1892. d, Murat Bay. d. Murray River. o, Peake, 23 Mar
1909. d, Scorpion Springs Conservation Pk, 5 km SW
of Nanams Well, 17 Dec 1983 (all SAM). Western Aus-
tralia. d, 29°35’S, 115°09’E, 27 km NNW of Eneabba,
24 Oct 1984. d, 33°18’S, 123°22’E, 18 km NNW of Mt
Ragged, 12 Nov 1969 (both ANIC).

Diagnosis. Eyes with lateral margins rounded,
margins of pronotum without distinct teeth,
anterior margin of fore coxa without large teeth,
inner face of fore coxa pale proximally, tegmina
of female covering first 3-4 abdominal seg-
ments, apical segment of cercus with rounded
margin, pa of male genitalia produced into
single spine.

Description. Body rather robust, brown, greyish
brown or greenish brown. Apical margin of head
broadly arched, eyes slightly protuberent anter-
iorly with rounded lateral margins, frontal
shield with distinct subantennal ridge. Protho-
rax moderately to very elongate, slight but dis-
tinct supracoxal expansion, lateral margins
smooth except in prozona of female where they
are finely tuberculate, metazona smooth dor-
sally with distinct mid dorsal keel, prozona with
few small scattered tubercles dorsally but with
denser and more pronounced tubercles beneath.
Fore coxa with 5 small sharply pointed teeth on
anterior edge, females often with few smaller
ones between, inner face not distinctly pat-
terned. Fore femur with all spines blackish. Fore
tibia with 14-15 inner and 8-1 1 outer spines, all
tipped blackish brown. Mid and hind femora
sometimes with genicular spine. Wings of male
covering first seven abdominal segments; teg-
men hyaline except for costal area and costal
margin of discoidal area which are opaque
brown above, costal area beneath glossy black in
proximal fifth with remainder whitish with
black cross veins, costal margin of discoidal area
intense orange-brown beneath; discoidal area
usually with dark spot at proximal and distal end
of stigma though distal spot sometimes absent,
venation slightly to distinctly darkly pigmented.
Wings of female just surpassing caudal margin
of third abdominal segment to just short of cau-
dal margin of fourth; tegmen similar in colour to
that of male except discoidal area completely
opaque. Hindwings of both sexes with costal
area orange-brown, remainder hyaline with
veins slightly to distinctly pigmented particu-
larly toward apex.

Abdomen without distinct white mid dorsal
stripe, anterior margin of sternites 3-5 coloured
black and red but only visible when abdomen
flexed. Cerci (fig. 73-74) as long as to about
twice as long as subgenital plate, apical segment
with margin rounded. Male genitalia (figs 78-
80) with dpr of vph strongly curved, broad at
base, narrow toward tip; anterior portion of vl of
Iph broadly rectangular, slightly produced at
junction with pa; apr swollen medially, tip
slightly mucronate; pa produced into single
spine directed laterally to posterio-laterally, sur-
face finely shagreened except at tip.

Measurements (mm). Body length, ¢ 78-97,
o 87-127. Head width, d 7.8-8.8, 9 9.0-11.6.
Head depth, s 4.6-7.4, 9 6.0-9.6. Pronotum
width, ¢ 3.8-4,6, o 5.4-7.2. Pronotum length,
d 25-32, 9 28-45. Fore coxa length, 9 12-16, 9
16-21. Fore femur length, d 14-18, 9 18-24.
Tegmen length, g 48-53, 9 32-42. Cercus length,
3 6.4-9.8, o 8.0-9.0.

Immature stages. Nymphs have a broken white
mid dorsal abdominal stripe and are often more
mottled in colour than adults. There is also a
green colour variant in nymphs however this
appears to become brownish in the final instar.
The ootheca (figs 75-77) is large and globular
with a thick spongy outer layer, eggs arranged in
a w-shaped formation and is pale brown or green
in colour. It is deposited on the stems of shrubs
and tall grasses, and is often parasitised by wasps
of the genus Podagrion.

Distribution and habits. Known from eastern
Australia, essentially along the Great Dividing
Range, from Cape York south through Victoria
to the south eastern corner of South Australia
(fig. 170). There are also two doubtful records
from Western Australia. Predominantly a shrub
dweller although nymphs may sometimes be
found in grasses. Adults are most commonly
encountered from December to April. The dis-
tinctive colours on the ventral costal margins of
the wings and on the anterior margins of
the abdominal sternites are seen during the
defensive display.

Remarks. Serville's syntypes of this species can-
not be located (Balderson, 1984). This is a
widely distributed and somewhat variable
species. Specimens become progressively larger
and the prothorax more elongate toward the
north. The cerci also become reduced in size at
the northern end of the range. Beier's A./.gigan-
tea falls within this range of variation.

32 G. A. MILLEDGE

Archimantis sobrina Saussure
Figures 81-92. 174

Archimantis sobrina Saussure, 1873; 26.
Archimantis minor Giglio-Tos. 1917: 43. Syn.
nov.

Material examined. Syntype female of Archimantis
minor, King George's Sound (Western Australia).
Captain Grey, 40 12-16 17b. Syntype female of Archi-
mantis minor. Swan River (Western Australia), 43 14
(both BMNH).

Other specimens examined (2529, 1509, 94 juv.).
ACT. 36, Black Mt, 29 Dec 1965. c, Black Mt, 16 Jan
1961, 3d, Black Mt. 22 Dec 1964. c, Black Mt, 25 Jan
1965. 149, Black Mt, 29 Dec 1961. 3c, Black Mt, 3 Jan
1962. 2c, Black Mt, 4 Jan 1962. 4c, Black Mt, 24 Dec
1962. g, Black Mt, 14 Jan 1963. 38, Black Mt, 2 Jan
1963. 3, Black Mt, 5 Jan 1962. 3g, Black Mt, 15 Jan
1962.2, Black Mt, 26-28 Jan 1963.6, Black Mt, 29 Jan
1963. c, Black Mt, 20 Jan 1950. 29, Black Mt, 13 Jan
1950. d, Black Mt, 30 Dec 1951, d, Black Mt, | Jan
1950. 20, Black Mt, 5 Jan 1952. c, Black Mt, 20 Jan
1953. 20, Black Mt, 26 Dec 1961. d, Black Mt, 4 Jan
1952. 28, Black Mt, 22 Dec 1952, g, Black mt, 19 Jan
1953. c, Black Mt, 24 Dec 1952. +. Black Mt, 31 Jan
1968. 3c, Black Mt, 12 Dec 1967, 39, Black Mt, 21 Dec
1969. c, Canberra, 18 Dec 1959. d, Canberra, 12 Jan
1965. d, Canberra, 6 Jan 1964. g, Canberra, 30 Dec
1950. d, Canberra, 5 Dec 1961. 28, Canberra, 20 Dec
1950. 29, Canberra. Dec 1949, g, Canberra, 26 Dec
1978. 9, Canberra, | Jan 1950. 9, Canberra, 4 Mar
1966. 2d, Canberra, 11 Jan 1972. 1 juv., Canberra, 1
May 1983. c, Cotter Dam, Dec 1930, 9, Pierces Creek,
21 Jan 1950.9, 8 km NNE of Tharwa, 17 Dec 1956 (all
ANIC).

Old. <. Hughenden, 21 Mar 1930. 7. Windorah, 5
Nov 1963 (both AM). 39,4 km SW of Birdsville, 4 Dec
1974. d, Blackall, 10 Apr 1972. 1 juv.. 48 km N of
Blackall, 27 Dec 1961.2, 23°02’S, 139°18’E, 62 km SW
of Boulia. 16 Oct 1978. d, 7 km N of Cunnamulla, 29
Jan 1971. 2 juv., 23 km W of Cunnamulla, 13 Jan
1965. d, Gilruth Plains Stn. E of Cunnamulla, 8 Jan
1964. d, Longreach, 20-31 Jan 1972. d, 2141'S,
14030'E. Selwyn Mine, 160 km SE of Mt Isa, 7 Oct
1991. d, St George, 19 Jan 1969. l Juy., 50 km NNE of
Wyandra, 25 Dec 1961. 9, 20°18’S, 139"03'E, 20 km
ESE of Yelvertoft HS, nr Camooweal, 9 Apr 1976 (all
ANIC). o, 25°48°S, 146735'E, Augathella, 18 Jan 1993
(NMV), I juv.. 17 km N of Ardmore Stn, 105 km SW
of Mt Isa, 27 Sep 1977. 9, Blackburn, 18 Sp 1905. 29,
Cunnamulla (all SAM).

NSW. c. Broken Hill, Mar 1943. 3, Calumet, 43 km
NE of Binnaway, 23 Dec 1932. 1 juv., Mootwingie,
142 km NE of Broken Hill, 15 Sep 1955, d, Stephens
Ck. 24 km ENE of Broken Hill, 29 Jan 1976, 20, War-
rumbungle Nat Pk, 13-18 Dec 1977 (all AM). 9, Bar-
rington Tops. 1945-6. 39, 18 km WNW of Bourke. 9
Feb 1972.8, Brewarrina, 1914. g. 6 km W of Cobar, 10
Dec 1971. l juv., 42 km S of Condobolin, LI Dec 1962.
d. Cowra, 15 Feb 1967. 9, Deniliquin. 2, Deniliquin, |

Aug 1961. 1 juv., 22 km SW of Dubbo, 21 Dec 1961. 9,
15 km W of Dubbo, 22 Dec 1961. d, Forbes, 14-15
Nov 1979, 9, Henty, 18 Jan 1962. | juv., 32"30'S,
140°20’E, Kinchega Nat Pk, Jan 1986. 9, Lake Cowal,
15 Dec 1970. 3, 30 km S of Lightning Ridge, 27 Dec
1978. 1 juv., 14 km WNW of Monia Gap, nr Hillston,
23 Mar 1972. 9, Mountain Gap, nr Hillston, 25 Oct
1971. 4, 12 km NNE of Parkes, 21 Dec 1961, l juv.,
Peak Hill, 14 Sep 1954. 3, 49, 1 juv., 5 km SSE of
Reefton, 20 Dec 1961. g, Tibooburra, 16 Nov 1949. 9,
7 km NW of Trangie, 14 Jan 1962. 20, Trangie Exp.
Stn., 6 km NW of Trangie, 13-21 Sep 1978. 1 juv., 87
km NE of Wentworth, 6 Nov 1962. 9, Yass, 28 Dec
1928. d, 30 km WNW of Wentworth, | Jan 1989. d,
Wittabrenna Ck, N of Tibooburra, 11 Nov 1971 (all
ANIC). o, Angora Stn, via Booligal, 20 Mar 1965. 1
juv., Broken Hill, Feb 1927. c, Broken Hill, 11 Jul
1904. ¢, Menindee Lakes Caravan Park, 26 Dec 1973.
(all SAM).

NT. 9, nr Tobermorey, Sep 1930 (AM). g. Alice
Springs, 5 Oct 1955. d, Alice Springs, 23 Oct 1955. 76,
36, 23418, 133752 E, Alice Springs, 3-8 Nov 1988. d,
23'4]'S, 133°52’E, Alice Springs, 27-28 Oct 1988. 4d,
23°41/S, 134915'E, 39 km E of Alice Springs, 25-26
Sep 1978. d, 23 4l'S, 134915'E, 39 km E of Alice
Springs, 5 Oct 1978. 4d, 19°24’S, 135°58’E, 15 km SW
of Alroy Downs HS, 10 Apr 1976. 1 juv., 25707'S,
135*30'E, 38 km NNE of Andado HS, 28 Sep 1972. g,
9, 257175, 130757 E, 10 km WNW of Ayers Rock, 6
Nov 1980. g, 21°40’S 133'45'E, 21 km SW of Barrow
Creek. 12 Oct 1972. d, Barrow Creek Telegraph Stn, 20
Mar 1955, d, 24°36’S, 133 12'E, 7 km SW of Henbury
HS, 2 Oct 1972, 48, 22'18'S. 137°52’E, Illugnarra
Waterhole, 90 km SSW of Uradangi, 15 Oct 1978. 9,
Junction Waterhole, Todd River, 10 Oct 1978. d,
25*50'S, 133°18’E, Kulgera, 17 Jan 1982. 9, Mt Ever-
ard, Jindalce Site, 45 km NW of Alice Springs, 27 Feb
1987, 60, 26°00’S, 131°25’E, 26 km WNW of Mulga
Park HS, 18 Jan 1982. d, | juv., 24°11’S, 134901'E,
Ooraminna Camp, 56 km SE of Alice Springs, 23 Sep
1987. g, 23°00’S, 136°08’E, Plenty River, 245 km ENE
of Alice Springs, 14 Oct 1978, d, 23'42'S, 134°12’E,
Ross River Rd, 33 km E of Alice Springs, 3 Nov 1988.
32, 20°53’S, 130°25’E, Sangsters Lake, 38 km SE of
The Granites Mine, Tanami Desert, 31 Oct 1988. c,
24*11'S, 133°31’E, Stuart Hwy, 65 km SSW of Alice
Springs, 4 Nov 1988. d, 9, 1 juv., 19*59'S, 129°427E,
Tanami Borehole, Jul-Sep 1971. l juv., Tennant
Creek, 5 Nov 1965, d' 9, 20751'S, 130°16’E, 35 km S of
The Granites Mine, Tanami Desert, 29 Oct-2 Nov
1988,26, 25°21°S, 131°03’E, Uluru Motel, Ayers Rock,
4, 6 Nov 1980 (all ANIC). 1 juv., 4 km W of Alice
Springs, 24 Sep 1987. g, 31 km S of Alice Springs, 29
Sep 1987. 2 juv., Ayers Rock, Sep 1948. 1 juv., Glen
Helen Gorge, Finke River, 14 Oct 1987. 9, Kings Can-
yon Rd, 30 km W of Stuart Hwy, 1 Oct 1987. 1 juv.,
20°52'S, 130°16’E, SE of The Granites, Tanami
Desert, 27-31 Oct 1987 (all NMV). 7. Alice Springs,
Horn Expd., 1894, d, Alice Springs, June 1894, 9,
Petermann Range. d, Tennant Creek (all SAM). d,
Prouse Gap, 150 km N of Alice Springs, 4 Oct 1969
(WAM).

33

REVISION OF THE ARCHIMANTINI (MANTIDAE)

Sè

ke la seats aga A a
ji Dan

81

re

Figures 81-82, Archimantis sobrina. 81, male; 82, female. Scale = 10 mm.

34 G. A. MILLEDGE

Figures 83-92, Archimantis sobrina. 83, male head; 84, female head; 85, male cercus; 86, female cercus; 87,
ootheca, lateral; 88, ootheca, dorsal; 89, ootheca, transverse; 90, male genitalia, dorsal; 91, male genitalia, ven-
tral; 92, male genitalia, right lateral. Scale = 2 mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE)

SA. 9, Alarinna, Musgrave Ranges. d, Arkaroola, Flin-
ders Ranges, 22 Jan 1976, d, Innamincka, Coopers
Creek, 24 Jan 1976 (all AM). 29, 26°08’S, 130°54’E, 25
km NW of Amata, Musgrave Ranges, 19 Jan 1982. l
juv., 5 km SE of Goolwa, 19 Jun 1951. 9, 23 km NE of
Hawker, 24 Jan 1959. d, 33°07’S, 136°38’E, 20 km NE
of Kimba, 5 Oct 1982. d, 12 km S of Kimpton, 7 Jan
1989. 1 juv., 38 km SSE of Marree, 24 Jul 1956. I juv.,
27km NNW of Padthaway, 13 Mar 1953, 9, 10km E of
Penong, 6 Jan 1972. 9, 1 juv., 35°24’S, 140°48’E, 18 km
SSW of Pinnaroo, 25 Oct 1983. 1 juv., 16 km ESE of
Poochera, 30 Oct 1969. 1 juv., 27 km NW of Port
Augusta, 12 Jul 1956, 2 juv., 13 km SW of Quorn, 27
Jul 1956. c, Roseworthy Agricultural College, nr
Gawler, Dec 1972. d, 32°20’S, 138°36’E, Slippery Dip
Camp, Brachina Creek, 9 Nov 1987. 1 juv., Urrbrae,
Aug 1946. d, Urrbrae, Jun 1956. 8, 6 km S of Wintinna
HS, 5 Feb 1984 (all ANIC). I juv., Abminga RS, 22 Sep
1987. 1 juv., Alberrie Creek RS, 40 km W of Marree,
17 Sep 1987. o, edge of Lake Frome, 16 May 1977. |
juv., 31 km NW of William Creek, 18 Sep 1987, 2 juv.,
0.5 km W of Yappala HS, N of Hawker, 16 Sep 1987
(all NMV). 125, 4 juv., Adelaide. 9, Adelaide, 21 Dec
1892. 9, Adelaide, 23 Feb 1896. 1 juv., Adelaide, 26 Jul
1886. 2, Adelaide, 6 Nov 1959. d, nr Agnes Creek Stn,
II Oct 1977. o, 30°22’S, 137°08’E, Andamooka Stn,
Apr 1981. 49, Anna Creek HS, 6 Dec 1974. 1 juv.,
Arcoona Creek, Gammon Ranges, 17 Sep 1956. 1 juv.,
Birkinhead. d, Black Swamp, 5 Dec 1952. ¢, Brachina
Gorge, Flinders Ranges, 13 Dec 1974. d, Brighton, 12
Nov 1901. d, Brinkworth. 1 juv., Buckleboo. c, Cara-
ppee Hill, Eyre Penninsula, 8 Nov 1964. 9, Ceduna. 9,
Ceduna, Jan 1924, | juv., Clayton Crossing, 29 Jul
1955. 9, Coomandook. d, Coopers Creek Ferry Cross-
ing, 30 Nov 1974. 1 juv., Cortina Stn, Jan 1968. g,
Eden Hills, 28 Jan 1956. o, nr Frasers Hut, Cariewer-
loo Stn, March 1956. 7. Dalhousie Springs, 6 Oct 1980.
g, Finniss Springs HS, 7 Dec 1964. 3, Frome River, nr
Marree, 25 Oct 1956. | juv., Gammon Range Nat Pk.
Arcoona Creek, 5 May 1989, d, Glenunga HS, 4 Dec
1932. 4 juv., Glenunga, 9 Sep 1932. 9, Goodwood, 14
Jan 1964. 9, Goodwood Park, 19 Nov 1888. d, Grange.
g, Hilton, 30 Dec 1956. o, Islington, 15 Nov 1886. g,
Kensington Park, 10 Nov 1980. 1 juv., Lake Arcoona,
3 Jan 1900. g, Lake Callabonna, 1893. o, Lake Mulli-
gan, 1893. 9, Lake Palankarinna, 3 Mar 1972. 26,
Leigh Creek, 18 Nov 1890. d, 28°19’S, 136°16’E, Levi
Creek, 8 km NW of Ely Perry Spring, 7 Dec 1974. o,
Loxton, 30 Dec 1955. 4, Laxton, 26 Nov 1956. 9, Lox-
ton, 26 Dec 1955. d, Lucindale. 1 juv., 29911'8,
134°19’E, 2 km SW of Mabel Creek HS, 28 Oct 1984. o,
Marrec. d, Marree Racecourse, | Dec 1974.2, 32°57’S,
137°24’E, Middleback Stn, 12 Nov 1983. o, 32°57’S,
137724'E, Middleback Stn, 12 Feb 1984. d, Mitcham,
16 Nov 1978. I juv., Mitcham, 23 Mar 1979. o, Mit-
cham, 30 Nov 1979. 3, Mt Davies, 18 Nov 1966. 9, Mt
Lofty Range. 9, Murat Bay. 9, Murray River. 9, Nor-
manville, Nov 1933. 9, Ooldea, 30 Apr 1898. 29, Ool-
dea. d, o. 2 juv., Orroroo, 1942-44. d, Paddington,
Outer Harbour, Nov 1933. d, Parkside. 1 juv., Poo-
chera, 13 Jun 1956. d, 2 juv., Purple Downs, Pimba, 12
Dec 1937. d, Port Lincoln district, 16 Oct 1957. g,

laj
Lu

Quorn, Dec 1893. 9, Thebarton, 2 Jan 1960. g, Unley,
Jan 1883. 9, Warradale. | juv., Wilpena Nat Pk, 10
May 1989. 1 juv., 7 km W of Wilson, 9 May 1956. 26.
Yeelana. 9, Yurgo (all SAM).

Vic. 1 juv., 12 km NNW of Annuello, 5 Nov 1962. d,
Kyabram, 26 Oct 1946. o, Melbourne, 1966. d.
35°04’S, 143°11’E, 11 km W of Piangil, 27 Oct 1973
(all ANIC). o, 34°46’S, 142°31’E, 30 km WNW of
Annuello, 20 Oct 1985. 9, Chelsea, 23 Jan 1918. d,
Cheltenham, 13 Dec 1905. 9, 2.8 km NNW of China-
mans Well, Big Desert, 12 Oct 1982. 9, Downshore Rd,
Elsternwick, 5 Jan 1908. o, Falls Creek Village, 15 Mar
1984.2, 9, Frankston. 3, Gippsland. d, Glenelg River, 7
km NNE of Nelson, 25 Nov 1966. d, Hattah Lakes,
Feb 1983. 39, o, Hattah Lakes Nat Pk, 21 Oct 1982. g,
34^54'S, 142*08'E, 19 km SW of Hattah, 17 Oct 1985.
l juv., 3446'S, 14220'E, 5 km E of Hattah, 22 Oct
1985. d, Laverton, Sep 1977. d, 59, Lurg, E of Benalla,
Mar 1977. 3,9, 3 juv., 0.2 km N of Milmead Rock, Big
Desert, 20 Mar 1987, 9, Mordialloc. 9, 35?24'S,
141°09’E, 15.9 km SSW of Murrayville, 19 Feb 1987.
d, Oakleigh, 1902. 9, Redcliffs, 19 Mar 1928. 9, 8.1 km
N of Round Swamp, Big Desert, 28 Mar 1985. o, San-
dringham, 1892. g, Tallangatta, 24 May 1904. 1 juv.,
Whipstick, 11 Apr 1948. 9, Old Tom Rd, Whipstick
Forest, N of Bendigo, 17 Feb 1985 (all NMV).

WA. c, 9, Bornholme, 8 Dec 1921. 28, Karratha, 25
Oct 1978. g, 50 km SW of Sanfire Flat, bn Broome and
Port Hedland, 29 Oct 1978 (all AM). d, Balgo Hills, 16
Oct 1985. I juv., Bohemia Ridge, E of Christmas
Creek HS, 22 Oct 1969. o, Broge Hill, 160 km S of Halls
Creek, 25 Sep 1985. 1 juv., 15 km SW of Cocklebiddy,
3 Nov 1969. 1 juv., 32°08’S, 126°18’E, 23 km ESE of
Cocklebiddy, 25 Oct 1977. d, 30°09’S, 115°07’E,
Cockleshell Gully, 19 km NNE of Jurien, 25 Oct 1984,
l juv., 3128'S, 120°50’E, 67 km SSW of Coolgardie, 6
Nov 1969. o, Cutler Rd, Jandakot, Mar-Apr 1982. d,
26°01’S, 113°35’E, 12 km SSE of Denham, 22 Oct
1984. 1 juv., 29°20’S, 115*01'E, 12 km SE of Dongara,
11 Sep 1981. d, 29°16’S, 114°55’E, 3 km SSW of Don-
gara, 15 Oct 1970. 9, Eneabba, 7 Oct 1981. d, 9, 1 juv.,
32°08’S, 126°17’E, Eyre Tower, Microwave Stn, 22 km
ESE of Cocklebiddy, 7 Oct 1982. d, 29, 33°42/S.
123'26'E, Fisheries Rd, 2 km ESE of Price Hill, Cape
Arid Nat Pk. 15 Oct 1982. 29, 27*50'S, 114°43’E, Four
Mile Pool, Murchison River, 23 Oct 1984. 1 juv., Fre-
mantle, 12 Aug 1934. | juv., 28°47’S, 114°34’E, Ger-
aldton, | May 1971. 2 juv., 27°49’S, 114°11'E, nr
Grandstand Rock, 12 km SE of Kalbarri, Kalbarri Nat
Pk, 19 Oct 1984. o, 31^53'S, 116*05'E, John Forrest
Nat Pk, Darling Ranges, 28 Oct 1985. d, 33/238,
123°24’E, Junana Rock. 9 km NW of Mt Ragged, 26
Oct 1977. d, 30°13’S, 115°18’E, 28 km ENE of Jurien,
26 Oct 1984. o, 30°54’S, 121°32’E, 21 km SSE of Kal-
goorlie, 17 Feb 1978. 9, 14 km W of Kitchener, 15 Jan
1974. 0, Lake Douglas, 12 km SW of Kalgoorlie, 13 Jan
1989. 9, 27734'S, 114°26°E, Loop Rd, 30 km ENE of
Kalbarri, Kalbarri Nat Pk, 17 Oct 1984. 2o, 2 juv.,
27739'S, L14*17'E, Loop Rd, 14 km ENE of Kalbarri,
Kalbarri Nat Pk, 20 Oct 1984. o, 21*34'S, 117903/E, 3
km WNW of Millstream HS, 23 Apr 1971. d, 21*35'S,
117°12’E, 15 km E of Millstream HS, 20 Oct 1970. d,

36 G. A. MILLEDGE

2210'S, 11502/E, 20 km S of Minderoo HS, 17 Oct
1970, 1 juv., 33°41’S, 1237 10'E, 8 km NW of Mt Bar-
ing. SW of Mt Ragged, 16 Oct 1982. 9, 3501'S,
117°54’E, Mt Clarence, Albany, 13 Feb 1980. 9, Mt
Hawthorn, 10 Nov 1953. d, 39, 1 juv.. 33°59’S,
122°08’E, | km NNW of Mt Le Grand, Cape Le Grand
Nat Pk, 11 Oct 1982, 49, 5 km SW of Mt Ragged, 12
Nov 1969. 9, 1 juv., 33°56’S, 119*59'E, Mylies Beach,
Fitzgerald Nat Pk, 31 Oct 1984. d, 25°11’S, 113°507E,
nr New Beach, 40 km SSE of Carnarvon, 21 Oct 1984.
| juv., Ninghan Stn, 20 Jul 1963. 1 juv., 327098,
12143 E, 7 km WNW of Norseman, 13 Oct 1984. |
juv., 34^ 14'S, 115712 E, Pinnacles/Cervantes, 2-5 Mar
1982. 1 juv., 33°37’S, 120°24’E, 33 km E of Ravens-
thorpe, 10 Feb 1980. d, 58 km N of Reid, Nullarbor, 4
Feb 1968. I juv., Rottnest Island, 22 Jun 1975. d, Scad-
den, 21 Oct 1977. g, 9, 347268, 117956/E, Stirling
Range Nat Pk, 50 km SSW of Borden, 12 Jan 1980. o.
33518, 12301'E, Thomas River, 23 km WNW of Mt
Arid, Cape Arid Nat Pk, 13 Oct 1982. d, l juv.,
32°35’S, 117°19’E, Tutanning Nature Reserve, 22 km
SE of Pingelly, 16-17 Sep 1981. 9, 23 13'S, 114726'E,
13 km WSW of Winning HS, 29 Apr 1971. d, 43 km
NW of Wittenoom, 22 Apr 1963 (all ANIC). 9, Mid-
dalya, 17 Aug 1904. d, 9, Wembley, 18 Nov 1948 (all
NMV). 9, 1 juv., Beverley. 9, Mt Giles Weather
Station, Nov 1966. 9, Mullewa. 20, Nanga, neck of
Peron Penninsula, 3 Aug 1972. 9, Roebourne, I Mar
1899. 3, 17 km SW of Three Springs, 7 Nov 1968 (all
SAM). d, Applecross, 28 Nov 1963. | juv.. Bickley, 2
Sep 1965. d, Deepdene, Karridale, 10 Aug 1962. 1 juv.,
Dumbleyung, 31 Mar 1963. 9, Dumbleyung, 19 Dec
1963. 1 juv.. Kalamunda, 7 Dec 1962. 1 juv., Kelm-
scott, 14 Apr 1971. I juv., Lake Brown. 9. Laverton, 26
Nov 1964. | juv., 3301/8, 120°44’E, McDermid Rock,
27 Sep-3 Oct 1978. c, 28 km NE of Millstream, 26 May
1965. c, Mt Hawthorn. 2 juv.. 30°59’S, 119°07’E, 13.6
km SSW of Mt Jackson, 5-11 Sep 1979.9, Mt Pleasant,
26 Oct 1963. o, Mt Pleasant, | Dec 1963. &, Mt Tom
Price, Mar 1966. 26, Mullewa, 1914. 9, Murchison dis-
trict. 9, Murdock, 27 Jan 1978. 9, Nedlands, Nov 1969.
9, Peringillup, Dec 1913. <. Perth, g, Perth, Oct 1914.
d, South Perth. 26 Dec 1972. 9, Walyhomoning Rock,
May 1972. ¢, Warburton Ranges, Dec 1962. c, Wem-
bley Downs, | Nov 1969. o, West Perth. 9, West
Subiaco. 9, Yandil (all WAM).

Diagnosis. Eyes with lateral margins rounded,
lateral margins of pronotum not strongly
toothed, fore coxa without large teeth on
anterior margin or distinctly patterned inner
face, tegmina of female covering first two
abdominal tergites, apical segment of cercus
with pointed to rounded margin, pa of male
genitalia compact.

Description. Body of moderate to large size,
colour brown, yellow brown or green. Apical
margin of head broadly arched, eyes slightly pro-
tuberent anteriorly with lateral margins
rounded, frontal shield with distinct subanten-

nal ridge. Prothorax elongate, rather broad in
larger specimens, supracoxal expansion slight
but distinct; lateral margins finely denticulate in
prozona and smooth in metazona of male, simi-
lar in female except anterior third of metazona
finely denticulate; prozona with small scattered
tubercles dorsally, denser more pronounced
tubercles ventrally; metazona smooth with dis-
tinct mid dorsal keel. Fore coxa with 6-7 very
small teeth on anterior margin, females with
smaller tubercles between teeth particularly
proximally; inner face without distinct pattern-
ing, pale proximally. Fore femur with all spines
tipped blackish brown. Fore tibia with 13-16
inner and 10-11 outer spines, all tipped blackish
brown. Mid and hind femora without genicular
spine. Tegmina and hindwings of male reaching
to caudal margin of eighth abdominal segment
to just beyond supraanal plate; tegmen hyaline
except for costal area and costal margin of dis-
coidal area which are opaque and of similar
colour to body above; beneath costal margin of
discoidal area intense orange-brown, costal area
glossy black in proximal fifth, remainder whitish
with black cross veins; discoidal area occasion-
ally with dark spot proximal of stigma dorsally,
venation unpigmented to slightly pigmented.
Tegmina and hind wings of female just reaching
caudal margin of second abdominal tergite; teg-
men of similar colour to male except that opaque
portion of discoidal area more extensive,
occasionally with blackish spot at distal end of
stigma and veins of discoidal area sometimes
darkly pigmented. Hindwings of both sexes with
costal margin orange-brown, remainder hyaline.
Abdomen usually with narrow, whitish mid dor-
sal stripe; anterior margin of abdominal ster-
nites 3-6 coloured black and red, only visible
when abdomen flexed dorsally; cerci (figs 85-
86) elongate, apical segment with pointed to
rounded margin. Male genitalia (figs 90-92)
with dpr of vp strongly curved, broad at base,
narrowing toward tip; anterior part of vl of Iph
broadly rectangular, narrowly produced at junc-
tion with pa; apr swollen medially, tip slightly
mucronate; pa compact and strongly sha-
greened, with two lateral projections, anterior
one knob like, posterior one similar but slightly
longer and more pointed with small tooth at
apex.

Measurements (mm). Body length, ¢ 70-100, 9
70-110. Head width, 3 6.5-10.2, o 9.0-11.4.
Head depth, d 3.8-5.1, 9 5.7-7.4. Pronotum
width, 7 3.5-5.7, 9 5.0-7.9. Pronotum length, d
22-31, 9 22-38. Fore coxa length, & 12-17,

REVISION OF THE ARCHIMANTINI (MANTIDAE) 37

9 13-21. Fore femur length, ¢ 14-20, 9 16-25.
Tegmen length, 9 48-60, o 16-24. Cercus length,
d 7.4-9.5, 9 7.0-11.2.

Immature stages. Nymphs occasionally with
more mottled colour pattern than adults.
Ootheca (figs 87-89) roughly globular oblong,
emergence area often protruding somewhat pos-
teriorly, spongy layer rather narrow so that
ootheca appears smaller than that of A. latistyla,
eggs arranged in W formation, colour whitish or
pale greenish; deposited on the stems of shrubs
and tall grasses, often parasitized by wasps ofthe
genus Podagrion.

Distribution and habits. Occurs over the drier
regions of mainland Australia, absent from the
monsoonal north and the eastern side of the
Great Dividing Range but extends into wetter
regions in the south east and south west (fig.
171). Found on shrubs and tall grasses, rests with
underside uppermost, adults most commonly
collected from October to January.

Remarks. Saussure's types for this species can-
not be located (Balderson, 1984). This is a
widely distributed and variable species. North-
ern specimens become larger and the prothorax
becomes broader, in contrast to that of A. /ati-
styla which becomes more elongate. Specimens
from the southwest are rather small, brown in
colouration, lack the dorsal abdominal stripe
and have the apical segment of the cercus with a
rounded margin. These are probably adap-
tations to a more shrub dominated habitat, simi-
lar though less pronounced characters are found
in specimens from the Big Desert region of Vic-
toria, a predominately shrubby region. The
southwestern form was described as a seperate
species, A. minor, by Giglio-Tos (1917) however
this form grades gradually into the more typical
form to the north and west and is considered a
mere variant.

Archimantis armata Wood-Mason
Figures 93-101, 172

Archimantis armatus Wood-Mason, 1877: 76.
A. armata Wood-Mason, 1878: 584.

Material examined. Holotype female of A. armatus,
North Australia, C. French, 587/1 (ZSIC, in
alcohol).

Other specimens examined (159, 109, 5 juv.). Qld. d,
Clermont, Apr 1921. g, Clermont, 7 Feb 1981 (both
AM). I juv.. 20 km W of Arcadia HS, 24 Oct 1967. 1
juv., 25 km N of Bell, 27 Oct 1967. 9. Blackall, 22 Apr
1972. 3, Black River, 20 km N of Townsville, 20 Jul
1990. | juv., Bundaberg, Apr 1971. d, Electra, nr

Bundaberg, 25 Apr 1978. d, Issacs River Crossing,
Dingo-Mt Flora Rd, 24 Jan 1982. d, Mackay, 4May
1971. 1 juv., 25°36’S, 151°44’E, Oaky Creek, 14 km N
of Gayndah, 19 Apr 1982. 2 juv,, Wild Horse Swamp,
25 km SW of Rolleston, 25 Oct 1967 (all ANIC). 9,
Emerald (QM). 29, Brisbane, 12 Oct 1915. d, 19°38’S,
146°50’E, 6.5 km N of Calcium, 18 Feb 1989. c, Roma,
9 Jan 1984. d, Townsville, 6 May 1975. 9, Townsville,
I Jul 1934 (all NMV). 9, Childers. 29, Mitchell, 3-4
Jan 1974. d, Somerset (all SAM). 9, Banana, 10 Dec
1961.9, Banana, 10 Feb 1964, d, Coalstoun Lakes, Feb
1961. c, Dalby, 25 May 1951. 9, Yuleba, Jan 1964 (all
UQ).

NSW. 9, Moree, 26 Dec 1951. d, Pallal Ck., 17 km
Sw of Bingara, 5-6 Jan 1956 (both ANIC).

Diagnosis. Eyes with lateral margins rounded,
pronotum with distinctly toothed lateral mar-
gins which are slightly lamellate anterior to coxal
insertion, fore coxae without large teeth on
anterior margin or distinctive pattern on inner
face, tegmina of female covering first 2%
abdominal tergites, apical segment of cerci with
rounded margin, pa of male genitalia compact
with single pointed projection.

Description. Body large, elongate and rather
robust, brown in colour. Apical margin of head
broadly arched, eyes slightly protuberent anter-
iorly with lateral margins rounded, frontal
shield with distinct subantennal ridge. Pro-
thorax elongate, slight but distinct supra coxal
expansion, lateral margins distinctly though
rather finely toothed and slightly lamellate
anterior to coxal insertion; prozona with few
scattered tubercles above, more numerous and
much more pronounced tubercles beneath
which are often paler than surrounding area;
ventral area between coxal insertion creamish
surrounded by mauvish grey; metazona with
distinct mid dorsal keel. Fore coxa with 6-8
small teeth on anterior edge, inner face not dis-
tinctly patterned. Fore femur with all spines
tipped blackish brown, shallow pit on ventral
face near fourth discoidal spine for reception of
terminal outer tibial spine. Fore tibia with 15—
17 inner spines and 10-13 outer spines, all
tipped blackish brown. Mid and hind femora
without genicular spine. Wings of male reaching
to about caudal margin of abdominal segment
seven. Tegmen hyaline except for costal area and
costal margin of discoidal area which are opaque
brown above; below costal area black in proxi-
mal fifth, remainder whitish with black cross
veins, costal margin of discoidal area orange
brown; stigma bounded proximally and distally
by dark spot on dorsal surface. Hindwing with
costal area orange brown, remainder hyaline.

G. A. MILLEDGE

38

L

T
IS ĈI L SIS EERE
Ee =.

93

EI

Se I

Figures 93-94, Archimantis armata. 93, male; 94, female. Scale = 10 mm

REVISION OF THE ARCHIMANTINI (MANTIDAE) 39

Figures 95-101, Archimantis armata. 95, male head; 96, female head; 97, female cercus; 98, male cercus; 99, male
genitalia, dorsal; 100, male genitalia, ventral; 101, male genitalia, right lateral. Scale = 2 mm.

40 G. A, MILLEDGE

Wings of female covering first 22 abdominal
tergites. Tegmen similar to male except discoi-
dal area completely opaque brown with veins
rather darker brown. Hindwing with marginal
coloured area slightly more extensive and veins
darker coloured, particularly apically. Abdomen
without distinct pale mid dorsal abdominal
band, anterior margin of abdominal sternites 3-
6 blackish, apical segment of cercus (figs 97-98)
with rounded margin. Male genitalia (figs 99-
101) with dpr of vp very broad at base, strongly
curved, narrow toward tip; anterior section of vl
oflph broadly rectangular, narrowly produced at
junction with pa; apr swollen medially, tip
slightly mucronate; pa compact, surface folded
and strongly shagreened with single large
glabrous tooth projecting posterio-laterally.

Measurements (mm). Body length, 9 85-94, 9
103-112. Head width, S 5.4-8.2, 9 9.2-10.2.
Head depth, 9 4.5-4.6, 9 6.0-7.4. Pronotum
width, g 4.3-5.0, 9 6.0-7.3. Pronotum length, g
28-29, 9 36-37. Fore coxa length, & 14-15, 9 19-
21. Fore femur Length, à 16-17, 9 21-24. Teg-
men length, 9 49, 9 29-30. Cercus length, d 9.5-
11.5, 9 11,0-12.0,

Immature stages. Nymphs have a pale mid dor-
sal stripe but it is not as prominent as in some
other members of the genus. The toothed mar-
gins of the pronotum are not apparent in very
early instars. The ootheca is unknown.

Distribution and habits. Recorded from far
north eastern New South Wales and eastern
Queensland as far north as Townsville (fig. 172).
Nothing is known ofthe habits of this species but
it is probably a shrub dweller.

Remarks. This species is not well represented in
collections considering its distibution covers
some fairly well populated areas and may not be
common in nature.

Archimantis monstrosa Wood-Mason

Figures 102-110, 172

Archimantis monsirosa Wood-Mason, 1878: 583.
Archimantis latizonata Sjóstedt, 1918: 21.

Material examined. Holotype female of A. monstrosa,
Victoria River, North Australia (Northern Territory),
10 Mar 1856, R.Elsey Esq., 57.134 1 (BMNH).
Other specimens examined (209, 3o, 38 juv.). Qld. c,
L7720'S, 14457'E, Emu Ck, 27 km SW of Dimbulah,
25-26 Nov 1981. d, 18°10’S, 140^ 1 5E. Flinders River,
59 km SW of Normanton, | Dec 1981. d, 1725/8,
145°05’E, 15 km W of Irvinebank, 27-28 Nov 1981
(all ANIC). g, 17°13’S, 145^17' E, 5.5 km N of Collins
Weir, W of Atherton, 10 Feb 1989. 20, 17°51/S,
141*08'E, Glenore Pumping Station, Norman River,

13 Jan 1993. | juv., 19°33’S, 140°51’E, Julia Creek
Rd., Dugald River crossing, 15 Jan 1993. d, Karumba,
7 Apr 1989. 9, 18°43’S, 144°38’E, Kennedy Develop-
ment Rd, Spring Creek HS turnoff, 17 Apr 1991. 29,21
km SSW of Mt Garnet, 11 Apr 1991. d, 17938'S,
141*09^E, 0.5 km S of Wills Ck., NE of Normanton, 11
Jan 1993 (all NMV). d, 89, 5 juv., Mornington Island
Mission, 20-21 May 1963 (SAM).

NT. 1 juv., MacArthur River, 18 Jun 1976. d, 9,
Nhulunbuy, Feb 1973. d 9, Nhulunbuy, Mar 1973. 9,
Nhulunbuy, Feb 1974. 1 juv., Port Keats, 20 Jun 1952
(all AM). l juv., 8 km NW of Adelaide River, 10 Jun
1972. d, 16°19’S, 136°05’E, 36 km SW of Borroloola, 4
Nov 1975. 1 juv., 1628/8, 136°08’E, Bukalara Range,
47 km SSW of Borroloola, 23 Apr 1976. d, 12°25’S,
132°59’E, 3km ENE of Cahills Crossing, East Alligator
River, 12 Nov 1972, | juv., 2 km NW of Cahills Cross-
ing, East Alligator River, 14 Sep 1982. 1 juv., 5 km W
of Jabiru, 7-10 Feb 1983. 1 juv., 16°23’S, 129°31’E, 13
km WNW of Kildurk HS, 29 Jul 1969. o, Koongarra,
15 km E of Mt Cahill, 6 Apr 1973. 1 juv., 12°50°S,
132°51’E, 16 km NE of Mt Cahill, 13 Jun 1973. 2 juv.,
15 km SW of Mudginberri HS, 31 Aug 1982. 1 juv.,
Patonga Rd, 12 km WSW of Mt Cahill, 14-15 Sep
1981. d, 11907'S, 132708 E’, Smith Point, Cobourg
Peninsula, 27 Jan 1977. 26, Tindal, 30 Nov 1967. 29,
14°31’S, 132°22’E, Tindal, 13 km ESE of Katherine, 2
Dec 1967. 4, 39, Tindal, 17 Dec 1967, 1 juv.”, 16°24’S,
131902/E, 3 km ENE of Victoria River Downs HS, 2
Jun 1969. 1 juv., 16°12’S, 130°26’E, 70 km WNW of
Victoria River Downs HS, 25 Jun 1968. 1 juv., Bor-
roloola, McArthur River, 16 Jun 1929, o, 11°08’S,
132°09’E, 2 km SE of Smith Point, Cobourg Peninsula,
5 Feb 1977 (all ANIC). 1 juv., Darwin, 7 Aug 1912
(NMV). 29, 4 juv,, Darwin, o, 1 juv., Groote Eylandt. 9,
Northern Territory. 4o, 1 juv., Roper River. d, Staple-
ton (all SAM).

WA. 1 juv., 17-25 km NNE of Bedford Downs HS,
1-8 Jul 1964. 1 juv., 17°19’S, 1227 10'E, 8 km S of Cape
Bertholet, West Kimberley District, 19 Apr 1977. 1
juv., 14°39’S, 126°57’E, Drysdale River, Kimberley
District, 18-21 Aug 1975, 2 juv., 18 km E of Gibb
River HS, Kimberley District, 21 Jun 1979, 1 juv.,
14*17'S, 126°39’E, Kalumburu, 13 Jun 1985. o, Kim-
berley Research Stn, 10 Dec 1951. d, Kimberley
Research Stn, 3 Mar 1962. 1 juv., 14^49'S, 125°42’E,
Mitchell River Falls, Kimberley District, 12 May
1983. 1 juv., 14°37’S, 125°48’E, 8 km SW of Walsh
Point, Admiralty Gulf, 17 May 1983. 9, Wyndham. 9,
Wyndham, 17 Dec 1930 (all ANIC). o, 3 juv., Wotju-
lum, Sep 1955 (WAM).

Diagnosis. Eyes with lateral margins rounded,
pronotum with strongly toothed margins which
are broadly lamellate anterior to coxal inser-
tions, fore coxa without large teeth on anterior
margin or distinctive pattern on inner face, teg-
mina of female covering first 3! tergites of
abdomen, apical segment of cercus with
rounded margin, pa of male genitalia compact
with double pointed projection.

41

REVISION OF THE ARCHIMANTINI (MANTIDAE)

—ryTy==>"
2 A e E E S rie SA

102

N

Figures 102-103, Archimantis monstrosa. 102, male; 103, female. Scale - 2 mm.

42 G. A. MILLEDGE

106

Figures 104-110, Archimantis monstrosa. 104, male head; 105, female head; 106, female cercus; 107, male
cercus; 108, male genitalia, dorsal; 109, male genitalia, ventral; 110, male genitalia, right lateral. Scale = 2
mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE) 43

Description. Body large elongate and robust,
brown in colour. Apical margin of head broadly
arched, eyes slightly protuberent anteriorly with
lateral margins rounded but slightly angular,
frontal shield with distinct subantennal ridge.
Prothorax elongate, marked supracoxal expan-
sion; lateral margins strongly toothed and dis-
tinctly lamellate anteriorly, strongly tuberculate
ventrally anterior to coxal insertion; metazona
with distinct mid dorsal keel; prozona with few
scattered tubercles dorsally, denser more pro-
nounced tubercles ventrally which are paler than
surrounding area. Fore coxa with 6-8 small
teeth on anterior margin, inner face not dis-
tinctly patterned. Fore femur with all spines
blackish. Fore tibia with 10-11 external and 14-
16 internal spines, all tipped blackish brown.
Mid and hind femora without genicular spine.
Wings of male not quite covering abdomen. Teg-
men hyaline except for costal area and costal
margin of discoidal area which are opaque
brown above; below costal margin of discoidal
area orange brown, costal area glossy black in
proximal fifth, remainder whitish with black
cross veins; discoidal area with dark spot dor-
sally at either end of stigma. Hindwing with
costal area orange brown, remainder hyaline.
Wings of female covering first 32 tergites of
abdomen. Tegmen similar to that of male but
discoidal area completely opaque with venation
darkly pigmented. Hindwing with orange brown
area more extensive and venation more darkly
coloured, particularly apically. Abdomen lack-
ing pale mid dorsal stripe, anterior margin of
abdominal sternites 3-6 blackish but only vis-
ible when abdomen flexed. Cercus (figs 106-
107) elongate, apical segment with rounded
margin, distal segments relatively broader than
those of A.armata. Male genitalia (figs 108—110)
with dpr of vph broad at base, strongly curved,
narrow toward tip; medial lobe strongly pro-
duced slightly mucronate; anterior section of vl
oflph broadly rectangular, narrowly produced at
junction with pa; apr swollen medially, tip
slightly mucronate; pa compact, strongly
shagreened with two short tooth like projections,
one directed posteriorly, the other posterio-
laterally.

Measurements (mm). Body length, d 90-97, 9
115-117. Head width, 3 8.8-9.5, o 11.0-11.6.
Head depth, d 4.5-5.6, 9 5.5-6.3. Pronotum
width, 3 7.0-8.4, 9 8.5-9.8. Pronotum length,
d 29-32, o 42-44. Fore coxa length, S 17-18,
9 22-23. Fore femur length, ¢ 19-21, 9 27-29.

Tegmen length, d 50-52, o 40-44. Cercus length,
d 8.7-10.0, 9 10.5-13.0.

Immature stages. Nymphs with pale mid dorsal
abdominal stripe but not as prominent as some
other members of the genus. Ootheca
unknown. i

Distribution and habits. Found across the trop-
ical north of Australia (fig. 172). I have collected
later instars of this species in shrubs and early
instar nymphs in low shrubs and grasses in open
woodland, in northern Queensland.

Remarks. It is possible that this species may be
conspecific with A. armata as the amount of
variation between the two species is no greater
than that found in several other species in this
genus. However at present all specimens exam-
ined can be assigned to one or other species.
Further collecting in the area between Charters
Towers and Mount Surprise in Queensland
should clarify this situation.

Archimantis quinquelobata (Tepper)
Figures 111-125, 169

Fischeria quinquelobata Tepper, 1905: 238.

Rheomantis quinquelobata (Tepper)-Giglio-Tos,
1917: 44,

Archimantis quinquelobata (Tepper)-Tindale, 1923:
441.

Material examined. Syntype female of A. quinquelo-
bata, Yactoo, South Australia, 19 May 1887, M.Craw-
ford. Syntype female of A. quinquelobata, north west
South Australia, Mar-Dec 1903, H.Basedow, I 14067.
Syntype female of A. quinquelobata, Broken Hill
(NSW), 3 Jan 1890, F.J. Burgess (all SAM). See Balder-
son (1984) for discussion of type material.

Other specimens examined (499, 509, 33 juv.). Old.
d, 2624'S, 146°15’E, Charleville, 7 Apr 1976. g,
21°41’S, 140°30’E, Selwyn Mine, 160 km SE of Mt Isa.
16 Oct 1990 (both ANIC). 2 juv., 25°48’S, 146°35’E,
Augathella, 18 Jan 1993. o, Charleville, 15 Mar 1921.
o, 21°13’S, 146°29’E, 140 km S of Charters Towers, 19
Feb 1989. 1 juv., 25°40’S, 146°30’E, Khyber Rd., 14
km N of Augathella, 18 Jan 1993 (all NMV). 9, Cun-
numulla (SAM ).

NSW. d, Bogan River, 40 km E of Coolabah, 8 Jan
1963. o, 23 km NNE of Bourke, 13 Mar 1959. 3c, 18
km WNW of Bourke, 9 Jan 1972. 1 juv., 47 km N of
Bourke, 23 Dec 1961. 9, 30°50’S, 146°33’E, 23 km SSE
of Byrock, 5 Apr 1976. d, 7 km W of Cobar, 20 Feb
1963.9, 6 km W of Cobar, 23 Mar 1972. I juv., 6km W
of Cobar, 10 Dec 1971. ljuv., 7 km W of Cobar, 1 Feb
1964. l juv., 25 km SW of Dubbo, 16 Jan 1961. 1 juv.,
20 km N of Dubbo, 19 Jan 1961. 1 juv., 12 km SE of
Girilambone, 23 Dec 1961. 1 juv., 12 km NW of
Goolma, 14-15 Jan 1961. 1 juv., 8 km W of Mendoo-
ran, 18 Jan 1961. I juv., 5 km SE of Merrygoen, 18 Jan

G. A. MILLEDGE

44

Archimantis quinquelobata. 111, male; 112, female. Scale = 10 mm.

Figures 111-112,

REVISION OF THE ARCHIMANTINI (MANTIDAE) 45

Figures 113-125, Archimantis quinquelobata. 113, male head; 114, female head; 115, male foreleg, inside; 116,
ootheca. lateral; 117, ootheca, dorsal; 118, ootheca, transverse; 1 19, male cercus; 120, female cercus; 121, mid
instar nymph, abdomen lateral; 122, mid instar nymph, mid left femur, dorsal; 123, male genitalia, dorsal; 124,
male genitalia, ventral; 125, male genitalia, right lateral. Scale = 2 mm.

46 G. A. MILLEDGE

1961.9, Trangie, 6 Apr 1956. (all ANIC). 69, 79, 22 km
NNW of Coombah, 29 Jan 1990 (NMV). 23, Broken
Hill, LI Dec 1906. 9, Carinda, nr Orange, Mar 1928. 1
juv., 32 km W of Euabalong, 28 Jan 1969. 9, Menindee
Lakes, 16 May 1969 (all SAM).

NT. g, 235418, 134^ 1 5'E, 39 km E of Alice Springs,
25-26 Sep 1978. d, 23°41’S, 133°52’E, Alice Springs,
3-4 Nov 1988, g, Ayers Rock, 7 Nov 1980. c, Ayers
Rock, 16 Feb 1967, 1 juv.. 12 km SE of Frewena, 23
Aug 1960. d, 23°48’S, 132°21’E, 5 km NE of Gosses
Bluff, 10 May 1969. o, 37 km NNW of Renner Springs,
24 Mar 1955. o, 20°53’S, 130°25’E, Sangsters Lake, 38
km SE of The Granites Mine, Tanami Desert, 31 Oct
1988. d, 25°21’S, 13103 E, Uluru Motel, Ayers Rock,
6 Nov 1980 (all ANIC). d, 55 km NNW of Alice
Springs, 27 Sep 1987. 29, 20752'S, 130°16’E, 10 km
WSW of Sangsters Bore, SE of The Granites, Tanami
Desert, 27-31 Oct 1987 (all NMV). 9, Alice Springs,
Jun 1894. | juv., Palm Creek (both SAM).

SA. 9, Marte, 6 Feb 1888 (AM). d, 26°09’S, 130°35’E,
56 km W of Amata. Musgrave Ranges, 19 Jan 1982. d,
5 km ESE of Bopeechee. | Jan 1969. 1 juv., 31917'S,
131°34’E, 27 km E of Ivy Tank Motel, 31 Oct 1970. 1
juy., Leigh Creek, 29 Sep 1965. 1 juv., Wilpena Pound,
25 Feb 1970 (all ANIC). 9, Overland Railway, 17 km E
of Golden Well. Jan-Mar 1909. 1 juv., 31 km NW of
William Creek, 18 Sep 1987 (both NMV). 1 juv., Cor-
tina Stn. c, Kingoonya. ljuv., 26°04’S, 135°05’E, Lake
Suprise, Simpson Desert. d, Macumba Creek, 22 Nov
1909. 69, Mt Painter, Flinders Range. 9, Murray
River. 9, Ooldea. 9, north South Australia, 18 Feb
1888. d, Tintara. 9, Wynbring (all SAM).

Vic. 9, 24 km NNE of Bendigo, 3 Jan 1987. 39, Hat-
tah Lakes Nat Pk, 10 Apr 1982. 26, 4o, Hattah Lakes
Nat Pk, 8 Apr 1982. 9, nr Murray River. l juv.,
3573285, 14229, Waithe Reserve, 21 Jan 1987 (all
NMV).

WA Australia. d, Balgo Hills, 19 Oct 1985. d, Brogo
Hill. 160 km S of Halls Creek, 14 Oct 1985. I juv., 27
km ENE of Cosmo Newbery Mission, 14 Oct 1970. 9,
27750'S, 114°43’E, Four Mile Pool, Murchison River,
23 Oct 1984, I juv., Gahnda Rockhole, 1-2 Feb 1967.
3.11 km N of Geraldton, 26 Jan 1973. d, 29°36’S,
L15*15'E. Gravel Scrape, 25 km N of Eneabba, 24 Oct
1984. c. 6 km W of Hayes Hill, nr Lake Cowan, 8 Feb
1983. 3, 30°09’S, 115*07'E, 19 km NNE of Jurien, 25
Oct 1984. 1 juv., 5 km NNW of Kalgoorlie, 11 Feb
1975. 3 juv., 27°34’S, 14?26' E, Loop Rd, 30 km ENE of
Kalbarri, Kalbarri Nat Pk, 17 Oct 1984. 4, d, 18 km E
of Meekatharra, 28 Apr 1963. I juv., 10 km W of Mia
Mia HS, nr Exmouth Gulf, 29 Jan 1972. 39, Mt Leon-
ora, nr Leonora, 13 Feb 1981. 2 juv., Mt Leonora, nr
Leonora, 20 Feb 1979. 1 juv., 26°31’S, 119953/E, 4 km
ESE of Mt Russell, NW of Wiluna, 15 Oct 1984, 8, 85
km ENE of Port Hedland, 18 Apr 1963. g, 60 km SE of
Sandstone, 23 Jan 1973. d, 1,5 km SE of Spargoville,
10 Feb 1980. 9, 3 km NW of Spargoville, 17 Feb 1982.
l juv., 28°32’S, 121°00”E, Victory Creek, 50 km NNW
of Leonora, 14 Oct 1984. 1 juv., 21*58'S, 118°03’E, 42
km NW of Wittenoom, 11 Nov 1970. 9, 48 km ESE of
Wittenoom, 25 Apr 1963. 1 juv., 28°26’S, 116°0S’E, 20
km W of Wurarga, NE of Mullewa, 16 Oct 1984 (all

ANIC). l juv., Middalya, 17 Aug 1904 (NMV). 9, Cool-
gardie, 1 May 1896. 29, Fraser Range, Oct 1891 (all
SAM). 9, 27°42’S, 121°37’E, 7.5 km SE of Banjiwarn
Stn, 22-28 Feb 1980. 9, 31712'S, 120°17’E, Boorabbin
Rock, 20-21 Jan 1982, 9, Charles Knob, 500 km NE of
Laverton, 20 Feb 1964. 9, 29°57’S, 121°07’E, 3.8 km
NE of Comet Vale Siding, 7-15 Mar 1979. c, Mt Eger-
ton. 9, Lake Side. 9, Landor Stn. 9, Narrogin. 9, Tardun,
l Jan 1963. d, Towrana Stn (all WAM).

Diagnosis. Lateral margin of eyes rounded, lat-
eral margins of pronotum with only very fine
denticles in anterior part, anterior margin of fore
coxa with four large and one or two smaller teeth
which are black on inner face, tegmina of female
covering first 2⁄2 abdominal tergites, apical seg-
ment of cercus with rounded margin; pa of male
genitalia with two lateral projections, the
anterior one short and flattened, the lower one
elongate unciform.

Description. Body of male slender, of female
more robust, considerable variation in size;
colour grey to brownish, sometimes with lighter
and darker mottlings. Apical margin of head
slightly arched; eyes anteriorly protuberent, with
rounded lateral margins; frontal shield with dis-
tinct subantennal ridge. Prothorax moderately
to extremely elongate, slight but distinct supra-
coxal expansion; lateral margins very finely
tuberculate in prozona and smooth in metazona
of male, more strongly tuberculate in prozona
and finely tuberculate in anterior half of meta-
zona of female; prozona with few small scattered
tubercles above, stronger tubercles below that
are cream coloured surrounded by dark back-
ground; metazona with distinct mid dorsal car-
ina, reddish black ventrally between bases of
fore coxae. Fore coxa (fig. 115) with four large
flat triangular teeth and usually smaller proxi-
mal one, occasionally small distal one, these
teeth coloured black ventrally and contrasting
with pale colouration of rest of inner face of
coxa. Fore femur with all spines tipped blackish
brown. Fore femur with 13-15 inner and 9-10
outer spines, all tipped blackish brown. Mid and
hind femora with or without genicular spine.
Tegmina of male covering first six abdominal
tergites; costal area opaque brownish above,
darker orange brown beneath in proximal fifth
with remainder pale semi opaque; anterior third
of discoidal area opaque brownish above and
darker orange brown beneath with large dark
spot at proximal end and smaller dark spot at
distal end of stigma dorsally, remainder hyaline,
venation slightly to distinctly dark coloured.
Hind wing of male with distal two thirds of

REVISION OF THE ARCHIMANTINI (MANTIDAE) 47

costal area and apical portion of discoidal area
brownish with veins darker, remainder hyaline.
Tegmina of female covering first 21% abdominal
tergites; costal and discoidal areas opaque
brownish or grey above, reddish brown beneath,
often with scattered semi opaque areas in discoi-
dal area; dark patches at ends of stigma exten-
sive, sometimes joining posteriorly, stigma paler
than surrounding area; venation more darkly
coloured. Hindwing of female very reduced,
only about half the length of tegmin; smokey
blackish brown in costal area and anterior
portion of discoidal area, particularly apically,
more orange brown beneath, remainder hyaline.
Abdomen without pale mid dorsal stripe,
anterior margin of sternites 3-6 blackish, pos-
terior margin of sternites 2-5 with fine creamy
yellow border; cerci (figs 119-120) slightly to
distinctly elongate, margin of apical segment
rounded. Male genitalia (figs 123-125) with dps
of vph strongly curved, narrow at base and
gradually narrowing to point distally; anterior
portion of vl of Iph narrow, slightly produced at
junction with pa; apr swollen proximally, tip
simple; pa finely shagreened with narrow projec-
tion at junction with vl and two prominent pro-
jections dextrad, anterior one short and slightly
flattened to spatulate with distal margin finely
serrate, posterior one longer and spiniform,
directed laterally then curving ventrally; small
sclerite in membrane anterior to pa.

Measurements (mm). Body length, à 80-103, e
71-132. Head width, 3 7.2-8.2, 9 8.0-11.4.
Head depth, d 4.3-4.6, 9 5.0-7.4. Pronotum
length, ¢ 23-34, 9 27-47. Pronotum width, l
3.7-4.1, 9 5.4-6.5. Fore coxa length, ¢ 14-16, 9
15-24. Fore femur length, 3 16-18, 9 18-27.
Tegmen length, g 48-52, 9 20-35. Cercus length,
d 4.5-9.0, o 3.9-10.0.

Immature stages. Nymphs often display a small
lobiform expansion apically on posterior edge of
mid and hind femora (fig. 122) and single, small
bulbous dorsal protrusions between abdominal
tergites 3-6 (fig. 121). Ootheca (figs 116-118)
rather depressed oblong with fairly thick spongy
layer, cells arranged in a shallow u-shaped for-
mation, colour pale grey, brown or greenish,
deposited near ground on shrubs, grasses, logs or
rocks, when on flat surfaces rather depressed in
appearance, often parasitised by wasps of the
genus Podagrion.

Distribution and habits. Recorded from the
more arid regions of mainland Australia (fig.
169). Usually found in woody shrubs though

occasionally in tall grasses. Adults most com-
monly collected from January to April.

Archimantis brunneriana Saussure
Figures 126-136, 173
Archimantis brunneriana Saussure, 1871: 277.

Material examined (109, 89, 10 juv.). Old. d, Kin-
bombi Falls, nr Goomeri, 19 Dec 1976, d, 20 km N of
Monto, 6 Dec 1979 (both AM). d, Armstrong Creek
Crossing, 13 km NNW of Guthalungra, 26 Jan 1982. 1
juv., 3 km N of Bundaberg, 23-30 Jun 1971. , Burnett
River Crossing, 10 km NE of Eidsvold, 9 Jan 1970. d,
9, Ceratodus, 31 Dec 1955. 4, 17°12’S, 144°34’E, 6 km
SE of Chillagoe, 26 Nov 1981. I juv., 13'56'S,
143°12’E, Coen, 18 Jul 1986. d, 2 juv., 1,7 km W of
Gayndah, 3 Jan 1965. 1 juv., 23°20’S, 150735'E, Mt
Archer (NW summit), Rockhampton, 30 Jun 1980. c,
Mt Garnet, Dec 1960. 1 juv., 257268, 152°56’E, Mt
Tibrogargan, Glasshouse Mountains, 15 Aug 1985. o,
10 km NE of Stanwell, 6 Jan 1965. 9, Tannum Beach,
nr Gladstone, 22-23 Dec 1955. 1 juv., Watalgan
Range, nr Bundaberg, 6 Nov 1971. 9, 20'10'S,
152933/E, Woodgate Nat Pk, 29 km ENE of Childers,
13 Aug 1985 (all ANIC). 1 juv., Bruce Hwy, 4 km S of
Bowen, l Apr 1991. d, 12°44’S, 143°16’E, Claudie
River, Iron Range Nat Pk, 25 km NW of Lockhart
River, 10 Nov 1988. d, 8 km E of Emuford, 30 Dec
1989. 39, 16°48’S, 145°22’E, 22 km NNW of Mareeba,
15 Feb 1989. 2 juv., 44 km SSW of Mundubbera, 31
Mar 1991 (all NMV). 9, Brisbane (QM).

Diagnosis. Lateral margin of eye angular, lateral
margin of pronotum without teeth, anterior
margin of fore coxa without large teeth, inner
face of fore coxa with pattern of small pale spots
on blackish background that peters out distally,
tegmina of female covering first two abdominal
tergites, apical segment of cercus with pointed
apex, pa of male genitalia with tip of projection
directed laterally and slightly anteriorly.

Description. Body slender, grey-brown, brown,
yellow-brown or pale green in colour. Head
strongly anterio-posteriorly compressed, apical
margin broadly arched in female, almost hori-
zontal in male; frontal shield with distinct
subantennal ridge in male, only very slight ridge
in female; eyes with angular lateral margins. Pro-
thorax slender with slight but distinct supra
coxal expansion; lateral margins entire in male,
minutely denticulate for whole length in female
but sparser posteriorly; prozona sparsely granu-
late above, more densely tuberculate below;
metazona with distinct mid dorsal carina, darkly
coloured ventrally between bases of fore coxae.
Fore coxa (fig. 130) with few very small blunt
teeth on anterior edge in male, more numerous
in female with 2 or 3 larger ones proximally;

G. A. MILLEDGE

48

Archimantis brunneriana. 126, male; 127, female. Scale = 10 mm.

-127,

Figures 126

REVISION OF THE ARCHIMANTINI (MANTIDAE) 49

Figures 128-136, Archimantis brunneriana. 128, male head; 129, female head; 130, male foreleg, inside; 131,
ootheca, lateral; 132, male cercus; 133, female cercus; 134, male genitalia, dorsal; 135, male genitalia, ventral;
136, male genitalia, right lateral. Scale - 2 mm.

50 G. A. MILLEDGE

inner face dark brown, becoming black proxi-
mally with numerous irregular pale spots. Fore
femur with all spines tipped black-brown, claw
groove slightly distal of mid point. Fore tibia
with 15-16 inner and 9-11 outer spines, all
more or less tipped black. Inner face of tarsal
segments sometimes blackish. Mid and hind
femora without genicular spine. Wings of male
reaching just beyond fifth abdominal tergite;
tegmen with costal area opaque white except for
posterior proximal third which is black, costal
margin of discoidal area orange-brown, colours
more intense below, remainder of tegmen hyal-
ine. Tegmina of female reaching just beyond
caudal margin of second abdominal tergite;
colouring similar to male except that pale costal
margin very narrow, black colour extends for
three quarters of the length of costal area and
orange-brown band broader, covering about one
third of discoidal area. Hindwings of both sexes
with costal and sub costal areas orange-brown,
remainder hyaline. Abdomen sometimes with
broad pale median dorsal stripe, anterior margin
of sternites 3-7 blackish; apical segment of cer-
cus (figs 132-133) with pointed apex. Male geni-
talia (figs 134-136) with dpr of vph narrow at
base, strongly curved; anterior section of vl of
Iph broadly triangular, broad projection at junc-
tion with pa; apr with medial bulge, distal end
short, tip slightly mucronate; pa with single unci-
nate spiniform projection, lightly shagreened,
slightly sinusoidal distad with sharp angle near
tip.

Measurements (mm). Body length, ¢ 72-78, 9
86-92. Head width, ¢ 6.5-7.3, o 8.4-9.0. Head
depth, 9 3.1-3.6, 9 4.8-5.1. Pronotum length, c
23-26, o 32-35. Pronotum width, d 2.9-3.4, o
4.3-4.8. Fore coxa length, € 13, 9 16-19. Fore
femur length, ¢ 15, o 19-21. Tegmen length, d
36-37, ş 20-22. Cercus length, $ 8.2-8.4, 0 10.9—
12.4.

Immature stages. Nymphs similar in colour to
adults; ootheca (fig. 131) rather small, brownish,
thin spongy layer, eggs in w-formation,
deposited on stems of shrubs and grasses.

Distribution and habits. Recorded from eastern
Queensland, from Iron Range in the north to
Brisbane in the south (fig. 173). Found in grasses
and shrubs in woodland habitats.

Archimantis straminea Sjòstedt
Figures 137-147, 173

Archimantis straminea Sjóstedt, 1918: 17,

Material examined. Lectotype d, here designated,
Noonkanbah, NW Austr., Dec, Mjóberg, (3) 271 82.
Paralectotype 9, here designated, NW Austr., Mjóberg,
(3) 272 82 (both NHRM).

Other specimens examined (206, 69, 21 juv.). Qld. d,
Butcher Creek, 20 km W of Cloncurry, 21 Jan 1977. d,
65 km E of Hughenden, 3 Feb 1981 (both AM). 3 juv.,
8 km E of Dunbar HS, 10 Oct 1965. 9, 25 km SE of Mt
Christison, Hughenden dist., 10-11 Jan 1965. 1 juv., 8
km ENE of Prairie, 10 Jan 1965 (all ANIC), d. 9,
19°14’S, 140°21’E, Burke and Wills Junction, 16 Jan
1993. o, 17°33’S, 141°09’E, Eleven Mile Ck., 16 km
NNE of Normanton, 10 Jan 1993. 4d, o, 1 juv.,
17°33’S, 141°09’E, Eleven Mile Ck., 16 km NNE of
Normanton, 5 Jan 1993, 1 juv., 6 km NE of George-
town, 15 Apr 1991.6, o, 1 juv., 12 km E of Georgetown,
12 Apr 1991. 29, 17°51’S, 141°08’E, Glenore Pumping
Station, Norman River, 13 Jan 1993. 1 juv., 17°38’S,
141°09’E, 9 km NE of Normanton, 4 Jan 1993. 39, 1
juv., 17°36’S, 141°09’E, 11 km NE of Normanton, 6
Jan 1993 (all NMV). ¢, Cloncurry, 8 Apr 1947
(UQ).

NT. d, Alpha Creek, 35 km W of Timber Creek
township, 8 Jan 1986. 1 juv., 16*08'S, 136706 E, 22 km
WSW of Borroloola, 2 Nov 1975. 1 juv., 17 km S of
Dunmarra, 4 Nov 1965. c, Keep River Crossing, Vic-
toria Hwy, 7 Jan 1986. 1 juv., 25 km NNW of Larri-
mah, 20 Nov 1966. 1 juv., 10 km NW of Larrimah, 4
Nov 1965. 1 juv., 30 km NW of Mataranka, 17 Aug
1960. 39, October Creek, Carpentaria Hwy, 180 km E
of Daly Waters, 11 Jan 1986. d, 11*01'S, 136°45’E,
Rimbija Island, Wessel Is, 12 Feb 1977. l juv., 35 km E
of Timber Creek, 29 Oct 1965. 3, Tindal, 6 Dec 1967.
9, 16 km W of West Barnes River Crossing, Victoria
Hwy, 8 Jan 1986 (all ANIC).

WA. 1 juv., 18°22’S, 122953'E, 85 km ESE of
Broome, 16 Aug 1976. 1 juv., 18 km E of Gibb River
HS, Kimberley dist., 21 Jun 1979, 1 juv., 14°13’S,
126°44’E, 13 km NE of Kalumburu, 17 Jun 1985. 29,
Lissadel Stn, nr Kununurra, Feb 1982. | juv.,
McSpeery Gap, Napier Range, SE of Kimberley
Downs HS, 27 Oct 1969. 1 juv., Wyndham East, 27
Oct 1965 (all ANIC),

Diagnosis. Lateral margin of eye distinctly angu-
lar, lateral margin of pronotum without teeth,
anterior margin of fore coxa without large teeth,
inner face of fore coxa with pattern of large pale
spots on blackish background which extends dis-
tally, tegmina of female covering only first
abdominal tergite, apical segment of cercus with
pointed apex, pa of male genitalia directed
laterally then caudally.

Description. Body moderately large, elongate
and slender; pale brown, yellow brown or green
in colour. Head strongly anterio-posteriorly
compressed, apical margin only slightly arched,
eyes with lateral margins distinctly angled,
frontal shield lacking subantennal ridge. Protho-
rax elongate and slender with slight but distinct

51

REVISION OF THE ARCHIMANTINI (MANTIDAE)

A

Figures 137-138, Archimantis straminea. 137, male; 138, female. Scale - 10 mm

LA
N

G. A. MILLEDGE

Figures 139-147, Archimantis straminea. 139, male head; 140, female head; 141, male foreleg, inside; 142, male
cercus; 143, female cercus; 144, ootheca, lateral; 145, male genitalia, dorsal; 146, male genitalia, ventral; 147,
male genitalia, right lateral. Scale = 2 mm.

REVISION OF THE ARCHIMANTINI (MANTIDAE) 53

supracoxal expansion; lateral margins entire in
male , virtually so in female; prozona sparsely
granulate above, more densely tuberculate
below; metazona with distinct mid dorsal keel,
darkly coloured between bases of fore coxae.
Fore coxa (fig. 141) with numerous small
tubercles on anterior edge; inner face dark
brown to blackish with pale spots which become
larger distally. Fore femur with all spines tipped
blackish brown, claw groove slightly distal of
mid point. Fore tibia with 14-16 inner and 10-
11 outer spines, all tipped blackish. Mid and
hind femora without genicular spine. Wings of
male reaching just beyond fifth abdominal ter-
gite; tegmen with costal area opaque white
except for proximal third which has posterior
black band, costal margin of discoidal area
orange-brown, colours more intense beneath,
remainder hyaline. Tegmina of female only
covering first abdominal tergite; colour of costal
area similar to male except inner black band
broader, extending almost to apex; discoidal
area completely opaque, pale pinkish above,
intense rose colour below. Hindwings of both
sexes with costal area orange-brown, remainder
hyaline, those of female very reduced. Abdomen
with broad, pale mid dorsal stripe, anterior mar-
gin of sternites 3-7 red and blackish; cerci (figs
142-143) very elongate, apical segment with
pointed apex. Male genitalia (figs 145-147) with
dpr narrow at base, with abrupt curve at mid
point; anterior section of vl of Iph broad with
rounded margins; apr with medial bulge, distal
end short, tip slightly mucronate; pa with single,
long spiniform projection, directed caudally,
laterally then dorso-caudally.

Measurements (mm). Body length, ¢ 78-91, 9
100-108. Head width, d 8.5-9.0, 9 10.0-10.8.
Head depth, d 4.0-5.0, 9 5.0-5.5. Pronotum
length, d 28.0-31.5, 9 39.0-42.0. Pronotum
width, d 3.8-4.0, 9 4.5-4.8. Fore coxa length, d
13.8-15.5, o 18.5-19.5. Fore femur length, d
15.5-18.0, 9 21.5-23.0. Tegmen length, ¢ 38.0-
41.0, o 14.0-14.5. Cercus length, ¢ 12.0-15.4, 9
16.5-18.5.

Immature stages. Nymphs similar in appearence
to adults. Ootheca (fig. 144) similar in appear-
ance to that of A.brunneriana but pale greenish
in colour with a slightly thicker spongy layer.

Distribution and habits. Found across the trop-
ical north of Australia, west of the Dividing
Range (fig. 173). I have collected this species in
long grasses in open woodland country in north
Queensland.

Archimantis gracilis sp. nov.
Figures 148-157, 174

Material examined (7d, 49, 5 juv.). Holotype: c.
12°50’S, 132^5I'E, 16 km E of Mt Cahill, Northern
Territory, 7 Mar 1973, K.H.L.Key (ANIC).

Paratypes. NT. d, 29, 127258, 132°58’E, 1 km N of
Cahills Crossing, East Alligator River, 30 May 1973,
K.H.L.Key. 9, 5 km NNW of Cahills Crossing, East
Alligator River, 25 May 1973, K.H.L.Key. 9, 12725'S,
132°58’E, 1 km N of Cahills Crossing, East Alligator
River, 29 May 1973, K.H.L.Key. g, 12731'S, 132°54’E,
9 km NE of Mudginbarry HS, 10 May 1973,
M.S.Upton & J.E.Feehan. d. 12°17’S, 133 14'E, 15 km
WSW of Nimbuwah Rock, 1 Jun 1973, K.H.L.Key. 34,
12°22’S, 133°01"E, 6 km WSW of Oenpelli, 30 May
1973, K.H.L.Key (all ANIC).

Other specimens examined. NT. 1 juv,, nr Cahills
Crossing, 18 km SW of Oenpelli, 20 Sep 1981. I juv.,
12*52'S, 132*50'E, 15 km E of Mt Cahill, 12 Jun 1973.
l juv., 12°22’S, 133°01’E, 6 km WSW of Oenpelli, 30
May 1973. 2 juv., 13°19’S, 132'47'E, Twin Falls
Gorge, 52 km S of Mt Cahill, 26 Mar 1982 (all
ANIC).

Diagnosis. Body slender, lateral margins of eyes
rounded, lateral margins of pronotum and
anterior margin of fore coxa not strongly armed,
interior face of fore coxa becoming blackish
proximally but without distinctive colour pat-
tern, tegmina of female covering first 2%
abdominal tergites, apical segment of cercus
with rounded margin, pa of male genitalia with
single elongate process.

Description. Body of moderate size, rather slen-
der in form particularly in the male; colour of
dry specimens yellow-brown to chocolate
brown, females often peppered with darker spots
on abdomen, dosal surface of pronotum and
outer face of fore legs. Head with slightly arched
apical margin, eyes somewhat protuberent
anteriorly with rounded lateral margins, frontal
shield with distinct subantennal ridge. Protho-
rax slender, especially in male, with slight but
distinct supracoxal expansion; prozona lightly
granulate above, more strongly tuberculate
below; metazona with fine mid dorsal keel, dark-
ish between bases of fore coxae. Fore coxa (fig.
152) with number of small tubercles on anterior
edge, more so in female, usually with two more
prominent ones in proximal third; inner face
pale distally, becoming blackish and faintly
tubercled proximally. Fore femur with discoi-
dal, outer, large inner and tips of small inner
spines blackish. Foretibia with 14 inner and 8-
10 outer spines, all tipped blackish brown. Mid
and hind femora usually with small genicular
spine. Wings of male extending just beyond sixth

G. A. MILLEDGE

54

Archimantis gracilis. 148, male; 149, female. Scale = 10 mm.

-149,

Figures 148

REVISION OF THE ARCHIMANTINI (MANTIDAE) 55

153

Figures 150—157, Archimantis gracilis. 150, male head; 151, female head; 152, male foreleg, inside; 153, female
cercus; 154, male cercus; 155, male genitalia, dorsal; 156, male genitalia, ventral; 157, male genitalia, right
lateral. Scale = 2 mm.

56 G. A. MILLEDGE

abdominal tergite. Tegmen of male hyaline
except for costal area which has opaque white
marginal band, followed by black band in proxi-
mal third, followed by brown band at costal
margin of discoidal area and and orange-brown
band in proximal third, these colours more
intense below; stigma obscure. Tegmen of
female extending to about middle of third
abdominal tergite, costal and discoidal areas
completely opaque, costal area dark blackish,
discoidal area dark brown with fuscous reticu-
late pattern more prominent distally and
roughly coinciding with veins; stigma usually
indicated by paler area with blackish spot at
proximal and distal end; colouring more intense
beneath. Hindwing of male with distal two
thirds of costal area flushed brown, remainder
hyaline. Hindwing of female about four fifths as
long as tegmen, costal area and longitudinal
veins brown, rest of wing hyaline except for
smokey arca apically. Abdominal sternites 3-6
with blackish anterior margins; cercus (figs 153-
154) elongate, terminal segment with rounded
apical margin. Male genitalia (figs 155-157)
with dpr of vph narrow at base, strongly
recurved; anterior section of vl of Iph rather
small in area and situated noticeably to the left,
with narrow elongate projection at junction with
pa; apr with only slight medial bulge, extending
well beyond dpr, tip simple; pa with single spin-
iform projection, initially directed posteriorly,
then curving laterally, slightly sinusoidal with
tip directed anterio-laterally, faintly shagreened
except toward tip, with small more strongly sha-
greened ventrally directed secondary projection
near elbow of main projection; apr very short.

Measurements (mm). Body length, 3 77-82, 9
90-98. Head width, & 6.9-7.1, 9 8.5-8.8. Head
depth, d 4.2, 9 5.4-5.7, Pronotum length, d 26-
28, 9 33-35. Pronotum width, d 3.1, o 4.3-4.8.
Fore coxa length, d 13, 9 17-18. Fore femur
length, 9 15, 9 20-21. Tegmen length, ¢ 41-43, Q
23-25. Cercus length, 9 7.3-8.0, o 8.4-9.6.

Immature stages. Dry specimens of nymphs are
more mottled than adults with indications of a
pale mid dorsal abdominal stripe of uneven
width. Ootheca unknown.

Etymology. Specific name derived from the
Latin gracilis meaning slender or thin.

Distribution and habits. Only recorded from a
small area in the East Alligator River region of
the Northern Territory (fig. 174). Habits
unknown but probably a shrub dweller.

Archimantis vittata sp. nov.
Figures 158-165, 174

Material examined. Holotype: d, 11°45’S, 142°35°E,
Heathlands HS, Queensland, 22 Jan 1992, T.Weir
(ANIC).

Paratypes: 9, 11°08°S, 142°29’E, Jardine River,
Cape York Peninsula, Queensland, 12 Oct 1979, M.S.
and B.J. Moulds (AM). d, Dulhunty River Crossing,
Cape York Peninsula, Queensland, 27-28 Sep 1974,
G.B. Monteith (QM).

Diagnosis. Body slender, lateral margins of eyes
rounded, lateral margins of pronotum and
anterior margin of fore coxa not strongly armed,
interior face of fore coxa becoming blackish
proximally but without distinctive colour pat-
tern, tegmen of male with diagonal bands in
discoidal area, terminal segment of cercus with
rounded apical margin, pa of male genitalia
compact.

Description (male only). Body rather small for
genus, elongate, slender, colour of dry specimens
brown with darker speckling on abdomen and
outer face of fore legs. Head with almost straight
apical margin; eyes slightly protruberent anter-
iorly, lateral margins rounded; frontal shield
with distinct subantennal ridge. Prothorax
elongate and very slender, with slight but dis-
tinct supracoxal expansion, lateral margins very
faintly tuberculate in prozona and anterior third
of metazona; prozona faintly granulate above,
more strongly tuberculate below; metazona with
fine mid dorsal keel, dark reddish between bases
of fore coxae. Fore coxa (fig. 160) with few, very
small, blunt teeth on anterior edge; inner face
pale distally, becoming blackish with few very
small pale tubercles proximally. Fore femur with
discoidal, outer, large inner and tips of small
inner spines blackish. Fore tibia with 16-17
inner and 8-9 outer spines, all more or less
tipped blackish brown. Mid and hind femora
without genicular spine. Tegmen (fig. 161)
reaching just beyond abdominal tergite 6; costal
area with opaque white marginal band, narrow-
ing distally and followed by darker band in
proximal third which is brown above and black
beneath; narrow zone between subcosta and
radius dark brown, further band posterior to
radius in proximal third which is cream above
and yellow-orange below; discoidal area with
major veins flushed brown giving banded
appearance except for area posterior to stigma
which, along with remainder of tegmen, is hyal-
ine. Wing with distal half of costal area and zone
between subcosta and radius flushed brown,

57

REVISION OF THE ARCHIMANTINI (MANTIDAE)

158

Figure 158, Archimantis vittata, male. Scale = 10 mm.

58 G. A. MILLEDGE

159

161

AL. LA
N A Oe

A N
Gi == JA e Uo
WIT C Se R Cao
~=

TT
E PC

5
SA
= S,
PES T MEAN (Naza

Figures 159-165, Archimantis vittata, male. 159, head; 160, foreleg, inside; 161, tegmen; 162, cercus; 163,
genitalia, dorsal; 164, genitalia, ventral; 165, genitalia, right lateral. Scale = 2 mm.

remainder hyaline. Cerci (fig. 162) elongate, ter-
minal segment with rounded apical margin.
Male (figs 163-165) genitalia with dpr of vph
compact, mandible like in appearance, with
small tooth at apex, dorsal surface lightly ridged
dextrad; anterior section of vl of Iph with short
broad projection at junction with pa; apr with-
out noticeable median bulge, tip simple; pa com-
pact and strongly shagreened anteriorly, with
short, abruptly curved, uncinate posterior pro-
jection.

Measurements (mm). Body length, 66-67.

Head width, 6.8. Head depth, 3.5. Pronotum
length, 22-23. Pronotum width, 2.8-2.9. Fore
coxa length, 10.8. Fore femur length, 13.5. Teg-
men length, 35-36. Cercus length, 6.2.

Immature stages. Unknown.

Etymology. Specific name from the Latin vitta-
tus, meaning decorated with bands.

Distribution and habits. Known only from three
records near the tip of Cape York Peninsula (fig.
174). Habits unknown but probably a shrub
dweller.

REVISION OF THE ARCHIMANTINI (MANTIDAE) 59

Acknowledgments

I would like to thank Dr T.Kronestedt (NHRM),
Dr E.Matthews (SAM) and Dr J.van Tol
(RNHL) for the loan of type material, Mr J.Bal-
derson (ANIC) for allowing me to examine
Archimantis type material in his care, Mr
E.C.Dahms (QM), Mr T.Houston (WAM), Mr
M.Moulds (AM), Dr D.Rentz (ANIC) and Ms
M.Schneider (UQ) for the loan of specimens.

References

Balderson, J., 1984. Catalogue of Australian Man-
todea. CSIRO Technical Paper No 23.

Beier, M., 1935. Mantodea: Fam. Mantidae: Subfam.
Mantinae. Genera Insectorum 203: 1-146.

Beier, M., 1963. Neue und bemerkenswerte Mantiden
verschiedener Herkunft. Sruttgarter Beitrage zur
Naturkunde 106: 1-11.

Beier, M., 1964. Blattopteroidea. Ordung Mantodea
Burmeister 1838 (Raptoriae Latrielle 1802; Man-
toidea Handlirsch 1903; Mantidae auct.) Bronn's
Klassen und Ordnung des Tierreichs (5) (3) 6: 849—
970. (Geest & Portig : Leipzig).

Giglio-Tos, E., 1912. Mantidi Esotici. V. Mantes,
Tenoderae, Hierodulae et Rhomboderae. Societa
entomologica Italiana, Bollettino 43: 1-167.

Giglio-Tos, E., 1917. Mantidi Esotici. Generi e specie
nouve. Societa entomologica Italiana, Bollettino
48: 43-108.

McCoy, F., 1886. Natural history of Victoria. Prodro-
mus of the zoology of Victoria, or, figures and
descriptions of all classes of the Victorian indigen-
ous animals. Vol. 2, Decade 13: 79-118. Trubner
and Co.: London.

Milledge, G., 1990. Revision of the genus Nesoxypilus
Beier (Mantodea: Amorphoscelidae: Paraoxypili-
nae). Memoirs of the Museum of Victoria 50(2):
347-355.

Paijmans, K., 1975. Explanatory notes and vegetation
map of Papua New Guinea. CSIRO. Land
Research Series No.35.

Rentz, D.C.F., 1985. Monograph of the Tettigoniidae
of Australia. Volume 1, The Tettigoniinae.
CSIRO: Canberra.

Saussure, H. de, 1869. Essai d'un systéme des Man-
tides. Mittheilungen der Schweizer entomolog-
ischen Gesellschaft 3: 49-73.

Saussure, H. de, 1871. Mélanges orthopterologiques.
Supplement au troisième fascicule. Mémoires de
la Société de Physique et d'Histoire Naturelle de
Geneve 21: 239-336.

Saussure, H. de, 1873. Mélanges orthopterologiques.
Fascicule 4. Mémoires de la Société de Physique et
d'Histoire Naturelle de Geneve 23: 1-160.

Serville, J.G.A., 1838. Histoire naturelle des insectes.
Orthoptéres. Libraire Encyclopedique de Roret:
Paris.

Sjòstedt, Y., 1918. Results of Dr E.Mjóberg's Swedish
scientific expeditions to Australia 1910-1913. 17.
Mantidae and Phasmidae. Arkiv for Zoologi
11(19): 1-60.

Tepper, J.G.O., 1905. Insects collected in the north-
western region of South Australia proper by H.
Basedow; with descriptions of new species of
Mantidae and Phasmidae. No.2. Transactions,
Proceedings and Reports of the Royal Society of
South Australia 29: 237-245.

Tindale, N.B., 1923. Review of Australian Mantidae.
Records of the South Australian Museum 2: 425-
457.

Werner, F., 1922. Zur Kenntnis der Manntodeenfauna
der Niederlandischen Kolonien. Zoologische
Mededelingen 7: 115-126.

Wood-Mason, J., 1877. On a small collection of
orthopterous insects of the families Phasmidae
and Mantidae from Australia and New Britain,
with descriptions of four new species. Annals and
Magazine of Natural History 20: 74—77.

Wood-Mason, J., 1878. On new and little known Man-
tidae. Proceedings of the Zoological Society of
London 38: 580-587.

G. A. MILLEDGE

60

onon

T---4

ylomantis baldersoni (0) and Coenomantis kraussiana (m).

Figure 166. Distributions of Austromantis albomarginata (e) and Nullabora flavoguttata (o).
Distributions of Corth

Figure 167.

61

REVISION OF THE ARCHIMANTINI (MANTIDAE)

y variegata (0) and A. papua (e).

Figure 168. Distributions of Austrovate:

lobata (9).

Figure 169. Distribution of Archimantis quinque

G. A. MILLEDGE

Figure 170. Distribution of Archimantis latistyla (9).

Figure 171. Distribution of Archimantis sobrina (m).

REVISION OF THE ARCHIMANTINI (MANTIDAE)

Figure 172. Distributions of Archimantis monstrosa (€) and A. armata (©).
Figure 173. Distributions of Archimantis straminea (9) and A. brunneriana (0).
Figure 174. Distributions of Archimantis gracilis (A) and A. vittata (A).

63

Fa a

=

miti ko n bk aran

— en aj n

Memoirs of the Museum of Victoria 56(1):65-68 (1997)

A NEW SPECIES OF PROGRADUNGULA FORSTER AND GRAY
(ARANEAE: GRADUNGULIDAE) FROM VICTORIA

G. A. MILLEDGE

Museum of Victoria, 71 Victoria Crescent. Abbotsford, Melbourne, Vic. 3067, Australia

Abstract

Milledge. G.A., 1997. A new species of Progradungula Forster and Gray (Araneae; Gradu-
ngulidac) from Victoria. Memoirs of the Museum of Victoria 56(1): 65-68.
Progradungula otwayensis, sp. nov., from the Otway Ranges in Victoria is described and

notes on its biology are given.

Introduction

Progradungula carraiensis Forster and Gray
from north-eastern New South Wales and
Macrogradungula moonya Gray from north-
eastern Queensland are the only two cribellate
species of the primitive araneomorph family
Gradungulidae currently known. P. carraiensis
is only known from the type locality, Carrai Bat
Cave in New South Wales (Gray, 1983),
although Gray (Forster et al., 1987) suggests it
may be slightly more widespread. M. moonya is
known from the Boulder Creek region, near
Tully in Queensland (Forster et al, 1987).
Recent collecting in the Nothofagus forests of
Otway Ranges in Victoria has revealed an
undescribed species of Progradungula occurring
at a number of sites.

Morphological descriptions are of the holo-
type male. For comparative purposes descrip-
tive terminology and form follow Forster et al.
(1987), except for leg spination which follows
Platnick and Shadab (1975). Moran (1985),
when describing Gradungula brindabella (now
placed in Kaiya), noted the asymmetrical distri-
bution of the leg spines in that species and
adopted a simplified notation as a result. As the
species described here displays a similar asym-
metrical spine distribution, numbers and pos-
itions of leg spines given are those of the right
legs only and should be taken as an approxi-
mation rather an absolute. Measurements are in
millimetres. All material is housed in the
Museum of Victoria (NMV).

Progradungula Forster and Gray

Progradungula Forster and Gray. 1979: 1053. Type
species Progradungula carraiensis Forster and Gray.
by original designation.

Diagnosis. See Forster et al. (1987) for diagnoses
of this and other gradungulid genera.

Progradungula otwayensis sp. nov.
Figures 1-3

Material examined. Holotype: d, Victoria, Otway
Ranges, Aire Crossing Track, 0.5 km N of Aire River
crossing, 38'42'S, 143°29’E, 31 Jan 1995, G. Milledge,
K3260 (NMV).

Paratypes: Vic. 3d juvs., l 9 juv. same data as holo-
type, K3261; 2 d juvs., Otway Ranges, Young Creek
Rd, 0.2 km NE of Ciancio Creek crossing, 38°42’S,
143929/E, 31 Jan 1995, G. Milledge, K3262; 3 juvs.,
Otway Ranges, Phillips Track, 0.5 km N of Triplet
Falls, 38 *40'S, 143°29’E, 30 Jan 1995, G., Milledge,
K3263; 3 juvs., as above except 20 Feb 1992, K2473-
5; 1 juv., as above except 17 Mar 1992, K2476; | juv.,
Otway Ranges, Maits Rest, 10 km W of Apollo Bay,
38°45’S, 143°19’E, 20 Feb 1992, G. Milledge, K2477
(all NMV).

Etymology. Specific name refers to the region
where this species has been collected.

Diagnosis. This species can be separated from P.
carraiensis by the presence a single parembolic
process on the male palpal embolus.

Description. As in P. carraiensis except as fol-
lows. Carapace 5.42 long, 3.98 wide. Abdomen
5.40 long, 4.75 wide. Colour: carapace pale fawn
except for orange-brown eye region and pale
grey patch anterior to fovea, chelicerae dark
brown; abdomen light mauvish grey with paler
mid-dorsal stripe on anterior half and 3 pairs of
pale dorsolateral chevrons on posterior half: legs
pale fawn proximally becoming darker brown
distally. Eye sizes and interdistances: AME 0.15,
ALE 0.29, PME 0.25, PLE 0.25, AME-AME
0.18, AME-ALE 0.26, PME-PME 0.26, PME-
PLE 0.31, ALE-PLE 0.08; MOQ length 0.69,
front width 0.48, rear width 0.76. Cheliceral
stridulatory area small, similar to that of Macro-
gradungula (Davies, 1993), also present in juv-
eniles but less well developed. Cheliceral groove
with 4 large promarginal teeth and 3-4 much
smaller retromarginal denticles. Labium 0.69

66 G. A. MILLEDGE

2

1mm

o BEECH FOREST

o LAVERS HILL

Figures 1-3. Right palp of holotype male of Progradungula otwayensis (hairs omitted). 1, prolateral; 2, ventral; 3,
retrolateral.
Figure 4. Distribution of Progradungula otwayensis.

NEW SPECIES OF SPIDER 67

long, 0.85 wide. Sternum 2.75 long, 1.81 wide.
Leg formula 1423. Spination: femora: I d1-0-1,
p3-4-4, r3-3-3, v0-1-0; II d2-2-1, p3-3-4,
r3-2-3, v0-3-0; II dl-1-1, pl-3-4, r3-3-4,
v2-2-0; IV dl1-4-1, pl-3-1, r4-1-3, v1-0-0;

tibiae: I di-0-1, p3-2-2, r2-1-2, vI-1-1; Il
d1-0-1, p2-2-2, rl-1-2, v2-2-3; III dl-1-1,
pl-1-2, r2-1-1, v3-2-2; IV di-1-0, p2-2-1,
rl-1-2,v4-3-1;tarsi1p2-1-0,r2-1-0,v3-3-1:
II p2-2-2, r2- 1-1, v4-4-3; III p3-2-2, r2-1-2,
v3-4-4; IV 3-2-2, r2-1-2, v4-3-4.

I II Ill IV Palp
Femur 10.64 8.07 7.41 $8.55 3.04
Patella 2:37 Abe N52" TOO 0.76
Tibia 1092 8.36 6.56 8.08 2.95
Metatarsus 11.40 9.12 7.60 931 —
Tarsus 2.56 2.28 1.80 2.25 1.80
Total 37.89 30.01 24.89 30.09 8.55

Palpal bulb (figs 1-3) with single parembolic
process on embolus, tip of embolus with mem-
branous area more expansive and indentatation
more pronounced than that shown in the illus-
tration of P. carraiensis by Gray (1983). Palpal
spination: femur: d0-1-2, pO-0-l, r0-1-1;
tibia: pO-2-0.

Distribution and habits. Known from several
localities in the Otway Ranges (fig. 4). So far this
species has only been collected from the vicinity
of Nothofagus cunninghamii trees and later
instars appear dependant on the hollows that are
often found in the bases of older trees for day-
time retreats (although one web has been seen in
the hollow stump of a nearby eucalypt). Web
structure and defensive behaviour are similar to
that described for P.carraiensis (Gray, 1983;
Forster et al., 1987) although the web structure
of P.otwayensis has not been as closely studied.
The catching ladder and supporting web is
sometimes constructed several metres from the
retreat, to which it is attached by a single sturdy
guyline, but more commonly it is constructed in
the vegetation surrounding the base ofthe tree or
in the sucker-like shoots growing from the tree
trunk. It may be attached to the ground but more
often is built amongst the vegetation, up to two
metres above ground. The spiders are only
active at night and assume their catching pos-
ition an hour or so after sunset. The egg case 1s
unknown.

Remarks. As noted in the introduction, the leg
spine count of the holotype for this species is
variable so, although the overall number appears
to be significantly lower than that of P. carraien-
sis, further adult specimens are required before
the usefulness of this character can be ascer-
tained, The lack of an adult female for the
description of this species means it may be mis-
placed in Progradungula. This is because Gray
(Forster et al., 1987) used female characters to
separate Macrogradungula from Progradungula.
as no male of Macrogradungula was available.
However, due to the similarities in dimensions
and form of the body, and of the cheliceral teeth,
of the male of this species and that of P. car-
raiensis, it is reasonable to place it in Prograd-
ungula.

The presence of the cheliceral stridulatory
area in this species and in Macrogradungula, but
apparently not in P. carraiensis, would appear
significant. However, it seems that this feature is
present in the male holotype of P. carraiensis
but was overlooked in the original descrip-
tion (M. Gray, pers. comm.). More specimens
are needed before the relationships between
Macrograndungula and Progradungula can
be clarified.

Although this species has been collected from
a number ofsitesin the Otway Ranges. its appar-
ent dependence on mature N.cunninghamii
trees for refuges may make its survival some-
what precarious unless areas where Myrtle
Beech occurs are preserved.

References

Davies, V.T., 1993. The spinning field and stridulating
apparatus of penultimate male Macrogradungula
moonya (Araneae: Austrochiloidea: Gradunguli-
dae). Memoirs of the Queensland Museum 33(1):
175-178.

Forster, R.R. and Gray, M.R., 1979. Progradungula, a
new cribellate genus of the spider family Gradu-
ngulidae (Araneae). Australian Journal of Zoology
27: 1051-1071.

Forster, R.R., Platnick, N.I, and Gray, M.R., 1987. A
review of the superfamilies Hypochiloidea and
Austrochiloidea (Araneae, Araneomorphae).
Bulletin of the American Museum of Natural His-
tory 185: 1-116.

Gray, M.R., 1983. The male of Progradungula car-
raiensis Forster and Gray (Araneae, Gradunguli-
dae) with observations on web and prey capture,
Proceedings of the Linnean Society of New South
Wales 107(1): 51-58.

68 G. A. MILLEDGE

Moran, R.J., 1985. Gradungula brindabella (Araneae: Platnick, N.I. and Shadab, 1975. A revision of the
Gradungulidae). a new species of hypochyloid spider genus Gnaphosa (Araneae, Gnaphosidae)
spider from eastern Australia. Bulletin of the Brit- in America. Bulletin of the American Museum of
ish Arachnological Society 6: 304-308. Natural History 155: 1-66.

Memoirs of the Museum of Victoria 56(1):69-82 (1997)

SUPPLEMENT TO A REVISION OF AUSTRALIAN MEMBERS OF THE BEE GENUS
HOMALICTUS (COCKERELL) (HYMENOPTERA: HALICTIDAE)

KENNETH WALKER

Department of Entomology, Museum of Victoria,
71 Victoria Crescent, Abbotsford. Vic. 3067, Australia

Abstract

Walker, K., 1997, Supplement to a revision of Australian members of the bee genus Homal-
ictus (Cockerell) (Hymenoptera: Halictidae). Memoirs of the Museum of Victoria 56: 69-82.

Five new species, Homalictus adiazetus, H. pilosignya, H. megagnathus, H. sicarius and
H, verticulus, the males of H. andrewsi (Kirby), H. atrus Walker and H. maitlandi (Cockerell)
and female of H. forrestae are described. The discovery of H. megagnathus extends the
known distribution of H. (Papualictus) Michener, previously New Guinea, into north-east-
ern Australia and requires the subgeneric diagnosis to be revised. Inferred species relation-
Ships, based on newly available character states, are revised.

Introduction

Walker (1986) revised and inferred sister-group
relationships for 39 known species of Australian
Homalictus Cockerell. A supplement to that
revision is now necessary. Five new species are
recognised, including the first Australian species
ofthe subgenus Papualictus Michener and males
of three and the female of one previously known
species are described and figured for the first
time. These species and additional character
states (both sexes: head, mesoscutum and pro-
podeum sculpture characteristics; and male
genitalia) are used to revise inferred species-
group relationships.

Terminology and Abbreviations

Terminology and methods follow Walker (1986)
with these exceptions: the gaster 1s called the
metasoma; the forewing length measurement is
taken from the base of the arcuate basal vein (M)
to the distal-most margin of the third submargi-
nal cell (2nd r-m); and, relative head measure-
ments are standardised to a head width of 100
units allowing them to be expressed as a percent-
age and therefore directly compared between
sexes and species. Sculpture definitions follow
Harris (1979) and punctation density rates are
defined as: dense — interspaces between punc-
tures less than diameter of a puncture; close —
interspaces between punctures equal to diam-
eter of puncture; open — interspaces between
punctures greater than one but less than twice
diameter of puncture; sparse — interspaces
between punctures equal to or greater than twice
diameter of puncture.

Institutions are abbreviated as follows: AM,
Australian Museum, Sydney, New South Wales;
ANIC, Australian National Insect Collection,

69

CSIRO, Canberra, Australian Capital Territory;
BCRI, Biological and Chemical Research Insti-
tute, Rydalmere, Sydney, New South Wales;
BMNH, Natural History Museum, London
(statutory name: British Museum (Natural His-
tory); NMV, Museum of Victoria, Melbourne,
Victoria.

Descriptive abbreviations are as follows:
AOD, antennocular distance; CL, clypeus
length; CW, clypeus width; EW, eye width in
side view; FL, flagellum length; GL, glossa
length; GW, maximum genal width in side view;
HL, head length; HW, head width; IAD, inter-
antennal distance; IOD, interocellar distance;
LID, lower interorbital distance; ML, mandible
length; OAD, ocellantennal distance; OOD,
ocellocular distance; S1-S8, metasomal sterna
1-8; SL, scape length; T1-T5, metasomal terga
1-5; UID, upper interorbital distance.

Homalictus Cockerell

Homalictus Cockerell, 1919: 13. Type species: Hal-
ictus taclobanensis Cockerell, 1915: 488 (by original
designation).

Subgenus Homalictus
Diagnosis. See Walker (1986: 115).

Homalictus (Homalictus) adiazetus sp. nov.

Material examined. Holotype. 9 Qld, Bunya Mts
(26°51’S, 151°34’E), 22 Jan 1938, N. Geary, 2000 ft
(610 m) (AM; Missing right hind tarsal segments.)
Paratype. lo NSW, Minnamurra Falls (34°38’S,
150°43’E), 10 Feb 1962, C.E. Chadwick (BCRI).

Diagnosis. A member of the “sphecodoides”
species-group (see Walker (1986) for species-
group definitions); female with frons striate
above antennal bases, pronotum dorsolateral

70 K. WALKER

angles sharply acute, mesoscutum mesially and
posteriorly densely punctate, parapsidal areas
scabrous, dorsal surface of propodeum striate
with a few interconnectives mesially.

Description of female. (male | unknown)
(measurements of holotype in bold). Body
length: 5.18—5.39 mm; Forewing length: 1.48-
1.51 mm; Head width: 1.47-1.49 mm. Relative
head measurements: HW: 100; HL: 81-82; UID:
58-59; LID: 54-55; AOD: 18-20; IAD: 9-10;
OAD: 33-34; IOD: 16-17; OOD: 15-16; CL:
18-19; GW: 16-17; EW: 24-25; SL: 44-45; FL:
80-82.

Structure. Head broad, inner orbits converging
below, median frontal carina almost absent,
extending just beyond supraclypeal area, eyes
with sparse cover of minute setae. Scape reaches
at least posterior margin of median ocellus. Cly-

peus weakly convex, in side view, anterior half

shining, densely to closely punctate with large,
deep punctures, posterior half dull, covered with
à fine reticulate pattern, sparsely punctate with
small, shallow punctures. Frons striate above

antennal bases to level of posterior margin of

median ocellus, sculpture laterally striate though
weakened to smoothly and sparsely punctate
along inner margin of eyes, lower paraocular
areas smooth, shining and glabrous; vertex
almost smooth, with a few transverse striac.
Pronotum dorsolateral angles sharply acute,
weakly projected. Mesoscutum surface dull,
anterior margin straight, anteriorly sparsely
punctate, mesially and posteriorly densely punc-
tate, parapsidal areas scabrous, punctures con-
tiguous with raised edges. Scutellum shining,
closely to densely punctate, scutellum length
equal to length of dorsal surface of propodeum.
Dorsal surface of propodeum not defined by car-
inae, sculpture broadly striate with a few inter-
connectives mesially, sculpture reaches lateral
margins only. Mesepisternum and metepister-
num smooth, covered with a fine reticulate pat-
tern. Fore basitarsal comb fan shaped, hind
basitibial plate apically rounded, inner hind tib-
ial spur coarsely serrate with apices of at least
five teeth, their bases fused.

Colour. Head, scutellum and propodeum
black, antennal scapes dark brown, flagellum
segments light brown; mesoscutum dull with a
dark blue hue; metasoma red-brown suffused
with dark brown apically; legs red-brown except
femora, trochanters and coxae a darker brown.
Vestiture. Body sparse, head and mesoscutum
with short erect, minutely branched hair, meta-
notum and lateral margins of scutellum densely

hirsute; metasomal TI-T2 glabrous, T3-T4
with sparse hair cover.

Distribution. South-eastern Queensland and
coastal central New South Wales.

Etymology. The epithet “adiazetus” means
unpolished and refers to the sculpture pattern on
the head and mesoscutum.

Remarks. Homalictus adiazetus body colour
patterns are similar to M. megastigmus (Cock-
erell) and M. niveifrons (Cockerell) but the coarse
vertical striae on the frons and the densely punc-
tate mesoscutum differs from both.

Homalictus (Homalictus) andrewsi
(Kirby) comb. nov.

Figures 1-3
Halictus andrewsi Kirby, 1900: 86.

Syntypes. 119, Christmas Island, north part of the
island, January 1898, Andrews (BMNH; not exam-
ined),

Material examined. Christmas I. 529 Ethel Beach
(10°28’S, 105°42’E), 15 Apr 1989, J.C. Cardale (col-
lected in either closed forest or yellow trays); 7o, 20 nr
Grants Well (10°29’S, 105°39’E), 15, 24 and 29 Apr
1989, J.C. Cardale (¢ collected at light); 59, 19 Central
Area Wkshp (10°29’S, 105*38'E), 14-15 Apr 1989, J.C.
Cardale; 19 Lily Beach Road (10°28’S, 105°42’E), 13-
28 Apr, J.C. Cardale, malaise trap/trough. (ANIC).
Diagnosis. A member of the urbanus species-
group; female with frons finely striate, pro-
notum dorsolaterally rounded, mesoscutum
posterior two thirds densely punctate, dorsal
surface of propodeum smooth except with sev-
eral striae on basal half; male with frons striate,
mesoscuium impunctate, genae with conspicu-
ous cover of long, plumose hair.

Description of female. Body length: 5.24-5.78
mm; Forewing length: 1.32-1.41 mm; Head
width: 1.55-1.62 mm. Relative head measure-
ments: HW: 100; HL: 88-91; UID: 48-49; LID:
51-52; AOD: 18-19; IAD: 09-10; OAD: 33-34;
IOD: 15-16; OOD: 09-10; CL: 20-21; GW: 17-
19; EW: 26-27; SL: 45-46; FL: 95-98.

Structure. Head elongate, inner orbits diverging
below, ocellocular distance markedly less than
interocellar distance. Frons finely striate, striae
with interconnectives giving appearance of a
widely spaced reticulate pattern, supraclypeal
area and clypeus dull, covered with minute
reticulate pattern, almost impunctate except for
several weak, sparse, shallow punctures. Prono-
tum dorsolaterally rounded, not well projected.
Mesoscutum and scutellum dull, covered with a

THE BEE GENUS HOMALICTUS 71

conspicuous minutely reticulate pattern,
anterior half of mesoscutum sparsely punctate,
posterior two thirds and scutellum densely
punctate with shallow punctures. Dorsal surface
of propodeum not defined by carinae, surface
dull, smooth except with several striae on basal
half. Metasomal terga impunctate.

Description of male. Body length: 4.69-4.80
mm; Forewing length: 1.19-1.22 mm; Head
width: 1.32-1.34 mm. Relative head measure-
ments: HW: 100; HL: 91-93; UID: 54-55; LID:
40-41; AOD: 13-14; IAD: 11-12; OAD: 31-33;
IOD: 14-15; OOD: 14-15; CL: 21-22; GW: 16-
17; EW: 29-30; SL: 36-37; FL: 114-116.

Structure. Head elongate, inner orbits con-
verging below, not strongly so, scape just reaches
anterior margin of median ocellus. Remainder
of body similar to female except: frons sculpture
distinctly striate, supraclypeal area, clypeus,
mesoscutum and scutellum impunctate, all dull
and covered with a conspicuous reticulate pat-
tern, dorsal surface of propodeum sculpture
restricted to basal margin.

Colour. Head and mesosoma dull metallic
green, head with a golden sheen, metasoma with
dark emerald green, surface with distinctive
sheen; legs with coxae and femora dark green
except mid femora suffused with red-brown,
tibiae and tarsi red-brown and variously suf-
fused with dark brown.

Vestiture. Body sparse, head and mesoscutum
with sparse cover of short, erect, branched hair,
genae with conspicuous cover of long, plumose
hair forming a beard, metasomal sternites with
moderate cover of erect, plumose hair, hair not
forming distinctive pattern.

Genitalia and associated sterna. (figs 1—3).

Distribution. Christmas Island.

Remarks. The male is described and figured here
for the first time. Kirby's description of the
female is adequate for species recognition, how-
ever, the partial redescription presented here
provides characters necessary for comparative
purposes. Homalictus andrewsi is most like H.
urbanus (Smith) but differs in both sexes by the
sculpture on the dorsal of the propodeum
(andrewsi- almost smooth; urbanus- coarsely
ruguloso-striate). The shape of the eyes in the
female (inner orbits diverging below) does not
occur in any other member of the urbanus
species-group. Kirby (1900) described two halic-
tids, Halictus andrewsi and H. binghami (the
latter now placed under Pachyhalictus Cock-
erell; see Michener (1978)), from the collections

made by Mr C. Andrews on Christmas Island in
1898. Ms J. Cardale collected extensively (direct
sweeping, yellow pan, malaise and light traps) on
Christmas Island in 1989 yet the only halictid
species collected was H. andrewsi (pers. comm.
J.C. Cardale). Pachyhalictus binghami may now
be extinct.

Homalictus (Homalictus) atrus Walker
Figures 4-6
Homalictus atrus Walker, 1986: 122.

Material examined. Holotype. 9 Qld, Moses Ck, 4 km
NE Mt Finnigan (15°47’S, 145*17^E), 14-16 Oct 1980,
J.C. Cardale (ANIC).

Other specimens examined: Qld: 139, 199 Mcll-
wraith Range, 30 km E of Coen (13°50’S, 143°17’E), 3
Nov 1988, K. Walker (NMV).

Diagnosis. A member of the “blackburni”
species-group; male with frons impunctate, cly-
peus black, pronotum dorsolateral angles acute,
mesoscutum sparsely to openly punctate, dorsal
surface of propodeum sculpture ruguloso-strio-
late, fore and mid leg tarsal segments flanged
laterally, genal hairs long, fore leg coxae and tro-
chanters and mid and hind coxae with dense
cover of long, plumose hair, lateral margins of
fore tarsi with long, simple hairs, rasp-like sculp-
ture on the apicoventral surface of the genitalia
volsellae.

Description of male. Body length: 4.24-4.47
mm; Forewing length: 1.01-1.06 mm; Head
width- 1.27-1.29 mm. Relative head measure-
ments: HW: 100; HL: 86-88; UID: 58-61; LID:
34-35; AOD: 12-13; IAD: 10-11; OAD: 35-37;
IOD: 18-20; OOD: 14-15; CL: 20-21; GW: 15-
16; EW: 32-33; SL: 28-30; FL: 175-179.

Structure. Head broad, sculpture smooth,
covered with a finely reticulate pattern, frons
and supraclypeal area impunctate, clypeus
sparsely punctate; inner orbits converging
strongly below; scape reaches well short of
anterior margin of median ocellus. Pronotum
dorsolateral angles acute, weakly produced.
Mesoscutum and scutellum dull, microtessel-
late, both sparsely to openly punctate with shal-
low punctures. Dorsal surface of propodeum not
defined by carinae, sculpture ruguloso-striolate,
extends to dorsal rim. Fore and mid leg tarsal
segments flanged laterally.

Colour. Body black except antennal flagella
dark brown; legs brown except tarsi light red-
brown.

Vestiture. Body sparse except genal hairs long,
forming a "beard", mesoventral area and fore

~
to

leg coxae and trochanters with dense cover of
long, plumose hair, mid and hind coxae and tro-
chanters with similar hair, forming a moderate
cover; lateral margins of fore tarsi with long,
simple hairs.

Genitalia and associated sterna. (ĥgs 4-6).

Distribution. Cape York Peninsula and north
Queensland.

Remarks. The male is described and figured here
for the first time. /Zomalictus atrus is most like
H. luteipes (Friese) but differs in both sexes by
colour and sculpture characters (atrus 9: tibiae
black, propodeum openly ruguloso-striolate, d:
clypeus black, propodeum ruguloso-striolate;
luteipes 9: tibiae light red-brown, propodeum
closely ruguloso-striolate, &: clypeus dull white
on lower one third, propodeum weakly strio-
late). Female characters alone inferred a sister-
group relationship of H. atrus with H. luteipes
(Friese), a Papua New Guinea species. However,
with additional male characters and new species
now available, two clades involving five species
scem apparent. Prominent projections on the
apicoventral surface of the genitalia volsellae
provide a synapomorphy for a clade containing
H. luteipes (fig. 4), H. cassiaefloris (Cockerell)
(see Walker, 1986: fig. 19a) and H. eurhodopus
(Cockerell) (see Walker, 1986: fig. 19d). Foreleg
vestiture (fore leg coxae and trochanters with
dense cover of long, plumose hair and lateral
margins of fore tarsi with long, simple hairs) and
the rasp-like sculpture on the apicoventral sur-
face of the genitalia volsellae provide synapo-
morphies for a second clade containing 71. atrus
and /1. pilosignya sp. nov.

Homalictus (Homalictus) forrestae Walker
Homalictus forrestae Walker, 1986: 139,

Material examined. Qld, Heathlands (11°45’S,
142735'E): 29 25 Jul-18 Aug; Id Jun-25 Jul; 29 18
Aug-18 Sep; 19 18 Sep-21 Oct; 1992, P. Zborowski,
J, Cardale, T. Weir, L. Miller and E. Nielsen, ex Mal-
aise trap, (ANIC)

Diagnosis. A member of the "forrestae" species-
group; female with frons weakly striate, prono-
tum dorsolaterally rounded, mesoscutum
mesially closely to densely punctate, openly to
closely punctate in parapsidal areas, dorsal sur-
face of propodeum sculpture coarsely ruguloso-
striate, anterior surface of hind tibia with hairs
nearly simple, bristlelike or with 1 or 2 branches
(no area with abundant, short, erect, plumose
hairs as is usual for the genus), hind tibiae
slender, under surface scarcely concave.

K. WALKER

Description of female. Body length: 4.62-4.85
mm; Forewing length: 0.87-0.96 mm; Head
width: 1.24-1.32 mm. Relative head measure-
ments: HW: 100; HL: 92-94; UID: 64-65; LID:
56-58: AOD: 18-19; IAD: 09-10; OAD: 36-37;
IOD: 18-19; OOD: 18-19; CL: 20-21; GW: 16-
17; EW: 27-28; SL: 40-41; FL: 70-72.

Structure. Head elongate, inner orbits con-
verging strongly below, median frontal carina
present, reaches less than halfway to median
ocellus; eyes with sparse cover of minute setae.
Scape elongate, just reaches anterior margin of
median ocellus. Clypeus with more than two-
thirds projected below lower margins of eyes,
weakly convex in side view, anterior margin
straight, anterior half with several, large punc-
ture, posterior half openly to sparsely punctate
with small, shallow puncture, supraclypeal area
sparelv punctate. Frons and genae weakly stri-
ate, striae reach anterior margin of lateral ocelli,
vertex smooth. Pronotum dorsolaterally
rounded, weakly projected. Mesoscutum surface
dull, covered with microtessellate pattern, anter-
iorly impunctate, mesially closely to densely
punctate, openly to closely punctate in parap-
sidal areas. Scutellum openly to closely punc-
tate, scutellum length equal to length of dorsal
surface of propodeum. Dorsal surface of pro-
podeum not defined by carinae, sculpture
coarsely ruguloso-striate, extends to dorsal rim.
Mesepisternum and metepisternum striate.
Anterior surface of hind tibia with hairs nearly
simple, bristlelike or with 1 or 2 branches (no
area with abundant, short, erect, plumose hairs
as is usual for the genus); hind tibiae slender,
under surface scarcely concave; hind basitibial
plate apically rounded; inner hind tibial spur
coarsely serrate with 2 teeth.

Colour. Head blue-green, though frons above
antennal bases, supralclypeal area and basal half
of clypeus with copper tinge, remainder of cly-
peus black; antennae dark brown except under-
neath of flagellar segments light brown; mesos-
cutum and scutellum green-blue suffused with
copper tinge, propodeum dark blue, metasomal
terga dark brown; legs with coxae, trochanters
and femora dark brown, apical end of femora,
tibiae and tarsi light red-brown.

Vestiture. Body sparse, except tomentose hair
on pronotum mesodorsal and dorsolateral areas,
lateral margins of propodeum with long plumose
hair; ventral metasomal scopae as usual for the
genus, hind leg scopae present though not as
pubescent as is usual for the genus.

THE BEE GENUS HOMALICTUS 73

Figures 1-9. Homalictus spp. male genitalia. H. andrewsi: figs 1-3: 9, Christmas Island, Grants Well, Apr 1989,
J.C. Cardale., ANIC. H. atrus: figs 4-6: 3, Qld, Mcllwriath Range, 30 km E of Coen, 3 Nov 1988, K. Walker,
NMV. H. maitlandi: figs. 7-9: 9, Qld, Claudie River, | mi W of Mt Lamond, 19 Dec 1971, D.K. McAlpine, G.A.
Holloway, D.P. Sands, AM. 1, 4, 7: ventral view (left half) and dorsal view (right half); 2, 5, 8: lateral view; 3, 6, 9:
metasomal S7 and S8. Scale lines (associated with each species) = 0.5 mm.

74 K. WALKER

Distribution. Cape York Peninsula and western
Queensland.

Remarks. The Heathlands population was ident-
ified as H. forrestae after comparison of the
genitalia of the single new male with that of the
male holotype from western Queensland. The
female is described and figured here for the first
time and its character suite confirms the
inferred sister-group relationship with H.
urbanus. Homalictus forrestae is most like H.
urbanus but differs with the female hind tibiae
slender, under surface scarcely concave and
anterior surface with hairs nearly simple, the
male flagellum length (urbanus Fg:UID > 1.5;
Jorrestae Fg: UID < 1.5)and male TI punctation
(urbanus, a sparsely punctures mesially; forres-
tae, closely punctate mesially). The female hind
tibial characters also occur on species of H.
(Papualictus) and H. (Quasilictus). However, I
consider Homalictus proper the correct place-
ment for H. forrestae as the hind tibial characters
are the only characters shared, in both sexes,
with the other two subgenera.

Homalictus (Homalictus) maitlandi (Cockerell)
Figures 7-9

Halictus maitlandi Cockerell, 1910: 223.
Homalictus maitlandi. — Michener, 1965: 180,
338. — Walker, 1986: 145.

Material examined. Holotype 9 Qld. Cairns, Kuranda,
Mar 1902, Turner (BMNH).

Other specimens examined: Qld. (69) same data as
holotype: lo Mossman Gorge (16°29’S, 145°16’E),
Daintree Nat. Park, 26 Oct 1988, K. Walker (NMV);
lọ Mcllwraith Range, c. 510 m (13°45’S, 143°21’E),
22-27 Jul 1977, R.W. Taylor, in rain forest (ANIC);
le, 3 km ENE Mt Tozer (12°44’S, 143°14’E), 28 Jun-4
Jul 1986, J.C. Cardale, ex. Malaise trap (ANIC); 29 11
km ENE Mt Tozer (12°43’S, 143°18’E), 11-16 Jul
1986, J.C. Cardale, ex. ethanol (ANIC); 18 Claudie
River, | mi W Mt Lamond (12°43’S, 143°17’E), 19 Dec
1971, D.K. McAlpine, G.A. Holloway, D.P, Sands
(AM).

Diagnosis. See Walker (1986).

Description of male. Body length: 6.16 mm;
Forewing length: 1.48 mm; Head width: 1.62
mm. Relative head measurements: HW: 100;
HL: 85; UID: 58; LID: 42; AOD: 15; IAD: 10;
OAD: 28; IOD: 16; OOD: 16; CL: 22; GW: 16;
EW: 31; SL: 34; FL: 142.

Structure. Head broad, inner orbits converg-
ing strongly below, eyes glabrous; median
frontal carina absent; scape reaches anterior
margin of median ocellus; clypeus convex, shin-
ing though covered with fine reticulate pattern.

Frons smooth though covered with distinct tess-
ellate pattern; vertex broad. Pronotum dorsola-
teral angles bluntly obtuse, well projected.
Mesoscutum and scutellum smooth. Dorsal sur-
face of propodeum not defined by carinae,
smooth except weakly ruguloso-striolate along
posterior margin. Metasomal terga impunctate
and smooth. Fore and mid tarsal segments
flanged laterally; hind basitibial plate complete,
bluntly acute apically.

Colour. Body black except metasoma dark
brown, anterior half of clypeus pale white-yel-
low, antennal scapes and pedicles red-brown,
legs and coxae, trochanters and basal half of fore
femora brown, remainder of legs light red-brown
except tarsal segments pale white.

Vestiture. Body sparse except lower frons and
paraocular areas with cover of adpressed, plu-
mose hair, genae with dense cover of long, plu-
mose hair forming a “beard”; anterolateral cor-
ners of mesoscutum, metanotum and lateral
margins of propodeum with dense cover of long,
plumose golden hair, posterior margin of mesos-
cutum with weak band of hair, fore coxae with
some long, plumose hair, fore trochanters with
dense cover of long, plumose hair, hair length
exceeds length of femur; fore and mid tarsi with
dense cover of simple and plumose hair, mid
and hind femora almost glabrous.

Genitalia and associated sterna. (figs 7-9).

Distribution. Cape York Peninsula and north
Queensland.

Remarks. Walker's (1986) comments on H.
maitlandi were based on the only available
material (types collected in 1902). Recently col-
lected specimens confirm the species is extant
and provide the male character suite described
and figured here. Examination of the male geni-
talia characters affirms the placement of mait-
landi within Homalictus and these characters,
together with flanged tarsal segment characters,
provide synapomorphies for the clade of H.
maitlandi, H. latitarsis (Friese) and H. grossope-
dalus Walker.

Homalictus (Homalictus) pilosignya sp. nov.
Figures 10-12
Material examined. Holotype. 9 Old, Mcllwraith
Range, 30 km E of Coen (13°50’S, 143°17’E), 3 Nov
1988. K. Walker (NMV, T-16621).
Paratypes. 69 (NMV, T-16622-T-16627), 38 (NMV,
T-16628-T-16630) same data as holotype.

Diagnosis. A member of the
species-group; female with

“blackburni”
frons smooth,

THE BEE GENUS HOMALICTUS 75

pronotum dorsolateral angles obtuse, mesoscu-
tum sparsely to openly punctate, dorsal surface
of propodeum mesially coarsely ruguloso-stri-
ate, laterally striate; male with mesoscutum
impunctate, dorsal surface of propodeum rugu-
loso-striolate, fore and mid leg tarsal segments
laterally flanged, genal hairs long, fore leg tro-
chanters with dense cover of long, plumose hair,
lateral margins of fore tarsi with long hairs,
chel on only the outer surface of the hair
shaft.

Description of female. (measurements of holo-
type in bold) Body length: 5.62-5.69 mm; Fore-
wing length: 1.41-1.43 mm; Head width: 1.74-
1.79 mm. Relative head measurements: HW:
100; HL: 81-83; UID: 50-52; LID: 46-47; AOD:
17-18; IAD: 8-09; OAD: 32-33; IOD: 13-14;
OOD: 12-13; CL: 17-18; GW: 15-16; EW: 25-
26; SL: 45-46; FL: 82-83.

Structure. Head broad, inner orbits converg-
ing below, median frontal carina absent; eyes
with sparse cover of minute setae. Scape
elongate, reaches at least posterior margin of
median ocellus. Clypeus weakly convex in side
view, anterior margin coarsely and irregularly
indented, remainder of surface dull, covered
with sparse, minute puncture, supraclypeal area
impunctate. Frons, vertex and paraocular areas
smooth, covered with a microtessellate pattern,
sparsely punctate with piliferous punctures.
Pronotum dorsolateral angles obtuse, weakly
projected. Mesoscutum surface dull, covered
with microtessellate pattern arranged as a circu-
lar pattern, openly to sparsely punctate with pil-
iferous punctures. Scutellum sculpture similar
to mesoscutum except close to openly punctate,
scutellum length equal to length of dorsal surface
of propodeum. Dorsal surface of propodeum not
defined by carinae, sculpture mesially coarsely
ruguloso-striate, laterally striate, sculpture
reaches dorsal rim. Mesepisternum and mete-
pisternum smooth, with fine reticulate pattern.
Fore basitarsal outer apicolateral comb absent,
hind basitibial plate apically obtuse, inner hind
tibial spur coarsely serrate with 3 teeth.

Colour. Body black except antennal flagella
brown, mesoscutum and scutellum sooty black,
metasoma tergum 1 with dark blue hue, legs with
tarsi brown.

Vestiture. Body sparse, except tomentose hair
on pronotum mesodorsal and dorsolateral areas,
mesoscutum anterolaterally and anterior
spiracle cover, lateral margins of propodeum
with long plumose hair; hind leg and ventral
metasomal scopae as usual for the genus.

Description of male. Body length: 3.92-4.62
mm; Forewing length: 3.85-4.01 mm; Head
width: 1.29-1.34 mm. Relative head measure-
ments: HW: 100; HL: 80-82; UID: 55-56; LID:
34-36; AOD: 09-10; IAD: 11-12; OAD: 33-34;
IOD: 15-16; OOD: 11-12; CL: 16-17; GW: 14-
15; EW: 32-34; SL: 28-30; FL: 160-163.

Structure. Head broad, sculpture smooth,
with a fine reticulate pattern, frons, clypeus and
supraclypeal area impunctate; inner orbits con-
verging strongly below; scape reaches well short
of anterior margin of median ocellus. Pronotum
dorsolateral angles broadly obtuse to rounded,
weakly produced. Mesoscutum and scutellum
dull, microtessellate, both impunctate. Dorsal
surface of propodeum not defined by carinae,
sculpture finely ruguloso-striolate, sculpture not
reaching dorsal rim. Fore and mid leg tarsal
segments laterally flanged.

Colour. Body black except antennal flagella
suffused with dark brown; legs dark brown
except tarsi light red-brown.

Vestiture. Body sparse except clypeus and
lower paraocular areas with some erect, plumose
hairs, genal hairs long, forming a “beard”, fore
leg trochanters with dense cover of long, plu-
mose hair, mid trochanters with some plumose
hair, though not as long or cover as dense as ves-
titure on fore trochanters, mesoventral area with
moderate cover of simple hair; lateral margins of
fore tarsi with long hairs, branched only on outer
surface of hair shaft.

Genitalia and associated sterna. (ĥgs 10-
12).

Distribution. Cape York Peninsula and north
Queensland.

Etymology. The epithet is from "pilosus" mean-
ing hairy and “ignya” referring to the upper
section of the leg.

Remarks. Homalictus pilosignya shares with H.
atrus fore leg vestiture (coxae and trochanters
with dense cover of long, plumose hair and lat-
eral margins of fore tarsi with long, simple hairs)
and the rasp-like sculpture on the apicoventral
surface of the genitalia volsellae but differs in the
female mesoscutum and scutellum colour (pilo-
signya sooty black; atrus black) and male vesti-
ture (pilosignya with plumose vestiture on fore
trochanters only (atrus plumose vestiture on fore
leg trochanters and all coxae), plumose hair
absent on the mesoventral area (atrus plumose
hair present) and the hairs branched on one side
only of the lateral margins of the fore tarsi (atrus
hairs simple).

76 K. WALKER

Homalicius pilosignya belongs to a species-
group (termed “hackburni” by Walker (1986)
for Australian species only and “buccinus” by
Pauly (1986) for Australian and non-Australian
species) which contains over 50 species that
occur in India, Sri Lanka, VietNam, Thailand,
Malaysia, Indonesia, Philippines, New Guinea,
Bismark Archipelago, Caroline, Bismarck and
Solomon Islands, the New Hebrides and Aus-
tralia (Pauly (1986)). This species-group forms a
monophyletic clade defined by several synapo-
morphies (both sexes with apex of marginal cell
terminating on wing margin, female lacking fore
basitarsal outer apicolateral comb absent, male
with genal vestiture long (forming a “beard”),
flanged tarsal segments, and coxae and/or tro-
chanters with plumose and elongated vestiture).
Australian members of this clade (H. atrus, H.
blackburni, H. cassiaefloris, H. dampieri, H. eur-
hodopus, H. grossopedalis, H. latitarsis, H. mait-
landi and H. pilosignya sp. nov.) occur through-
out the “Torresian” faunal province as defined
by Main (1981), with the greatest species diver-
sity found in north Queensland and Cape York
Peninsula.

Homalictus (Homalictus) verticulus sp. nov.
Figures 13-14

Material examined. Holotype. 9 Northern Territory,
12 km NNE of Borroloola (15°58’S, 136°21’E), | Nov
1975, J.C, Cardale, on Terminalia volucris R.Br. ex
Brenth. (ANIC).

Paratype. 19, NT, 22 km WSW of Borroloola
(16 08S, 136'06'E), 2 Nov 1975, J.C. Cardale, caught
in malaise trap. (ANIC),

Diagnosis. A member of the “urbanus” species-
group; female with frons striate, inner orbits par-
allel to weakly diverging below, vertex long and
broad (IOD = vertex length), genae enlarged
(GW 1.3 x EW), pronotum dorsolateral angles
large and acute, mesoscutum posterior half
closely punctate except parapsidal areas densely
punctate, dorsal surface of propodeum striate
with several basal interconnectives.

Description of female. (measurements of holo-
type in bold) Body length: 4.85-5.01 mm; Fore-
wing length: 1.05-1.06 mm; Head width: 1.53-
1.65 mm, Relative head measurements: HW:
100: HL: 80-81; UID: 61-62; LID: 62: AOD:
22-23; IAD: 9-10; OAD: 35; IOD: 17-18: OOD:
18-19; CL: 15-16; GW: 26-27; EW: 20: SL: 39-
40; FL: 72-74.

Structure. Head broad (fig. 13), inner orbits
parallel to weakly diverging (in holotype) below,
median frontal present, extends less than half-

way to median ocellus; eyes with sparse cover of
minute setae. Scape not reaching anterior mar-
gin of median ocellus. Clypeus flat in side view,
anterior margin straight, surface dull, anterior
half with. densely punctate with large, shallow
punctures, remainder openly punctate with
small puncture, supraclypeal area impunctate.
Fronsstriate to level of anterior margin of lateral
ocelli, areas lateral to median ocelli smooth, ver-
tex long and broad (IOD= vertex length), striate,
striae continue onto enlarged genae (fig. 14).
Labrum basal area with 2 large tubercles on
either side of midline. Pronotum dorsolateral
angles large and acute. Mesoscutum surface dull,

covered with a fine reticulate pattern, impunc-
tate to sparsely punctate, remainder closely

punctate except parapsidal areas densely punc-
tate. Scutellum shining, sparely to openly punc-

tate, scutellum length longer than length of
dorsal surface of propodeum. Dorsal surface of
propodeum not defined by carinae, dorsal rim

rounded and shining, sculpture striate with sev-

eral interconnectives basally, lateral striae con-

tinue onto vertical surface. Mesepisternum and

metepisternum coarsely striate. Hind basitibial

plate apically rounded, inner hind tibial spur

coarsely serrate with 3 large teeth.

Colour. Head dark blue/green, antennae
brown, mandible amber, mesoscutum semi-
metallic blue/green, scutellum blue, propodeum
and metasoma black, legs brown.

Vestiture. Body sparse, except tomentose hair
on pronotum dorsolateral areas, and mesoscu-
tum anterolaterally, lateral margins of propod-
eum with long plumose hair; hind leg and ventral
metasomal scopae as usual for the genus.
Distribution. Borroloola region, Northern Terri-
tory.

Etymology. The epithet is from "vertex", mean-
ing top, and refers to the unusual nature of the
vertex,

Remarks. Homalictus verticulus shares enlarged
head characters with H. ctenander Michener but
differs markedly in head, mesoscutum and pro-
podeum sculpture patterns and body colour. It is
most like H. holochlorus (Cockerell) in head
sculpture characteristics, body size and colour
but differs in the female by enlarged vertex and
genae, prominent pronotum dorsolateral angles
(acute distally), mesoscutum punctation (verti-
culus, mesially closely punctate, parapsidal
areas densely punctate; holochlorus mesially
openly punctate, parapsidal areas closely punc-
tate) and sculpture on the propodeum (verticu-
lus, striate; holochlorus, ruguloso-striolate).

THE BEE GENUS HOMALICTUS TI

Subgenus Papualictus

Homalictus (Papualictus) Michener, 1980: 8. Type
species: Homalictus megalochilus Michener, 1980: 8
(by original designation) = Homalictus lorentzi
(Friese). synonymy by Pauly (1986).

Diagnosis revised from Michener (1980). Both
sexes with frons, vertex and genal area coarsely
striate; frontal carina present between antennal
bases; females with hind tibiae slender, not
scarcely concave beneath, with hair of outer sur-
face sparse and simple; males with head much
broader than long (HW at least 1.2 x HL), cly-
peus low and transverse, about five times as
wide as long, apical truncation concave, episto-
mal suture lateral to tentorial pit horizontal,
mandibles enlarged and sickle shaped, shifted
slightly posteriorly so that a triangular malar
space is formed, pre-episternum elevated to
form a rough vertical ridge.

Homalictus (Papualictus) megagnathus sp. nov.
Figures 15-19

Material examined. Holotype. d, Qld, Mt Webb Nat.
Pk (15°04’S, 145°07’E), 27-30 Apr 1981, I.D. Nau-
mann (ANIC; Genitalia removed and placed in vial on
pin).

Paratypes. Qld: 18, 9 km ENE Mt Tozer (12°43’S,
143°17’E), 5-10 Jul 1986, J.C. Cardale, at MV light
(ANIC); le 3km ENE Mt Tozer(12"44'S, 143°14’E), 28
Jun-4 Jul 1986, J.C. Cardale, ex. Malaise trap/ethanol
(ANIC).

Diagnosis. Female with frons, vertex and genae
coarsely striate, anterior surface of hind tibia
with hairs nearly simple, bristlelike or with 1 or 2
branches (no area with abundant, short, erect,
plumose hairs as is usual for the genus); hind
tibiae slender, under surface scarcely concave;
male with head broader than long, clypeus short
and transverse, apical truncation concave, ante-
romesial margin elevated as a large boss, under-
neath boss clypeus strongly concave, mandibles
elongated and sickle shaped, shifted posteriorly;
each gena with a large lamella-shaped process.

Description of male. (measurements of holotype
in bold) Body length: 3.93-4.24 mm; Forewing
length: 0.66-0.68 mm; Head width: 1.27-1.29
mm. Relative head measurements: HW: 100;
HL: 70-72; UID: 62-64; LID: 61-62; AOD: 20-
21; IAD: 11-12; OAD: 31-32; IOD: 16-18;
OOD: 20-21; CL: 10-12; CW: 54-56; GW: 21-
22; EW: 23-24; SL: 37-38; FL: 92-94; ML: 62-
64.

Structure, Head broader than long (fig. 15),
inner orbits weakly converging below to almost

parallel below, median frontal carina well devel-
oped, reaches half way to median ocellus. Cly-
peus smooth and shining, short and transverse,
almost all of clypeus positioned above lower
margins of eyes, apical truncation concave, only
about one-fifth as wide as clypeus, anteromesial
margin elevated as a large, rounded, forwardly
projected rounded smooth shining boss, under-
neath boss clypeus strongly concave, epistomal
suture lateral to tentorial pit horizontal. Supra-
clypeal area weakly elevated, covered with a fine
reticulate pattern. Frons coarsely striate above
antennal bases, sculpture continues to level of
anterior margin of lateral ocelli, laterally con-
tinues almost to inner margin of eye. Mandibles
shifted posteriorly so that anterior articulation is
well behind lower inner margins of eyes, pos-
terior articulation of mandibles shifted poster-
iorly to well behind lower outer margins of eyes,
the shifted posterior mandibular articulation
forms a small triangular malar space; mandibles
elongated and sickle-shaped, longer than lower
interorbital distance, apex of mandible rounded
and weakly upturned, pollex absent. Labrum
with rounded boss across the entire basal area,
distally labrum rounded. Genal width, in side
view, subequal to eye width, each gena with a
single large, transversely broad, lamella-shaped
process (fig. 16), processes originate on under-
side of genae slightly behind level of posterior
mandibular articulation. Vertex broad and stri-
ate, slightly wider than interocellar distance.
Pronotum dorsolaterally rounded, in side view
lateral margin with distinct carina extending to
dorsolateral angle. Pre-episternum, forward of
pre-episternal groove, elevated to form rough
vertical ridge; mesepisternum, above pleural sig-
num, rounded and smooth; venter of mesepis-
ternum without transverse ridge behind front
coxae. Mesoscutum anterolateral margins raised
forming a lip, mesially lip absent, anteriorly dull
and impunctate, covered with a fine reticulate
pattern, mesially and laterally shining though
mesial area finely striate and openly punctate,
parapsidal areas with a fine broadly reticulate
pattern, sparsely punctate. Scutellum smooth
and highly polished, sparsely punctate with min-
ute punctures. Dorsal surface of propodeum not
defined by carinae, smooth and polished except
a few weak, short striae basally. Metasomal TI
highly polished and impunctate except for a few
minute punctures, T2-T5 shining and impunc-
tate except basally each tergite with several weak
transverse lines. Fore basitarsal comb fan
shaped; hind basitibial plate weakly present and
apically acute.

78

K. WALKER

Figures 10-16. Homalictus spp. H. pilosignya (paratype): male genitalia, fig. 10, ventral view (left half) and dorsal
view (right half); fig. 1 1, lateral view; fig. 12, metasomal S7 and S8. H. verticulus (holotype): figs 13-14, head front
and lateral views respectively. H. megagnathus (holotype): figs 15-16, head front and lateral views respectively.
Scale lines (associated with each species) = 0.5 mm.

THE BEE GENUS HOMALICTUS 79

Colour. Body black except, clypeus, mandibles.
| antennal scapes and legs light red-brown, coxae
black, femora suffused with dark brown, anten-
nal flagellum brown.

Vestiture. Body sparse, frons and mesoscutum
with sparse cover of erect hair, lower paraocular
areas with similar, though adpressed, hair, meta-
somal sternites with sparse cover of erect long
and short plumose hair.

Genitalia and associated sterna. (figs 17-19).

Description of female. Body length: 4.47 mm;
Forewing length: 1.08 mm; Head width: 1.36
mm. Relative head measurements: HW: 100;
HL: 82; UID: 58; LID: 52; AOD: 18; IAD: 09;
OAD: 34; IOD: 16; OOD: 16; CL: 18; GW: 14;
EW: 26; SL: 42; FL: 82; ML: 36.

Structure. Agrees with description of male
except as follows: Head with inner orbits con-
verging below. Clypeus convex, sculpture openly
punctate with shallow punctures, surface dull,
covered with fine reticulate pattern except
anterior margin smooth and shining. Supracly-
peal area sculpture and punctation as on cly-
peus. Frons coarsely striate above antennal
bases, ridges more prominent than in male.
Mandibles less shifted posteriorly relative to eye
than in male, malar area short, mandibles of
ordinary size and form. Genal width, in side
view, less than eye width, sculpture striate. Ver-
tex striate, slightly narrower than interocellar
distance. Pronotum dorsolateral angle acute, in
side view lateral margin of pronotum with a
ridge extending to dorsolateral angle, ridge less
prominent than in male. Pre-episternum, for-
ward of pre-episternal groove, weakly elevated
to form a vertical ridge. Mesoscutum anterola-
teral margins as in male, surface dull, covered
with a fine reticulate pattern, sculpture mesially
closely punctate, in parapsidal areas openly
punctate. Scutellum surface dull as on mesoscu-
tum, sculpture openly to sparsely punctate. Dor-
sal surface of propodeum as in male except basal
striae rugulose. Metasomal tergites as in male.
Hind basitibial plate well developed, apically
acute. Anterior surface of hind tibia with hairs
nearly simple, bristlelike or with 1 or 2 branches
(no area with abundant, short, erect, plumose
hairs as is usual for the genus); hind tibiae slen-
der, under surface scarcely concave. Inner mar-
gin of inner hind tibial spur with 2 or 3 apically
rounded teeth.

Colour. As in male but with antennal scapes
and flagella light red-brown infuscated with dark
brown, mesoscutum dull coppery-green, scutel-
lum with blue hue.

Vestiture. As in male but with metanotum
with tomentum of short white hair; femoral and
sternal scopa well developed, similar colour to
legs.

Distribution. Cape York Peninsula and north
Queensland. E

Etymology. The epithet is from “mega” and
“gnathos” and refers to the enlarged mandibles.

Remarks. Homalictus (Papualictus) was erected
for five species found in the moderate to high
altitudes of New Guinea, New Britain and New
Ireland (Michener, 1980). The discovery of H.
megagnathus now extends its distribution into
northern-east Australia. Michener produced a
diagnosis for H. (Papualictus) but noted that
since males were not known for all described
species, the male diagnostic characters *may not
all actually be subgeneric attributes" (p. 8).
Comparisons between the diagnostic characters
of H. (Papualictus) and character states of H.
megagnathus revealed the following differences.
Body length was described as large for both sexes
in H. (Papualictus), but H. megagnathus is best
termed small (9: 8.5-11 mm, 10-11 mm;pok l <
4.5 mm, respectively). Body length is the only
variable female subgeneric character. Male sub-
generic characters differ as: Head ratio (width
1.2 X length; width 1.4 X length respectively);
genal width (GW 2 X EW; GW almost equal to
EW, respectively); pronotum dorsolateral angle
large and distally acute versus small and
rounded; and the following two characters do
not occur on H. megagnathus: mesepisternum,
above pleural signum, elevated to form a rough
prominence; and dorsal surface of propodeum
strongly elevated to form a shining, longitudi-
nally elongate boss.

Subgeneric placement not resolved
Homalictus sicarius sp. nov.
Figures 20-24

Material examined. Holotype. €. Qld, 11 km NW of
Bald Hill. McIlwraith Range (13°44’S, 143°20’E), 26
Jun-13 Jul, I.D. Naumann, at light, search party
campsite (ANIC; genitalia removed and placed in vial
on pin).

Diagnosis. Male with head and vertex broad
(HW 1.5 X HL; vertex width 0.8 X IOD),
median frontal carina present, clypeus low and
transverse (CW 4.5 X CL), anteromesial margin
concave, epistomal suture lateral to tentorial pit
horizontal, frons weakly striate, genae area and
vertex smooth, mandibles elongated (ML 0.9 X

80 K. WALKER

Figures 17-24. Homalictus spp. H. megagnathus (holotype): male genitalia, fig. 17, ventral view (left half), dorsal
view (right half); fig. 18, lateral view; fig. 19, metasomal S7 and S8. H. sicarius (holotype): figs 20-21, head, front
and lateral views respectively. Male genitalia: fig. 22, ventral view (left half) and dorsal view (right half); fig. 23,
lateral view; fig. 24, metasomal S7 and S8. Scale lines (associated with each species) = 0.5 mm except figs 22-24 =
0.25mm.

THE BEE GENUS HOMALICTUS 81

LID), shifted posteriorly so that forming a small
triangular malar space, GW subequal EW, pron-
otum dorsolateral angles obtuse, mesoscutum
impunctate, dorsal surface of propodeum
smooth.

Description of male. (female unknown)
(measurements of holotype in bold) Body
length: 4.16 mm; Forewing length: 0.96 mm;
Head width: 1.25 mm. Relative head measure-
ments: HW: 100; HL: 74; UID: 62; LID: 60;
AOD: 18; IAD: 14; OAD: 30; IOD: 18; OOD: 18;
CL: 15; CW: 68; GW: 22; EW: 24; SL: 38; FL:
116; ML: 56.

Structure. Head broader than long (figs 20-
21) inner orbits weakly converging below,
median frontal carina present, reaches one third
to median ocellus. Clypeus with a dull sheen,
covered with a fine reticulate pattern, impunc-
tate, short and transverse (CW 4.5 X CL),
almost half of clypeus positioned above lower
margins of eyes, apical margin not truncate
though anteromesial margin concave with lat-
eral points forming small processes, epistomal
suture lateral to tentorial pit horizontal. Supra-
clypeal area weakly elevated, covered with a fine
reticulate pattern. Frons appears smooth though
with weak striae above antennal bases, striae not
reaching anterior margin of lateral ocelli, lateral
margin smooth. Mandibles shifted posteriorly
so that anterior articulation is behind lower
inner margins of eyes, posterior articulation of
mandibles shifted posteriorly to well behind
lower outer margins of eyes, the shifted posterior
mandibular articulation forms a small triangular
malar space (fig. 21); mandibles elongate,
weakly sickle-shaped, almost as long as lower
interorbital distance, apex of mandible rounded,
pollex absent. Genal width, in side view, sube-
qual to eye width. Vertex broad (Vertex width
0.8 X IOD) and smooth. Pronotum dorsolateral
angles obtuse, in side view weakly elevated. Pre-
episternum, forward of pre-episternal groove,
not elevated to form rough vertical ridge; mese-
pisternum, above pleural signum, rounded and
smooth; venter of mesepisternum without trans-
verse ridge behind front coxae. Mesoscutum
anterior margin rounded, entire surface dull and
impunctate, covered with a fine reticulate pat-
tern. Scutellum smooth, polished and impunc-
tate. Dorsal surface of propodeum not defined
by carinae, smooth and polished except a few
weak, short striae basally. Metasomal terga pol-
ished and impunctate. Fore basitarsal comb fan
shaped; hind basitibial plate well developed,
apically acute.

Colour. Body black except, apical half of cly-
peus with metallic blue/red tinge, mandibles and
basal one quarter of antennal scapes amber,
antennal flagellar segment brown, coxae and
femora brown, tibiae and tibiae and tarsi light
red-brown suffused with dark brown.

Vestiture. Body sparse, frons and mesoscutum
with sparse cover of erect hair, lower paraocular
areas and clypeus with moderate cover of short
plumose hair, metasomal sternites with sparse
cover of erect long and short plumose hair.

Genitalia and associated sterna. (figs 22-
24).

Distribution. Cape York Peninsula.

Etymology. The epithet is from “sica” meaning
dagger and refers to the shape of the mandi-
bles.

Remarks. Subgeneric placement of H. sicarius
will require association of the female and exam-
ination of the full character suite. Several
character states of the male suggest subgeneric
placement in H. (Papualictus). In particular the
head ratio (HW 1.4 X HL), clypeus low and
transverse (clypeus width 4 X CL), anteromesial
margin concave, mandibles elongated, weakly
sickle-shaped and with the posterior articulation
slightly shifted posteriorly to form a small tri-
angular malar space. However, comparison with
my diagnosis of H. (Papualictus) (Walker, 1986)
highlighted the following differences: frons, ver-
tex and genae not striate, the clypeus apical
margin not truncate and concave beneath, not
forming an elevated boss, mandible length less
than LID and pre-episternum not elevated to
form a rough vertical ridge. I considered this
single male specimen warranted description to
flag a second Australian species that possesses a
number of subgeneric male head characters of
H. (Papualictus) and may eventually be placed in
that subgenus.

Acknowledgments

I wish to thank the curators of the institutions
listed for the loan of their specimens and Pro-
fessor C.D. Michener, Drs Glynn Maynard and
Gary Poore for their constructive comments.

References

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Michener, C.D., 1980. The large species of Homalictus
and related Halictinae from the New Guinea area
(Hymenoptera, Apoidea). American Museum
Novitates 2693: 1-21.

Pauly, A., 1986. Les abeilles de la sous-famille des Hal-
ictinae en Nouvelle-Guinée et dans l'Archipel Bis-
mark (Hymenoptera: Apoidea: Halictidae). Zool-
ogische Verhandelingen 227: 1-58.

Walker, K.L., 1986. Revision of the Australian species
of the genus Homalictus (Hymenoptera: Halicti-
dae). Memoirs of the Museum of Victoria 47(2):
105-200.

Memoirs of the Museum of Victoria 56(1):83-89 (1997)

DESCRIPTIONS OF NEW LEPTOPHLEBIIDAE (INSECTA: EPHEMEROPTERA)
FROM AUSTRALIA. I. TILL YARDOPHLEBIA GEN. NOV.

JOHN C. DEAN

Environment Protection Authority, Catchment and Marine Studies,
27 Francis Street, Melbourne, Vic. 3000, Australia

Abstract

Dean, J.C., 1997. Descriptions of new leptophlebiidae (Insecta: Ephemeroptera) from
Australia. I. Tillyardophlebia gen. nov, Memoirs of the Museum of Victoria 56: 83-89,
The genus Tillyardophlebia gen. nov. is established to accommodate a group of leptoph-
lebiid mayflies from eastern Australia. Adults and nymphs of T.rufosa sp. nov. from Victoria
and New South Wales and T.alpina sp. nov. from New South Wales are described and

figured.

Introduction

The family Leptophlebiidae is the dominant
mayfly family in Australia. Despite recent
descriptions of new genera (Dean, 1987, 1988;
Campbell and Suter, 1988; Campbell, 1993), the
family remains poorly known, with many undes-
cribed genera held in various collections (Dean
and Suter,1996). The present paper is the first in
a series, and describes a new genus and two new
species from south-eastern Australia.
Additional taxa will be described as nymphs are
associated with adults and sufficient adult
material becomes available. Examined material
has been preserved in alcohol, with parts of some
specimens mounted on microscope slides. All
material is lodged in the Museum of Victoria
(NMV).
Tillyardophlebia gen. nov.

Diagnosis. Imago. Forewing with membrane

| hyaline (Fig. 1); costal and subcostal cells either

uniformly washed with reddish-brown or hyal-
ine with cells in apical third of wing translucent,
whitish. Length-width ratio 2.8-3.0. 4-9 costal
crossveins basal to the bulla, 10—19 distal to the
bulla. MA forked at 0.38-0.41 wing length. MP,
attached by crossvein to MP, at 0.17-0.18 wing
length. ICu, not linked to CuA-CuP crossvein ;
base of ICu, linked by crossvein to CuA; ICu,
and ICu, moderately diverging as wing margin
approached. Hindwing 0.24—0.28 length of fore-
wing. Costal margin with shallow concavity just
beyond midlength, costal space relatively broad
both basal and distal to this (Fig. 2). Vein Sc

| joining costal margin at about 0.95 wing length.

Hindwing with 8-13 costal crossveins and 8-10
subcostal crossveins. All legs with tarsal claws
dissimilar, one with an apical hook and oppos-
ing ventral flange, the other obtuse, pad-like
(Fig. 6). Forelegs of male with ratios of segment

83

lengths 0.71-0.89:1.00 (3.6-3.9 mm):0.04-
0.05:0.37-0.43:0.37-0.45:0.28-0.36:0.10-0.11.
Male genitalia with claspers 3-segmented, nar-
rowing abruptly at about midlength (Fig. 4).
Penes long, narrow, widely separated for most of
length, fused at base only; with pair of stout,
curved ventral spines a little basal to where the
penes fuse (Figs 21, 22, 24, 25). Female ninth
sternum with apical margin either flattened or
very shallowly concave (Fig. 5).

Subimago. Wings uniformly
brown.

Mature nymph. Head prognathous, antennae
about twice width of head. Mouthparts as in Fig-
ures 14-20. Clypeus with lateral margins slightly
diverging to anterior. Labrum 1.1-1.4 times
broader than clypeus; width 1.8-2.3 times maxi-
mum length; 2 setal fringes close to anterior
margin, each fringe extending across more than
half width of labrum; anterior margin with cen-
tral notch, the base of which is concealed by
frontal setal fringe, which lies along an over-
hanging canopy. Mandible with outer margin
rounded, dense tuft of long setae at midlength,
and series of shorter setae along margin between
tuft and outer incisor; incisors slender, with | or
more subapical denticles; prostheca large.
Lingua of hypopharynx with well developed lat-
eral processes, anterior margin with deep
incision; superlingua with dense setal fringe
along anterior margin. Maxilla somewhat squat,
subapical row of about 25 pectinate setae; palp
moderately short, terminal segment about same
length as middle segment, middle segment bear-
ing simple setae only. Labium with glossae not
turned under ventrally, lying in same plane as
paraglossae; palp 3-segmented, terminal seg-
ment about half length of middle segment. Legs
relatively large, apex of hind femur reaching
beyond midlength of abdomen (Fig. 7); all

yellowish-

84 J. C. DEAN

segments of leg with fringe of setae along outer
margin (Fig. 9); tarsal claws with ventral teeth,
progressively larger apically (Fig. 10). Abdomi-
nal segments without setae on lateral margins,
strongly developed posterolateral spines on seg-
ments 2-9 (Fig. 8); posterior margins of abdomi-
nal terga with series of stout spines, longer spines
interspersed with shorter (Fig. 11). Gills present
on abdominal segments 1-7, each gill lanceolate,
with upper and lower lamellae equally devel-
oped (Fig. 12). Caudal filaments with apical
whorl of stout spines on each segment, fine setae
sparse (Fig. 13).
Type species. Tillyardophlebia rufosa sp. nov.
Etymology. The genus is named for R.J.Tillyard,
in recognition of his pioneering work on Aus-
tralian mayflies.
Remarks. Brief descriptions of Tillyardophlebia
have previously been provided by Dean and
Suter (1996) under the designation ‘Genus D’.
More species than those described below are
known from Tasmania, Victoria and Queens-
land, but a shortage of material precludes their
description in the present work.

The genus Tillyardophlebia can be distin-

guished from all other leptophlebiid genera by
the following combination of characters.

Imago: 1, forewing with ICu, not attached to
CuA-CuP crossvein; 2, forewing approximately
4 times length of hindwing; 3, hindwing with Sc
joining wing margin at about 0.95 wing length; 4,
tarsal claws dissimilar; 5, male genitalia with
penes long, narrow, fused near base only, with a
pair of stout ventral spines near base; 6, ninth
sternum of female with apex either flattened or
with very shallow concavity.

Nymph. 1, labrum slightly broader than cly-
peus and with notch in anterior margin; 2, width
oflabrum 1.8-2.3 times maximum length; 3, tar-
sal claws with ventral teeth; 4, abdominal seg-
ments without lateral fringe of setae; 5, postero-
lateralspinespresentonabdominalsegments2-9.

Tillyardophlebia is a member of the Meridial-
aris lineage as defined by Pescador and Peters
(1980), which also includes the Australian genus
Austrophlebioides. While the two Australian
genera are closely related, Tillyardophlebia is
clearly distinguished by the form of the male
genitalia and, in the nymph, by the form of the
labrum and the absence of setae on the lateral
margins of the abdominal segments.

Imagos
l.

Nymphs
l

Keys to described species of Tillyardophlebia

Legs with all segments yellow, brown pigmentation restricted to
small area around apices of femora; male genitalia with ventral,
triangular flange at about mid length of each penis lobe (Figs
ZII) 5 etu Sel ume ume rti Tillyardophlebia rufosa
Legs with femora dark brown, all other segments pale brown:
male genitalia without ventral flange at mid length of each penis
lobe (Figs2425),...,................ Tillyardophlebia alpina

Gills with membranes heavily pigmented, purple; femora of all
legs dark reddish-brown ............. Tillyardophlebia alpina
Gills with membranes pale, yellow-white; femora of all legs light

brown with several contrasting pale patches

Tillyardophlebia rufosa sp. nov.
Figures 1-23
lype material. Holotype: Victoria. Badger Creek, d/s
weir, 23 Feb 1984, J. Dean, NMV T-16709, male
imago reared from nymph.

Paratypes: same location, 31 Jan 1985, J. Dean,
NMV T-16710, male imago reared from nymph; same
location, 21 Feb 1980, J. Dean, NMV T-16711, male
imago reared from nymph (wings, genitalia, legs,
assorted nymphal parts mounted on slides); same
location. 20 Mar 1980, J. Dean, NMV T-16712,
female 1mago reared from nymph: same location, 21
Feb 1980, J. Dean, NMV T-16713. female imago

HITS RENATE Tillyardophlebia rufosa

reared from nymph (wings, ninth sternum, legs,
assorted nymphal parts mounted on slides).

Other material examined. Vic. Badger Creek, d/s
weir, various dates, J. Dean, 14 nymphs; same
location, various dates, J. Dean, 5 female subimagos
reared from nymphs; Watts River, Fernshaw, 4 Mar
1976. J. Dean, female imago reared from nymph; same
location, 24 Feb 1977, J. Dean, 1 nymph; O'Shannassy
River, u/s reservoir, 17 Feb 1977, J.Dean, 1 nymph;
Thomson River, u/s Easton Portal. 22 Jan 1975,
J.Dean, 1 nymph. NSW. Thredbo River, 1.5 km u/s
Deadhorse Gap, 21 Jan 1984, J. Dean, 2 nymphs;
Leather Barrel Creek, Alpine Way, 6 Feb 1985, J.
Dean, 3 nymphs.

A NEW GENUS OF MAYFLY 85

Description. Imago. Length of male: body 10.6-
11.6 mm, forewing 10.0-10.3 mm; Length of
female: body 11.7 mm, forewing 11.7 mm.
Upper lobes of male eyes orange-brown, in
contact dorsally. Thorax golden. Legs with all
segments yellowish, apex of femur brown. Fore-
wings with costal and subcostal cells washed
with reddish-brown (Fig. 1). Male abdomen pre-
dominantly paler brown, with pair of thin, dark
brown, longitudinal stripes along midline, an
adjacent paler macula on each side of midline
close to anterior margin, and further lateral to
this an area of dark brown pigmentation (Fig.
23). Female abdomen similar, although with
pale maculae less obvious and darker anterola-
teral region less contrasting with the rest of the
segment. Penes lobes with ventral, triangular
flange at about midlength, and triangular ventral
process immediately distal to the fused basal
section (Figs 21 22).

Subimago. Wings yellowish-brown, forewing
costal and subcostal cells with brown pigmen-
tation of imago showing through. Female abdo-
men reddish-brown.

Nymph. Labrum width 2.0-2.1 times maxi-
mum length; width of frontal setal fringe 0.5-0.6
times width of labrum, narrower than secondary
fringe; anterior notch broad (Fig. 14). Femora
with upper surface contrasting pale brown and
yellow (Fig. 9). Gills with membrane pale, yel-
low-white. Abdominal terga contrasting brown
and yellow colour pattern (Fig. 7).

Comments. Association of adult and nymph by
rearing. Nymphs occur in small to medium sized
forest streams, where they have been collected
from the surface of large cobbles and small
boulders in moderate currents.

Tillyardophlebia alpina sp. nov.
Figures 24-26

Type material: Holotype: New South Wales. Ram-
shead Creek, the Cascades, Merritt's Track, 27 Jan
1984, J. Dean, NMV T-16714, male imago. Paratype:
same location and date, NMV T-16715, female subi-
mago.

Other material examined: NSW. Ramshead creek,
The Cascades, Merritt's track, 27 Jan 1984, J. Dean, 2
nymphs: Ramshead creek, u/s Mt Kosciusko summit
track, 27 Jan 1984, J. Dean, 6 nymphs; Lake Coota-
patamba, 24 Jan 1984, J. Dean, | nymph; Lake Albina
inflowing stream, 9 Feb 1985, J. Dean, 24 nymphs;
creek near top of Crackenback chair lift, 15 Dec 1978,
J. Dean, 6 nymphs.

Description. Imago. Length of male, body 12
mm, forewing 11 mm. Upper lobes of male eyes

pinkish-tan, in contact on meson of head. Tho-
rax golden. All legs with femur dark brown,
other segments pale brown. Forewings with
costal and subcostal cells washed with pale
brown. Abdomen predominantly darker brown,
somewhat blotchy, each segment paler in pos-
terior half; terga with thin, dark brown, longi-
tudinal stripes along the midline, ill defined pair
of paler maculae near anterior margin, l on
either side of the midline (Fig. 26). Penes lobes
without ventral flange at midlength; inner mar-
gin of each lobe with ventral process immedi-
ately distal to fused basal section (Figs 24, 25).
Lobes in ventral view relatively long and nar-
row.

Subimago. Wings yellowish-brown.

Nymph. Labrum width about 2.2 times maxi-
mum length; width of frontal setal fringe 0.5-0.6
times width of labrum, narrower than secondary
fringe; anterior notch narrow. Femora with
upper surface strongly pigmented, reddish-
brown. Gills with membrane darkly pigmented,
purplish. Abdominal terga uniformly dark red-
dish-brown.

Comment. Association of adult and nymph is
based on dissection of the male genitalia from a
ripe nymph. The species is known only from the
Kosciusko region, and nymphs have been col-
lected from stony substrates of small streams.
both above and just below the treeline.

Acknowledgments

Dr Ken Walker, Curator of Entomology,
Museum of Victoria, is thanked for making
available material held in the collections of the
museum. Part ofthe work on the immatures has
been funded by the Land and Water Resources
Research and Development Corporation as an
MRHI R and D project.

References

Campbell, I.C., 1993. A new genus and species of lep-
tophlebiid mayfly (Ephemeroptera: Leptophlebii-
dae: Atalophlebiinac) from tropical Australia.
Aquatic Insects 15: 159-167,

Campbell, I.C. and Suter, P.J., 1988, Three new gen-
era, a new subgenus and a new species of Leptoph-
lebiidae (Ephemeroptera) from Australia. Journal
of the Australian Entomological Society 27; 259—
2758:

Dean, J.C., 1987, Two new genera of Leptophlebiidae
(Insecta: Ephemeroptera) from south-western
Australia. Memoirs of the Museum of Victoria 48:
91-100.

86 J. C. DEAN

Dean, J.C., 1988. Description of a new genus of lep-
tophlebiid mayfly from Australia (Ephemerop-
tera: Leptophlebiidae: Atalophlebiinae). Proceed-
ings of the Royal Society of Victoria 100: 39-45.

Dean, J.C. and Suter, P.J., 1996. Mayfly nymphs of
Australia. A guide to genera. Identification Guide
No.7, Co-operative research Centre for Freshwater

Ecology. Murray-Darling Freshwater Research
Centre: Albury, New South Wales.

Pescador, M.L. and Peters, W.L., 1980. Phylogenetic
relationships and zoogeography of cool-adapted
Leptophlebiidae (Ephemeroptera) in southern
South America. Pp 43-56 in: Flannagan, J.F. and
Marshall, K.E. (eds), Advances in Ephemeroptera
Biology. Plenum Press: New York

Figures 1-6. Tillyardophlebia rufosa. Imago. 1, wings, male; 2, hind wing, male; 3, male genitalia, lateral; 4, male
genitalia, ventral; 5, female ninth sternum, ventral; 6, tarsal claws, male. d

A NEW GENUS OF MAYFLY 87

"n v. v v
KANA SA Vv viv, VV.
VI Nw vvv ne NUI vi ANN VWA v.

MA

11

Figures 7-13. Tillyardophlebia rufosa. Nymph. 7, whole nymph, dorsal; 8, postero lateral spines of abdomen; 9,
fore-leg; 10, tarsal claw; 11, spines on posterior margin of abdominal tergum 6; 12, gill from abdominal segment
4; 13, terminal filament, midlength.

88 J. C. DEAN

Figures 14-20. Tillyardophlebia rufosa. Nymphal mouth parts. 14, labrum; 15, left maxilla, ventral; 16, left
mandible, dorsal; 17, right mandible, dorsal; 18, hypopharynx; 19, labium, dorsal (left of midline) and ventral
(right of midline); 20, terminal segment of labial palp, dorsal.

A NEW GENUS OF MAYFLY 89

Figures 21-26. Male imagos. Tillyardophlebia rufosa. 21, penes, ventral; 22, penes, lateral; 23, abdominal terga
4-6, dorsal; T. alpina 24, penes, ventral; 25, penes, lateral; 26, abdominal terga 4-6, dorsal.

Memoirs of the Museum of Victoria 56(1):91-123 (1997)

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN GENERA OF
CHLOROCYSTINI (INSECTA: HOMOPTERA: TIBICINIDAE)

A.J. DE BoER

Instituut voor Systematiek en Populatie Biologie (Zoólogisch Museum). University of Amsterdam,
PO Box 94766, 1090 GT Amsterdam, The Netherlands

Abstract

Boer. A.J. de, 1997. Phylogeny and biogeography of Australian genera of Chlorocystini
(Insecta: Homoptera: Tibicinidae). Memoirs of Museum of Victoria 56(1): 91-123.

Six cicada genera belonging to the Chlorocystini are endemic to Australia. These (Chlor-
ocysta Westwood, 1851; Cystopsaltria Goding and Froggatt, 1904; Cystosoma Westwood,
1842; Glaucopsaltria Goding and Froggatt, 1904; Owra Ashton, 1912: and Venustria Goding
and Froggatt, 1904) can be divided into three monophyletic groups. The phylogeny and
biogeography of these groups is discussed, descriptions are given ofthe groups and all species
concerned. A key to males is presented for all species of Chlorocystini in Australia.

Introduction

The “Baeturia and related genera complex” was
defined as a supposedly monophyletic group for
which aedeagal characters are regarded synapo-
morphic (De Boer, 1990). Recently this complex
was identified as the tribe Chlorocystini (sensu
stricto) (De Boer, 1995d) comprising about 150
species attributed to 14 genera. Most species
occur in New Guinea but the distribution of the
tribe includes Maluku and Timor, the Bismarck
Archipelago, Solomon Islands, Vanuatu,
Samoa, Tonga and parts of northern and eastern
Australia.

Of the 13 species in Australia two (Thaumas-
topsaltria globosa Distant, 1897 and Guineap-
saltria flava (Goding and Froggatt, 1904)) also
occur in New Guinea. All others are endemic to
Australia and, apart from Gymnotympana rufa
(Ashton, 1914) and Gymnotympana varicolor
(Distant, 1907), belong to eridemic Australian
genera.

The present publication forms part of a phy-
logenetic and biogeographic study of the Chlor-
ocystini (sensu stricto) and deals with the
endemic Australian genera: Chlorocysta West-
wood, 1851; Cystopsaltria Goding and Froggatt,
1904; Cystosoma Westwood, 1842; Glaucopsal-
tria Goding and Froggatt, 1904; Owra Ashton,
1912; and Venustria Goding and Froggatt, 1904.
Cystosoma has two species, Chlorocysta three
and the others are monotypic.

All nine species were recently discussed
(Moulds, 1990) but descriptions and drawings of
male genitalia are given for the first time here.
The redescriptions recount the characters used
in a phylogenetic reconstruction of the Chloro-
cystini as a whole (De Boer, 1995d). Here, the

91

relationships of and between Australian genera

are discussed separately. A computer analysis of

the distribution of shared characters of all 148

species of the Chlorocystini (sensu stricto) has

shown that the Australian genera can be subdiv-

ided into three groups:

l. Cystopsaltria and Cystosoma, as mono-
phyletic group;

2. Chlorocysta, Glaucopsaltria and Owra, as
monophyletic group: and

3. Venustria.

The latter takes a somewhat isolated position,

but is presumably closely related to Gymnotym-

pana Stàl, 1861 and shares several characters

with the Australian species (G. rufa and G. var-

icolor) (De Boer, 1995a; 19954).

The results of phylogenetic analysis (De Boer,
1995d)are summarised and relationships within
and between the Australian groups is treated in
more detail. The groups are diagnosed and all
species are described. A key to males of all
Australian species of the Chlorocystini is
presented.

Methods

The material examined for this study is pre-
served in the following collections: BMNH,
Natural History Museum (formerly: British
Museum (Natural History) London; BPBM,
Bernice P. Bishop Museum, Honolulu; CSIRO,

Commonwealth Scientific. and Industrial
Research Organisation, Canberra; DEI,
Deutsches Entomologisches Institut, Eber-

swalde; IZW, Polska Akademia Nauk, Instytut
Zoologii, Warszawa; KBIN, Koninklijk Belgisch
Instituut voor Natuurwetenschappen, Brussels;
Moul, personal collection Mr M.S. Moulds,

92 A, J. DE BOER

Sydney; RMNH, Nationaal Natuurhistorisch
Museum (formerly Rijksmuseum van Natuur-
lijke Historie), Leiden; SEM, Snow Entomologi-
cal Museum, Lawrence, Kansas; SMD, Staat-
liches Museum für Tierkunde, Dresden; SMF,
Natur Museum und Forschungs Institut “Sen-
kenberg", Frankfurt am Main; SMN, Staatliches
Museum für Naturkunde, Stuttgart; ZIM, Zool-
ogisches Institut und Zoologisches Museum,
Hamburg; ZMA, Instituut voor Systematick
en Populatie Biologie (Zoólogisch Museum),
Amsterdam; ZMB, Institut für Spezielle Zoo-
logie und Zoologisches Museum der Humboldt-
Universitit, Berlin; and ZMH, Zoological Mu-
seum of the University of Helsinki, Helsinki.

Some of the terms used in the descriptions are
explained in figs 5 and 13. To examine the male
genitalia the pygofer was pulled out after over-
night softening, with a sharp needle inserted
between the pygofer and the 8th abdominal seg-
ment. The aedeagus was pulled out at the same
time by inserting the needle between the
claspers. Descriptions were made from dried
museum material. This drying often affects
colour. Bright green becomes yellowish brown,
but colour marks like blackish stripes and spots
remain intact. Measurements are based on all
available specimens. Only some of the most
important and the most recent systematic litera-
ture concerning the genera and species is cited.
For more complete lists of literature is referred
to the catalogues of Metcalf (1963) and Duffels
and Van der Laan (1985).

Phylogeny

In the following discussion figures in parenth-
eses refer to apomorphies in the cladogram (fig.
1). The Chlorocystini (sensu stricto) form a
monophyletic group for which an S- curved aed-
eagus with winged lateral crests is the supposed
apomorphy (1). The phylogenetic relationships
between the 148 described species of this tribe
were analysed with the aid ofthe program PAUP
(Swofford, 1993) using a data matrix of 154
characters and 409 character states (De Boer,
1995d). The tribe Prasiini was used as the sister
group and the genus Muda as outgroup. The
result of this analysis showed that on a generic
level the support of some of the proposed phy-
logenetic relationships is very weak.

Two major subdivisions can be made. Chlor-
ocysta, Glaucopsaltria, Owra and Venustria can
be grouped with Baeturia, Guineapsaltria, Gym-
notympana, Papuapsaltria and Scottotympana
based on the following synapomorphies: proxi-
mal spine of fore femur erect (3), a smoothly

vaulted pronotum without distinct medial fis-
sure (4) and a distinct hyaline border along the
hind margin of tegmen (5). Other genera have a
somewhat wrinkled head and pronotum with a
fairly distinct medial fissure on the pronotum
and a very narrow border along the hind margin
of tegmen; these character states also occur in
most species of the Prasiini, the presumed sister
group of the tribe.

Baeturia
Scottotympana
Gymnotympana

Venustria

Chlorocysta
Glaucopsaltria
Owra

Guineapsaltria

Papuapsaltría

? Ew Aedeastria

? Thaumastopsaltria
Mirabilopsaltria

Cystopsaltria
Cystosoma

Prasiini

Figure 1. Tentative cladogram of the Chlorocystini.
Numbers refer to apomorphies discussed in the text.

Another subdivision can be made based on male
operculum size. Venustria forms a monophyletic
group with Baeturia, Gymnotympana and Scot-
totympana. These genera share a fairly large
operculum as apomorphy (7) in which the
medial margin lies medial to the meracanthus.

In other genera of Chlorocystini, in Prasiini and

in Mudathe medial margin ofthe operculum lies

generally lateral to the meracanthus.

Several characters of Venustria are also found
in Scottotympana and many species of Gymno-
tympana and indicate a close relationship with
these two genera:

l. tegmina with reddish venation and a broad
hyaline border along the hind margin;

2. along proximal spineon the fore femur, often
longer than the distance to middle spine;
and

3. a very short meracanthus.

However, Baeturia and Scottotympana presum-

ably form a monophyletic group; claspers which

are not fused at the base are a presumed synap-
omorphy for these two genera (8). V. superba
shares a very similarly shaped basal part of the
operculum with Gymnotympana rufa and

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 93

G. varicolor while the female of V. superba shares
a similar thorn-shaped protuberance on the
pygofer with G. rufa. (De Boer, 1995a; 1995d).
The last two characters suggest that Venustria
should be included in and be synonymised with
Gymnotympana but no unambiguous synapo-
morphy for Venustria and Gymnotympana
together has been found. The phylogenetic
analysis showed that Venustria as the sister
group of Gymnotympana is the most parsimoni-
ous solution (De Boer, 19954).

The phylogenetic relationships between
Chlorocysta, Glaucopsaltria and Owra and the
group formed by Baeturia, Guineapsaltria,
Gymnotympana, Papuapsaltria, Scottotympana
and Venustria are not clear, as indicated by the
polytomy in fig. 1. Guineapsaltria and Papuap-
saltria are possibly sister groups, based on a very
small and rounded male operculum in many of
their species (6). Chlorocysta, Glaucopsaltria
and Owra share a very broad vertex with the
small ocelli wide apart with most species of
Gymnotympana and all but one species of Gui-
neapsaltria; the distance between the lateral
ocelli often exceeds three times the width of the
frontal ocellus. The fairly short meracanthus of
the species of Chlorocysta, Glaucopsaltria and
Owra and the vague colour pattern found on the
head and thorax of two of the Chlorocysta
species, might indicate a close relationship with
Gymnotympana, Scottotympana and Venustria.
The phylogenetic analysis showed that a recon-
struction with Chlorocysta, Glaucopsaltria and
Owra as sister group of Baeturia, Gymnotym-
pana, Scottotympana and Venustria is slightly
more parsimonious than the option with Gui-
neapsaltria and Papuapsaltria as sister group of
that group. On a tree with a total length of 906
steps the latter option is only one step longer (De
Boer, 1995d).

Cystosoma and Cystopsaltria share a strongly
furrowed pronotum and a very narrow border
along the hind margin of tegmen with Aedeas-
tria, Mirabilopsaltria and Thaumastopsaltria, a
strongly bent proximal spine on the fore femur
with Mirabilopsaltria and Thaumastopsaltria
and an angularly protruding postclypeus with
Thaumastopsaltria. All these characters are also
widely distributed in Prasiini and are presumed
plesiomorphic. Furthermore, Cystosoma and
Cystopsaltria have small male opercula which
are regarded plesiomorphic; the opercula of Cys-
tosoma schultzi and Cystopsaltria immaculata,
though relatively small, do slightly extend medi-
ally of the meracanthus but the operculum of
Cystosoma saundersii does not. The two species

of Cystosoma and most species of Thaumastop-
saltria and Mirabilopsaltria have a narrow tym-
bal cavity; the tergite part between the auditory
capsule and the 2nd sternite is very short. This
character is presumably apomorphic for Cystos-
oma, Cystopsaltria, Mirabilopsaltria and Thau-
mastopsaltria together but then lost in some of
the species (2). It is not clear whether or not Aed-
eastria should also be included in this group;
though Aedeastria has a wide tymbal cavity it
shares distinct diverging fissures on the vertex as
a possible synapomorphy with Mirabilopsaltria
and Thaumastopsaltria. The erect fore femoral
spine of Aedeastria, however, suggests a mono-
phyletic origin with the group containing Baetu-
rid, Chlorocysta, Glaucopsaltria, Gymnotym-
pana, Guineapsaltria, Owra, Papuapsaltria,
Scottotympana and Venustria (see fig. 1 ques-
tionmarks)

The coloured tegmina of Cystosoma and Cys-
topsaltria which are also found in Mirabilopsal-
triaand Thaumastopsaltria might also indicate a
relationship between these genera though this
character is also found in some genera of
Prasiini.

Biogeography

Biogeographic patterns of Chlorocystini and a
similarly distributed group of cicadas, the sub-
tribe Cosmopsaltriaria, were recently compared
and analysed (De Boer, 1995c). It appears that
most of the species of these groups are found on
terranes derived from a historic oceanic island
arc, which developed since about 40 million
years ago as a result of subduction of the former
Tethys Sea under the Pacific tectonic plate (De
Boer, 1995c). This arc is known as the Outer
Melanesian Arc (Hamilton 1979; Holloway,
1979, 1984; Duffels, 1986; Rangin et al., 19902,
1990b; Daly et al., 1991), Due to the continuous
northward movement of Australia and the west-
ward movement of the Pacific this volcanic
island arc broke up and fragments collided with
the northern craton of Australia where they now
form part of New Guinea (Pigram and Davies,
1987). It is supposed that the Outer Melanesian
Arc, before breaking up, formed an important
route of dispersal for south-east Asian biota
invading the Pacific (Duffels 1986, Duffels and
De Boer, 1990, De Boer, 1995c).

The distribution of Chlorocystini over the arc
terranes, the fact that their presumed sister
group (the oriental Prasiini as defined by De
Jong, 1985) is most speciose in Sulawesi and that
the sister group of these groups combined (the
genus Muda) occurs in south-east Asia and the

94 A. J. DE BOER

greater Sunda Islands, suggest that the ancestor
of that tribe also used the island arc as route of
dispersal. In fact. the distribution and phylogeny
of these cicadas reflect the geotectonic history in
that the main vicariant speciation events in
the cladograms correspond with presumed
sequences of fragmentation of the island arc
(De Boer, 1995c). This means that the ancestor
of Chlorocystini originates from Asia and that
all its Australian species must have invaded Aus-
tralia by dispersal presumably through New
Guinea (De Boer, 1995c).

Two patterns of distribution of Australian
Chlorocystini can be recognized. It appears that
species not belonging to endemic genera but
belongingto genera found in New Guinea as well
(Guineapsaltria flava, Gymnotympana | rufa,
Gymnotympana varicolor and Thaumastopsal-
tria globosa) are restricted to the northern and
eastern parts of the Cape York Peninsula; the
only exception (7. globosa) is also recorded from
Groote Eylandt, NT (Moulds, 1990). This same
pattern is found in several species that do not
belong to Chlorocystini: Diceropyga subapicalis
(Walker, 1868) (see Duffels, 1977), Lembeja
paradoxa (Karsch, 1890) and Lembeja vitticollis
(Ashton, 1912) (see De Jong, 1982) belong to
non-endemic genera and are restricted to the
Cape York Peninsula (Moulds, 1990).

Venustria, an endemic Australian genus that
should possibly be included in Gymnotympana
(see discussion above; De Boer, 1995a; 19954),
is also restricted to the Cape York Peninsula.
The two groups of endemic Australian genera
havea much wider distribution, reaching farther
southward. These have a strikingly similar dis-
tribution in northern and eastern Qld, including
the Cape York Peninsula and north-eastern
NSW.

The distribution patterns of the species of
endemic and non-endemic Australian genera
and their relative positions in the cladogram
suggest different ages. There must have been at
least two periods of dispersal from New Guinea
to Australia. For the non- Australian genera a
dispersal during Pliocene-Pleistocene low sea-
level stands has been proposed (De Boer, 1992b;
1995c). The two groups of endemic Australian
genera branch off lower down in the cladogram
and must therefore be older. Their ancestors
must have reached Australia earlier, possibly
following the first collision between Australia
and a fragment of the Outer Melanesian Arc,
which is dated at about 25 mya (Pigram and
Davies, 1987; De Boer, 1995c).

For maps of distribution of the Australian
species described here see Moulds (1990) and De
Boer (1995c).

2(1)

Key to males

Tegmen with variable venation, often differing between left and
right tegmen of individuals, forming 9 or more apical areas and
several subapical areas between apical and ulnar areas ..... 2
Tegmen venation regular, with 8 or 9 apical areas, but without
RADAMI ACAS e, riarmo e rre te doda Feta etre ole co 10
Tegmen hyaline, with fairly broad hyaline border along hind mar-
gin and broad costal area. Proximal spine of fore femur erect (fig.
20). Postclypeus not swollen ventrally (figs 4, 45). Pronotum
smooth, without distinct medial furrow .................. 3
Tegmen opaque or weakly reddish or greenish tinged, with nar-
row border along hind margin and narrow costal area. Proximal
spine of fore femur strongly bent, adjacent to femur (fig. 60).
Postclypeus distinctly swollen ventrally (fig. 70). Pronotum with
distinct tedial furrows: ceto acute ss e sa cube pr MG 7
Aedeagus Z-curved in apical part, proximal part straight (cf. fig.
12). Tymbal with 9 or more ridges .................ssse. 4
Aedeagus S-curved (figs 31, 42). Tymbal with 6-8 ridges ... 6
Head and thorax without colour markings. Tymbal with 9-10
ridges. Clasper base forming a low collar around base of anal
valves, which is sloping proximally of aedeagus (fig. 6). Aedeagus
not incised at apex (figs 11, 12) ....... Chlorocysta vitripennis

5(4)

603)

7(2)

8(7)

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

Head and thorax with pattern of dark brown spots and stripes.
Tymbal with 11-12 ridges. Clasper base forming a high collar
around base of anal valves, which is erect proximally of aedeagus
(figs 14, 25). Aedeagus incised at apex (figs 18, 27) ........ 5
Distomedial corner of operculum not reaching medially of mer-
acanthus (fig. 24). Ventral part of postclypeus darkly suffused.
Tymbal with 12 ridges, 4 of which do not reach ventral tymbal
EMOTES gi do 22554 E Chlorocysta suffusa
Distomedial part of operculum almost pointed, elongate, reach-
ing medially of meracanthus (fig. 15). Ventral part of postclypeus
not darkly suffused. Tymbal with 11 ridges, 4 of which do not
reach ventral tymbal margin ............. Chlorocysta fumea
Body length over 28 mm. Abdomen strongly inflated. Seventh
tergite wedge- shaped in lateral view, long dorsally and short at
ventral margin; pygofer turned to a ventral position. Distal part of
operculum longer than basal part and reaching beyond apex of
meracanthus (fig. 44). Wing with 6 apical areas .............

TU SG NUT T SM APES. Ate SUA Ao inj Glaucopsaltria viridis
Body length under 18 mm. Abdomen weakly inflated. Seventh
tergite not wedge- shaped in lateral view; pygofer directed pos-
teriad. Distal part of operculum much shorter than basal part and
not reaching to apex of meracanthus (fig. 33). Wing with 5 apical
BIL 1958 E D rer de TIE T Owra insignis
Tegmina rounded at apex, venation not reticulate. Wings with 6
apical areas. Pronotum not forming a sharply edged lateral crest.
Head and pronotum not triangle-shaped; anterior margins of
postclypeus and vertex lobes not forming a continuous line with
lateral margins of pronotum. Cruciform elevation of mesonotum
broader than long across its centre. First and second sternites not
adjacent. Clasper straight, with broad domed clasper hollow and
with laminiform and sharply pointed medial protrusion. Tymbal
WIEN ORISSA nl ONE V LONE iter less Thaumastopsaltria globosa
Tegmina pointed at apex, venation reticulate in distal half. Wings
with more than 6 apical areas. Pronotum forming a sharply edged
lateral crest. Head and pronotum triangle-shaped; anterior mar-
gins of postclypeus and vertex lobes forming an almost continu-
ous line with lateral crest of pronotum. Cruciform elevation of
mesonotum narrower than long across its centre. First and second
sternites adjacent. Clasper hook-shaped, with narrow clasper hol-
low in downwardly directed apical part and without medial pro-
trusion. Tymbal with 7 or more ridges ................... 8
Veins CuA and M not fused at corner of basal area (fig. 72). First
radial area of tegmen divided by longitudinal vein. Venation of
apical and ulnar areas reticulate. Pygofer opening narrow, V-
shaped (fig. 78). Aedeagus with dorsal appendages near its apex
(figs 85, 86). Tymbal with 10 ridges .Cystopsaltria immaculata
Veins CuA and M fused at corner of basal area (fig. 81). First
radial area of tegmen not divided. Venation of apical areas reticu-
late, ulnar areas normal. Pygofer opening broad, U-shaped (fig
57). Aedeagus without dorsal appendages (figs 62, 77). Tymbal
LE ete muss ER HD E Cs uz SE m cui 9

Body length over 40 mm. Abdomen globularly inflated. Medial
margin of operculum not reaching medially of meracanthus (fig.
nJ uiuo Sed kk a a o's her eR exter des Cystosoma saundersii
Body length under 30 mm. Abdomen distinctly but not globularly
inflated. Medial margin of operculum reaching medially of mer-
acanthus-(fig. M) ess iS mas dle debis Cystosoma schmeltzi
Wing with extremely slender anal fields; 2nd anal field rudimen-
tary, not hyaline. Abdomen with reddish ventral band. Tergites
sharply folded towards sternites ventrolaterally. Ventral part of
pygofer not incurved to its margins; ventral and basal margins of
pygofer forming a continuous, sharply edged, ridge ....... Il
Wing with broad anal fields; 2nd anal field distinct and hyaline.
Abdomen without reddish ventral band. Tergites gradually
curved towards sternites ventrolaterally. Ventral part of pygofer
gradually incurved to its margins; ventral and basal margins of
pygofer not forming a sharply edged ridge ............... 12
Tymbal with 5 ridges. Tymbal and lateral part of 2nd tergite sep-
arated by broad wedge-shaped cavity. Second tergite strongly
curved between auditory capsule and 2nd sternite. Caudodorsal
beak erect and conically shaped; its lateral margins curving
inwards and fusing. Aedeagus short and almost straight, directed

AA eet aed atr euh Peu acie MÈ n Sra fh eR Gymnotympana rufa
Tymbal with 4 ridges. Tymbal and lateral part of 2nd tergite sep-
arated by narrow gap. Second tergite straight between auditory
capsule and 2nd sternite. Caudodorsal beak strongly curved, but
not conically shaped, its lateral margins not fusing. Aedeagus long
and strongly curved down, reaching beyond ventral margin of
DysOfere 58 342 so aa eius ea le ack Gymnotympana varicolor

96 A. J. DE BOER
9(8)
10(1)
11(10)
upwards, not reaching beyond ventral margin of pygofer
12(10)

Body length over 20 mm. Operculum large, covering most of
tymbal cavity and reaching medially of meracanthus. Basal part
of operculum medially distinctly longer than laterally (fig. 55).
Tegmina slightly bronzed, with 9 apical areas. Clasper broad and
lobate, with large clasper hollow and strongly curved down, not
forming a shaft around aedeagus (fig. 51). Aedeagus erect, its apex
adjacent to base of anal valves. Aedeagus broad, swollen in proxi-
mal part and without subapical lobe ....... Venustria superba
Body length under 16 mm. Operculum small, not covering tymbal
cavity and not reaching medially of meracanthus. Basal part of
operculum oblong. Tegmina clear hyaline, with 8 apical areas.
Clasper long, slender and straight, without distinct clasper hol-
low, directed posteriad and forming a shaft around aedeagus.
Aedeagus directed posteriad between claspers, its apex near apex
of clasper. Aedeagus very slender, with distinct subapical lobe.
yon prise RÀ Fue een hn ENTE ATI USER SENE Guineapsaltria flava

Chlorocysta, Glaucopsaltria and Owra

Description. The species of Chlorocysta and
Glaucopsaltria are large compared to other
Chlorocystini, body length 20-33 mm; only
Owra is distinctly smaller, under 16 mm. Body
reddish brown or greenish, generally uniformly
coloured, but abdomen often with reddish seg-
mental hind margins and a row of dark coloured
ventrolateral spots. Male abdomen strongly

inflated 1.4-2.3 X as long as head and thorax
together, in females 0.9-1.2 X. Head broad and
short (fig. 3), 2.4-3.0 X as wide as long and 1.8-
2.2 X as wide as distance between eyes. Postcly-
peus broad and bluntly rounded anteriorly, 2.3-
3.3 X as wide as long. Anterior margin of post-
clypeus continuous with anterior margins of ver-
tex lobes. Sides of postclypeus with about 8-10
distinct furrows, ending in short rows of short
parallel ridges, which form a narrow band along

PHYLOGENY AND BIOGEOGRAPH'* OF AUSTRALIAN CICADAS 97

lorum. Postclypeus not swollen ventrally;
anterior margin (lateral view) straight or weakly
convex (fig. 4). Vertex very broad and smooth;
diverging fissures from centre of head to corners
of postclypeus weakly developed. Vertex lobes
sometimes with some weak longitudinal
wrinkles. Vertex 1.7-2.4 X as wide as long; 1.3-
1.6 X as wide as postclypeus and 1.6-2.4 X as
wide as eye. Ocelli small and far apart. Distance
between lateral ocelli 2.4—4.1 X width of frontal
ocellus and 0.8-1.2 X distance between lateral
ocellus and eye. Pronotum 2.3-2.6 X as wide as
long and fairly smooth, without medial furrow.
Pronotal collar laterally weakly amplified and
slightly curving down at anterior margin of lat-
eral amplifications. Tegmina and wings hyaline,
venation reddish or sometimes ochraceous. Teg-
men venation variable with 9-15 apical areas
and a more or less continuous band of subapical
areas. Costal area hyaline and very distinct,
widening towards tegmen apex. A distinct hyal-
ine border along hind margin of tegmen (fairly
narrow in Owra). Wings with 5, in Glaucopsal-
tria 6, apical areas and a distinctly broader hyal-
ine border than in tegmina. Legs ochraceous and
unmarked. Fore femur (fig. 20) with row of 3
erect and sharply pointed spines, diminishing in
length towards tibia. Tymbal with 6-12 parallel
sclerotized ridges. The most proximal ridges
often not reaching ventral margin of tymbal.
Short intercalary ridges forming a lateral band at
half- width across tymbal. Opercula very small.
Basal part of operculum slightly vaulted with
two rounded elevations and weakly wedge-
shaped; longest medially and gradually tapering
towards lateral margin. Basal part with distinct
crest around its distolateral corner, lateral part
of this crest very short and in males often glo-
bularly swollen. Distal part of operculum in
males angularly oblong and longer than basal
part (though extremely short in Owra) and not,
or only partly covering tymbal cavity in ventral
view; often leaving most of folded membrane
exposed. Lateral margin of male operculum
straight or weakly convex, slightly directed
mesiad and forming a distinct and obtuse angle
with crest of basal part and with straight distal
margin. The straight medial margin lies laterally
of meracanthus. Meracanthus fairly short, but
generally reaching beyond operculum. Female
operculum shorter than in male and sickle-
shaped. Male abdomen very delicate and dis-
tinctly inflated. First tergite in male medially
often more than half as long as 2nd tergite (but
Glaucopsaltria with extremely long 2nd tergite)
and not hidden under metanotum. Middorsal

part of second tergite hardly longer than lateral
parts, proximal margin of second tergite weakly
convex medially. Lateral parts of 2nd tergite
weakly swollen at anterior margins and adjacent
to tymbals. Tergite part between auditory cap-
sules and sternite 2 almost straight, forming a
distinct ridge along tymbal cavity. Sternites 1
and 2 not adjacent (fig. 13 arrow). Auditory cap-
sules not swollen, hardly elevated relative to
connecting bar between abdomen and tymbal.
Female abdomen shorter and more robust than
that of male, with short and broad pygofer (fig.
22). Ovipositor sheaths almost reaching to apex
of sharply pointed caudodorsal beak. Male
pygofer short and rounded, with convex distal
and ventral margins. Caudodorsal beak short
and erect; not curving over basal part of claspers
or anal valves and bluntly rounded, truncate, or
rectangular at apex. Lateral lobes of pygofer
weakly curved inwards with small, weakly
inflated, protuberances. Ventral margins of
pygofer converging, generally forming a sharp
angle at base of pygofer opening. Claspers fairly
slender and hook-shaped; sharply curving down
at half-length. Apical part of clasper with large,
sharply edged, clasper hollow (in Owra without
clasper hollow). Claspers weakly diverging
towards rounded apices. Basal part of clasper
forming a continuous ring-shaped collar around
base of anal valves. Aedeagus S-curved (in
Chlorocysta more Z-curved in apical part), with
very narrow lateral crests along proximal part.

Monophyly of the three genera. In the follow-
ing discussion figures in parentheses refer to
apomorphies in the cladogram (fig. 2). Chloro-
cysta, Glaucopsaltria and Owra form a mono-
phyletic group based on the following synapo-
morphies: 1, a continuous band of subapical
areas in tegmen; 2, a wedge shaped basal part of
operculum; and 3, small male auditory capsules
(De Boer, 1995d).

C. fumea

7 C. suffusa
C. vitripennis
O. insignis
G. viridis

Figure 2. Cladogram of the species of Chlorocysta,
Glaucopsaltria and Owra. Numbers refer to apomor-
phies discussed in the text.

$5 2*3

The tegmen venation tends to be variable and
often differs between left and right tegmen
within individuals. The number of apical areas

98 ^. J. DE BOER

varies from 9-10 in Owra to 11-13 in Chloro-
cysta and to 13-15 in Glaucopsaltria but 1s
alwavs more than 8, the number most common
in related genera and in most cicadas. Further-
more, the tegmina have a more or less continu-
ous band sometimes of a double row of subapical
areas between apical and ulnar areas. Similarly
high numbers of apical areas in the tegmina are
also found in Baeturia inconstans De Boer
(De Boer, 1994c), Gymnotympana (4 species, De
Boer. 1995a), Mirabilopsaltria (2 species, De
Boer, 1996) and all but one species of Thaumas-
topsaltria (De Boer, 1992b). A continuous band
of subapical areas is regarded apomorphic for
Chlorocysta, Glaucopsaltria and Owra (1 1n fig.
2). Although in the tegmina of the species of
Gymnotympana, Mirabilopsaltria and Thau-
mastopsaltria noted above some subapical areas
occur, these never form a continuous band.
Subapical areas do not occur in Baeturia.

The basal part of the male operculum is
slightly wedge-shaped, longest medially and
slightly tapering to its lateral margin. The oper-
culum base is generally oblong in species of
Chlorocystini but in most species of Gymnotym-
pana the operculum base has its greatest length
laterally (De Boer, 1995a). Only in Gymnotym-
pana rufa, G. varicolor and Venustria superba the
basal part of operculum is longest medially but
narrows more abruptly at about one third of its
width. The weakly wedge-shaped operculum
base is regarded apomorphic for Chlorocysta,
Glaucopsaltria and Owra (2).

Chlorocysta, Glaucopsaltria and Owra have
small and weakly protruding auditory capsules.
Although similarly unswollen capsules are pres-
ent in Thaumastopsaltria (De Boer, 1992b) they
presumably represent a parallel development
and can be regarded as apomorphic for this
group (3).

Phylogenetic relationships. Chlorocysta and
Owra share wings with 5 apical areas synapo-
morphic for these genera (4). Wings with 5 apical
areas occur in several genera of cicadas but only
sporadically in other genera of the Chlorocys-
tini.

The three species of Chlorocysta share a some-
what Z-curved aedeagus (a modification of the
S-curved aedeagus, an apomorphy of the Chlor-
ocystini) and a high number (9-12) of tymbal
ridges (5, 6) as apomorphies. Owra insignis
shares an incised aedeagal apex with C. fumea
and C. suffusa but similarly incised aedeagal
apices occur in several related genera (e.g., Aed-
eastria, Guineapsaltria and Papuapsaltria). C.
fumea and C. suffusa are supposed to be sister

species sharing a large tymbal with 11-12 ridges
(7). Furthermore, these species share a wide
pygofer opening and a distinct colour pattern on
head and thorax but the phylogenetic value of
these characters is not clear since they also occur
in several related genera.

Chlorocysta Westwood

Cystosoma (Chlorocysta) Westwood, 1851: 208. —
Walker. 1852: 1133.

Chlorocysta.-Stal, 1863: 575. — Goding and Frog-
gatt, 1904: 566, 596, 658. — Distant, 1906: 153, 159.
— Boulard, 1979: 35. — Duffels and Van der Laan,
1985: 249, — Moulds, 1990: 185-186, — De Boer,
1990: 64. — De Boer, 1991: 2-3. — De Boer, 1992a:
164. — De Boer, 1993a: 16-17. — De Boer, 1993b:
142. — De Boer, 1994a: 3. — De Boer, 1995a: 4, 8, 24.
— De Boer, 1995b: 6.

Glaucocysta Goding and Froggatt, 1904: 566.

Mardalana Distant, 1905: 213, 215. — Distant,
1906: 154, 159. — Metcalf, 1963: 257. — Duffels and
Van der Laan, 1985: 249.

Mardarana [sic] Kato, 1932: 185. — Kato, 1956:
70.

Mardalena [sic] Boulard, 1979: 46.

Type species. Cystosoma vitripennis Westwood,
1851.

Diagnosis. Green body. Basal part of operculum
wedge shaped. Male abdomen strongly inflated.
Male auditory capsules weakly inflated. Teg-
mina with more than eight apical areas and con-
tinuous band of subapical areas. Tymbal with
9-12 ridges. Aedeagus in lateral view Z-shaped
near apex.

Remarks. Chlorocysta, originally described as a
subgenus of Cystosoma, is the type genus of
Chlorocystini Distant, 1905. The genus contains
three species but several undescribed species
have been distinguished (Moulds, pers. comm.).
The peculiarly Z-curved aedeagus is a supposed
apomorphy for Chlorocysta,

Chlorocysta vitripennis (Westwood)
Figures 3-13

Cystosoma | (Chlorocysta)
1851: 208.

Chlorocysta. vitripennis. — Goding and Froggatt,
1904: 566, 659. pl. xix fig. 6. — Distant, 1906: 159. —
Burns, [957: 643. — Metcalf, 1963: 256-257. — Duf-
fels and Van der Laan, 1985: 249, — Moulds, 1990:
188-189, pl. 22 figs 2, 2a-b. — De Boer, 1995a: 16, 77.
— De Boer, 1995b; 5.

Cicada congrua Walker. 1862: 303-304. — Goding
and Froggatt. 1904: 611, 657.

Chlorocysta macrula Stàl, 1863: 575. — Goding and
Froggatt. 1904: 566, 659, 660.

vitripennis Westwood,

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 99

Figures 3-13. Chlorocysta vitripennis Westwood, 1851: 3, male head in dorsal view, Mt Tamborine; 4, male
postclypeus in lateral view, Mt Tamborine; 5, pygofer in lateral view, Mt Tamborine; 6, claspers and aedeagus,
Mt Tamborine: 7, pygofer in oblique view, Mt Tamborine; 8, male caudodorsal beak in dorsal view, Mt Tam-
borine; 9, female operculum, Dorrigo; 10, female caudodorsal beak in dorsal view, Dorrigo; 11, aedeagus in
lateral view, Mt Tamborine; 12, aedeagus in lateral view, ex. coll. v Voixem; 13, male operculum, Mt Tamborine,
arrow indicating gap between sternites 1 and 2.

bp = basal part of operculum: c = crest around distolateral corner of basal part of operculum; cb = caudodorsal
beak; dm = distal margin of operculum; di = distal margin of pygofer; do = dorsal margin of pygofer; dp = distal
part of operculum: ho = clasper hollow; la = lateral margin of operculum, m = medial margin of operculum; me =
meracanthus; mc = medial corner of operculum: pr = protuberance on lateral lobe of pygofer; ve = ventral margin

of pygofer.

102 A. J. DE BOER

Figures 14-24. 14-19, Chlorocysta fumea Ashton, 1914: 14, pygofer in lateral view; 15, operculum; 16, cau-
dodorsal beak in dorsal view: 17, aedeagus in lateral view; 18, aedeagal apex; 19, claspers. 20-24 Chlorocysta
suffusa Distant, 1907: 20, male fore femur in lateral view, Cairns; 21, female operculum, Cooktown; 22, female
genital segment in lateral view, Cooktown; 23, female caudodorsal beak in dorsal view, Cooktown; 24, male
operculum, Julatten.

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

an almost right angle with slightly longer distal
margin. Distal margin straight, but weakly con-
vex near angular distomedial corner. Medial
margin straight. Distal part of operculum
elongate at distomedial corner, reaching mesiad
beyond apex of meracanthus. Meracanthus
short, not reaching distal margin of oper-
culum.

Abdomen. Strongly inflated, almost uni-
formly brown coloured, but with darkened
segmental hind margins and a lateral row of
darkened spots.

Genitalia. Pygofer in lateral view (fig. 14)
short and globularly swollen. Dorsal margin
weakly concave, convexly bending into erect
caudodorsal beak. Distal margin slightly convex
between beak and lateral protuberance. Ventral
margin angularly convex. Ventral margins con-
verge to a bluntly rounded angle at base of
pygofer opening. Caudodorsal beak in dorsal
view (fig. 16) with subapical swelling and
pointed apex. Lateral lobe of pygofer with small
weakly swollen lateral protuberance. Claspers
(fig. 19) fused at base to a continuous and broad
collar around base of anal valves. Dorsal margin
of clasper ending in right angle on surface of this
collar and not forming a lateral crest on lateral
surface of clasper base. Collar erect, not domed,
between claspers. Distal corner of clasper bend-
ing mesiad around aedeagus, supporting aedea-
gus in upright position. Claspers sharply curving
down and strongly diverging towards apices.
Apical part of clasper with sharply edged hollow
at its inwardly directed side. Aedeagus (fig. 17)
slender and apically rounded in lateral view,
strongly Z-curved in apical third, basal two-
thirds almost straight. Aedeagus with very slen-
der lateral crests and without dorsal swellings.
Aedeagal apex with dorsoventral incision; aed-
eagus ending in two small and almost pointed
lateral lobes (fig. 18).

Measurements: Body length: 29.8 mm; teg-
men length: 31.7 mm; head length: 2.3 mm;
pronotum length: 2.8 mm; mesonotum length:
6.2 mm; head width: 6.5 mm; width of pronotal
collar: 7.2 mm.

Distribution. Endemic to the Cape York Penin-
sula, Qld. Moulds (1990) recorded the species
only known from along the old Leo Creek track
at the southern end of the Mcllwraith Range, at
an altitude of around 300 m.

Remarks. C. fumea is the largest species of this
genus and is easily recognized by its distome-
dially elongated male operculum. C. fumea is
closely related to C. suffusa, described next,

103

sharing a similar colour pattern and incised aed-
eagal apex. Only one male was available for
study.

Chlorocysta suffusa (Distant)
Figures 20-29

Mardalana suffusa Distant, 1907: 418. — Burns,
1957: 644. — Metcalf, 1963: 258. — Duffels and Van
der Laan, 1985: 249,

Chlorocysta suffusa. — Moulds, 1990: 186-187, pl.
22 figs 7, 7a.

Material examined. Australia. S. Qld, Nauklér. coll.
Dodd, d, ZMH. Cairns 9 km W, ii-iv. 1985, d, ZMA.
Cairns, A.P. Dodd d Det. Chlorocysta vitripennis G+F.
BPBM. Cairns, 1920, J.F. Illingworth, d, 9 Det. Chlor-
ocysta vitripennis G+F, BPBM. Clohesi River, S.F. SW
of Kuranda, 18.1.1984, M.S. and B.J. Moulds, ¢ C. suf-

fusa det. M.S. Moulds, o C. suffusa det. M.S. Moulds,

Moul. Gordonvale, J.F. Illingworth, 3, o, BPBM. Julat-
ten, 10.xi.1979, M.S, and B.J. Moulds, o C. suffusa det.
M.S. Moulds, Moul. Same data but 20.x1.1979, d C.
suffusa det. M.S. Moulds, Moul. 15.1.1981, ¢ Mardal-
ana suffusa det, M.S. Moulds, ZMA. 17.1.1981, ¢ C.
suffusa det. M.S. Moulds, Moul. 24,1.1981,9C. suffusa
det. M.S. Moulds, Moul. 13,111.1982, d C. suffusa det.
M.S. Moulds, Moul. Kamerunga, nr Cairns, 14.1.1977,
M.S. and B.J. Moulds, g C. suffusa det. M.S. Moulds,
Moul. Kuranda, 200m, 12.11.1956, J.L. Gressitt, d,
BPBM. Kuranda, 14.11.1988, J. Hasenpusch, & C. suf-

fusa det. M.S. Moulds, Moul. Quarantine Bay, nr

Cooktown, 17.11.1982, M.S. and B.J. Moulds, 9 C. suf-

fusa det. M.S. Moulds, Moul. Tully Falls, S of Raven-

shoe, 11.1.1977, M.S. and B.J. Moulds, 9 C. suffusa det.
M.S. Moulds, Moul. Upper Smithfield, nr Cairns,
7.1.1984, M.S. and B.J. Moulds, 9 C. suffusa det. M.S.
Moulds, Moul. Windsor Tableland, NW of Moss-
mann, 10.1.1984, M.S. and B.J. Moulds, € C. suffusa
det. M.S. Moulds, Moul. Woobadda R., S of Blooms-
field, 13.1.1984, M.S. and B.J. Moulds, & C. suffusa det.
M.S. Moulds, 9 C. suffusa det. M.S. Moulds, Moul.

Description. Body ochraceous to brown, prob-
ably green when alive and covered with short
brown hairs (for photographs see Moulds, 1990:
pl. 22 figs 7, 7a). Head and thorax with indistinct
pattern of brown stripes. Females with conspicu-
ously striped abdomen. Abdomen in males 1.4-
1.6 x head and thorax together, in females 1.0—
1.2 x. Tegmina of males 1.1—1.2 x body length, of
females 1.3-1.4 x.

Head. With similar colour pattern as in
C. fumea, though strongly varying in intensity.
Vertex light brown, darker brown around lateral
ocelli and sometimes around frontal ocellus and
with dark brown spots bordering lateral corners
of postclypeus. Vertex lobe with 2 dark brown
spots between lateral ocellus and eye and a dark
spot bordering proximomedial corner of eye.

104 A. J. DE BOER

Figures 25-37. 25-29, Chlorocysta suffusa Distant, 1907: 25, pygofer in lateral view, Julatten; 26, pygofer in
oblique view, Julatten; 27, claspers and aedeagus, Julatten; 28, male caudodorsal beak in dorsal view, Julatten;
29, aedeagus in lateral view, Julatten.

30-37 Owra insignis Ashton, 1912: 30, pygofer in lateral view; 31, aedeagus in lateral view; 32, male caudodorsal
beak in dorsal view; 33, male operculum; 34, female caudodorsal beak in dorsal view; 35, pygofer in oblique view;
36, clasper; 37, female operculum.

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

Vertex lobes slightly darkened on anterior parts.
Postclypeus in dorsal view dark brown at lateral
corners and along anterior margin. Ventral part
of postclypeus with large dark medial spot. Lat-
eral surface of postclypeus with 9-10 irregular
rows of short parallel ridges continuing into ten
parallel streaks running towards central
fissure.

Thorax. Pronotum ochraceous brown, with a
broad lighter ochraceous coloured medial band
that is not, or only slightly, dilating towards dis-
tal pronotal margin. Proximal end of this band
marked by a pair of small dark brown parame-
dian spots at pronotal collar. Pronotum light
brown along oblique fissures and towards
medial band. Mesonotum greyish brown, only
slightly darkened in paramedian spots at prono-
tal margin and in vague reticulate pattern of
angular brown spots, forming lateral streaks
from pronotal margin to corners of cruciform
elevation. Cruciform elevation greenish
ochraceous, with brown medial band. 2 black
spots in front of elevation.

Tegmina and wings: Hyaline. Tegmen ven-
ation variable, differing even between left and
right tegmen of individuals. Tegmina with 12-
13 apical areas and a, at some places double,
band of 6-8 subapical areas. Wings with 5 apical
areas.

Tymbals. 12 dark brown parallel sclerotized
ridges; 8 ridges spanning the tymbal from dorsal
to ventral margin, the 4 most proximal ridges do
not reach ventral margin and successively short-
ening, with most proximal ridge shortest. 12
short and lighter coloured intercalary ridges
seem to form a lateral band across tymbal.

Operculum. Male operculum (fig. 24) closely
resembling that of C. fumea in size, but with a
more rectangular shaped distal part, not
elongated at distomedial corner. Basal part of
operculum slighly vaulted and wedge-shaped;
longest medially and tapering towards its lateral
margin. Basal part forming a distinct, though
very short crest around rectangular distolateral
corner. Distal part of operculum almost rec-
tangular, adjacent to body and not covering tym-
bal cavity in ventral view, though, as in C.
fumea, covering most of folded membrane. The
straight and almost parallel lateral and medial
margins both making an almost right angle with
straight distal margin. Meracanthus short, not
reaching to distal margin of operculum. Female
operculum (fig. 21) short, sickle-shaped and
erect.

Abdomen. Male abdomen strongly inflated,
almost uniformly brown to green coloured, seg-

105

mental hind margins often darker coloured.
Ventrolateral row of slightly darkened spots on
segments 3-7. Female abdomen with pattern of
broad longitudinal lateral streaks similar as in
Gymnotympana strepitans (De Boer, 1995a) and
a distinct dark brown ventromedial band.
Female caudodorsal beak reaching to apex of
ovipositor sheaths (fig. 22) and triangular in dor-
sal view, somewhat swollen to its base and
sharply pointed at apex (fig. 23).

Male genitalia. Pygofer in lateral view (fig. 25)
short and globularly rounded. Dorsal margin
slightly concave, continuous with straight and
slender caudodorsal beak. Distal margin slightly
convex between beak and lateral protuberance.
Ventral margin straight, but forming a rounded
corner under lateral protuberance. Ventral mar-
gins converging and forming a sharp angle at
base of pygofer opening (fig. 26). Caudodorsal
beak in dorsal view (fig. 28) short, triangular and
pointed or narrowly rounded at apex. Lateral
lobe of pygofer curving inwards, forming a tri-
angularly swollen protuberance, which does not
extend beyond distal margin of pygofer.
Claspers (fig. 27) very similar to C. fumea.
Clasper base fused to a high and continuous col-
lar around base of anal valves. Dorsal margin of
clasper ending in right angle on this collar and
not forming a crest on lateral surface of clasper
base. Clasper base forming collar around base of
anal valves, which abruptly, but slightly, curves
inwards between claspers. Claspers forming
diverging ridges from dorsal margins to base of
anal valves. Distal corner of clasper bending
mesiad around aedeagus, supporting aedeagus
in upright position. Claspers curving down and
strongly diverging towards apices. Apical part of
clasper with sharply edged hollow at its inwardly
directed side. Aedeagus apically rounded in lat-
eral view (fig. 29), slender and Z-curved in apical
third. Basal two-thirds of aedeagus almost
straight, with very slender lateral crests and
slightly swollen dorsally, at distal ends of lateral
crests. Aedeagal apex with dorsoventral
incision; aedeagus ending in two small, rounded,
lateral lobes (fig. 27).

Measurements (mean + sd): Body length c:
24.2-27.9 mm (26.2 mm + 1.1), 9: 20.0-21.5
mm (20.5 mm + 0.6); tegmen length c: 28.2-
31.2 mm (30.0 mm + 1.1), 9: 26.3-28.3 mm
(27.5 mm + 0.7); head length 4: 2.1-2.5 mm (2.3
mm), 9: 1.9-2.5 mm (2.2 mm); pronotum length
d: 2.5-2.8 mm (2.7 mm), 9: 2.7-2.8 mm; meson-
otum length d: 5.4-6.4 mm (5.8 mm), o: 4.9-5.6
mm (5.3 mm); head width e: 6.0-6.7 mm (6.4
mm), 9: 6.0-6.6 mm (6.3 mm); width of pronotal

106 A. J. DE BOER

collar d: 6.7-7.5 mm (7.0 mm), 9: 6.6-7.3 mm
(6.9 mm).

Distribution. Along the eastern coast of the Cape
York Peninsula, Qld. Moulds (1990) recorded
the species from Iron Range, the Mcllwraith
Range near Coen and Cooktown south to the
Paluma Range.

Remarks, C. suffusa closely resembles C. fumea
in tymbal shape and colour pattern but can be
easily separated from that species by a dark
brown suffused spot on the ventral side of the
postclypeus. Males of C. suffusa are easily sep-
arated from C. Jumea by their rectangular oper-
culum.

Owra Ashton

Owra Ashton, 1912: 224. — Metcalf, 1963: 252. —
Duffels and Van der Laan, 1985: 248. — Moulds,
1990: 184-185. — De Boer, 1992b: 18, 19, 20. — De
Boer. 1993a: 16-17. De Boer, 1993b: 142. — De
Boer, 1995a: 8. — De Boer, 1995b: 6.

Owra

Type species. insignis Ashton, 1912

(monotypic),
Owra insignis Ashton
Figures 30-37

Owra insignis Ashton, 1912: 224, pl. Ll figs 6-6a. —
Burns. 1957, 642, — Metcalf, 1963: 252. — Duffels
and Van der Laan, 1985: 248. — Moulds, 1990: 185,
pl. 22 figs 6, 6a,

Material examined. Australia. Lake Barine, 530 m,
31.i-1.ii.1964. J. Sedlacek, 9, BPBM; Cairns, A.P.
Dodd, Det. Chlorocrsta macrula Stal, BPBM.
Description. Body reddish brown but according
to Moulds (1990) olive green when alive (for
photographs see Moulds, 1990: pl. 22 figs 6, 6a).
Abdomen in male 1.5 x head and thorax
together, in female 1.0 x. Tegmina of male 1,1 x
body length. of female 1.4 x.

Tegmina and wings. Hyaline, venation red-
brown and costa bright red. Tegmina with 10
(according to Moulds, 1990, sometimes 9) apical
areas and a regular band of 3 or 4 subapical
areas. Costal area very broad. Tegmen with very
narrow hyaline border along hind margin. Wing
with 5 apical areas and broad hyaline border
along hind margin.

Tymbals. 5 parallel sclerotized ridges span-
ning the tymbal from dorsal to ventral margin
and a 6th, most proximal ridge, almost reaching
ventral margin. 5 distinct intercalary ridges
seem to form a lateral band across tymbal.

Opercula. Male operculum (fig. 33) extremely
small, not covering tymbal cavity in ventral view
and leaving folded membrane completely

exposed. Basal part of operculum about 4 x as
long as distal part, slightly vaulted and with dis-
tinct crest around rectangular distolateral cor-
ner. Basal part wedge-shaped; medially dis-
tinctly longer than laterally. Distal part of male
operculum very short, sickle-shaped and erect,
hardly more than a continuation of the crest
around distolateral corner of basal part. Mera-
canthus reaching well beyond operculum.
Female operculum (fig. 37) very similar to that
of male, with extremely short and sickle-shaped
distal part,

Abdomen. Male abdomen slender and weakly
inflated, almost unicoloured ochraceous, but
segmental hind margins orange-brown. Second
tergite (fig. 33) weakly curved along tymbal cav-
ity. Female abdomen light brown with reddish
segmental hind margins and ventrolateral row of
brown spots on segments 3-8. Ovipositor
sheaths not reaching to apex of caudodorsal
beak. Female caudodorsal beak in dorsal view
(fig. 34) broad and bluntly rounded at apex.

Male genitalia. Pygofer in lateral view as in fig.
30. Dorsal margin of pygofer straight, strongly
concave to base and angularly bending into
straight, slender and erect caudodorsal beak.
Distal margin slightly convex, angularly bending
into margin of beak. Lateral lobe of pygofer
strongly curving inwards towards end of distal
margin and forming an angularly rounded and
swollen lateral protrusion. Ventral margin of
pygofer strongly convex near this protrusion,
but almost straight towards base of pygofer.
Ventral margins converging to short and straight
basal margin at base of pygofer opening (fig. 35).
Caudodorsal beak in dorsal view (fig. 32) short
and broad, truncate at apex. Claspers very dif-
ferent from those of related species, missing a
clasper hollow. Clasper (fig. 36) broad, globu-
larly swollen near base, strongly curved down to
very slender and elongate, almost laminiform
apical part. Claspers strongly diverging towards
their truncate apices. Clasper base forming a low
collar around base of anal valves. Aedeagus (fig.
31) strongly upcurved at half-length and with
slender lateral crests. Aedeagal apex incised, as
in C. fumea and C. suffusa, but ending in two
more sharply pointed lateral lobes (fig. 35).

Measurements. Body length g: 15.1 mm, 9: 13.3
mm; tegmen length d: 17.0 mm, 9: 19.0 mm;
head length s; 1.6 mm, 9: 1.7 mm; pronotum
length d: 1.6 mm, 9: 1.8 mm; mesonotum length
d: 3.0 mm, 9: 3.3 mm; head width d: 4.1 mm, 9:
4.6 mm; width of pronotal collar d: 4.0 mm, 9:
4.6 mm.

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

Distribution. Only known from the south-east-
ern corner of the Cape York Peninsula, north-
eastern Qld. Moulds (1990) recorded the species
from Thornton Peak range north of Daintree to
Mission Beach and widespread on the Atherton
Tableland.

Remarks. Owra insignis is a very small
ochraceous brown species, at first sight closely
resembling Guineapsaltria flava and Gymno-
tympana rufa. Owra, however, is closely related
to Chlorocysta and Glaucopsaltria, sharing the
presumed synapomorphic tegmen venation.
Owra shares the 5 apical areas in the wing with
Chlorocysta. Only one male and one female have
been examined.

Glaucopsaltria Goding and Froggatt

Glaucopsaltria Goding and Froggatt, 1904: 657,
659. — Moulds, 1990: 189 — De Boer, 1992b: 18, 19,
20. — De Boer, 1993a: 16-17, — De Boer, 1993b: 142.
— De Boer, 1995a: 8. — De Boer, 1995b: 6.

Glaucopsaliria in synonymy of Chlorocysta. — Dis-
tant. 1906: 159. — Metcalf, 1963: 255-256, 473.

Type species. Glaucopsaltria viridis Goding and
Froggatt (monotypic).

Glaucopsaltria viridis Goding and Froggatt
Figures 38-44

Glaucopsaltria viridis Goding and Froggatt, 1904:
566.658. — Kirkaldy, 1907: 308 (6). — Moulds, 1990:
190, pl. 22 fig 3. — De Boer, 1995a: 16.

Chlorocysta viridis. — Distant, 1906: 159. — Frog-
gatt. 1907; 352. — Burns, 1957, 643. — Metcalf, 1963:
256. — Duffels and Van der Laan, 1985: 249.

Material examined. Australia. Qld. 24.11.1946, d,
SEM, Brisbane. 1973. J.B. Vogel, 9, ZMA. Qld, J.A.
Grant, BM- CSIRO Expedition, BM 1973-346, 7.
BMNH. Rockhampton. Museum Godefroy No 4836,
g det. C. viridis. ZIM.

Description. Body olive green or yellow-green
(for photograph see Moulds, 1990: pl. 22 fig 3).
Tegmina slightly shorter than body. Abdomen
1.9-2.2 x head and thorax.

Tegmina and wings. Hyaline. Tegmen ven-
ation variable and differing even between left
and right tegmen of individuals. Tegmina with
13-15 apical areas and a continuous band. at
some places concisting of a double row, of 7-10
subapical areas. Costal area very distinct, broad-
ening towards tegmen apex. Tegmen with dis-
tinct hyaline border along hind margin, though
narrower than in wing. Wing with 6 apical
areas.

Legs. Fore femur with row of 3 erect spines
(fig. 43).

107

Tymbals. 6 transverse sclerotized ridges span-
ning the tymbal from dorsal to ventral margin
and a 7th, most proximal, ridge almost reaching
ventral margin. 7 short intercalary ridges seem
to form a lateral band across tymbal.

Operculum (fig. 44). Basal part of operculum
slighly vaulted, wedge-shaped; longest medially
and tapering towards lateral margin. Basal part
forming an irregularly protruding crest around
distolateral corner. Distal part of operculum
longer than basal part and reaching beyond apex
of meracanthus. Distal part only partly covering
tymbal cavity in ventral view, though covering
most of folded membrane. Lateral margin mak-
ing an almost right angle with basal part of oper-
culum and initially almost straihgt, but angu-
larly bending mesiad where operculum curves
towards body. Distal margin straight, making a
sharp angle with straight medial margin. Disto-
lateral corner broadly rounded. Operculum
strongly curved-up along proximal part of its lat-
eral margin (especially in Brisbane specimen),
forming a small lobe (fig. 44 arrow).

Abdomen. Strongly inflated, uniformly green
or discoloured yellow, with traces of green. Sec-
ond tergite medially about 3 x Ist tergite and
laterally gradually tapering to auditory capsules.
Seventh tergite wedge-shaped in lateral view:
very long and convex dorsally (lateral view),
short ventrally.

Genitalia. Pygofer in lateral view as in fig. 39.
Dorsal margin weakly concave, continuous with
erect caudodorsal beak. Distal margin weakly
convex between beak and lateral protuberance.
Ventral margin straight. forming an almost rec-
tangular corner just under lateral protuberance,
concavely bent near base of pygofer opening.
Ventral margins converge and form a sharp
angle at base of pygofer opening (fig. 38). Cau-
dodorsal beak in dorsal view (fig. 41) broad and
triangular, almost rectangular at apex. Lateral
lobe of pygofer bending inwards towards distal
margin, forming an angularly rounded and lat-
erally flattened protuberance. Clasper as in
Chlorocysta, but with longer and more slender
apical part (fig. 40). Apical part of clasper
strongly bent down towards smoothly rounded
apex and with a sharply edged hollow at
inwardly directed side. Dorsodistal corner of
clasper bending mesiad around aedeagus, sup-
porting aedeagus in upright position. Claspers
almost parallel, only slightly diverging towards
apices and fused at base to a fairly low and con-
tinuous collar around base of anal valves.
Medial part of collar curving inwards, slightly
dome-shaped proximally of aedeagus, between

108 A. J. DE BOER

Figures 38-46. 38-44, Glaucopsaltria viridis Goding and Froggatt, 1904: 38, pygofer in oblique view, Australia;
39, pygofer in lateral view, Brisbane; 40, claspers, Australia; 41, caudodorsal beak in dorsal view, Australia; 42,
aedeagus in lateral view, Australia; 43, male fore femur in lateral view, Australia; 44, operculum, Brisbane, arrow
indicating upcurving lateral lobe. 45-46 Venustria superba Goding and Froggatt, 1904: 45, male postclypeus in
lateral view, Kuranda; 46, male head in dorsal view, Kuranda.

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

dorsal crests of claspers. Dorsal margin of
clasper forming a sharply edged crest, continu-
ing on lateral surface of clasper base. Aedeagus
(fig. 42) weakly S-curved, with very short and
narrow lateral crests along proximal half. Aed-
eagal pore oval.

Measurements. (mean): Body length: 29.8-
33.8 mm (32.1 mm); tegmen length: 29.5-31.3
mm (30.0 mm); head length: 2.1-2.4 mm (2.3
mm); pronotum length: 2.7-2.9 mm (2.8 mm);
mesonotum length: 5.1-6.0 mm (5.5 mm); head
width: 6.4-6.5 mm; width of pronotal collar:
6.8—7.0 mm (6.9 mm).

Distribution. Along the eastern coast of Qld and
in north-eastern NSW. Moulds (1990) recorded
the species from Daintree River punt crossing
and Cairns in northern Qld, from Mackay, Car-

lisle Island, Middle Percy Island and Eungella in
central Qld and from the Kolan River north of
Gin Gin to Dorrigo.

Remarks. Glaucopsaltria viridis is larger than the
Chlorocysta species with a strongly inflated
abdomen. G/aucopsaltria is easily separated
from Chlorocysta and Owra, since it has six
(instead of five) apical areas in the wings. Males
of G. viridis can be recognized by the shape ofthe
Tth tergite which is very long middorsally but
short ventrally so that the pygofer is curved to a
ventral position. Females have not been exam-
ined. The local name of this species is Bottle
Cicada (Moulds, 1990).

Venustria Goding and Froggat

Venustria Goding and Froggatt, 1904: 565, 596. —
Distant, 1906: 128, 129. — Metcalf, 1963: 203. —
Duffels and Van der Laan, 1985: 233. — Moulds,
1990: 32, 180. — De Boer, 1990: 64. — De Boer, 1991:
2-3. — De Boer, 1992a: 164. — De Boer, 1992b: 19.
— De Boer, 1993a: 16-17. — De Boer, 1993b: 142. —
De Boer, 1994a: 3. — De Boer, 1994b: 130. — De
Boer, 1995a: 1, 3, 4, 8, 11. — De Boer, 1995b: 5.

Venustra [sic]. — Kato, 1932: 181.

Type species. Venustria superba Goding and
Froggatt, 1904 (monotypic).

Remarks. Distant (1906) included Venustria in
his division Taphuriaria (now the tribe Taphu-
rini, which has as principal characters: “Eyes
projecting beyond anterior margin of pronotum;
pronotum subquadrate, not distinctly narrowed
anteriorly; abdomen about as long as space
between apex of head and base of cruciform
elevation; a more or less distinct posterior
metasternal process visible in males between or
at the base of the opercula" (Distant, 1906).

109

Recently Moulds (1990) transferred the genus to
Chlorocystini since several characters (e.g., fore-
wing vein Sc separated from costa; apical cells
mostly longer than ulnar cells; and an inflated
male abdomen) are “associated with that
tribe".

/enustria has a weakly S-curved aedeagus,
with very narrow lateral crests and should there-
fore be placed in Chlorocystini. The width ofthe
head and shape ofthe pronotum of Venustria are
very similar to those of Chlorocysta and its
metasternum, though more protruding than in
Chlorocysta, does not differ greatly from that of
other genera of the tribe. Males of Venustria,
however, have a very short and rather solid look-
ing abdomen, often shorter than head and tho-
rax together. The eyes are large and angular with
inwardly directed proxomedial corners and the
tegmina are relatively long so that in general
aspect Venustria looks fairly different from all
other species of the tribe.

Venustria superba Goding and Froggatt
Figures 45-55

Venustria superba Goding and Froggatt, 1904: 565,
597, 603, pl. xix figs 7, 7a. — Distant, 1906: 628. —
Burns, 1957: 634. — Metcalf, 1963: 203-204. — Duf-
fels and Van der Laan, 1985: 233. — Moulds, 1990:
180-181, pl. 16, fig. 11. — De Boer, 1995a: 3, 4, 9, 13,
15, 74, 77.

Material. Australia. Australién, 1888, v Müller, 9,
SMN. Etty Bay nr Innisfail, 23.11.1982, M.S. and B.J.
Moulds, à Venustria superba det. M.S. Moulds, ZMA.
Cairns, J.F. Illingworth, 1920, 59, 9 det. Venustria
superba, BPBM. Kuranda 1 ml. E, 11.iii.1964, LF.B.
Common and M.S. Upton, d, CSIRO.

Description. Body light brown (for photograph
see Moulds, 1990: pl. 16, fig. 11). Females about
as long as males, but with more robust head and
thorax and longer tegmina. Abdomen of males
relatively short, 0.8-1.1 X head and thorax
together, of females 0.9-1.0 X. Tegmina fairly
long in males and females, being 1.3-1.5 X body
length.

Head (fig. 46): Ochraceous brown, darkened
towards anterior margins of vertex lobes, with
row of 3 dark spots on vertex lobe between lat-
eral ocellus and eye. Head slightly broader than
anterior part of pronotum, 2.4-3.1 X as broad
as long and 2.1-2.4 X as broad as distance
between eyes. Head distinctly shorter than dis-
tance between eyes. Postclypeus 2.6-3.2 X as
wide as long and smoothly rounded anteriorly,
its anterior margin convex and almost continu-
ous with anterior margins of vertex lobes.

110 A. J. DE BOER

Figures 47-55. Venustria superba Goding and Froggatt, 1904: 47, female genital segment, Cairns; 48, female
operculum. Cairns; 49, female caudodorsal beak in dorsal view, Cairns; 50, pygofer in lateral view, Kuranda; 51,
pygofer in oblique view, Kuranda; 52, aedeagus in lateral view, Etty Bay; 53, male caudodorsal beak in dorsal
view, Kuranda: 54, male fore femur in lateral view, Kuranda; 55, male operculum, Etty Bay.

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS LI

Postclypeus (fig. 45) weakly swollen ventrally,
| anterior margin (lateral view) weakly convex.
Lateral sides of postclypeus with about 12 fur-
rows and 5-6 very short and irregular rows of
short parallel ridges on weakly inflated crest
along the lorum. Vertex smooth with weak
medial furrow and without diverging furrows.
Vertex in males 1.8-2.2 x, in female 2.3-2.4 X
as wide as long and, in both sexes, 1.4-1.8 X as
wide as width of eye. Eyes large and angular;
medial margins of eyes strongly converging:
shortest distance between eyes at their proxo-
medial corners. Ocelli small and wide apart.
Distance between lateral ocelli 0.8-1.0 X dis-
tance between lateral ocellus and eye and 2.8—
3.4 X width of frontal ocellus.

Thorax. Pronotum ochraceous, with dark
streaks in and along oblique fissures and a nar-
row light ochraceous or greenish coloured
medial band bordered by vaguely darkened
streaks. Pronotum without medial fissure, 2.3-
2.7 X as wide as long and, across the collar, 1.1—
1.2 X as wide as head. Amplified lateral corners
of pronotal collar larger than in Chlorocysta.
Mesonotum ochraceous brown, with reticulate
pattern of ochraceous stripes and brown specks,
forming a pair of paramedian semicircular spots
at pronotal margin and a pair of converging lat-
eral bands from pronotal margin to corners of
cruciform elevation. 2 dark spots in front of
cruciform elevation.

Legs. Fore femur (fig. 54) with row of 3 erect
spines, diminishing in length towards tibia.
Most proximal spine very long, longer than
distance to middle spine.

Tegmina and wings. Hyaline, but slightly
bronzed in apical areas. Venation reddish
brown. Tegmen with 9 apical areas and distinct
costal area. Veins CuA and M adjacent, but not
fused near basal area. Wing with 6 apical
areas.

Tymbals. 4 transverse sclerotized ridges span-
ning the tymbal from dorsal to ventral margin, a
5th ridge almost reaching ventral margin and a
6th, most proximal, ridge spanning about three-
quarters of tymbal width. 5 short intercalary
ridges seem to form a lateral band across
tymbal. ]

Opercula. Male operculum (fig. 55) covering
greater part of tymbal cavity in ventral view and
extending medially of meracanthus. Basal part
of operculum medially distinctly longer than lat-
erally, abruptly broadening at about third of its
width and strongly vaulted. Distal part angularly
oblong. Lateral margin long and straight, angu-
larly bent into weakly convex distal margin. Dis-

tomedial and medial margins straight, disto-
medial corner broadly rounded, medial corner
more narrowly rounded, almost rectangular.
Meracanthus very short, hardly reaching beyond
basal part of operculum. Opercula widely separ-
ated medially. Female operculum (fig. 48) very
small. Basal part as in males, medially distinctly
longer than laterally. Distal part shorter than
basal part, its straight lateral margin forming a
broadly rounded angle with weakly convex
distal margin.

Abdomen. Light brown, silvery pilose and
with dark brown midventral band. Males with
darkened 8th and sometimes 7th, tergite and
ventrolateral row of slightly darkened spots on
segments 3-7. Male abdomen more solid in
aspect than in other species of Chlorocystini,
hardly swollen and almost without fold between
tergites and sternites; nearly circular in cross sec-
tion. First tergite very short and partly hidden
under metanotum. Anterior margin of 2nd ter-
gite concave medially. Lateral parts of 2nd
tergite swollen anteriorly and almost adjacent to
tymbal, leaving only a narrow gap between tym-
bal and 2nd tergite. Auditory capsules globularly
swollen and distinctly elevated relative to con-
necting bar between abdomen and tymbal. Ter-
gite part between auditory capsule and sternite 2
almost straight, with distinct crest along tymbal
cavity. First sternite globularly swollen between
opercula. Female abdomen about as large as that
of male, but more strongly tapering towards
apex. Female pygofer (fig. 47) long and slender
with distinct thorn-shaped protuberance at ven-
tral margin. Ovipositor sheaths reaching to apex
of caudodorsal beak. Female caudodorsal beak
in dorsal view (fig. 49) triangular and sharply
pointed at apex.

Male genitalia. Pygofer in lateral view as in fig.
50. Dorsal margin almost straight, but concave
to base and weakly convexly bent into stout and
slightly posteriorly curved caudodorsal beak.
Distal margin angularly convex, concavely bent
into caudodorsal beak and ending in almost
right angle on broad and angular lateral pro-
trusion, Ventral margin strongly concave. Lat-
eral lobe of pygofer with very small angularly
swollen lateral protuberance, distinctly reaching
beyond distal margin of pygofer. Pygofer open-
ing broad (fig. 51); ventral margins ending wide
apart at straight basal margin. Caudodorsal beak
in dorsal view (fig. 53) triangular and sharply
pointed at apex. Clasper (cf. fig. 51) very short
and stout, with rounded lobate and downwardly
directed apical part and broad, sharply edged
inwardly directed, clasper hollow. Dorsal

112 A. J, DE BOER

margin of clasper sharply curving upwards
proximally, merging with broad ring-shaped
clasper base. Basal part of clasper very broad in
lateral view, distinctly swollen around base of
anal valves, especially mid-between claspers,
possibly representing the remnants of a medial
uncus lobe. Aedeagus (fig. 52) weakly S-curved,
strongly swollen in proximal half and curving
down towards apex. Aedeagus strongly widening
near base, forming 2, somewhat lobate, basola-
teral protuberances flanking a large hollow at the
fold between aedeagus and its basal plate. These
protuberances continue into short lateral crests
along proximal part of aedeagus. Aedeagus with
very slender, almost membranous, laminiform
ventrolateral crests, just medially of these lateral
crests, which curve into the ventral hollow
between the basolateral protuberances. Aed-
eagal pore almost round, weakly incised
ventrally.

Measurements (mean + sd): Body length d:
20.6-25.2 mm (x23.4 mm + 1.3), 9: 23.4 and
26.2 mm; tegmen length g: 31.0-34.6 mm (x33.5
mm + 1.1), 9: 35.1 and 35.3 mm; head length c:
2.4-3.1 mm (x2.6 mm), 9: 2.5 and 2.6 mm; pron-
otum length d: 2.9-3.6 mm (x3.3 mm), 9: 3.4 and
3.7 mm; mesonotum length d: 6.0-7.8 mm (x7.0
mm), 9: 6.8 and 8.0 mm; head width g: 6.7-7.7
mm (x7.5 mm), 9: 8.0 and 8.1 mm; width of
pronotal collar 3; 7.6-8.8 mm (8.5 mm), 9: 8.8
and 9.2 mm.

Distribution. Endemic to the south-eastern cor-
ner of the Cape York Peninsula, north-eastern
Old. Moulds (1990) recorded the species from
Mt Hartley approximately 30 km south of
Cooktown to the Kirrama Range north-west of
Cardwell.

Remarks. The species can be recognized by
slightly bronzed tegmina with 9 apical areas.
Males are characterised by short lobate claspers
and females are casily recognized by the sharp,
thorn-shaped, protuberance at the ventral mar-
gins of the pygofer (fig. 47). The local name of
this species is Frog Cicada (Moulds, 1990).

Cystosoma and Cystopsaltria

Description. Body reddish brown, without any
distinct colour markings. Surface of head and
pronotum roughly wrinkled and pitted. Head
and pronotum conically shaped; anterior mar-
gins of postclypeus and vertex lobes forming an
almost continuous straight line with lateral mar-
gin of pronotum, though interrupted by eyes.
Females smaller than males. Abdomen of males
strongly inflated, being 1.3-2.0 X head and tho-

rax, in females 0.9-1.1 X. Male tegmen in Cys-
tosoma 0.8—1.1 X, in Cystopsaltria 1.4 X body
length, in females 1.2-1.4 X. Head (fig. 69) nar-
rower than anterior part of pronotum, with long
and angularly protruding postclypeus. Head
1.9-2.3 X as broad as long. Postclypeus dis-
tinctly protruding, its sharply edged anterior
margin forming a right angle at apex. Postcly-
peus 1.6-2.1 X as broad as long, distinctly
swollen ventrally (fig. 70), its anterior margin
(lateral view) convex or forming an obtuse angle
at half length. Lateral sides of postclypeus some-
times slightly bulbous, with about 9-12 weak
furrows and a weakly inflated crest along lorum.
Vertex wrinkled, with distinct medial fissure,
but without diverging fissures between ocelli.
Central part of vertex only slightly elevated,
often even somewhat pressed down and con-
cave. Vertex in Cystopsaltria narrower than in
Cystosoma. Vertex 1.6-2.1 X as broad as long.
Distance between eyes 0.9- 1.1 X length of head
and 1.1-1.4 X postclypeus width. Ocelli wide
apart in Cystosoma, more closely together in
Cystopsaltria. Distance between lateral ocelli
0.8-1,1 X distance between lateral ocellus and
eye, and, in Cystosoma 2.4-3.7 X, in Cystopsal-
tria 1.9-2.2 X width of frontal ocellus. Prono-
tum with grooved and pitted surface and distinct
medial furrow. Amplified lateral corners of
pronotal collar forming an inflated crest along
anterior margin, which is continuous with the
sharply edged anterolateral margin of prono-
tum. Mesonotum with a more smooth surface
and with very narrow cruciform elevation, nar-
rower than long across its centre. Fore femur
(fig. 60) with row of 3 spines, diminishing in
length towards tibia, most proximal spine
strongly bent and adjacent to femur, reaching to
about half-way the distance to middle spine (see
arrow). Tegmina green, opaque rusty brown or
greenish tinged in museum material, fairly slen-
der and pointed at apex. Venation reticulate in
distal half of tegmen. This reticulation includes
ulnar areas in Cystopsaltria, but Cystosoma has
normally developed ulnar areas. Costal area and
border along hind margin of tegmen very nar-
row, Wings hyaline, in Cystosoma with 7-10, in
Cystopsaltria with 12-14 apical areas, often
somewhat reticulate near wing apex and with
fairly broad hyaline border along hind margin.
Tymbal with 6-10 transverse sclerotized ridges
from dorsal to ventral margin. Male operculum
fairly small. Basal part of operculum hardly
vaulted, but in males forming a large, often glo-
bularly swollen, protuberance at distolateral cor-
ner. Distal part of male operculum angularly

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

oblong, curved to close against the body and
covering most of tymbal cavity in ventral view.
Opercula widely separated medially, by broad
and rounded first sternite, Meracanthus short,
not reaching distal margin of operculum.
Female operculum shorter than that of male,
with a more elongate lateral crest of basal part.
Male abdomen very delicate and strongly
inflated, without distinct folds between tergites
and sternites; almost circular in cross section.
First tergite in male fairly long, not hidden under
metanotum. Proximal margin of second tergite
convex medially and almost straight between
auditory capsules and sternite 2 in Cystosoma
(fig. 68), though weakly curved in Cystopsaltria
(fig. 82) and forming a fairly distinct ridge along
tymbal cavity. Lateral parts of 2nd tergite
weakly swollen at anterior margins and adjacent
to tymbals. Sternites 1 and 2 adjacent. Auditory
capsules in males weakly developed, hardly pro-
truding, but distinctly elevated relative to con-
necting bar between tymbal and abdomen.
Female abdomen more robust than that of male,
but much smaller with short and broad genital
segment. Ovipositor sheaths not reaching to
apex of bluntly rounded caudodorsal beak (fig.
63). Male pygofer globularly rounded, with short
caudodorsal beak. Beak not curved over basal
part of claspers or anal valves. Lateral lobes of
pygofer curved inwards and forming bluntly
rounded lateral protuberances. Claspers broad
at base and hook-shaped; sharply curving down
at half-length. Apical part of clasper with large,
sharply edged, clasper hollow. Claspers weakly
diverging towards rounded apices. Basal parts of
claspers forming a continuous but very low ring-
shaped collar around base of anal valves. Aed-
eagus weakly S-curved. Aedeagal pore round.

Tribal placement. Distant (1905) included Cys-
tosoma in his division Hemidictyaria (presently
tribe Hemidictyini) which has as its principal
character a narrow head. This division also con-
tained Hemidictya Burmeister, 1835, Hovana
Distant, 1905 and seven genera that were later
brought into the tribe Prasiini (e.g., Arfaka
Distant, 1905; Iruana Distant, 1905; Jacatra
Distant, 1905; Lacetas Karsch, 1890; Lembeja
Distant, 1905; Prasia Stal, 1863 and Sapantanga
Distant, 1905). Recently Moulds (1990) trans-
ferred Cystopsaltria from Chlorocystini to Hem-
idictyini based on the narrow head, inflated
male abdomen and reticulate tegmina, charac-
ters very similar to Cystosoma.

A study of male genitalia, however, shows that
Cystosoma and Cystopsaltria should be attri-

113

buted to Chlorocystini (sensu stricto). Cystos-
oma and Cystopsaltria have a weakly S- curved
aedeagus with winged lateral crests, the synapo-
morphy of that tribe. The shapes of pygofer and
claspers also agree with such an allocation. Fur-
thermore, for the discriminating characters used
by Moulds, the inflated male abdomen of Cys-
tosoma and Cystopsaltria is larger but otherwise
very similar to that of most species of Chloro-
cystini and different from that of Hemidictya,
and the head of Cystosoma and Cystopsaltria 1s
not as notably different as that of several other
genera of Chlorocystini.

Hemidictya frondosa Burmeister, 1835 and
Hovana distanti (Brancsik, 1893), the two
remaining species of the Hemidictyini, are
undoubtedly closely related and probably sister
species, interesting from a biogeographical point
of view, since the former comes from Brazil and
the latter from Madagascar. These two are
characterised by (1) very broad, opaque yellow-
green and apically pointed tegmina with reticu-
late venation in the distal halves, (2) a costa
strongly widened and flattened in its proximal
half, (3) a distally elongate and sharply pointed
mesonotum reaching to the 2nd abdominal seg-
ment, and (4) a strongly streamlined head and
pronotum, with anterior margins of postclypeus
and vertex lobes forming a nearly straight and
almost continuous line with margins of the eyes
and lateral edges of pronotum. The last charac-
ter is also found in the African genus Lacetas
indicating that Lacetas should possibly be
included in Hemidictyini.

Cystosoma and Cystopsaltria share opaque
apically pointed and reticulate tegmina and a
sharp lateral edge of the pronotum with Hemid-
ictya and Hovana. Whether reticulate tegmen
venation can be regarded synapomorphic is dif-
ficult to decide since its extent varies between
the four genera. However, in general aspect the
tegmina of Hemidictya and Hovana are very dif-
ferent from those of Cystosoma and Cystopsal-
tria. Tegmina of the former two are broader,
more squarish and have a widened costa, while
the veins CuA and M fuse well before reaching
the basal area.

The sharp lateral edge of pronotum is defi-
nitely different, sharper, in Hemidictya and
Hovana. Head and pronotum of these two
species are more strongly streamlined than in
Cystosoma and Cystopsaltria; even the eyes are
flattened and contribute to this streamline. A
lateral pronotum edge similar as in Cystosoma
and Cystopsaltria was found in several other
species of Chlorocystini (e.g., Mirabilopsaltria

114

viridicata (Distant, 1897), M. humilis (Blóte,
1960), Gymnotympana montana De Boer, 1995,
G. olivacea Distant, 1905 and G. verlaani De
Boer, 1995; see De Boer, 1995a; De Boer, 1995b,
1996),

Study of male genitalia of Hemidictya fron-
dosa (unpublished) does not suggest relationship
to Chlorocystini since there are considerable dif-
ferences in shapes of pygofer, clasper and aedea-
gus between species of that tribe.

Monophyly. Cystosoma and Cystopsaltria are
easily recognized by the following synapomor-
phies: apically pointed tegmina and venation
reticulate in the distal halves of tegmina (l and 2
in fig. 56). The hyaline wings too, tend to be
somewhat reticulate towards their apices and
have more than 6 apical areas. The species
further share: 1, a very stout and angularly pro-
truding postclypeus, in lateral view not unlike
that of Thaumastopsaltria, but more strongly
protruding and often with convexly swollen
sides; 2, a sharp lateral edge of the pronotum in
dorsal view, though interrupted by the eyes,
almost continuous with the anterior margins of
postclypeus and vertex lobes; and 3, a very slen-
der cruciform elevation on the mesonotum, nar-
rower than long across the centre. These charac-
ters also occur in Prasiini, the presumed sister
group of Chlorocystini and might therefore be
plesiomorphic.

C. saundersii

3 C. schultzi

12 .
C. immaculata

Figure 56. Cladogram of the species of Cysrosoma and
Cystopsaltria. Numbers refer to apomorphies dis-
cussed in the text.

Phylogeny. Although Cystosoma schultzi and
Cystopsaltria immaculata share a fairly large
male operculum, other characters indicate that
Cystosoma schultzi and Cystosoma saundersii
form a monophyletic group. The Cystosoma
species have. compared to Cystopsaltria, a dis-
tinctly broader pygofer opening and a similar
pattern of reticulation in the tegmina more
restricted to the apical parts of tegmen. Further-
more, the veins CuA and M fuse at the corner of
the basal area in Cystosoma (fig. 72), a probable
synapomorphy for the two species of that genus
(3 in fig. 56).

Cystosoma Westwood

Cicada Westwood. 1842: 118.

Cystosoinga. Westwood. 1842: 118. — Goding and
Froggatt, 1904: 566, 595, 662. — Distant, 1906: 182.
185, — Metcalf, 1963: 433-434. — Boulard. 1979: 46.

A. J. DE BOER

— De Jong, 1982: 182. — Duffels and Van der Laan..
1985: 315. — Moulds, 1990: 192. — De Boer, 1990:
64. — De Boer, 1992a: 164. — De Boer, 1992b: 18, 19.
— De Boer, 1993a: 16-17. — De Boer, 1993b: 142. —
De Boer, 1995a: 8, 12, 16. — De Boer, 1995b: 3.

Type species. Cystosoma saundersii Westwood,
1842.

Diagnosis. Head with flattened vertex and large,
angularly protruding, postclypeus. Pronotum
with distinct medial fissure and rough, pitted,
surface. Tegmina opaque green, apically
pointed. Tegmen venation reticulate, with many
cross veins in apical areas, ulnar areas of normal
shape. Veins CuA and M fusing at corner of
basal area. Male abdomen strongly inflated.

Remarks. Cystosoma is presumed to be mono-
phyletic, the fusion of the veins CuA and M at
the corner of the basal area is the supposed apo-
morphy for the genus.

Cystosoma saundersii Westwood
Figures 57-68

Cicada saundersii Westwood, 1842: 118.

C'ystosoma saundersii. — Westwood, 1842: 118. —
Goding and Froggatt, 1904: 566, 662, 663. — Distant,
1906: 185. — Burns, 1957: 670. — Metcalf, 1963: 435.
— Duffels and Van der Laan, 1985: 315. — Moulds,
1990: 193-196, pl. 23 figs 1, la-b. — De Boer, 1995b:
3.

Cytosoma [sic] saundersii. — Musgrave, 1953: 13, |
fig.

Cystosoma laundersii [sic]. — Schremmer, 1957:
19, Fig. 11, 46.
Material examined. Australia. Australia, Saunders, c,
e Cystosoma saundersii det. Edm. Schmidt, IZW. Bris-
bane, 1973, J.B. Vogel, 39, ZMA; Brisbane, Kenmore,
xi. 1972, J.B. Vogel, 39, 9, ZMA. Cairns, 9 km W., ii-
1v.1985, 9. ZMA. Cunninghams Gap, 700-750 m, 20—
30.x1.1963, J.L. Gressitt, e, o, BPBM. N. Holland,
Koch. d, d det. Cpstoyoma saundersii Westwood,
ZMB. Nov. Holl., (Koch), v. Heyden, 7 Cystosoma
saundersii det, R.M. de Jong, SMF. NS Wales, 1908,
coll. A. Jacobi, 9, SMD; Qld, E. H.F. Walter, 9, BPBM.
Richmond river, NS Wales, 1908, Coll. A. Jacobi, c.
SMD. Tully, New Holland, Camille van Voixem, d det
Cystosoma saundersii Westw., KBIN. Without locality
label: Coll. Camille van Voixem, g, KBIN. Jocoli det.
Coll. Breddin, &, DEI; $ Cysrosoma saundersii det.
R.M, de Jong. SMF. Without labels: 9, ZMB,
Description. Body yellowish brown but green
when alive, unmarked (for photograph see
Moulds, 1990: plate 23 Figs l, la-b). Abdomen
in males 1.7-2.0 X head and thorax together, in
females 0.9-1.1 X. Tegmina of males 0.8-1.0 X
body length, of females 1.2-1.3 X.

Head. Greyish brown. Medial part of vertex
hardly elevated, ocelli somewhat pressed

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 115

Figures 57-68. Cystosoma saundersii Westwood, 1842: 57. pygofer in oblique view, Tully; 58, pygofer in lateral
view, Tully; 59, male caudodorsal beak in dorsal view, Tully; 60, male fore femur in lateral view, Tully, arrow
indicating strongly bent proximal spine; 61, aedeagus in dorsal view, Brisbane; 62, aedeagus in lateral view,
Brisbane; 63, female genital segment in lateral view, Cunningham's Gap; 64, female caudodorsal beak in dorsal
view, Cunningham's Gap; 65, female caudodorsal beak in dorsal view, Brisbane; 66, clasper, Tully; 67, female
operculum, Brisbane; 68, male operculum, Tully.

116

inward. Surfaces of vertex and dorsal part of
postclypeus very rough, covered with small pits,
anterior parts of vertex lobes wrinkled. Vertex
and postclypeus with distinct medial fissure.
Vertex with semicircular furrow around frontal
ocellus, but without diverging fissures between
ocelli. Postclypeus distinctly swollen ventrally,
anterior margin (lateral view) angularly convex
at about half-length. Sides of postclypeus with
11 weak furrows and a smooth and narrow,
weakly inflated, crest along lorum.

Thorax, Pronotum pitted, smooth in medial
furrow and with long wrinkles across collar.
Mesonotum smooth and unmarked.

Tegmina and wings: Tegmina opaque green-
brown and pointed at apex. Basal half oftegmen,
including ulnar areas, with regular venation, dis-
tal half of tegmen reticulate, with many cross
veins. Veins M and CuA fused at, or very close
to, corner of basal area (cf. fig. 81). Wings
hyaline, with irregular venation, 7 or 8 apical
areas and 2 or 3 subapical areas. Apex of wing
tending to be reticulate, with several cross
veins in Ist, 2nd and sometimes 3rd apical
arca.

Legs. Fore femur (fig. 60) with 3 spines, most
proximal spine strongly bent, adjacent to
femur.

Tymbals, Large, in lateral view covering more
than half of body width. 5 transverse sclerotized
ridges spanning the tymbal from dorsal to ven-
tral margin and a 6th most proximal ridge span-
ning about three-quarters of tymbal width. 6
distinct intercalary ridges form a lateral band
across tymbal.

Opercula, Male operculum (fig. 68). Basal part
of operculum with a knobby lateral protuber-
ance at distolateral corner and weakly vaulted,
lateral vaulting almost absent. Distal part of
operculum rather large relative to basal part,
almost as long as wide and almost completely
covering tymbal cavity in ventral view. Disto-
medial corner of operculum extending medially
beyond meracanthus. Lateral margin long. run-
ning straight to base of protuberance at distola-
teral corner of basal part and convexiy bent into
straight and about equallv long, distal margin.
Distomedial corner blunt, medial margin
straight and slightly directed mesiad. Meracan-
thus reaching to about three-quarters of opercu-
lum length. Female operculum (fig. 67) narrower
and much shorter than in male. Basal part of
female operculum about as long as in male, but
more distinctly vaulted and with long and dis-
tinct crest along rectangular distolateral corner.
Distal part very short and erect, oblong, with

A. J. DE BOER

almost rectangular distomedial corner and
broadly rounded distolateral corner.

Abdomen. Ochraceous or green and
unmarked. Male abdomen strongly, almost glo-
bularly inflated, distinctly broader than thorax
and strongly convex dorsally; higher than thorax
in lateral view. First tergite not extending under
metanotum, more than half as long as 2nd tergite
middorsally. Auditory capsules weakly inflated
but distinctly elevated relative to connecting bar
between tymbal and abdomen. Tymbal cavity
very narrow; auditory capsules fairly close
together and tergite part between auditory cap-
sule and 2nd sternite very short. Second tergite
straight between auditory capsule and 2nd
sternite and forming a fairly distinct crest along
tymbal cavity. Female abdomen long and slen-
der, with conically protruding auditory capsules.
Female pygofer (fig. 63) short and weakly
swollen, in lateral view with weakly convex dor-
sal margin. Ovipositor sheaths not reaching to
apex of caudodorsal beak. Female caudodorsal
beak in dorsal view swollen triangular and
pointed at apex (fig. 65) but rounded in speci-
men from Cunningham's Gap, southern Qld
(fig. 64).

Male genitalia. Pygofer in lateral view as in fig.
58. Dorsal margin weakly convex, almost
straight, continuous with straight caudodorsal
beak. Distal margin weakly convex, concavely
bent into straight margin of beak. Lateral lobe of
pygofer slightly curving inwards and forming a
weakly developed, slightly swollen and rounded
lateral protuberance. Ventral margin forming a
broad and rounded corner just below this pro-
tuberance and angularly convex, but concave
towards pygofer base. Pygofer globularly
swollen; pygofer opening very broad, broadest
between lateral protuberances of pygofer lobes,
ventral margins of pygofer converging to a con-
vex basal margin at base of pygofer opening;
ventral part of pygofer opening broadly U-
shaped (fig. 57). Caudodorsal beak in dorsal
view (fig. 59) very stout and broadly rounded at
apex. Clasper (fig. 66) broad, square-shaped at
base, with stout triangular, downwardly directed
apical part, curving inwards distally of aedeagus
and forming an angular dorsodistal corner sup-
porting aedeagus in upright position. Claspers
strongly diverging towards rounded apices. Api-
cal part of clasper with distinct, sharply edged
clasper hollow. Aedeagus (fig. 62) very stout and
almost straight, slightly swollen and weakly
upcurved at three-quarters of its length, but
recurving towards apex. Aedeagus with very
slender lateral crests and a pair of broad and

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

rounded dorsal ridges (fig. 61). Aedeagal pore
broad and rounded.

Measurements (mean + sd): Body length 3:
42.0-51.0 mm (47.2 mm + 2.6), o: 30.0-33.8
mm (31.3 mm + 1.4); tegmen length d: 40.3-
47.8 mm (43.9 mm + 2.4), o: 37.7-42.2 mm
(40.2 mm + 1.7); head length à: 3.2-3.9 mm (3.5
mm), 9: 3.4-3.9 mm (3.6 mm); pronotum length
d: 4.2-4.8 mm (4.6 mm), 9: 4.2-4.5 mm (4.4
mm); mesonotum length c: 7.0-8.3 mm (7.8
mm), 9: 6.6-8.1 mm (7.4 mm); head width g:
7.1-7.8 mm (7.5 mm), 9: 6.8-7.8 mm (7.4 mm);
width of pronotal collar d: 10.8-11.9 mm (11.4
mm), 9: 9.7-11.5 mm (10.7 mm).

Distribution. Eastern Qld and north-eastern
NSW. Moulds (1990) recorded the species from
the Atherton Tableland in northern Qld, the
Clarke Range, Eungella plateau and Mackay in
Central Old, inland at Carnarvon Range and
from Kroombit Tops in southern Qld to Syd-
ney.

Remarks. Cystosoma saundersii is the largest
species of Chlorocystini, easily recognized by its
enormously inflated abdomen, hence the local
name “Bladder Cicada" (Moulds, 1990).

Cystosoma schmeltzi Distant
Figures 69-71, 73-75, 81

Cystosoma schmeltzi Distant, 1882: 32, pl, vii figs
II, lla-b. — Goding and Froggatt, 1904: 566: 662,
664. — Distant, 1906: 185. — Burns, 1957; 670. —
Metcalf, 1963: 436. — Duffels and Van der Laan,
1985: 316. — Moulds, 1990: 192-193, pl. 23 figs 2,
2a.

Material. Australia. Dama, Cape York, 9, ZMB: Gayn-
dah, Mus Godefroy No 17630, o paratype, ZIM.
Wacol, 28.11.1970, H. Sas, 9, RMNH.

Description. Body yellow-green but green when
alive and unmarked (for photographs see
Moulds, 1990: pl. 23 figs 2, 2a). Male abdomen
1.4 X head and thorax together, in females 0.9—
1.0 x. Tegmina of male 1.0 X body length, of
females 1.4 x.

Head (fig. 69): Ochraceous with traces of olive
green, vertex lobe with blackish spot between
eye and lateral ocellus. Head heavily wrinkled
between ocelli and on anterior parts of vertex
lobes, vertex with distinct medial fissure. Post-
clypeus distinctly swollen ventrally, anterior
margin (lateral view) angularly convex at about
half-length. Lateral parts of postclypeus with 11
weak furrows and a smooth and narrow, weakly
inflated, crest along lorum (fig. 70). da

Thorax. Pronotum pitted as in C. saundersii,

117

smooth in medial furrow and with long wrinkles
on collar. Mesonotum smooth and unmarked.

Tegmina and wings: Tegmina opaque green-
brown and pointed at apex. Basal halfoftegmen,
including ulnarareas, with regular venation, api-
cal half reticulate, with many cross veins. Veins
M and CuA fused at corner of basal area (fig. 81).
Wings hyaline, with almost regular venation and
7 apical areas. A fairly narrow hyaline border
along hind margin of wing, though distinctly
broader than opaque border of tegmen.

Tymbals. Large, in lateral view covering more
than half of body width. 6 transverse sclerotized
ridges spanning the tymbal from dorsal to ven-
tral margin and a 7th most proximal ridge span-
ning about three-quarters of tymbal width. 7
distinct intercalary ridges form a lateral band
across tymbal.

Opercula. Male operculum (fig. 71) with
weakly vaulted basal part as in C. saundersii.
Distal part of operculum very different from
that of C. saundersii, oblong and rather large
relative to basal part, completely covering tym-
bal cavity in ventral view and distinctly extend-
ing medially beyond meracanthus. Lateral mar-
gin long, gradually and concavely bending into
crest around distolateral corner of basal part and
convexly bending into long and weakly convex
distal margin. Distomedial corner broadly
rounded, medial margin straight. Meracanthus
reaching to about two-thirds of operculum
length. Female operculum as in C. saundersii
with erect and oblong distal part (cf. fig. 67).

Abdomen. Male abdomen ochraceous to
green and unmarked, distinctly inflated, but not
as globular as in C. saundersii; with dorsal mar-
gin (lateral view) not distinctly elevated relative
to dorsal margin of thorax. First tergite not hid-
den under metanotum and medially only slightly
shorter than 2nd tergite. Auditory capsules
weakly developed, hardly protruding and not
visible in dorsal view, but distinctly elevated
relative to connecting bar between tymbal and
abdomen. Tymbal cavity very narrow; auditory
capsules fairly close together and tergite part
between auditory capsule and 2nd sternite very
short. Second tergite straight between auditory
capsule and 2nd sternite and forming a fairly
distinct crest along tymbal cavity. Female geni-
tal segment short and weakly swollen as in C.
saundersii. Ovipositor sheaths not reaching to
apex of caudodorsal beak. Female caudodorsal
beak in dorsal view swollen triangular and
pointed at apex.

Male genitalia. Pygofer in lateral view as in fig.
73. Dorsal margin concave near base, but

118 A. J. DE BOER

|

Figures 69-77. Cystosoma schmeltzi Distant, 1882: 69, male head in dorsal view; 70, male postclypeus in lateral
view; 71, male operculum; 72, base of right tegmen, male (Cystopsaltria immaculata); 73, pygofer in lateral view;
74. male caudodorsal beak in dorsal view; 75, pygofer in oblique view; 76, clasper; 77, aedeagus in lateral
view.

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS

strongly convex to apex of stout and short cau-
dodorsal beak. Distal margin straight, forming
an obtuse angle with margin of beak, angularly
bending outwards at distal end, into weakly
developed, slightly swollen and rounded lateral
protuberance. Ventral margin weakly convex,
but forming a broad and rounded inwardly
curved corner just below this protuberance.
Pygofer globularly swollen; pygofer opening
very broad, broadest between lateral protuber-
ances of pygofer lobes. Ventral margins converg-
ing to a very short and convex basal margin at
base of pygofer opening; ventral part of pygofer
opening U- shaped (fig. 75). Caudodorsal beak
in dorsal view (fig. 74) very stout and broadly
rounded at apex. Claspers (fig. 76) parallel to
rounded apices, broad, square-shaped at base,
with long and slender, downwardly directed api-
cal part, curving inwards at angular dorsodistal
corner and supporting aedeagus in upright pos-
ition. Claspers fused at base to low collar around
base of anal valves. Apical part of clasper with
distinct but slender and sharply edged clasper
hollow. Aedeagus (fig. 77) slightly upcurved, but
recurving near apex, with very slender lateral
crests and pair of broad rounded dorsal ridges.
Aedeagal pore broad and oval.

Measurements. Body length d: 27.6 mm, o:
21.0 and 23.5 mm; tegmen length g: 28.2 mm, 9:
29.0 and 32.3 mm; head length d: 2.7 mm, 9: 2.7
and 2.8 mm; pronotum length c: 3.6 mm, 9: 2.8
and 3.0 mm; mesonotum length d: 6.4 mm, 9: 6.2
and 6.3 mm; head width: 5.4 mm, 9: 5.3 and 5.5
mm; width of pronotal collar d: 8.0 mm, o: 7.5
and 7.9 mm.

Distribution. Eastern Qld and northern NSW.
Moulds (1990) recorded the species from Moss-
man Gorge and Forty Mile Scrub, to inland
northern NSW south to Gunnedah.

Remarks. C. schmeltzi is distinctly smaller than
C. saundersii and has a less strongly inflated
abdomen. The local name is “Lesser Bladder
Cicada" (Moulds, 1990).

Cystopsaltria Goding and Froggatt

Cystopsaltria Goding and Froggatt, 1904: 559, 566,
661. — Distant, 1906: 154, 160. — Metcalf, 1963:
260. — Duffels and Van der Laan, 1985: 250. —
Moulds, 1990: 196-197. — De Boer, 1992b: 18-19, —
De Boer, 19932: 16-17. — De Boer, 1993b: 142. DG
Boer, 1995a: 8. — De Boer, 1995b: 3.

Type species. Cystopsaltria immaculata Goding
and Froggatt, 1904.

Remarks. Cystopsaltria is a monotypic genus,
closely related to Cystosoma.

119

Cystopsaltria immaculata
Goding and Froggatt

Figures 72, 78-80, 82-87

Cystopsaltria immaculata Goding and Froggatt.
1904: 566, 661. pl. xvii figs l. la. — Distant, 1906:
160. — Burns, 1957: 644-645. — Metcalf. 1963: 260.
— Duffels and Van der Laan, 1985, 250. — Moulds.
1990: 197-198, pl. 23 figs 3, 3a.

Material. Australia. Cairns, 9 km W., ii-iv.1985, 3.
ZMA. Kamerunga near Cairns, N. Qld., 10.1.1977,
M.S. and B.J. Moulds, d, ZMA. Mt Windsor Table-
land. NW of Mossman, 30.xii.1980, M.S. and B.J.
Moulds, 9 Cystopsaltria immaculata G. and F. det.
M.S. Moulds, ZMA.

Description. Body light brown (for photographs
see Moulds, 1990: pl. 23 figs 3, 3a). Male abdo-
men strongly inflated 1.3 X head and thorax
together, in females 1.0-1.1 X. Tegmina of male
1.4 X body length, in females 1.3-1.4 X.

Head. As in Cystosoma, but with somewhat
narrower vertex; ocelli more closely together.
Distance between lateral ocelli 1.9-2.2 X width
of frontal ocellus.

Thorax: Pronotum less distinctly pitted than
in Cystosoma, in male smooth, but with a dis-
tinct medial furrow.

Tegmina and wings. Tegmina opaque gree-
nish brown and reticulate as in the 2 foregoing
species, but with very different pattern of veins.
Distinctly more cross veins and larger part of
tegmen reticulate; no regular ulnar areas. Veins
M and CuA reach basal area separately. Radial
area divided by extra vein, parallel to costa and
fusing with vein M near corner of basal area (fig.
72). Tegmen with distinct, almost hyaline, cor-
dial fold. Wings hyaline, with irregular venation
asin C. saundersii, but with more (12-14) apical
areas. Apex of wing tending to be reticulate, with
several cross veins, especially near apex of first 4
or 5 apical areas.

Tymbals. Large, covering more than half of
body width in lateral view. 8 transverse sclero-
tized ridges spanning the tymbal from dorsal to
ventral margin, a 9th ridge spanning about
three-quarters of tymbal width and a 10th most
proximal ridge reaching to about half of tymbal
width. 9 weak intercalary ridges form a lateral
band across tymbal.

Opercula. Male operculum (fig. 82) resem-
bling that of C. schmeltzi. Distal part of opercu-
lum oblong and rather large relative to basal
part, covering greater part of tymbal cavity in
ventral view and distinctly extended medially
beyond meracanthus. Lateral margin long,
forming an obtuse angle at about third its length,

120 A. J. DE BOER

Figures 78-87. Cystopsaltria immaculata Goding and Froggatt, 1904: 78, pygofer in oblique view; 79, pygofer in
lateral view; 80, male caudodorsal beak in dorsal view; 81, base of right tegmen, male (Cystosoma schmeltzi); 82,
male operculum; 83, female caudodorsal beak in dorsal view, Cairns; 84, clasper; 85, aedeagus in lateral view; 86,
aedeagus in oblique view; 87, female operculum, Cairns.

PHYLOGENY AND BIOGEOGRAPHY OF AUSTRALIAN CICADAS 121

angularly bending into crest around distolateral
corner of basal part and angularly bending into
long and weakly convex distal margin. Distom-
edial corner angularly rounded, medial margin
weakly convex. Meracanthus reaching to about
3/4 of operculum length. Female operculum (fig.
87) much shorter than that of male. Distal part
sickle-shaped, with angularly convex distal mar-
gin.

Abdomen. Ochraceous brown and unmarked.
Male abdomen distinctly inflated, with its dorsal
margin (lateral view) higher than dorsal margin
of thorax. First tergite fairly short, middorsally
less than quarter as long as 2nd tergite. Auditory
capsules weakly developed, hardly protruding
and not visible in dorsal view. Tymbal cavity
much wider than in Cystosoma; tergite part
between auditory capsule and 2nd sternite much
longer than in that genus. Second tergite straight
between auditory capsule and 2nd sternite and
forming a distinct crest along tymbal cavity.
Female abdomen as in C. saundersii. Ovipositor
sheaths not reaching to apex of caudodorsal
beak. Female caudodorsal beak in dorsal view
(fig. 83) stout, bluntly rounded at apex.

Male genitalia: Pygofer in lateral view as in fig.
79. Dorsal margin almost straight and continu-
ous with short caudodorsal beak. Distal margin
straight, concavely bent into margin of beak,
angularly bending outwards at distal end, into
distinct and angularly swollen lateral protuber-
ance. Ventral margin straight and forming a very
small corner just below this protuberance.
Pygofer opening much narrower than in Cystos-
oma. Ventral margins converging to a sharp
angle at base of pygofer opening; ventral part of
pygofer opening V-shaped (fig. 78). Caudodorsal
beak in dorsal view (fig. 80) very stout and short,
broadly rounded at apex. Claspers (fig. 84)
almost parallel to sharply pointed apices.
Clasper broad, square-shaped at base, with long
and very slender, downwardly directed apical
part, curving inwards at angular dorsodistal cor-
ner, supporting aedeagus in upright position.
Claspers fused at base to a low collar around base
of anal valves. Apical part of clasper with dis-
tinct but slender and sharply edged clasper hol-
low. Aedeagus (figs 85, 86) very stout, swollen
and slightly upcurved in proximal half, but more
slender in distal half and recurving near apex.
Aedeagus without distinct lateral crests, but with
a pair of broad rounded dorsal ridges, ending in
weakly outcurving subapical appendages. Aed-
eagal apex incised.

Measurements (mean + sd): Body length c:
35.0 mm, 9: 31.5-35.1 mm (33.7 mm + 1.4);

tegmen length d: 40.0 mm, o: 42.3-46.6 mm
(45.1 mm + 1.7); head length d: 3.4 mm, 9: 3.2-
3.9 mm (3.7 mm); pronotum length d: 4.6 mm, 9:
4.6-5.0 mm (4.8 mm); mesonotum length d: 7.9
mm, o: 7.6-8.5 mm (8.2 mm); head width d: 7.4
mm, o: 7.4-7.8 mm (7.6 mm); width of pronotal
collar d: 10.6 mm, 9: 10.4-11.5 mm (11.1
mm).

Distribution. North-eastern Qld. Moulds (1990)
recorded the species from Gap Creek south of
Cooktown to Paluma.

Remarks. C. immaculata is in size intermediate
between Cystosoma schmeltzi and C. saundersii.
The species is easily recognized by the reticulate
vein pattern of tegmina, extending over a larger
part ofthe tegmen than in Cystosoma. Males can
be recognized by their aedeagus with distinct
dorsal appendages.The local name is ‘Rare
Bladder Cicada" (Moulds, 1990).

Acknowledgements

For the loan of material I am indebted to: Dr
W.J. Knight and Mr M.D. Webb (BMNH); Mr
G.M. Nishida and Mr K. Arakaki (BPBM); Dr
B.P. Moore (CSIRO); Dr H. Gadike (DEI); Dr E.
Kierich (IZW); Mr J. van Stalle (KBIN); Mr J.
van Tol (RMNH); Dr V.P. Gapud (SEM); Dr R.
Emmrich (SMD); Dr H. Schróder (SMF); Mr F.
Heller (SMN); Dr H. Strümpel (ZIM); Dr U.
Góllner-Scheiding and Dr J. Deckert (ZMB); Dr
A. Jansson (ZMH). Mr M.S. Moulds is thanked
for his help and cooperation and the loan of
specimens from his private coliection, he kindly
donated some specimens of C. immaculata to
our museum.

I would like to thank Mr G. Verlaan for tech-
nical assistance. I am indebted to Dr J.P. Duffels
and Prof. Dr F.R. Schram (Instituut voor Sys-
tematiek en Populatie Biologie, Amsterdam) for
their critical reading and comments on the
manuscript.

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Pigram, C. J. and Davies, P.J., 1987. Terranes and the
accretion history of the New Guinea orogen.
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193-212.

Rangin, G., Jolivet, L. and Pubellier, M.,1990a. A
simple model for the tectonic evolution of the
southeast Asia and Indonesia region for the past
43 m.y. Bulletin de la Société géologique de France
(8) 6 (6): 889-905.

| Rangin, G., Pubellier, M., Azema, J., Briais, A., Cho-
tin, P., Fontaine, H., Huchon, P., Jolivet, L.,
Maury, R., Muller, C., Rampnoux, J.-P., Stephan,
J.-F., Tournon, J., Cottereau, N., Dercourt, J. and
Ricou, L.E., 1990b. The quest for Tethys in the
western Pacific. 8 paleogeodynamic maps for

LI

Cenozoic time. Bulletin de la Société géologique
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Schremmer, F., 1957.Singzikaden Neue Brehmbii-
cherei 193. A. Ziemsen Verlag: Wittenberg-
Lutherstadt. 47 pp.

Stál, C., 1863. Hemipterorum exoticorum generum et
specierum nonnullarum novarum descriptiones.
Transactions of the Royal Entomological Society
of London (3) 1: 571-603.

Swofford, D.L., 1993. PAUP: Phylogenetic Analysis
Using Parsimony, version 3.1.1. Illinois Natural
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Walker, F., 1852. Supplement. List of the specimens of
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Walker, F., 1862. Characters of undescribed species of
Homoptera in the collection of F.P. Pasco, F.L.S.
Journal of Entomology 1 (5): 303-319, pl. 15.

Westwood, 1842. Illustrations of some genera belong-
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Westwood, J.O., 1851. Descriptions of some new
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Memoirs of the Museum of Victoria 56(1):125-128 (1997)

THE TYPE SPECIES OF ALEURODICUS DOUGLAS, A WHITEFLY GENUS
OF ECONOMIC IMPORTANCE (HOMOPTERA: ALEYRODIDAE)

J.H. MARTIN
Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK

Abstract

Martin, J.J.. 1997. The type species of Aleurodicus Douglas, a whitefly genus of economic
importance (Homoptera: Aleyrodidae). Memoirs of the Museum of Victoria 56(1): 125-128.

Slides prepared from type specimens of Aleurodicus cocois (Curtis), held in the Museum of
Victoria, Australia, have allowed their microscopic examination, possibly for the first time.
A lectotype is designated and an updated species diagnosis is presented. Its statusas the type
species of A/eurodicus is confirmed through validation of its synonymy with A. anonae

Morgan.

Introduction

Aleurodicus Douglas (in Morgan, 1892) is the
type genus of the mostly Neotropical whitefly
subfamily Aleurodicinae. It includes several
agricultural pest species, most notably A. disper-
sus Russell (1965), a polyphagous and extremely
fecund species. This insect has gained particular
notoriety by extending its native New World dis-
tribution to encompass much of the Pacific,
southern Asia and Africa over a period of only
15-20 years, arriving in northern Queensland by
1995 (Martin, 1996). A few Aleurodicus species
are natives of southern Asia and the western
Pacific; A. destructor Mackie (1912) is one of this
group, occasionally becoming a pest in tropical
Australia.

As part of a study of economically important
members of the Aleurodicinae in the Caribbean
region, it was discovered that Mound and Hal-
sey (1978) had mistakenly credited the Natural
History Museum, London (BMNH) as the
depository of syntypic specimens of Aleurodicus
cocois (Curtis, 1846). A. cocois has been regarded
as the type species of A/eurodicus through syn-
onymy (see below) and the purpose of this inves-
tigation has been to locate and examine type
material in order to assess this status. A. cocois
was described from coconut in Barbados but
material with the relevant data was absent from
BMNH. However five slides from Demerara
[Guyana] in BMNH, identified as A. cocois, bear
red labels as if of type status. This Guyanese
material may have mistakenly been used as the
basis for proposing the synonymy of A. anonae
Morgan with A. cocois (see Discussion, below).
Thus it was necessary to locate and examine the
true type material of A. cocois in order to reap-
praise this important synonymy.

Depositories. BMNH — The Natural History
Museum, London SW7 5BD, UK

NMV — Museum of Victoria, Abbotsford, Mel-
bourne, Vic. 3067, Australia

USNM — United States National Museum of
Natural History, Washington DC 20560, USA

Aleurodicus cocois (Curtis)
Figures 1-2

Aleyrodes cocois Curtis, 1846: 284-285. Lectotype
here designated

Aleurodicus anonae Morgan, 1892: 32 (Lectotype
designated and synonymised with A. cocois by Mound
and Halsey, 1978: 229).

Aleurodicus cocois (Curtis) Morgan, 1892:32.

Type material examined. Lectotype puparium of A/ey-
rodes cocois Curtis (here designated): Barbados, on
coconut tree [Cocos nucifera], Sir R Schomburgk,
January 1845 (NMV). Paralectotypes of A. cocois: 12
puparia, 2 third instar larvae, Barbados, same data as
lectotype (NMV, BMNH, USNM) Additional
material of A. cocois: 1 male, 1 female adult (on 1
slide), several dry adults in rather poor condition and
further puparia and third instar larvae on leaf frag-
ments, Barbados, same data as lectotype (all NMV).

Lectotype and paralectotype of Aleurodicus anonae
Morgan: 2 puparia, Guyana, Demerara, on Annona
muricata, S.J. McIntire (Douglas collection 1236).
1891 (BMNH).

Diagnosis. Pupal case. Rather evenly oval out-
line, widest at abdominal segment II, usually
1.00-1.40 mm long, 0.70-1.00 mm wide. Sub-
margin with a broad band of wide-rimmed
simple pores (terminology of Russell, 1965);
inner margin of this band is very characteristic,
on meso- and meta-thorax almost straight and
parallel to longitudinal moulting suture (Fig. 1).
but on abdomen curling around the large com-
pound pores (Figs 1, 2). Dorsal surface mesad of
wide-rimmed pore band smooth, punctuated
only by scattered septate pores (Fig. 2).

125

126 J. H. MARTIN

G
a”

2 9, 9 9090 GO
099° 919
JE DAO]
0 Ç Ba © 5 O)

Figures 1,2, A/eurodicus cocots, puparium (after Martin, 1987). 1, whole puparium, dorsal detail to right of line,
boundary of submarginal wide-rimmed pore band shown as dashed line; 2, dorsal detail of abdominal segments
HII to show submarginal wide-rimmed pores, scattered subdorsal septate pores and large compound pore in
lateral aspect.

WHITEFLY GENUS OF ECONOMIC IMPORTANCE

Cephalic and anteriormost 4 pairs of abdominal
compound pores usually 25-30 um in diameter,
posteriormost 2 abdominal pairs no more than
12 um (in all cases measured as the width of the
cylinder when pores in lateral aspect).

Distribution and host plants. Widely distributed
in New World tropics. Usually colonising coco-
nut but host records from 14 plant families are
quoted by Mound and Halsey (1978). Material
in BMNH from hosts other than Palmae com-
prises samples from Anacardiaceae, Annona-
ceae and Lauraceae.

Discussion

Horn et al. (1990) indicated that the collection of
John Curtis (who published entomological notes
in Gardener's Chronicle under the pseudonym
"Ruricola") had been deposited largely in the
Museum of Victoria, Melbourne. Ken Walker
(pers. comm.) confirmed that the dry material
sent to Curtis from Barbados was indeed present
in Melbourne, but that no slides appeared to
have been made. This dry material, the syntypes
of Aleyrodes cocois, was kindly loaned to the
author. Slides were prepared which have veri-
fied that the considerable numbers of specimens
in BMNH, identified as A. cocois over the years,
are conspecific with the type material.

The red-labelled Guyanese specimens in
BMNH are also from coconut. They bear J W
Douglas's collection number 1246/27, and one
slide bears the following note, in Laurence A.
Mound's handwriting: “The specimens referred
to in Morgan 1892 as from JWD ex-Demerara
— see Douglas diary". Consultation of Dou-
glas's diary (BMNH archive) reveals that batch
1246 comprised a number of samples received
from Mr S J McIntire in Demerara, 12 October
1891. Sample 27 bears the note “small fly and
fluff found on a cocoanut tree, alive when sent’.
Aleurodes cocois, Curt. true and diff. from that
on Anona (sic), no.1236. Those sent alive in a
bottle, to Mr Morgan”.

Morgan's (1892). paper is confusing and ram-
bling but is of great importance because it was
the vehicle for establishment of the genus Aleu-
rodicus, which contains several species of great
economic significance. “Aleurodicus Douglas
n.g." was proposed in the middle of Morgan's
paper, with an extremely brief diagnosis as a
footnote, initialled “J W D”. Immediately under
the generic heading is Morgan's own description
of his new species A/eurodicus anonae, although
this was only subsequently designated as the

[27

type-species of Aleurodicus, by Quaintance
(1908).

A. cocois is discussed in several places in Mor-
gan's 1892 paper, and the coconut sample
detailed in the Douglas diary is mentioned
twice, most particularly “Habitat: cocoa-nut
palm only, Demerara". However, the fact that
Curtis had described. A cocois from Barbados is
nowhere mentioned, even though Morgan
acknowledged McIntire for providing him with
a copy of Curtis’s descriptive article, sections of
which Morgan quoted. | believe that this has
been the source of major confusion. Quaintance
(1908) quoted A. cocois as “also [described] from
Demerara” (along with anonae), and yet he
included Barbados in its distribution data.
Although no particular status is claimed on the
red labels of the BMNH slides. it is probable that
Mound and Halsey. too, attributed unwarranted
significance to this Guyanese coconut material
from the Douglas collection.

Conclusions

From examination of the type material of A.
cocois and A. anonae the author concludes that
A. anonae is correctly placed as a junior syn-
onym of A. cocois, which is thus the type species
of Aleurodicus Douglas. Specimens on red-
labelled BMNH slides of A. cocois from coconut
in Demerara, which had been sent by McIntire
to Douglas and forwarded to Morgan, were cer-
tainly compared by Morgan with other Guya-
nese material he subsequently described as A.
anonae: however, they have no type status nor
particular significance taxonomically, having
only been identified as A. cocois through com-
parison with Curtis's written description.

Acknowledgements

The assistance of Ken Walker (NMV), who
swiftly located the Curtis dry material and
kindly arranged for its loan to BMNH, is grate-
fully acknowledged. The author's thanks are also
extended to Doug Williams and Gillian Watson
(CAB International Institute of Entomology).
for helpful suggestions made during this investi-
gation.

References

Curtis. J. (as “Ruricola”), 1846. Aleyrodes cocois (the
cocoa-nut Aleyrodes). Gardener's Chronicle 1846:
284-285.

Horn. W., Kahle, L, Friese, G. and Gaedike. R.. 1990,
Collectiones Entomologicae. Berlin. 573 pp.

128 J. H. MARTIN

Mackie, D.B., 1912. A new coconut pest. Philippine
Agricultural Review 5: 142-143.

Martin, J.H., 1987. An identification guide to com-
mon whitefly pest species of the world. Tropical
Pest Management 33(4): 298-322.

Martin, J.H., 1996. Neotropical whiteflies of the sub-
family Aleurodicinae established in the western

Palaearctic (Homoptera: Aleyrodidae). Journal of

Natural History 30(12): 1849-1859.

Morgan, A.C.F., 1892. A new genus and species of
Aleurodidae. Entomologist's Monthly Magazine
28: 29-33.

Mound, L.A. and Halsey, S.H., 1978. Whitefly of the
world. Chichester. 340 pp.

Quaintance, A.L., 1908. Homoptera, Family Aleyrod-
idae. In Wytsman, P. [ed.], Genera Insectorum 87:
1-11.

Russell, L.M., 1965. A new species of Aleurodicus
Douglas and two close relatives (Homoptera:
Aleyrodidae). Florida Entomologist 48: 47-55.

Memoirs of the Museum of Victoria 56(1):129-143 (1997)

ADDITIONS TO THE TAXONOMY OF THE LIMNORIIDAE
(CRUSTACEA: ISOPODA)

L.J. COOKSON

l CSIRO, Forestry and Forest Products,
Private Bag 10, Clayton South MDC, Vic. 3169, Australia

Abstract

Cookson, L.J., 1997, Additions to the taxonomy of the Limnoriidae (Crustacea: Isopoda).
Memoirs of the Museum of Victoria 56(1): 129-143,

The crustacean marine wood borers Paralimnoria andrewsi (Calman) and Limnoriasa-
seboensis Menzies are described from Australia for the first time. New characters are listed
that will aid in the separation of L. saseboensis, L. indica Becker and Kampf, L. simulata
Menzies and L. foveolata Menzies. L. indica and L. simulata are considered to be separate
species. Limnoria carptora sp. nov. is described from algal holdfasts growing at Heard Island
in the subantarctic. Additional distribution and habitat records are provided for L. agrostisa
Cookson, L. convexa Cookson, L. loricata Cookson, L. platycauda Menzies, L. quadripunc-
tata Holthuis, L. rugossisima Menzies, L. unicornis Menzies, and Lynseia annae Cookson

and Poore.

Introduction

This paper adds to an earlier study (Cookson,
1991) on the taxonomy of the Limnoriidae, an
isopod crustacean family of wood-, algal- and
seagrass-boring species. Since that publication
the Lynseiidae, a supposedly related family, has
been dismantled and its species of Lynseia trans-
ferred to Limnoriidae (Cookson and Poore,
1994), so that with Limnoria and Paralimnoria
there are now three genera in the family. Exten-
sive ecological research in Western Australia has
shown that the seagrass-boring species of
Lynseia and Limnoria agrostisa Cookson can
heavily colonize seagrass meadows, with
possibly important implications for the health of
those ecosystems (Brearley and Walker,
1993).

The present contribution re-examines the
species Limnoria indica, L. saseboensis, L. simu-
lata and L. foveolata in an effort to clarify dis-
tinguishing features. All four are very similar in
appearance, and indeed, Miiller (1988) synony-
mised L. indica and L, simulata. Also, a new
species is described from the subantarctic, and
the distribution or habitat records for a further
nine species extended (Table 1).

Material for this study was obtained from the
Museum of Victoria, Melbourne (NMV); Aus-
tralian Museum, Sydney (AM); United States
National Museum of Natural History,
Washington D.C. (USNM); Zoologisk Museum,
Copenhagen (ZMUC); Canterbury Museum,
Christchurch (CM); and the Bundesanstalt für
Materialforschung und -Prüfung (BAM).

Limnoria agrostisa Cookson

Limnoria agrostisa Cookson, 1991: 166-167, 169,
fig 10. — Cookson, 1990: 6, 15. — Brearley and
Walker, 1993: 415-428, fig la.

Material examined. SA, Port Pirie (33'12'S,
138°00’E), 4.1 m, subtidal on Posidonia and Amphi-
bolis spp: core sample 795-A4/5, T.J. Ward, Aug 1979,
AM P38958 (male, 3.75 mm), core sample 801-A4/8,
T.J. Ward, Mar 1980, AM P38986 (female, 2.9 mm).

WA, Bramble Point, Princess Royal Harbour
(35°03’S, 117°53’E), on Posidonia sinuosa, airlifted
sample epifauna, 0.1 m?, Hutchings, Wells and
Walker, 20 Jan 1988 (stn P. sinuosa 10), AM P39019
(ovig. female 4.0 mm). Green Island, Rottnest, 1-2 m,
from Amphibolis griffithii, D.I. Walker, 7 Feb 1989,
NMV J37418 (female, with | slide, 3.0 mm), NMV
J37420 (4 females, 1 male, 2 juv), 6 Feb 1989, NMV
J37419 (male, 1.7 mm). Seven Mile Beach. near Don-
gara, G. Edgar, 1985-1986, NMV J37421 (male, with
I slide, 2.0 mm).

Previous distribution records. South Australia and
southern Western Australia.

Remarks. L. agrostisa was originally described
on the basis of just two specimens. In this
additional material, one large male from Port
Pirie (AM P38958) has a pair of relatively large
anterior puncta on the pleotelson, similar in
comparative size to that found on L. unicornis
males, but not recurved. The male sometimes
has pereopod 7 poorly developed (NMV J37420,
J37421).

While L. agrostisa has been collected from
Posidonia, its preferred substrate is Amphibolis
griffithii (Brearley and Walker, 1993).

129

130

Table 1. New distribution records or substrates for the Limnoriidae.

L.J. COOKSON

Species

Location
*new record

Existing distribution
record

Paralimnoria

andrewsi (Calman)

Lynseta annae

Cookson & Poore

Limnoria convexa

Cookson

Limnoria indica
Becker & Kampf

Limnoria loricata

Cookson
Limnoria

platycauda Menzies

Limnoria
quadripunctata
Holthuis
Limnoria
rugosissima
Menzies

Limnoria unicornis

Menzies

Christmas Islands. Samoa.
Hawaii. Japan. Florida.
Caribbean. Philippines.
Puerto Rico. Ghana. Papua
New Guinea. Cocos Islands.

Geraldton to Dunsborough, Corner Inlet,
Western Australia. Ceduna, Victoria*
South Australia.

The Snares, New Zealand Campbell Island“,

New Zealand

Northern Queensland, Gladstone*,
Australia. Papua New Queensland
Guinea. Admiralty Islands. (southern limit).
Mandapam Camp, Madras*, India
Andaman Islands, India.

Hong Kong. Manila,

Philippines. Koniya, Japan.

Penang, Malaysia. Belize.

Tobago.

The Snares, New Zealand Campbell Island*,

New Zealand

West Indies. Puerto Rico. Bombay“, India

Belize. Cuba. Andaman

Islands. Aldabra Atoll.

Koniya, Japan. Satta Hip,

Thailand. Admiralty

Islands. Queensland,

Australia. Karwar, India

(NMV J37436, see Karande

et al., 1993).

Widespread cool temperate Cook Strait, New

distribution, including New Zealand (see

Zealand. Chilton, 1916).

NSW, SA, Victoria, Ninepin Point*,

Tasmania, Australia. The — Bicheno*,

Snares, New Zealand. Tasmania. Gabo
Island*, Victoria.

Caroline Islands. Andaman Port Douglas,

Islands. Palau. Huahine Queensland.

Island. San Salvador. Belize.

Papua New Guinea.

Northern Australia.

Broome*, Australia

Substrate Museum
*new record material
Jarrah NMV J17259
heartwood*
Posidonia NMV J41054
australis
Durvillaea NMV J37422
holdfasts NMV J37423
CM 5.37
Pine AM P35418
Wood test NMV J37426
panels
NMV J37424
NMV J37425

Himalayan NMV J37435
fir

Gutta- CM
percha cable

Macrocystis NMV J37434

holdfasts* NMV J37433
Ecklonia NMV J37417
holdfasts NMV J37416
Dead NMV J37454
mangrove

branch lying

on mud

amongst

mangroves*

ADDITIONS TO THE LIMNORIIDAE (CRUSTACEA) 131

Limnoria carptora sp. nov.
Figures 1, 2

Material examined. Holotype: Heard Island, Atlas
Cove (53900'S, 74°00’E). lower eulittoral, Durvillaea
antarctica holdfasts, G. Edgar, 21 Feb 1988 (stn HI CI
6). NMV 717255 (male, 4.4 mm, 1.2 mm wide pleo-
telson, with | slide).

Paratypes: Type locality, NMV 117256 (female, 4.9
mm, with | slide), NMV J17257 (male, 4.2 mm, with |
slide), NMV 117258 (2 males, 3.0, 4.8 mm, 2 non-ovig.
females, 3.9, 4.1 mm, 6 ovig. females, 4.6, 4.8, 5.0, 5.1.
5.2, 5.2 mm),

Diagnosis. Pleonite 5 convex dorsomedially,
without carinae. Pleotelson flattened, with
weakly raised lateral crests, with 3 pair of flat-
tened longitudinal carinae, anteromedial pair
broad. Pleonite 5 0.5 times as long as pleotelson.
Dorsal surface of pleotelson with scales fused,
covered with solitary scale spikes, without pits.
Posterior margin of pleotelson with dorsal row
of scale spikes; margin fringed with 4 stout setae
between which are short unsheathed setae and
scale spikes.

Antenna l with 4 flagellar articles; second
article with about 11 aesthetascs, third article
narrow. Flagellum of antenna 2 with 4 articles.
Mandibular palp with 3 articles. Mandibular
incisors without rasp and file. Lacinia mobilis of
right mandible unbranched, apically serrated.
Epipod of maxilliped strap-like, 4 times as long
as wide, reaching articulation of palp articles |
and 2; epipod with simple true setae.

Secondary unguis of pereopod 1 bifid. Ventral
comb seta present on merus of pereopod 7 and
carpus of pereopods 6, 7 and sometimes 5. Uro-
pod peduncle with short lateral spike setae, with-
out prominent tubercles; endopod 0.65 times as
long as peduncle.

Pleopod 2 with plumose setae up to 0.8 times
length of exopod. Appendix masculina long.
reaching beyond endopod tip, articulating
proximal to midlength of endopod. Endopod of
pleopod 5 anterior to exopod, oval, 0.8 times as
long as endopod of pleopod 2; peduncle of pleo-
pod 5 with simple seta laterally.

Additional characters. Body length up to 5.2
mm. Colour in alcohol light pink for fresh
material, becoming pale yellow upon prolonged
storage. Article 2 of mandibular palp with more
than 1 simple seta.

Etymology. From the Latin for carver (of hold-
fasts).

Distribution. Heard Island, Southern Ocean.
Lower eulittoral.

Substrates. Durvillaea antarctica holdfasts.

Remarks. This species is similar to L. stephen-
seni and L. antarctica. However, L. stephenseni
lacks carinae (other than lateral crests) and
puncta on the pleotelson, whereas L. carptora
has six flattened but distinct longitudinal cari-
nae. Also, the ventral branch of the secondary
unguis of pereopod | on L. stephenseni is greatly
reduced (see figures by Menzies, 1957; Wolff,
1990; Cookson, 1991), while for L. carptora this
ventral branch is much larger, and usually of
similar proportion to that found in L. antarctica.
The smallest ventral branch seen in the material
examined is illustrated (Fig. 2e). L. carptora has
four large stout setae on the posterior margin of
the pleotelson, whereas L. stephenseni has many
positioned around the entire hind perimeter of
the pleotelson. Furthermore, L. stephenseni
grows up to 9.8 mm long, while the longest L,
carptora specimen found so far is 5.2 mm. The
appearance of the lacinia mobilis of the right
mandible may not be a reliable difference
between the two species. It is simple in L. carp-
tora, while in L. stephenseni it may be simple
(Menzies, 1957; Wolff, 1990) or bifid (Cookson,
1991).

L. carptora differs from L. antarctica mainly
in the sculpturing found on pleonite 5 and the
pleotelson. L. carptora lacks a transverse carina
on pleonite 5. On the pleotelson of L. carptora,
the dorsomedial carinae are flatter and broader
than for L. antarctica, and anteromedial puncta
absent. Both L. carptora and L. stephenseni dif-
fer from L. antarctica in that they have less
defined and raised lateral crests on the pleotel-
son, the dorsal surface between the lateral crests
on the pleotelson is flatter, and article 3 of
antenna l is more narrow. L. carptora can reach
5.2 mm, while the largest L. antarctica specimen
found is 4.7 mm (Cookson, 1991).

In the key to the species of Limnoria (Cook-
son, 1991). L. carptora should be inserted
between L. antarctica and L. stephenseni at step
35:

— Pleotelson with carinae, pleonite 5 without
carinae dorsomedially. Secondary unguis of
pereopod | bifid. Posterior margin of pleo-
telson with 4 stout setae. Substrate algae

A re ra EET IERI. tere ts L. carptora

The distribution of L. carptora and L. antarctica
may overlap, as L. antarctica has been found at
Kerguelen Island (Menzies, 1957), which is near
Heard Island. However, L. stephenseni has not
been found in the waters off either island. Its
distribution is the Auckland Islands, Macquarie

L.J. COOKSON

[SS]
lo

; Ad HU 4, A400 das pea!
NT n "

Figure 1. Limnoria carptora sp. nov. A-F, male, NMV J17255, holotype: A, pleonite 5 and pleotelson, dorsal
view; B, posterior margin of pleotelson; C, dorsal surface of pleotelson; D, uropod, ventral view; E, flagellum of
antenna l: F, peduncle article 5 and flagellum of antenna 2. G, male, NMV J17257, paratype, maxilliped.

ADDITIONS TO THE LIMNORIIDAE (CRUSTACEA)

Figure 2. Limnoria carptora sp. nov. A-D, male, NMV J17255, holotype: A, pleopod 2; B, pleopod 5; C, right
mandible; D, lacinia mobilis of right mandible. E, female, NMV J17256, paratype, distal articles of pereopod 1,
lateral view.

Island and Marion Island (Wolff, 1990; Cook-
son, 1991). L. stephenseni and L. carptora have
been collected from Durvillaea (Wolff, 1990;
Cookson, 1991), and the former from Macrocys-
tis as well (Hale, 1937). L. antarctica has been
collected only from or near Macrocystis (Cook-
son, 1991).

Limnoria foveolata Menzies

Figure 3
Limnoria (Limnoria) foveolata Menzies, 1957: 175,
fig 33.
Limnoria foveolata. — Cookson, 1991: 142. —

Cookson, 1990: 2.

Material examined. Holotype, Kai Islands (6°5’S,
105°42’E), 52 m, Sigsbee trawl, sand, shells, Danish
Expedition, 4 Aug 1922, ZMUC (non-ovig. female,
with 2 slides).

Diagnosis (female). Pleonite 5 dorsomedially
with 2 subparallel longitudinal weak carinae
converging slightly posteriorly. Pleotelson with
1 pair of anteromedial puncta, with carinae pos-
teriorly, with pair of weak anterolateral carinae.
Pleonite 5 0.6 times as long as pleotelson. Dorsal
surface of pleotelson with scales fused, covered

with many solitary scale spikes. Lateral crests
and posterior margin of pleotelson without dor-
sal row of tubercles; posterior margin fringed
with 4 large stout setae between which are scale
spikes and short-sheathed setae.

Antenna 1 with 4 flagellar articles; second
article with 9 aesthetascs arising from 2 tufts.
Flagellum of antenna 2 with 5 articles. Mandibu-
lar palp with 3 articles. Mandibular incisors with
rasp and file. Lacinia mobilis of right mandible
with several teeth at apex. Epipod of maxilliped
subtriangular, about 3.3 times as long as wide,
just short of palp articulation; epipod with true
setae.

Secondary unguis of pereopod 1 bifid. Ventral
comb seta present on merus of pereopod 7 and
carpus of pereopods 2-7. Uropod peduncle
without tubercles; endopod 0.9 times as long as

peduncle.
Pleopod 2 with plumose setae up to 0.7 times
length of exopod. Appendix masculina

unknown. Endopod of pleopod 5 anterior to
exopod, oval, 0.75 times as long as endopod of
pleopod 2.

Additional characters. Body length 2.2 mm.
Colour in alcohol pale yellow.

134 L.J. COOKSON

Figure 3. Limnoria foveolata Menzies. A-J, female, ZMUC, holotype: A, pleonite 5 and pleotelson, dorsal view;
B. posterior margin of pleotelson. stout setae broken; C, dorsal surface of pleotelson; D, uropod, ventral view; E,
pleopod 2: F. pleopod 5; G, distal portion of right mandible; H, epipod and basis of endopod of maxilliped; I,
flagellum of antenna 1; J, peduncle article 5 and flagellum of antenna 2.

ADDITIONS TO THE LIMNORIIDAE (CRUSTACEA)

Distribution record. Known only from near the
Kai Islands, Indonesia. 52 m depth.

Substrates. Unknown. The presence of rasp and
file incisors on the mandibles suggests that this
species is a wood or seagrass borer.

Remarks. Only the female holotype of L. foveo-
latais known, looking similarto female L. indica
and L. saseboensis specimens. The differences
that separate L. foveolata from these species
remains unconvincing. Unfortunately, no male
specimens of L. foveolata are known, which
within this group of species is the sex that
appears to offer most distinctive features. The
pitted structure on the fifth pleonite and pleotel-
son must have been clearer when Menzies
(1957) named this species for its foveolate
appearance. However, when I examined the
holotype the pitting could not be seen with
reflected light at 80 times magnification, and
was only slight at 800 times magnification using
transmitted light. Perhaps some change in
prominence of the pitting can occur upon
storage. Such change was noted for the puncta of
Limnoria echidna Cookson, although for this
species the sculpturing became clearer upon
storage (Cookson, 1991). I have not noticed dra-
matic loss in clarity of pitting in other species. At
80X magnification, an absence of pitting is
characteristic of L. indica, while pitting is often
clearly visible in the typical L. saseboensis
specimen.

Menzies (1957) noted that L. foveolata dif-
fered from L. saseboensis in the lack of marginal
tubercles on the pleotelson, and longer maxilli-
pedal epipod. However, the latter character is
variable, as some specimens of L. saseboensis
have an epipod as long as that found in L. fov-
eolata. The feature that distinguishes L. foveo-
lata most easily from L. saseboensis and L.
indica, is the lack of tubercles on the lateral
crests of the pleotelson. L. foveolata also lacks
tubercles on the posterior margin of the pleotel-
son, but this condition can be found in some
specimens of both L. saseboensis and L. indica
also. L. foveolata has a weak pair of anterolateral
carinae on the pleotelson, which is normally
absent in L. indica but present in L. sase-
boensis. However, the difference again is not
impressive.

L. foveolata may become synonymised with
either L. saseboensis or L. indica on the basis of
future studies. The type locality lies within the
distribution range of L. indica, not L. sase-
boensis.

135

Limnoria saseboensis Menzies
Figures 4, 5

Limnoria (Limnoria) saseboensis Menzies, 1957:
141. 144, fig 18. — Menzies, 1959: 22.

Limnoria saseboensis. — Kühne, 1976: 546-547, fig
5.-Jones et al., 1976: 134, — Kensley and Schotte,
1989: 198, fig 87D. — Cookson, 1991: 143 155. —
Cookson, 1990: 4. — Cookson et al., 1991: 45.

Limnoria indica. — Gookson, 1987: 85-89, figs 1-8
(Goat Island material). — Cookson and Barnacle,
1987a: 143. — Cookson and Barnacle, 1987b: 287-
293 (Goat Island material).

Material examined. Paratypes, Japan, Sasebo, W.F.
Clapp, 22 Aug 1949 and 24 Jan 1950, USNM 91749 (5
males, | female; non-ovig female, 3.5 mm, with | slide:
specimens in poor condition).

Japan, Sasebo (3377'26”N, 129°44'16”E), LO m,
wood test panel submerged 6 Jul 1966-7 Dec 1966,
BAM (5 females 2.5-3.6 mm, male 2.9 mm), Koniya,
1968, BAM (male 2.9 mm, female 3.0 mm),

NSW, Sydney, pine bait blocks, R.D. Turner and
J.V. Marshall: Watsons Bay. 26 Nov 1971, AM
P35378 (5), Watsons Bay, 4 Aug 1971, AM P37046,
Cammeray, 10 Feb 1972, AM P35384 (1 male, 2
females up to 3.9 mm), Cabarita, 30 Jun 1971, AM
P38269 (1), AM P37044 (2), Goat Island, 27 May
1970. AM P35389 (33). Goat Island, 1971, AM
P35390(24), Goat Island, 4 Feb 1972, AM P37042 (3),
Goat Island, 0.5-1 m. E. pilularis timber, J. Beesley,
15 Sep 1975, NMV 137438 (2), various test timbers,
L.J. Cookson and J.E. Barnacle, | Mar 1985, NMV
J15258 (8), NMV 115263 (21), NMV J15257 (20),
NMV 715259 (16), NMV J15260 (male, 3.1 mm, with
1 slide), NMV 715268 (3), NMV 1715274 (20), NMV
J15276 (9), L.J. Cookson, 18 Mar 1983, NMV J15264
(11) NMV 715265 (20) NMV J15266 (14). Coffs Har-
bour, Police Jetty Marina. P. radiata block placed 14
Jan 1988, R. Livesay and L.J. Cookson, 4 Jul 1988,
NMV 115251 (5).

WA, Geraldton, wharf piles, R.D. Turner and J.V.
Marshall, 7 Nov 1970, AM P37045 (2), CCA-treated
jarrah piles pulled up on beach for several months, J.E.
Barnacle, 1 May 1989, NMV 1737443 (34), NMV
J37442 (male, 3.7 mm), NMV 737444 (male 3.3 mm,
with 1 side), NMV J37445 (male. 3.0 mm, with I
slide), NMV J37446 (female, 3.4 mm, with | slide),
NMV J37447 (intersex, 3.7 mm. with | slide), Gerald-
ton, Fishing Boat Harbour, 1 m below low tide, jarrah
piles, J. Boys, 5 May 1989, NMV J37440 (17), 28 Sep
1989, NMV J37439 (70), Roebourne, pine bait block,
R. Howlett, Nov 1960, NMV 714934 (male), Point
Samson, jetty, CCA-treated eucalypt pile, J.E. Barna-
cle, 15 Sep 1985, NMV 714962 (3) untreated eucalypt,
NMV J14961 (3, with | slide), jarrah piles washed onto
beach. J.E. Barnacle, 23 Apr 1989, NMV J37441 (8).

Qld. Wreck Reef, off Gladstone, (22911'S,
155°15’E), 12 m, in Casuarina from 'Cato' shipwreck,
W. Delaney, 18 May 1988, NMV J15210 (4 pleotel-
sons). Urangan Boat Harbour, Maryborough, low tide.
turpentine pile. J.E. Barnacle, 10 Feb 1986, NMV
J14995 (3). NMV J14996 (13).

136

L.J. COOKSON

Figure 4. Limnoria saseboensis Menzies. A, female from Sasebo, USNM 91749, paratype, pleonite 5 and pleo-
telson, dorsal view. B, male from Geraldton, NMV J37442, pleonite 5 and pleotelson.

Diagnosis (male). Pleonite 5 dorsomedially with
2 subparallel longitudinal carinae which may
converge slightly posteriorly. Pleotelson with 1
or 2 pairs of anteromedial puncta, if two pairs,
then one is directly behind other at a distance
egual to or longer than distance between
anterior puncta, with carinae behind posterior
pair of puncta, with an anterolateral pair of
puncta or long setae each anterior to short lateral
carina. Pleonite 5 0.5 times as long as pleotelson.
Dorsal surface of pleotelson with scales fused,
covered with many solitary scale spikes, some-
times strongly pitted anteriorly. Dorsal row of
tubercles extend from lateral crests to posterior
margin of pleotelson; posterior margin fringed
with 4 large stout setae between which are scale
spikes and short-sheathed setae.

Antenna 1 with 4 flagellar articles; second
article with about 15 aesthetascs arising from 2
tufts. Flagellum of antenna 2 with 5 articles.
Mandibular palp with 3 articles. Mandibular
incisors with rasp and file. Lacinia mobilis of
right mandible straight, apex with several teeth.
Epipod of maxilliped subtriangular, about 3
times as long as wide, not reaching palp articu-
lation; epipod with true setae.

Secondary unguis of pereopod | bifid. Ventral
comb seta present on merus of pereopod 7 and
carpus of pereopods 2-7. Uropod peduncle lat-
erally with short simple setae; with small
tubercles between plumose setae; endopod 0.85
times as long as peduncle.

Pleopod 2 with plumose setae up to 0.5 times
length of exopod. Appendix masculina reaching
apex of endopod of pleopod 2, articulating just
proximal to midlength of endopod. Endopod of
pleopod 5 anterior to exopod, broadly oval to
circular, 0.7 times as long as endopod of pleo-
pod 2; peduncle of pleopod 5 with simple seta
laterally.

Additional characters. With sexual dimorphism
of pleotelson sculpturing; female with pair of
anteromedial puncta followed posteriorly by
long carinae, pleotelson without other puncta.
Body length up to 4.1 mm. Colour in alcohol
pale yellow.

Distribution. Sasebo, Japan; Florida (Menzies,
1957); Japan (Kühne, 1976). Eastern Australia
from Sydney to Maryborough. Western Aus-
tralia from Geraldton to Point Samson near
Roebourne (current study). Depth range 0-12 m.

Substrates. From a causeway, presumably in
wood (Menzies, 1957). P. radiata and various
eucalypt timbers which were either untreated or
preservative treated, Syncarpia glomulifera (tur-
pentine), Casuarina (current study).

Remarks. For some time I wrongly identified
certain populations of this species as the very
similar L. indica (Cookson, 1987, Cookson and
Barnacle, 1987b). However, all references to L.
indica and L. saseboensis in Cookson (1991) are
correct.

ADDITIONS TO THE LIMNORIIDAE (CRUSTACEA) 137

Figure 5. Limnoria saseboensis Menzies. A-B, male from Sasebo, USNM 91749, paratype: A, pleopod 2, in situ;
B, pleopod 5, in situ. C-F, female from Sasebo, USNM 91749, paratype: C, antenna 2; D, right mandible; E,
lacinia mobilis of right mandible; F, maxilliped. G-H, male from Geraldton, NMV J37442: G, dorsal surface of
pleotelson; H, posterior margin of pleotelson. I, male from Point Samson, NMV J14961, flagellum of antenna 1.

138

Often, there is no difference between L. sase-
boensis and L. indica in shape or length of the
maxillipedal epipod (Kühne, 1976; current
study). although this character was used by Men-
zies (1957). Another character that cannot be
relied upon is the slight posterior convergence of
the longitudinal parallel carinae found dorsally
on pleonite 5 (Kühne. 1976: current study figs 4a
and 4b). The degree of pitting found on pleonite
5 and the pleotelson, as exemplified in Fig. 18D
given by Menzies (1957), also varies. This pit-
ting was well developed in the paratypes of L.
sasebocnsis. However, many specimens from
Australia lack the large carinated pits found on
the typical L. saseboensis specimen, so resemble
L. indica where carinated pits are not noticeable
at low magnification (80x). These pits are
noticeable on L. indica at higher magnification
(Kensley and Schotte, 1987). Menzies (1957) did
not give the number of puncta found on the two
parallel carinae on the dorsal surface of the pleo-
telson: "carinae tuberculate anteriorly and less
so posteriorly”. The male however has either
two or four puncta, and the female two puncta.
Few specimens are without puncta. Also, somc
male specimens from Australia and Japan have
an anterolateral pair of puncta on the pleotelson.
similar to that sometimes found on L. indica
males (Kühne, 1976; current study). These were
not to be found in the type material, although for
some, a large seta at the equivalent position was
noted. Therefore, the sexual dimorphism found
in L. indica can also occur in some specimens of
L. saseboensis. To further complicate the mat-
ter. the lacinia mobilis on the right mandible of
L. saseboensis was said to have two medially
curved tecth at the apex (Menzies, 1957). How-
ever. the number of teeth found can be greater
than this (Fig. 5e: Cookson, 1987, Figs. 6-8).
and so is similar to the lacinia mobilis found on
L. indica.

The population of L. saseboensis on the west
coast of Australia often differs from the popu-
lation on the east coast. While some males from
both coasts have the pair of anterolateral puncta
on the pleotelson, most males from eastern Aus-
tralia have two pairs of anteromedial puncta,
and most males in Geraldton have just one pair
of anteromedial puncta. However, this is not a
constant difference that could be used to separ-
ate the two populations as species. One male in
the Geraldton material (Fig. 4b) had all six
puncta. Also, the single male from Roebourne
(NMV 714934) had six prominent puncta (Roe-
bourne is an inland location, so the point of
collection was probably Point Samson). The six

L.J. COOKSON

puncta (four anteromedial, two anterolateral)
often found in male L. saseboensis from eastern
Australia is clearly shown in the Fig. 2 provided
by Cookson (1987). In this regard, L. saseboensis
males from eastern Australia can look very simi-
lar to L. indica males.

The most reliable difference to be found
between male L. indica and male L, saseboensis
specimens is that L. indica always has two pair of
anteromedial puncta, while L. saseboensis may
be without puncta or have just one pair. If L.
saseboensis has two pairs of puncta, then the two
puncta born on the same longitudinal carina will
be further apart or equal in distance to each
other, than they are with their paired partner. In
other words, the four puncta are longitudinally
more separated than laterally separated. For L.
indica, the two pair of puncta are always wider
apart than longer apart. Sometimes, the pos-
terior pair are especially wide, as was drawn in
the original description by Becker and Kampf
(1958), and shown in SEM photographs by
Kensley and Schotte (1987). Also, male L. sase-
boensis have carinae posterior to the hind pair of
anteromedial puncta, while L. indica males do
not. The key given by Cookson (1991) correctly
separates male L. saseboensis and L. indica,
based upon this character of carina presence or
absence behind the pleotelson puncta.

Other characters if present and clear, can
assist in identification. Lateral carinae (those
medial of lateral crests) are usually present in
both sexes of L. saseboensis. Such lateral carinae
are absent, or very slightly developed in L.
indica. The surface pitting of pleonite 5 and the
pleotelson can also be useful if developed. In
Australia, pitting was rare in the west coast
specimens, but more common in those from
eastern Australia, where the level of pitting was
also variable between individuals from the same
population. Another related character, found in
most but not all specimens of L. saseboensis, was
the pair of large circular carinae adjoining the
anterior end of the lateral crests. This circular
carina is constructed from several portions: the
anterior portion of the lateral crest, the antero-
lateral carina and puncta, and two bridging car-
inae (Figs. 4a, b). L. indica has the lateral crest
portion, and often the anterolateral puncta, but
the remaining components are essentially
absent. The endopod of pleopod 5, another
variable character, tends to be rounder in
L. saseboensis than L. indica.

It can at times be difficult or impossible, based
upon known characters, to separate female L.
indica and L. saseboensis. This is especially true

ADDITIONS TO THE LIMNORIIDAE (CRUSTACEA)

if pitting and lateral carinae on the pleotelson
are absent or ill defined. One further notable dif-
ference is that L. saseboensis is often larger. The
longest L. indica specimen known is 3.3 mm,
whereas the longest L. saseboensis specimen is
4.] mm.

In Australia, L. saseboensis has a warm tem-
perate distribution, while L. indica is tropical
and subtropical in range. Overlap of the two
species has not been observed. In eastern Aus-
tralia L. saseboensis extends from Sydney to as
far north as Maryborough, while L. indica can be
found from Gladstone to Papua New Guinea. In
Western Australia L. indica has not been found.
This absence might explain why L. saseboensis
can be found further north than on the east
coast, from Geraldton to Point Samson. L. sase-
boensis and L. indica appear to have similar
substrate preferences, in that, within their distri-
bution range on the east coast, they are the lim-
noriid species most likely to be found on
untreated turpentine timbers (Cookson, 1987;
further unpublished data).

Limnoria simulata Menzies
Figures 6, 7

Limnoria (Limnoria) simulata Menzies, 1957: 144,
fig 19. — Menzies, 1959: 20.
Limnoria simulata. — Carvacho, 1977: 17-18, figs

Müller, 1988: 397-403, figs 1-23 (Colombian
material). — Kensley and Schotte, 1989: 198. —
Cookson, 1991: 143, 192-193. — Cookson, 1990: 6.

Not Limnoria simulata. — Müller, 1988: 398
(Indian material). — McKoy-Hill, 1964: 46.

Material examined. Holotype, West Indies. Virgin
Islands, 16 Feb 1914, ZMUC (male, 3.0 mm, with 1
slide). Paratype, type locality, ZMUC (male).

USA, Florida, north of North Key, Tarpon Springs.
L m, found on Thalassia on mud flat. water tempera-
ture 13.3°C , salinity 23.6, Phillips, Greeley, Mann, 19
Dec 1958, USNM 103005 (leaf with burrow, 2 females
missing cephalon, male 2.5 mm. with | slide, ident. by
R.J. Menzies).

Colombia, Bahia de Nenguangue, near Santa Marta,
0.5 m, Thalassia, H.-G. Müller, 17 Jan 1986. ZMUC
(12 males, 2 females), 0-1 m, 5 Aug 1985, ZMUC (6
males, 4 females, | juvenile)

Diagnosis (male). Pleonite 5 dorsomedially with
a faint longitudinal sulcus. Pleotelson with l or 2
pairs of anteromedial puncta, if two pairs, then
puncta wider apart than longitudinally apart,
without carinae; with an anterolateral pair of
puncta or setae. Pleonite 5 0.5-0.6 times as long
as pleotelson. Dorsal surface of pleotelson with

139

scales partially fused, bordered posteriorly with
several short scale spikes, sometimes slightly pit-
ted anteriorly. Dorsal row of tubercles on lateral
crests of pleotelson, some bearing several short
Scale spikes; posterior margin fringed with 4-6
large stout setae between which are scale spikes
and short-sheathed setae, lacking posterior row
of tubercles, sometimes with irregular row of
short scale spikes in groups of 2-5.

Antenna 1 with 4 flagellar articles; second
article with about 6-1 1 aesthetascs. Flagellum of
antenna 2 with 5, sometimes 4 articles. Man-
dibular palp with 3 articles. Mandibular incisors
with rasp and file, rasp confined to distal half of
incisor. Lacinia mobilis of right mandible
straight, apex with several long or short teeth.
Epipod of maxilliped subtriangular, about 3
times as long as wide, reaching or just short of
palp articulation; epipod with true setae.

Secondary unguis of pereopod | bifid. Ventral
comb seta present or absent on merus of pereo-
pod 7, present on carpus of pereopods 2-7. Uro-
pod peduncle laterally with or without small
tubercles between plumose setae; endopod
0.75-1 times as long as peduncle.

Pleopod 2 with plumose setae up to same
length of exopod. Appendix masculina reaching
beyond apex of endopod of pleopod 2, articulat-
ing near midlength of endopod. Endopod of
pleopod 5 anterior to exopod, oval, 0.8 times as
long as endopod of pleopod 2.

Additional characters. Sculpturing on pleotelson
often sexually dimorphic, as female has just one
anteromedial pair of puncta, followed poster-
iorly by weak parallel carinae; female without
anterolateral puncta. Body length up to 3.2 mm
(Müller, 1988). Colour in alcohol pale yellow.

Distribution. Virgin Islands, West Indies (Men-
zies, 1957); Caribbean Sea of north Colombia
(Müller, 1988); Tarpon Springs, Florida (current
study). Depth: 0-4 m (Müller, 1988).

Substrates. Washed from the seagrass Thalassia
testudinum (Müller, 1988); leaves of the seagrass
Thalassia (current study).

Remarks. Some variations in the specimens
were noted. The ventral comb seta on the merus
of pereopod 7 was present in the specimens from
Tarpon Springs, and in some specimens from
Colombia, but absent in both types. The flagel-
lum of antenna 2 was drawn to have four articles
by Menzies (1957). However, this was difficult
to confirm from the slide that he had prepared,
as the tip was hidden by setae and debris. The
male paratype examined in situ had five articles,

140 L.J. COOKSON

Figure 6. Limnoria simulata Menzies. A-L, male, USNM 103005: A, pleonite 5 and pleotelson, dorsal view: B,
anterodorsal surface of pleotelson; C, posterodorsal surface of pleotelson; D, posterior margin of pleotelson; E,
pleopod 2; F, pleopod 5; G, maxilliped; H, flagellum of antenna 1; I, peduncle article 5 and flagellum of antenna
2: J, incisor of left mandible; K, right mandible; L, lacinia mobilis of right mandible.

ADDITIONS TO THE LIMNORIIDAE (CRUSTACEA) 141

C

man ke

TTIEITTTTIT IN TAREA AA
AN AN AN |

T

Figure 7. Limnoria simulata Menzies. A-C, male, ZMUC, paratype: A, pleonite 5 and pleotelson, dorsal view; B,
posterodorsal surface of pleotelson; C, posterior margin of pleotelson.

142 L.J. COOKSON

although the last was very small. Müller (1988)
also noted five articles in his specimens. The
posterior margin of the pleotelson bore six stout
setae in both type specimens, and in both
females from Tarpon Springs, whereas the male
from Tarpon Springs had four stout setae.

L. simulata, L. indica, L. saseboensis, and L.
hicksi Schotte, are all known to share, for some
specimens at least, the pattern on the pleotelson
of two anteromedial and one anterolateral pairs
of puncta.

L. simulata is most similar to L. indica, and
Müller (1988) synonymised L. indica with L.
simulata. However, I propose that the two
should remain separate species. L. simulata has
carinae on pleonite 5 much less defined than for
L. indica. In the specimens of L. simulata exam-
ined, there was a faint longitudinal sulcus on
pleonite 5. What might have been carinae on
either side of the sulcus, were so gradually
merged with the general surface, that I would
prefer not to call them carinae. Sulcus isthe term
used by Menzies (1957). L. simulata generally
has longer and more numerous setae dorsally on
pleonite 5 than L. indica. The anteromedial
puncta on the pleotelson of L. simulata are much
less defined than for L. indica. Often there is just
one anteromedial pair of puncta on the pleotel-
son of L. simulata, not two as always occurs for
L. indica. L. simulata lacks tubercles on the pos-
terior margin of the pleotelson, while the
majority of L. indica have these. Both species
have tubercles on the lateral crests, though often
less developed in L. simulata than L. indica. L.
simulata often has six stout setae on the pos-
terior margin of the pleotelson, while L. indica
has just four. Both the plumose setae on pleopod
2, and the appendix masculina, are longer in L.
simulata than L. indica. The rasp of the right
mandible is much reduced compared to that
found in L. indica.

Many of these differences are differences of

gradation. However, the dorsal structure of the
surface of the pleotelson is quite different. L.
indica has large solitary scale spikes, whereas L.
simulata has short teeth-like scale spikes
arranged in rows as groups of mostly three to
five. For L. simulata, some of these scale spikes
can also be seen on the tubercles on the lateral
crests. Another important ecological difference
is that L. simulata is a seagrass borer, while L.
indica has been found only in wood.

Acknowledgements

Material loaned for inclusion in this study was
generously provided by Dr Gary Poore (NMV),
Penny Berents and Dr Jim Lowry (AM), Dr
Torben Wolff and Dr Niel Bruce (ZMUC), Dr
H. Hertel (BAM), Dr lan Mannering (CM), Mar-
ilyn Schotte and Dr Brian Kensley (USNM),
B.S. Swami, Naval Chemical & Metallurgical
Laboratory, and Leela Devi. I would like to
thank both John Barnacle for the important col-
lections he made from various localities around
Australia, and Jan Cookson for drafting the
majority of figures.

References

Becker, G. and Kampf, W.-D., 1958. Funde der holz-
zerstórenden Isopodengattung Limnoria an der
Fesilandküste Indiens und Neubeschreibung von
Limnoria indica. Zeitschrift fiir Angewandte Zoo-
logie 45: 1-9,

Brearley, A. and Walker, D.I., 1993. Isopod borers in
scagrass species from south-western Australia.
Pp. 415-428 in: Wells, F.E., Walker, D.I., Kirk-
man, H., and Lethbridge, R. (eds), Proceedings of
the Fifth International Marine Biological Work-
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Memoirs of the Museum of Victoria 56(1):145-234 (1997)

A NEW GENUS OF MARINE ISOPOD (CRUSTACEA: FLABELLIFERA:
SPHAEROMATIDAE) FROM AUSTRALIA AND THE INDO-PACIFIC REGION

NIEL L. BRUCE
Zoologisk Museum, University of Copenhagen, Universitetsparken 15, DK 2100, Copenhagen , Denmark.

Abstract

Bruce, N.L., 1997. A new genus of marine isopod (Crustacea: Flabellifera: Sphaeromatidae)
s Australia and the Indo-Pacific Ocean. Memoirs of the Museum of Victoria 56: 145—

Oxinasphaera gen. nov. is established for a group of sphaeromatid species previously
placed in the genus Cymodoce Leach, 1818. Apomorphic characters allowing easy recog-
nition of genus are (in the male), hardened prominent spikes on the anterior margin of the
antennule peduncle, on the epistome (or with a transverse blade), and on the posterior of the
pereonites, maxilliped palp articles 3 and 4 with greatly elongated medial margin, article 5
also greatly elongated, all with an obliquely truncate densely setose apex, uropod endopod
with deeply bifid toothed apex and uropod endopod with hardened prominent spikes at
ventrodistal apex. Additionally, the pleon and pleotelson are dorsally denticulate, the pleo-
telson usually with a divided posterior margin but without a distinct exit channel. Species
transferred from Cymodoce are C. tuberculosa Stebbing, 1873, C. tripartita Richardson,
1910, C. multidens Richardson, 1910, C. bispinosa Baker, 1910, and C. australis Baker,
1929, The following new species are described: Oxinasphaera bisubula sp. nov. (type
species), O. aylostera sp. nov., O. copiapoa sp. nov., O. corypantha sp. nov., O. denmoza sp.
nov., O. epostoa sp. nov., O. frailea sp. nov., O. islaya sp. nov., O. kensleyi sp. nov., O. lobivia
sp. nov., O. lowryi sp. nov., O. matucana sp. nov., O. obregonia sp. nov., O. parodia sp. nov.,
O. poorei sp. nov., O. rebutia sp. nov., O. thetisae sp. nov., O. tual sp. nov.

The genus is recorded from South Africa, the Western Indian Ocean, around the Aus-
tralian coastline, New Caledonia, Indonesia, Papua New Guinea and the Philippines. All
species but one are recorded from shallow water at depths from the intertidal generally to
within 100 m; one species is recorded from a depth of 400 m. Many specimens and most
species are recorded as having been collected from sponges. A phylogenetic analysis of the
species is undertaken and brief comments given on the relationships within the genus and on
the biogeography of the genus. A key to the named species is given.

Contents

Introduction 146
Material and Methods 146
Sphaeromatidae 150
Oxinasphaera gen. nov. 151
Characters of taxonomic utility and general morphology 153
Relationships 154

Character discussion 154

Character list 155

Results 156

Discussion of trees 156
Key to the world species 157
Oxinasphaera bisubula sp. nov. 160
O. parodia sp. nov. 170
O. lobivia sp. nov. 173
O. frailea sp. nov. 176
O. denmoza sp. nov. 178
O. copiapoa sp. nov. 181
O. multidens (Richardson, 1910) 184
O. corypantha sp. nov. 186
O. obregonia sp. nov. 188
O. tual sp. nov 188

145

146

. rebutia sp. nov.

. aylostera sp. nov.

. tuberculosa (Stebbing, 1873)
. epostoa sp. nov.

. lowryi sp. nov.

. thetisae sp. nov.

. bispinosa (Baker, 1910)

. tripartita (Richardson, 1910)
kensleyi sp. nov.

australis (Baker, 1929).
matucana sp. nov.

poorei sp. nov.

O. islaya sp. nov.

Undescribed species
Acknowledgements

References

OODODEGSOGOGOD

Introduction

The genus Cymodoce Leach, 1814 is large, con-
taining a wide array of morphologically diverse
species. The genus and the European species,
including the type species, were reviewed by
Dumay (1972a, b, c, d). The most detailed recent
description to the genus is that of Harrison and
Holdich (1984), who gave an account of the Aus-
tralian species, together with extensive com-
ments on the confused taxonomic state of the
genus. Harrison and Holdich (1984, appendix 2)
also listed the known species and gave further
discussion on the status of the numerous mis-
placed species that, according to their percep-
tion, did not appear to belong to the genus.
Among the species that Harrison and Holdich
(1984) discussed was a group of distinctive
species related to Cymodoce tuberculosa Steb-
bing, 1873. Harrison and Holdich (1984), with-
out giving any reasons or descriptive comments
relating to morphological criteria, suggested that
these species would "require the formation of a
new genus to house them." This group of often
common and very distinctive Indo-West Pacific
isopods, here defined as a new genus, is easily
recognizable by the males having the antennule
peduncle article 1 with comb-like arrangement
of white prominent downwardly directed spikes,
similar prominent spikes on the epistome, and
prominent spikes arranged in transverse bands
across the dorsum of pereonites 2, 3, or 4 -7. The
generally very nodular and setose body surfaces
and the deeply bifid short uropod exopod and
cylindrical spiked uropod endopod also serve to
distinguish the genus. Although the new genus
can be clearly defined with several unambiguous
autapomorphies, the remaining species of
Cymodoce still present a no less confused assem-

N. L. BRUCE

191
193
195
199
202
205
208
214
218
218
222
225
227
231
232
232

blage, in reality containing species that should
be placed in several genera. The arguments and
discussion concerning Cymodoce have been pre-
sented by Harrison and Holdich (1984) and are
not repeated here.

The new genus is here defined, and the species
described together with an identification key.
The genus is can be clearly delimited by several
autapomorphies, and as monophyly can there-
fore reasonably be assumed, a cladistic analysis
was performed to examine the relationships
between the species, to more objectivley dis-
criminate the species groups, and to assess
character distribution and apparently homopla-
sious characters.

Material and methods

Material for study was obtained from Australian
state museums, by far the largest proportion
being that held by the Museum of Victoria, Mel-
bourne and the Australian Museum, Sydney.
Additional material was borrowed from the
other institutions listed, but the only unreported
material obtained other than from Australia
institutions was that which had been collected
by Th. Mortensen early this century and held at
the Zoologisk Museum, University of Copen-
hagen.

Methodology follows that of Bruce (1994b),
except that dissected appendages were drawn
using Nomarski illumination, and the scanning
electron microscope used was a Jeol JSM 840.
The cladistic analysis was performed using the
computer programme PAUP 3.1.1. The data set
was run using the heuristic search option (set-
tings tree-bisection-reconnection [TRR] and
MULPARS option). Trees and characters were
examined using the computer package

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

MacClade 3.03, and the cladogram figures (Figs
3 and 4) generated using MacDraw II.

Material for each species is categorized
according to type status; for named taxa the des-
ignation “non-type” refers to the principal study
material, and the designation “additional
material" refers to specimens that were merely
identified as belonging to the taxon after the
description had been finalized. The designation
"non-type" for new taxa refers to material
specifically excluded from type status because of
morphological differences, disparate geographic
location, or that the specimens were merely
identified as belonging to the taxon after the
description had been finalized.

Etymologies. Except where otherwise stated,
epithets are generic names of cacti and allude to
the spinose body surfaces of these isopods; these
names are to be treated as nouns in apposition.
The names were obtained from Riha and Subik
(1987).

Terminology. The species of the genus are in
many cases distinguished by the ornamentation
of the antennule, epistome, and dorsal body sur-
faces. Particular terms have been coined to
facilitate the description of these features (Fig.
1). Spikes — are hardened, usually acute cuti-
cular processes, which occur on antennule

147

peduncle articles 1 and occasionally 2, anterior
margin of the cephalon, epistome, dorsal surface
of pereonites, pleon and pleotelson, and also on
the uropods; tubercles — are low rounded or
acute processes, not always as prominent or
hardened as spikes, restricted to the pleon and
pleotelson; pleonal boss — the posteriorly
expanded medial part of the pleon posterior
margin. Antennule peduncle spikes are ident-
ified as (Fig. 1): anterior, posteroproximal, pos-
teromedial and posterodistal .

Descriptions. The type species has been
described in full detail, while descriptions of the
remaining species contain the principal charac-
ters distinguishing species. A representative
species typical of each species group has, where
sufficient material was available, also been
described in more than minimal detail. These
species are: O. tuberculosa and O. tripartita. Oxi-
nasphaera bispinosa and O. islaya sp. nov. Other
distinctive species are similarly treated.

Abbreviations. AM, Australian Museum, Syd-
ney; AMSBS, Australian Museum Shelf Benthic
Survey; NMV, Museum of Victoria, Melbourne;
TM, Tasmanian Museum and Art Gallery,
Hobart; Qld, Queensland, Australia; QM,
Queensland Museum, Brisbane; NTM, North-
ern Territory Museum, Darwin; SAM, South

Figure 1. Terminology. A, epistome: as, anterior spikes (or anterior blade when a single transverse ridge); lo,
lateral lobe. B, antennule: as, anterior spikes; pp. posterproximal spikes; pm, posteromedial spikes; pd, poste-

rodistal spikes. C, antennule: anterior spikes irregular; pb, posterior blade; D, pleon and pleotelson: pb, posterior
boss; ps, pleonal spike; as, anterior spike; tn, telson nodules; If, lateral flange.

148 N. L. BRUCE

Figure 2. World-wide distribution of Oxinasphaera. Dots represent named species, triangles (western Indian
Ocean) undescribed species.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 149

© Dv D
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IGS eg e? d PLÈ Se ROMS EEG SF WW HM PY LE IRAS Lè

; inas 4 i from 84 trees, length, 131; consist-
. Cladograms of Oxinasphaera. A, strict consensus tree, unordered
DUNS 0 260: retention index 0.349; homoplasy index 0.740; B, strict consensus tree, character 9 ordered,
from 68 trees length, 108: consistency index 0.315; retention index 0.503; homoplasy index 0.685.

150

C27

ga:

S 10.11

20

N. L. BRUCE

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e .
& No} e N o
^. S gè. L L Va AN
ERO PSE FK SOI
TÈS L'E ILL YP
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21,24
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Figure 4. Cladogram of Oxinasphaera using successive character weighting. Length, 90; consistency index 0.378;
retention index 0.624; homoplasy index 0.622. Solid bars = apomorphy, grey bar = homoplasy, white bar =

reversal.

Australian Museum, Adelaide; SAfM, South
African Museum, Durban; USNM, National
Museum of Natural History, Smithsonian Insti-
tution, Washington, DC, USA; WAM, Western
Australian Museum; ZMUC, Zoologisk
Museum, University of Copenhagen, Den-
mark.

PMS, plumose marginal setae; CP, circumplu-
mose.

NSW, New South Wales; NT, Northern Ter-
ritory, Qld, Queensland; SA. South Australia;
Vic., Victoria; WA, Western Australia.

Sphaeromatidae Latreille
Sphaeromatinae Latreille

Recent publications have called into question
the validity of characters on which the sphaero-
matid subfamilies are diagnosed and the difh-
culty of their interpretation, in particular the
presence and absence and development of pleo-
podal ridges (Bruce, 1993, 1994a, 1994b, 1995;
Brusca and Wilson, 1991; Kussakin and Malyu-
tina, 1993; Wägele, 1989). The subfamilies can
currently only be considered to be delimited by

their constituent genera. It is currently prema-
ture to offer a new diagnosis, and a thorough
revision of the Sphaeromatidae will necessitate
the relocation of many genera, and the develop-
ment of new concepts for genera and of generic
relationships within the family. Those com-
ments notwithstanding, the new genus described
here unequivocally belongs together with that
group of genera that form the core of the sub-
family Sphaeromatinae and includes, for
example, the genera Sphaeroma Bosc, 1802,
Neosphaeroma Baker, 1926, and Cymodoce.
Two species 1n Oxinasphaera, O. islaya sp. nov.,
and Oxinasphaera sp. 1, lack pleopodal ridges.

The development of the ornamentation of the
posterior margin of the pleon and pleotelson is
as equally variable as the degree of development
of the pleopodal ridges. Most species described
here have the posterior margin with two sub-
median excavations, which may also be
described as a median excavation in which lies a
process (in some species either interpretation is
valid), but one species (O. islaya) has an almost
entire posterior margin, and two (O. tripartita
and O. kensleyi) have a well developed dorsal

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 151

lobe overriding a median excision. In the O.
tuberculosa clade of species the lobe appears dor-
sal to the posterior notch, but it does not over-
ride the notch.

In sphaeromatid taxonomy, the presence of
posteriorly directed processes, single or paired,
on the posterior margin of the pleon (or pereon-
ite 6 or 7) has long been axiomatically con-
sidered to be of generic merit. The generic pairs
of Dynoides Barnard, 1914 and Clianella Boone,
1923 or Isocladus Miers, 1876 and Exosphae-
roma Stebbing, 1900, for example, are dis-
tinguished solely by the presence or absence of
such a process which, as can be readily demon-
strated, show intermediate conditions. Further-
more, such processes can readily be recognized
as homoplasious, also occurring in apparently
unrelated genera. In the genera Dynoides and
Clianella two species, D. brevispina Bruce, 1980
and D. brevicornis Kussakin and Malyutina,
1987, have greatly reduced pleonal processes,
which then leaves no distinguishing characters
between these two genera. The recognition of
autapomorphies should allow for a more mean-
ingful interpretation of these variable and
homoplasious characters. Such is the case for
Oxinasphaera, in which the pleonal boss ranges
from absent to with a pair of conspicuous
elongate processes, but recognition of the diag-
nostic autapomorphies as the characters of sig-
nificance allows the retention of those species
within one genus.

Oxinasphaera gen. nov.

Type species, Oxinasphaera bisubula sp. nov., here
designated.

Diagnosis of male. Pereonites 2-7 with 1 or 2
rows of distinct, usually acute spikes, segments
3-6 usually with 2 rows, anterior row of which is
larger than posterior row. Pleotelson posterior
margin medially excavate, with median process
set within or above indentation, occasionally
entire; without shallow open exit channel.
Antennule peduncle article 1 anteriorly with row
of prominent ventrally projecting spikes, poster-
iorly with 0-2 spikes. Epistome anterior margin
with 1, or 1 pair of prominent ventrally projec-
ting round acute or flat truncate spikes; postero-
lateral lobes with or without short spikes. Maxil-
liped palp article 2 medial margin strongly
produced, articles 3-4 with medial margins
greatly elongated (3-4 times as long as medial
width) and finger-like, article 5 elongate, lying
parallel to process of article 4; distolateral mar-
gins of articles 2-5 provided with abundant long

setae which are distally finely pectinate. Pleopod
2 appendix masculina about as long as endopod,
robust (4-9 times as long as maximum width),
not extending beyond or only slightly beyond
distal margin of ramus. Uropod attached mid-
laterally on pleon; exopod short, about half
length of exopod, slender, apex deeply bifid,
with 2 blade-like points; endopod projecting
beyond posterior of pleotelson, round in section,
not lamellar, apex with acute point and 1-3
prominent acute spikes, often with additional
abundant tubercles.

Description of male. Dorsal surfaces of body
granular and nodulose, often pilose. Cephalon
anterior margin often with anterolateral row of
acute spikes; laterally with distinct subocular
groove, a continuation of an anterior ridge, sec-
ond deep groove below the ridge; eyes large,
facets distinct, with prominent posterior lobe.
Coxae not distinctly demarcated. Pleon of 4 seg-
ments, segment 1 largely concealed by pereonite
7. segments 2-4 indicated by 2 distinct suture
lines running to posterolateral margin of pieon:
posteromedial margin of pleon with or without
boss, with or without posteriorly directed pro-
cesses. Pleotelson with anterolateral flange.
Pleonal sternite present, weakly developed.

Antennule peduncle article 1 more than twice
as long as 2, robust, article 3 slender, all articles
collinear; flagellum about as long or slightly
longer than peduncle, extending to about pos-
terior of pereonite 1. Antenna peduncle article 1
short, articles 2 and 3 shorter than 4-5 which are
longest; flagellum short, extending to about pos-
terior of pereonite l.

Epistome anteriorly acute, narrowly rounded
or truncate, medially constricted. Labrum unor-
namented, may be nodulose or setose. Mandible
incisor multicuspid; molar process prominent,
crushing, provided with lateral scale teeth; left
mandible with prominent lacinia mobilis; both
mandibles with spine row of 5 or 6 spines; palp
3-articled, article | longest, 3 shortest. Maxillule
lateral lobe with about 13 spines on gnathal sur-
face, most of which are prominently serrate,
with further 2 biserrate spines and distal surface;
medial lobe with 4 prominently serrate and plu-
mose spines, with further 2 short simple spines.
Maxilla with all articles well developed, with
prominent setae on lateral and middle lobes,
medial lobe with several robust CP spines and
additional slender spines. Maxilliped endite dis-
tal margin with 4 long laterally curving and 4
short CP spines, dorsal distolateral margin with
2-3 CP spines.

152 N. L. BRUCE

Pereopods all ambulatory, pereopods 1-3 sub-
similar, and more robust than 4-7; posterior
margins of merus and carpus with biserrate
spines, without prominent setulose fringe; dac-
tylus posterior margin with cuticular scale row,
with prominent simple secondary unguis and 1
flattened seta arising at lateral base of secondary
unguis; lateral margin with single seta (‘propodal
organ’), rarely with 2. Pereopods 6 and 7 pos-
terior and distal margins of carpus with con-
spicuous biserrate and trifid spines.

Penes paired, unfused, medially adjacent at
posterior of sternite 7, moderate to elongate in
length, extending at least to pleopod peduncles;
not conspicuously narrowed or acute.

Pleopods 1-3 both rami with PMS. Pleopod 1
endopod extending beyond exopod; exopod axis
oblique to peduncle, with simple spine at prox-
imolateral angle; peduncle with 3 coupling
hooks on medial margin. Pleopod 2 with appen-
dix masculina subbasally attached. Pleopods 3-
5 exopods with entire transverse suture. Pleo-
pods 4 and 5 exopods with thickened ridges or
folds (rarely without), with lateral margin
thickened, with row of short simple setae; endo-
pods without thickened ridges or folds. Pleopod
5 exopod with 3 or 4 scale patches.

Female, ovigerous: Body with nodules weakly
developed or absent; coxae visibly demarcated.
Pleotelson posterior margin entire, upturned,
with shallow ventral median depression, with-
out exit channel. Mouthparts metamorphosed.
Antennule peduncle article 1 anterior margin
finely denticulate, without spikes. Pereopod 7
with spines simple or finely serrate, relatively
longer and more slender than those of male.
Uropod endopod flat, with apical point.

Brood pouch made up of short oostegites,
overlapping slightly at mid-line, and arising
from sternites 1—4; eggs held in internal pouches
within body.

Female, non-ovigerous: Generally similar to ovi-
gerous female, but may retain trace of male
pleonal ornamentation, ventral margin of pleo-
telson posterior margin with ventrally flat rim,
with median depression not reaching posterior
margin, without exit channel; dorsal posterior
margin not upturned. Mouthpart, pereopodal
and pleopod morphology generally similar to
that of male.

Remarks. The defining apomorphies for the
genus are in the male: antennule peduncular
article | with hardened prominent spikes on the
anterior margin, epistome anteriorly with l or 2

hardened prominent spikes or transverse ridge;
maxilliped palp articles 3 and 4 each with a
greatly elongated medial margin, article 5 also
greatly elongated, all with an obliquely truncate
densely setose apex; posterodorsal margin of
pereonites 2-7 with 1 or 2 rows of conspicuous
spikes, uropod exopod with deeply bifid toothed
apex and uropod endopod with hardened
prominent spikes at ventrodistal apex.
Additionally, the pleon and pleotelson are dor-
sally denticulate, the pleotelson usually with a
divided posterior margin but without a distinct
exit channel. The uropod rami are both round in
section with the exopod about half as long as the
endopod.

Species of this genus are easily recognizable by
the males having the antennule peduncle article
1 with comb-like arrangement of prominent
white downwardly directed spikes, similar
prominent spikes on the epistome, and promi-
nent spikes arranged in transverse bands across
the dorsum of pereonites 2, 3, or 4-7. Other
characters include the very nodular and setose
body surfaces, deeply bifid short uropod exopod
and cylindrical spiked uropod endopod.

The recognition ofthe autapomorphies for the
genus allows recognition that the presence or
absence of pleonal processes, and differences in
morphology of the penial processes, appendix
masculina, pleopodal ridges and pleotelson pos-
terior margin are not necessarily of generic
merit.

Distribution. Species of Oxinasphaera are found
throughout the Indo-Pacific Ocean, with
recorded localities from the south-western
Indian Ocean at South Africa [Oxinasphaera
kenslevi sp. nov.], to the Philippines in the
north-cast [Oxinasphaera multidens (Richard-
son, 1910) and Oxinasphaera tripartita
(Richardson, 1910)], and to New Caledonia in
the east (Oxinasphaera corypantha sp. nov.)
(Fig. 2). The genus has been widely and com-
monly collected around the entire Australian
coastline with the exception of some remote and
ditficult to collect localities. Usually collected
within the continental shelf, but with one species
(Oxinasphaera parodia sp.nov.) recorded from a
depth of 400 metres. Several undescribed
species exist in the North-western Indian Ocean,
and are currently under study at the USNM (by
B. Kensley and M. Schotte). Other Indo-Pacific
regions may well yield further species once col-
lections have been made, and at present I regard
the distribution of the genus as incompletely
known. The relatively well recorded sphaero-

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

matid fauna of New Zealand (Hurley and
Jansen, 1977) apparently lacks representatives
of this genus.

Thesingle most commonly recorded or associ-
ated habitat for species of this genus is from, or
with, sponges. It is probable that much of the
collected material that has not been noted as
being taken from sponges is the result of these
animals exiting from their host once it has been
collected. Large samples taken by trawl or epi-
benthic sled, such as some of those of Oxinas-
phaera bispinosa, may indicate that some
species are also free living.

Etymology. The name is a combination of the
Greek oxina (harrow or rake, alluding to the
antennule peduncle morphology), with the end-
ing -sphaera, indicating familial affinity (femi-
nine).

Characters of taxonomic utility and general mor-
phology

Cephalon. The cephalon is frequently orna-
mented, and may be posteriorly granular or
pitted (e.g., Figs 11A-C) or smooth (Fig. 35A).
The absence or presence, distribution and form
of the spikes along the anterior margin, arrange-
ment of rostral spikes (absent, single or double)
and medial nodules should all be noted, the first
two characters being critical in species evalu-
ation.

Pereonites. Characters to note are: the arrange-
ment of pereonal spikes, presence of prominent
spikes on pereonite 1 or 2, and also which per-
eonites (e.g., 2-7, 3-7) have spikes. Spike rows
are usually double (Fig. 11F), rarely single or
triple, with the anterior row more prominent
and smaller nodules between the spike rows (Fig.
43B); note also the shape of spikes.

Coxae. Coxae vary little between species, but
differences can be seen in {he posterolateral mar-
gins of 5 and 6, some species being rounded,
others straight.

Pleon. There are several important and easily
observed characters to note, particularly the
presence or absence of posteriorly directed
paired processes (Figs 43C, 47C), the presence of
a posterior boss with paired lateral spikes (Fig.
11D), and the arrangement of the larger
tubercles. The pleonal boss is usually distinct
and armed with posteriorly directed spikes at
each posterolateral corner. In some species the
boss is altogether absent, while others lack a boss
but retain thé spikes, and in some the boss
approaches the state of processes.

153

Pleotelson. There are a number of important
characters to note, including the arrangement of
anterior spikes and of the larger tubercles; most
species have the dorsal surface covered with
papillose nodules (Figs 11E, 14E, 43D),
occasionally not evident (Fig. 16D); some
species have longitudinal rows of prominent
rounded nodules running from the posterior
margin and median lobe (Fig. 35C). The shape of
the posterior margin is usually with a median
excavation in which lies a coplanar lobe. Other
conditions of the pleotelson posterior margin
are: posterior margin flat and produced with two
submedian excavations; almost entire; with dis-
tinct dorsal lobe overlying median excavation;
median lobe large, laterally flanked by deep
grooves. The degree of setation should be
noted.

Cuticle. The cuticle is variously polished,
smooth or pitted, and ornamented in some
form. Some species have papillose nodules (Figs
11C, E, 14F), while in others the nodules are
simple (Fig. 21F) or absent (Figs 16F). Oxinas-
phaera islava has large shallowly domed
tubercles interspersed with smaller mushroom-
shaped nodules (Fig. 58D). In most species the
cuticle surface has scattered globular structures
(Fig. 12H), also found in Discidina Bruce, 1994b
(figs 54A, B). In only Oxinasphaera bispinosa
were these observed to occur in distinct clearly
defined pits (Fig. 43D).

Antennule. The morphology of antennular ped-
uncle article 1 is critical in species discrimi-
nation. Characters to note are the number of
anterior spikes, number and position of pos-
terior spikes, presence of a posterior blade and
whether spikes are regular or irregular. the pres-
ence or absence of spikes on antennule pedun-
cular article 2 is useful to note. The relative
length and the number of flagellar articles varies
slightly between species.

Antenna. Details of the antennal peduncle and
flagellum scarcely vary between species. The
degree of setosity of peduncular articles 4 and 5,
and number of flagellars article may vary.

Epistome. This character is of critical import-
ance in discriminating species. It should be
noted whether the spikes are round in section or
antero-posteriorly flattened, close set or set
apart, how many spikes are present, and whether
the spikes are set on or form a transverse ridge;
the presence of secondary spikes on lateral lobes
should be noted.

154 N. L. BRUCE

Mouthparts. Mouthparts are consistently uni-
form throughout the genus and of little use in
species discrimination. The mandible (Figs
11G, H, 43D, H) is of the typical generalized
sphaeromatid form with a prominent lacinia
mobilis on the left mandible, spine rows on both
mandibles and a peripherally toothed molar
with a flat unornamented surface. The maxilli-
ped palp setae are terminally plumose (Fig.
I2B). The endite dorsal surface has abundant
sctae, laterally with 3 large serrate setae (Fig.
44C).

Pereopods. Pereopods are generally very similar
throughout the genus. Differences can be
observed in the general robustness, number of
spines on the propodal palm, and the relative
length of both the carpal spines. Minor differ-
ences can be observed in the relative dimensions
of the pereopods. The posterior margin of per-
eopods 1-3 is provided with biserrate spines
(Fig. 12E), and distal margins of the carpus of
pereopods 6 and 7 with biserrate and trifid
spines (Fig. 44E). The dactylus has a robust sec-
ondary unguis with 2 associated setae (Figs 12C,
44D), the distolateral margin of the dactylus
with a single seta (Fig. 21 D) except for O. lowryi
which has 2.

Penes. These generally extend to the pleopod
peduncles. Some species have slender and
elongate penes that fit into a groove on the
medial margin of pleopod 1 endopod.

Pleopods. Pleopods are very uniform throughout
the genus. Pleopod | endopod medial margin
may be regular or with a ventral groove (when
receiving elongate penial process), or with dis-
tomedial lobe. Pleopod 2 appendix masculina
may be of even width or basally dilated, distally
narrowed, straight or curving laterally; the apex
varies from apically blunt to acute, usually
slightly longer than exopod, in some species not
extending beyond distal margin of endopod.
Pleopod 5 has 3 patches of distally fringed scales,
the distomedial patch being lobate (Fig. 44F,
44G).

Uropods. Often nearly identical among related
species; characters to note are the relative length
afthe exopod in relation to the endopod, the size
of the exopod spikes, the number and size of the
distal exopod spikes, and the setosity of the
rami. Uropodal setae are terminally roughened
(Fig. 12F), a character which encourages silt to
adhere to the animal. The uropod cuticle is simi-
lar to that of the pleotelson, the only appendage
for which this is the case.

Relationships

Sister group relationships have yet to be clearly
established in the Sphaeromatidae, and the
monophyly of many of the larger genera is open
to question. Cymodoce is one such genus, but the
type species and a group of closely related
species have been described in detail (Dumay,
1972a, b, c, d) allowing for a clear concept of the
genus (sensu stricto, based on the type species) to
be used when making outgroup comparisons.
Sphaeroma was chosen as the second outgroup
as it is a generally considered to be a monophy-
letic plesiomorphic genus, and was used to
polarise pleon and pleotelsonic character
states.

Cymodoce (sensu stricto) is regarded here as
the sister group to Oxinasphaera. The genus
Cymodoce has not been assessed in terms of cla-
distic criteria, and unambiguous synapomor-
phies with Oxinasphaera are difficult to ident-
ify, in part because the distribution of these
characters beyond the genera in question is not
known with certainty. Both genera have a simi-
lar mouthparts (particularly the elongate lobes
of the maxilliped palp), pereopodal and pleopo-
dal morphology, both commonly have some
form of pleonal boss, and both have a median
telsonic notch in which usually lies a simple
coplanar lobe, all of which are potential syapo-
morphies. The two genera can immediately and
easily be separated by the apomorphies ident-
ified (above) for Oxinasphaera, and also that
Cymodoce (sensu strictu) has a lamellar and lan-
ceolate uropodal exopod. Possible apomorphies
for Cymodoce (sensu stricto) include the pleotel-
son with 2 subparallel longitudinal ridges, pleo-
telson apex with two submedian notches, pleo-
telson with a prominent hardened hemispheri-
cal dome anterior to the posterior pleotelsonic
notches, uropod endopod flat and thickened
(eastern Atlantic species) or cylindrical in sec-
tion with a single distal apical spike (Australian
species). Detailed discussion of characters states
for Cymodoce, a large genus in desperate need of
revision, is beyond the scope of the present
study.

Character discussion

A generalized illustration of critical characters
of the antennules and pleon is given in Fig. 1.

Cephalic spikes (characters | and 2) the
anterior margin spikes are present in various
degrees of prominence, and are scored 0 only
when absent or so minute as to be indistinguish-
able from tubercles; O. poorei lacks spikes but

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

has a distinct anterior flange, and was recorded
as ornamented (1) rather than unornamented
(0) presence of 3 small rostral spikes was
recorded as |.

Pereonites (characters 3-8): records the spike
configuration. Very small spikes are recorded as
0 for characters 5 and 6, although they are not
absent. Character 8 is ambiguous, and not
clearly definable.

Pleon (characters 9-11): character 9, poster-
iorly directed processes, was ordered, the paired
processes being regarded as derived from a
pleonal boss; the plesiomorphic state (smooth,
no boss) is supported by outgroup comparison
and also developmental evidence as postem-
bryos, mancas and immature specimens lack the
pleonal boss; paired processes cannot be derived
directly from the plesiomorphic state, but the
possibility of reversals from state 2 to state l is
recognized. Posteriorly directed spikes (10) are
the spikes at the anterolateral angles of the
pleonal boss; lateral acute tubercles (11) are the
small tubercles commonly present on the lateral
margins of the pleonites 2, 3 and the lateral
flange.

Pleotelson (characters 12 -18): anterior spikes
(12) are those that oppose the pleonal spikes or
processes; the presence of a dorsal lobe (15)
precludes characters 16 and 17, as the notch is
without a median lobe ventrally.

Antennule (characters 19-23) and epistome
(characters 24-26) record the spike configur-
ation. The distinction between epistome spikes
and a transverse blade is not always clear: in
those species with paired acute spikes the spikes
are often basally united and flattened; in those
species with a blade the transition between a
transverse series of tubercles, small spikes and a
blade can be equally unclear.

Penes (character 27): penial processes in
Cymodoce are elongate, but are short in Sphae-
roma. The plesiomorphic condition is regarded
as short and unfused.

Pleopods (characters 28-30): The plesiomor-
phic condition for pleopod | (28) is with a
simple medial margin to the endopod as in
Sphaeroma; in many genera and species the
presence of a grooved medial margin is associ-
ated with elongate penial processes (e.g.. Cymo-
doce), the penes fitting into the groove. This not
always so, as shown by O. obregonia which has
elongate penial processes but no pleopodal
groove. The appendix masculina (29 and 30) in
both outgroups is elongate and evenly tapered
with a slender or narrowly rounded apex; being
abruptly narrowed and having a blunt apex are

155

both regarded as independent plesiomorphic
states.

Uropod (character 31): Sphaeroma has lamel-
lar uropods which lack terminal spikes. Indo-
Pacific species of Cymodoce have the uropodal
endopod similar to that of Oxinasphaera, but
with a single terminal spike, and have a longer
lamellar exopod that is not apically bifid.

Character list

Outgroups: Cymodoce, the probable sister
group; Sphaeroma, a generalized plesiomorphic
genus with regard to the pleon, pleotelson and
uropods. * = characters with assumed polarity.

1. Cephalon anterior margin: unornamented
(0); with distinct spikes or ridge (1).

2. Rostral spike: absent (0); single (1), double
(2).

3. Pereonite 1: unornamented (0); nodular
(1).

4. Pereonite 2: without prominent median
nodule(s) (0): with prominent median nod-
ule(s) (1).

5. Pereonite 2: without distinct spike rows (0);
with distinct spike rows (1).

6. Pereonites 4-7: without distinct spike rows
(0); with distinct spike rows (1).

7. Pereonites 4-7: spike rows subequal in size
(0); anterior spike row larger than posterior
(1).

8. Pereonite 7: posterior margin even (0); pos-
terior margin weakly bilobed (1).

9. Pleon posterior margin: regular, even, not
produced (0); posterior margin with medial
boss (1), with posteriorly directed processes
(2).

10. Pleon posterior dorsal surface: without
spikes (0); with spikes (1).

11. Pleon posterolateral surface: without acute
tubercles (0); with acute tubercles (1).

12. Pleotelson anterior margin: without pair of
submedian spikes (0); with pair of sub-
median spikes (1).

13. Pleotelson posterior margin: not posteriorly
flattened or extended (0); posteriorly flat-
tened and extended (1).

14. Pleotelson posterior margin: entire (0); with
simple shallow submedian notches (1), with
median notch with deep submedian grooves
(2).

15. Pleotelson posterior margin: without dorsal
lobe (0); with dorsal lobe projecting over
notch (1).

16. Pleotelson posterior margin: median lobe
slender, extending to posterior margin (0);

156 N. L. BRUCE

median lobe short, distinctly rounded, not
extending to posterior margin (1).

17. Pleotelson posterior margin: median lobe
without distinctly rounded tubercle(s) (0);
with such tubercles(s) (1).

18. Pleotelson posterior margin: shallow, flat in
lateral view (0); deep-sided in lateral view
(1).

19. Antennule peduncle article 1*: anterior
spikes not markedly flat, distally acute (0);
markedly flat, distally blunt or truncate
(1).

20. Antennule peduncle article 1: prominent
posterior spikes absent (0); prominent pos-
terior spikes present (1).

2]. Antennule peduncle article 1*: anterior
spikes regular in size (0); anterior spikes
irregular in size (1).

22. Antennule peduncle article 1: without pos-
terior blade (0); with posterior blade (1).

23. Antennule peduncle article 2: without small
anterior spike(s) (0); with small anterior
spike(s) (1).

24. Epistome anteriorly*: with spikes (0); with
transverse blade (1).

25. Epistome anteriorly*: with two spikes (0);
with single spike (1).

26. Epistome anterior spikes*: acute, round in
section (0); flattened, distallv truncate (1).

27. Penial processes: short, robust (0); elongate,
distally slender, extending to pleopod | ped-
uncle (1).

28. Pleopod | endopod: medial margin simple
(0); medial margin with dorsal groove (1).

29. Pleopod 2 appendix masculina: evenly
tapered (0); distally abruptly narrowed
(1).

30. Pleopod 2 appendix masculina*: distally
acute (0); distally bluntly rounded (1).

31. Uropod endopod: with single prominent
distal spike (0); with 2 or more prominent
distal spikes (1).

Results of analysis

The matrix of 25 taxa by 31 characters was
treated using the programme PAUP 3.1.1.
Character transformations were investigated
using the apolist option and examined using
MacClade 3.03. The tree data is given in the cap-
tions to figures 3 and 4. Initially 84 trees were
obtained, these being used to generate the strict
consensus tree (Fig. 3A); with character 9
ordered weighted 68 trees were obtained, the
strict consensus tree shown in Fig. 3B. The con-
sensus trees are largely not dichotomously
resolved, but the succesivley weighted tree (Fig.

3B) does demonstrate three distinct clades of the
O. bisubula polychotomy, the O. tripartita clade
and the O. tuberculosa clade. The successively
weighted tree (Fig. 4, character 9 ordered) main-
tains the principal clades shown by the consen-
sus tree, but the O, tripartita clade is shown as
part of a larger clade with the O. tuberculosa
clade. In both cases the level of homoplasy is
high, with a homoplasy index of 0.685 and 0.622
respectively.

Discussion of trees

The strict consensus tree (Fig. 3B) demon-
strates six dichotomously unresolved clades.
The O. bisubula clade is defined by two apomor-
phies: prominent posterior antennule spikes
(character 20) and the posterior margin with a
boss (character 9). Both of these characters occur
as homoplasies in other clades (O. lowryi,
character 9) or are reversed within the clade (O.
bispinosa, character 20, O. obregonia, character
9). Within this polychotomy there is a distinct
clade consisting of the species O. bispinosa, O.
tripartita and O. kensleyi. This clade is defined
by the following apomorphies: pleon with pos-
teriorly directed processes (character 9),
elongate penial processes (character 27), and
pleopod 1 endopod with a grooved medial mar-
gin (character 28). The latter two character states
are homoplasious, character 27 occurring in 0.
multidens, O. tual and O. obregonia, and charac-
ter 28 in O. corypantha and O. tual.

The O. tuberculosa clade is defined by the
autapomorphies of the deeply grooved pleotel-
son posterior margin (character 14), flattened
antennule spikes (character 19) and flattened
epistome spikes (character 26). The latter two
characters are homoplasious, also occurring in
O. bispinosa.

Of the remaining smaller clades, the O. aylos-
tera/O. rebutia clade is discussed below in
relation to the successivley weighted tree, as is
the O. australis/O. matucana clade. Both the
single taxon clades of O. poorei and O. islayi
inevitably lack unique apomorphies, although
both species have a number of defining autapo-
morphies

The successively weighted cladogram (Fig. 4)
supports the principal clades identified in the
consensus tree, but differs in placing the O. bis-
pinosa clade as sister group to the O. tuberculosa
clade. The O. bisubula clade is defined by the
unique appearance of posterior spikes on the
antennule peduncle (character 20). The clade O.
aylostera and O. rebutia, sister group to the
remaining O. bisubula clade, is defined by

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

several autapomorphies: prominent nodules on
pereonite 2 (character 4), pleotelson median
lobe short (character 16), pleotelson median
lobe with rounded tubercles (character 17), and
the telson being deep in lateral view (character
18). The last two characters are somewhat weak
and subjective, and the distribution and distinc-
tion of tubercles between species is equally sub-
jective. Although the remaining species of the O.
bisubula clade are dichotomously resolved, the
relationships between these species is far from
clear as the nodes are principally defined by
homoplasies and reversals, while some of the
characters (e.g., character 8) demonstrably show
intermediate states.

The O. tuberculosa clade is defined by the
unique appearance of flattened antennular
spikes, although this character reverses in O. tri-
partita and O. kensleyi. The clade containing
both the O. tuberculosa clade and O. bisubula
clade is defined by the unique appearance of flat-
tened spikes on epistome, again being reversed
in O. tripartita and O. kensleyi.

The clade Oxinasphaera australis and O.
matucana is defined by the posterior margin of
the pleotelson being flattened and extended
(character 13).

Relationships of the O. bispinosa clade. Oxinas-
phaera tripartita and O. kensleyi form a closely
related species pair defined by the unique occur-
rence of a posterior blade on the antennule ped-
uncle and the prominent pleotelsonic lobe that
overrides the telsonic sinus. In common with O.
bispinosa, the two species also have elongate
penes, and pleopod 1 endopod with a grooved
medial margin. All three species have elongate
pleonal processes, a critical synapomorphic
character. Oxinasphaera bispinosa retains the
plesiomorphic condition for the pleotelson pos-
terior margin, while the antennule and
epistome spikes (characters 19 and 26) are apo-
morphic.

Oxinasphaera tripartita and O. kensleyi could
not be scored for characters 14, 16, 17, 25, and
26. This species pair is also plesiomorphic for

157

character 19, and therefore lacks the two defin-
ing apomorphies (characters 19 and 26) for the
O. tuberculosa clade. Although the clade appears
united by the apparent unique appearance of
pleonal processes, close scrutiny of the mor-
phology of these processes suggests that their
occurrence in O. bispinosa and in the O. kens-
leyi/tripartita pair may also be homplasious.
The shape of the processes in O. bispinosa is
elongate and the processes are narrow and close-
set, with a deep and narrow separation; and the
apex is not hardened or spike-like (Figs 40A,
43C). In the O. kensleyi/tripartita pair the pro-
cesses are short, narrowing rapidly to the apical
point, widely separate, with a distally hardened
(i.e., spike-like) apex (Figs 45A, 47B,C, 48 A,B).
This, together with the change in position of the
clade shown in the two trees (figs 3B, 4), suggests
both that the position of this clade and of the
species within it are yet to be fully resolved. It is
of interest to note that all the western Indian
Ocean species (USNM specimens, personal
observation) have elongate pleonal processes.

Distribution of the major clades. The O. tubercu-
losa clade is restricted to Australian coastal
waters with the species pair of O. tuberculosa
and O. epostoa occurring on southern and north-
ern tropical coasts respectively. The remaining
species in this clade occur in south-eastern Aus-
tralia,

The O. bispinosa clade has a disparate distri-
bution. The species pair O. tripartita and O.
kensleyi occur in the western Pacific and south-
eastern South Africa respectively; O. bispinosa
occurs in south-eastern Australia. Several other
undescribed species occur in the western Indian
Ocean but the relationships of these species
require examination before any comments can
be made on their distribution.

The O. bisubula clade occurs in Indo-Malayan
region and the south-western Pacific. This group
of species is absent from the western Indian
Ocean and is also absent from south-western
Australia.

Key to world species of Oxinasphaera
Notes: 1, the key identifies only males; females can be reliably identified only
by association with males. 2, antennule, except where otherwise stated, refers

to peduncular article 1.

l. Pleotelson posterior margin median lobe in dorsal position, set

above posterior notch ..

AE ERN e E ed 2

— Pleotelson posterior margin median lobe in coplanar position, set
within posterior notch or lobe absent .................... 7

158

to

9

LI.

N. L. BRUCE

Pleotelson posterior margin median lobe not extending beyond
posterior margin, flanked by deep grooves; antennule article 1
without posterior blade or spikes; penes short ............. 3
Pleotelson posterior margin median lobe extending beyond pos-
terior margin, not flanked by deep grooves; antennule article 1
with posterior blade; penes elongate ..................... 6
Pleonal boss present; pereonites 6 and 7 with single row of
SPI KES 005 2234502) n i r Troas en O. lowryi, p. 202.
Pleonal boss absent; only pereonite 6 with single row of spikes
Cephalon and pereonite 1 nodulose; pleon and pleotelson with
numerous large spikes; epistome with single short truncate
SPIKO" a LA N a RM e Moria Ea cel jo O. thetisae, p. 205.
Cephalon and pereonite | unornamented; pleon and pleotelson
without numerous large spikes; epistome with 2 truncate
SPIRES ett „A e kon ca ka Song ch ta ked biz aton 5
Pereonites 2 and 3 densely nodulose; pereonite 7 with 2 spike
rows; pleon and pleotelson sparsely setose O. tuberculosa, p. 195.
Pereonites 2 and 3 weakly nodulose; pereonite 7 with 1 spike row;
pleon and pleotelson densely setose ...... O. epostoa, p. 199.
Antennule with 5-7 irregular anterior spikes, elongate posterior
blade; pleon posterior margin distinctly concave, processes with-
out ventral spike ..................... O. tripartita, p. 214.
Antennule with 5 irregular anterior spikes, narrow posterior
blade; pleon posterior margin weakly concave, processes with
ventral Spike cy 6 + Â x er etr rael a kid O. kensleyi, p. 218.
Pleotelson posterior margin entire, or medial lobe indistinct,
very weak; uropod exopod with 3-4 prominent serrations.........
stade Ve bona m scar v an A O. islaya, p. 237.
Pleotelson posterior margin with distinct medial lobe; uropod
exopod with ventral nodules only .................eesso. 8
Pereonal spikes flattened; cephalon anterior margin with distinct
flange on either side of rostrum; antennule with 3 flat anterior and
| large flat posteromedial spikes ........... O. poorei, p. 225.
Pereonal spikes rounded; cephalon anterior margin with or with-
out spikes on either side of rostrum; antennule with more than 3
anterior SPIKES. 131. Ra pit pini, dare ad 9
Pleon posterior margin with paired processes; epistome with 2
flat truncate spikes . a.nn anannanunnn. O. bispinosa, p. 208.
Pleon posterior margin without processes, with or without boss;
epistome spikes rounded, distally acute ................. 10
Pleon posterior margin with boss; pleotelson posterior margin
flattened, somewhat produced; antennule without posterior
SBIKeS: tr Roc LE TN AMA oa ne ta R Il
Pleon posterior margin regular, without boss; pleotelson posterior
margin not flattened or produced; antennule with posterior
SDIKeST toe a ne poke leeds eme, le Dia 12
Cephalon anterior margin with 3 prominent spikes, median one
rostral; epistome with 3 short spikes; pereonite 1 unornamented,
pereonite 2 weakly nodulose ............ O. australis, p. 218.
Cephalon anterior margin unornamented; epistome with trans-
verse blade; pereonite | with anterior pair of submedian nodules,
pereonite 2 with 2 spike rows ......... O. matucana, p. 222.
Pleotelson medial lobe not extending to posterior margin; per-
eonites l or l and 2 with prominent median nodules; pleonal boss
Very WEAK o, eT vu nutu d ets beum Mrl oko a, 13

21.

22.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

Pleotelson medial lobe extending to posterior margin; pereonites
1 and 2 without prominent median nodules; pleonal boss very

COT CRS MM LAU a ELATI RNC 14
Rostrum with bifid spike; pereonite 1 unornamented; appendix
masculina distally blunt ............... O. aylostera, p. 193.

Rostrum without spike; pereonite 1 with prominent median
rounded spike, 2 smaller nodules on either side; appendix mas-
culina distally acute < K e K Re Ee anan ea O. rebutia, p. 191.
Cephalon rostral spike distinct and single or absent ....... 5
Cephalon with distinct bifid, or indistinct trifid rostral spike 18
Cephalon without rostral spike; cephalon anterior margin unor-

ERI AE ee wm. R LENA nni BIO: 16
Cephalon with single rostral spike; cephalon anterior margin with
SSRI KESA TCI Des aerei EE ant sarti ME tue ala a ALES. 17

Pleotelson with 6 large posterolateral tubercles on either side;
antennule article 2 with spike; appendix masculina not extending
beyond distal margin of endopod ...... O. multidens, p. 184.
Pleotelson without large posterolateral tubercles; antennule
article 2 without spike; appendix masculina extending beyond
distal margin of endopod ............ O. corypantha, p. 186.
Pleonal boss well developed, each angle with 2-3 distinct
tubercles; antennule article 2 with single spike; pleopod endopod
medial margin with distinct distomedial lobe; appendix mascu-
lina distal half abruptly narrowed ...... O. obregonia, p. 188.
Pleonal boss weak, each angle with 1 tubercle; antennule article 2
without spike; pleopod endopod medial margin straight; appen-

dix masculina not abruptly narrowed ........ O. tual, p. 188.
Rostrum with 3 indistinct small spikes; pleotelson posteriorly
somewhat flattened ................... O. copiapoa, p. 181.
Rostrum with 2 distinct spikes; pleotelson not posteriorly some-
SV hratetiattened a pu c mU Me se da 19
Anterior margin of cephalon unornamented; pleotelson without
prominent posterior tubercles .......... O. denmoza, p. 178.
Anterior margin of cephalon with spikes; pleotelson with or with-
out prominent posterior tubercles ...................... 20

Anterior margin of cephalon with 4 prominent, discrete, acute
spikes on either side of rostrum; appendix masculina apically
irregularly truncated; medial margin of pleopod 1 endopod with
ati tk dense AP SA LR o ee MS. ade vg O. frailea, p. 176.
Anterior margin of cephalon with more than 4 spikes on either
side of rostrum; appendix masculina apically rounded; medial
margin of pleopod 1 endopod PMS not stiff ............. 21
Anterior margin of cephalon with continuous row of 6-7 spikes
on either side of rostrum; appendix masculina basally wide;
pleonal boss with 2 prominent tubercles anterior to posterolateral
ASI SEE Eo Va RR EE LP qs yn O. lobivia, p. 173.
Anterior margin of cephalon with spikes most prominent lat-
erally; appendix masculina not basally wide; pleonal boss without
prominent tubercles anterior to posterolateral angles ...... 22
Anterior margin of cephalon with spikes obvious; antennule with
6-7 anterior spikes; appendix masculina apically narrowly
rounded; pleotelson with 2 prominent tubercles posterior to those
opposing pleonal spikes ................. O. bisubula, p. 160.
Anterior margin of cephalon with spikes weak; antennule with 8-
9 anterior spikes; appendix masculina apically bluntly rounded
pleotelson without prominent tubercles posterior to those oppos-
ing pleonal spikes............---s----»- O. parodia, p. 170.

159

160

Oxinasphaera bisubula sp. nov.
Figures 5-12

Material examined. Holotype. d (4.7 mm), eastern
Bass Strait, 100 km NE of North Point, Flinders Is.,
Tas., 3852.6'S, 148°25.2’E, 15 Nov 1981, 130 m, fine
sand, R. Wilson (NMV J40489).

Paratypes. NSW. 169 (3.8-5.1 mm, mean = 4.3
mm). 89 (ovig 4,3, 4.3, 4.0, 3,9, 3.8, non-ovig 4.3, 4.1,
3.5 mm), 3 mancas (2.5, 2.0, 1.8 mm), off Moona
Moona Creek, Jervis Bay, 35°03.5’S, 150°41.0’E, 19
Jun 1982, 3 m, bivalves encrusted with sponges, J.K.
Lowry (AM P41171. slide AM P44214). 30 (5.1, 3.9
mm), off Moona Moona Creek, Jervis Bay, 350°35’S,
150°41.0°E, 3 Mar 1982, 4.5 m, on ascidian, Hermania
momus, in Ecklonia, P.B. Berents (AM P44199).

Bass Strait. 118 (5.0, 4.9, 4.9 SEM dissected, 4.8
SEM, 4.7, 4.5 dissected, 4.0, 4.0, 3.8, 3.5, 3.5 mm), 209
(3.8-4.5 mm), topotypes, same data as holotype
(NMV 140490, 2d, 2e ZMUC CRU1378). 38 (4.3, 4.1,
4.0 mm). 29 (ovig 4.5 damaged, dissected, 3.8 mm), 2
mancas ( 2.5, 1.1 mm), 28 km SSW of Marlo, Vic.,
37°59.0°S, 148°27.0’E, 30 Jul 1983, 51 m, muddy sand
and fine shell, M.F. Gomon and R.S. Wilson (NMV
126292). 26 (4.7, 4.6 mm), 65 km E of Cape Rochon,
Three Hummock Is, Tas., 40°10.9’S, 145°23.0’E, 13
Nov 1981, 75 m, shelly sand, R.S. Wilson (NMV
140491). 8 (4.4 mm), 20 km NNE of North Point, Tas.,
40738.0'S, 144°20.9"E, 4 Nov 1980, 37 m, muddy shell
grit, M. Gomon and G.C.B. Poore (NMV J26282). 6d
(4.6, 4.4, 4.3, 4.3, 4.2, 4.0 mm), 100 km SSE of Cape
Liptrap, Vic., 3945.9'S, 145°33.3’E, 13 Nov 1981, 74
m, muddy fine sand, R. Wilson (NMV 126257). 28
(4.2, 4.0 mm), 36 km SSW of Stokes Point, King Is,
Tas., 22 Nov 1981, 85 m, medium sand, dredged, R.S.
Wilson (NMV 14097).

Vic. 36 (4.8, 4.4, 4.0 mm), 69 ( 4.8, 4.4, 4.0, 3.7, 3.5,
3.3 mm), 8 mancas (2.8-0.9 mm), NE shore of
Cape Wellington, Wilsons Promontory, 3993,5'S,
146°28.7’E, 9 Feb 1982, 0 m, various SCUBA samples,
G. Smith and L. Rubleman (NMV J26317). 38 (4.9,
4.9, 4.8 mm ), female? (2.0 mm), Bastion Point, Mal-
lacoota, 37°34.3’S, 149°46.2’E, 6 Apr 1989, 5 m, reef
300 m offshore, hydroids, sponges, bryozoans and red
algae, G.C.B. Poore and R.S. Wilson (NMV 126366).
4d (4.6, 4.3, 4.3, 4.0 mm), Bastion Point, Mallacoota,
37°34.3’S, 149°46.2’E, 6 Apr 1989, 5 m, reef 300 m
offshore, sponges, G.C.B. Poore and R.S.. Wilson
(NMV J26402).

Tas. 99 (4.1-3.5, mm), 4o (4.2, 4.0, 4.0, 4.0 mm),
eastern side of Waubs Bay, Bicheno, 41953'S,
147"18'E. 23 Mar 1988, 7 m, reef, sponges on vertical
rock face, SCUBA, G.C.B. Poore and H.L. LewTon
(NMV J26238).

Non-paratypic material: Qld. 26, off Moreton Bay,
27°27’S, 153"39'E, 29 Mar 1969, 76.8 m, W.F. Ponder
on HMAS Kimbla (AM P44200).

NSW. cd, Nelson Head, Port Stephens, 32°43’S,
152°09’E, 27 Oct 1980, 24 m, rubble stones in channel,
J. Hall (AM P41173). d, 59, N of Fly Point, Nelson,
32°43’S, 152°09’E, 8 Nov 1981, 20 m, orange sponge
on dead mussel, R.T. Springthorpe and D. Stracy (AM
P44211). d, 29, 6 mancas, Fly Point, Nelson Bay,

N. L. BRUCE

32*43'S, 152°09’E, 28 Oct 1980, 15 m, algae, J. Halll
(AM P41194). 20, 69, 1 manca, Nelson Head, Porti
Stephens, 32°43’S, 152910'E, 27 Oct 1980, 18 m, sand
and shell grit, J. Hall and I. Loch (AM P41178). 26, 1 lq
(3 ovig), 7 mancas, Nelson Head, Port Stephens,
32*43'S, 152°09’E, 27 Oct 1980, 18 m, tufted bryo-
zoans and hydroids, J. Hall (AM P41190). 69, 69(3
ovig), 6 mancas, inside Box Head, Broken Bay,
33'33 8, 151°21’E, 22 Nov 1982, two species of
sponges, J.K. Lowry and R.T. Springthorpe (AM
P41174). d, inside Box Head, Broken Bay, 33°33’S,
151°21’E, 21 Nov 1982, from Echinoclathria sp., J.K.
Lowry and R.T. Springthorpe (AM P44209). 5 c, E of
North Head, Port Jackson, Sydney, 33°49’S, 151°18°E,
19 Feb 1973, 19 m, host Teichonella labrinthica,
AMSBS (AM P22191-P22194). 20, E of North Head,

Port Jackson, Sydney. 33°49’S, 151°18’E, 1973, 42 m,

host Polymastea craticia, AMSBS (AM P22197). 28,

same dala as previous except 19 m, 19 Feb 1973 (AM

P22190). 3d same data as previous except 32.9 m, 23

May 1973 (AM P22198). 9d, E of North Head, Port

Jackson, Sydney, 33'49'S, 151°18"E, AMSBS (AM

P24307). d, ovig 9, Port Hacking. Sydney, 34°03.9’S,

151707.6'E, 11 Jul 1971, 18 m, rock face, P.A. Hutch-

ings (AM P41203). 20, 2 ovigo, Port Hacking, Sydney,

34°05’S, 151"10'E, 13 Aug 1981, 15 m, sponge, J.K.

Lowry and R.T. Springthorpe (AM P41167).c, 49, off
Moona Moona Creek, Jervis Bay, 35°03’S, 150°41’E,

13 May 1981, 17.7 m, from sponge in scallop beds,

P.B. Berents (AM P41 192), 5c, 2 ovig 9, off Moona

Moona Creek, Jervis Bay, 35°03.5’S, 150°41.0’E, 19

Jun 1982. 8 m, mussels, epizoic algae and sponges on

sand covered rocks, J.K. Lowry (AM P41169). 8, 9. 8

immature and mancas, off Moona Moona Creck, Jer-

vis Bay, 35°03’S, 150°41’E, 19 Jun 1982, 3 m, from

sponges, J.K. Lowry (AM P44194). d, 89, off Moona

Moona Creek, Jervis Bay, 35°03.5’S, 15094 1.0/E, 19

Jun 1982, 3 m, encrusting sponge, J.K. Lowry (AM

P41170). 2 d, ocean side of Bowen Is., Jervis Bay,

35°07’S, 15046" E, 27 Apr 1971, 36 m, sponges, ascid-

ians, bryozoans and algae from large boulders, P.A.

Hutchings and P.B. Weate (AM P41195), 59, ovig 9,

ocean side of Bowen Is., Jervis Bay, 35*07'S, 150°46’E,

29 Nov 1971, 36.5 m, marine growth on boulders, P.A.

Hutchings (AM P44205). 30+ d, o, Darling Point, Jer-

vis Bay, 35°07.6’S, 150“45.6'E, 23 Jan 1973, 18 m,

scallops and mussels on sandy bottom, P.A. Hutchings

(AM P41204). g, several o and imm, 3705'S,

150°05’E, 30 Nov 1914, 55-91 m, sand, Dr Th. Mor-

tensen's Pacific Expedition 1914-1916, Endevour

(ZMUC CRU1379).

Vic. 49, 99 , 50 nm SE of Gabo Is., 37°22.3’S,
150°02.2’E, 19 Jun 1962, 75 m, HMAS Gascoyne (AM
P41181). 9, Gabo Is., 37°34’S, 159°55’E, 19 Feb 1973.
28 m, sponge community, J.E.Watson and S.A.
Shepherd (NMV 740496). d, Gabo Is, 37°34’S,
159°55‘E, 17 Feb 1973 (NMV 126216). 29, 2 mancas,
20 miles SW of Cape Everard, 37°48’S, 149°16’E, 27
Aug 1972 (NMV J26219).

Bass Strait. d, 31 km SSW of Cape Otway, Vic.,
39"08'S, 14324'E, 8 Oct 1980, 77 m, medium sand,
G.C.B. Poore (NMV J26253). 28, 23 km E of Cape
Rochon, Three Hummock Is, Tas., 40722.2'S,
145°17.0°E, 3 Nov 1980, 40 m, mainly sand,

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 161

M. Gomon and G.C,B.Poore (NMV 126266). 3,47 km
E of Cape Rochon, Three Hummock Is.. Tas.,
40723.8'S, 145*32.0'E, 3 Nov 1980, 66 m, muddy sand.
M. Gomon and G.C.B.Poore (NMV 126289). `

Tas. d, 29, 32 km NW of Devenport, 40°56.04’S.
146/39.00'E, 4 Feb 1980, 66 m, muddy sand, M.
Gomon and G.C.B. Poore (NMV 140482). 8d, 169 and
imm, 39 km NNE of Devenport, 40°49.75’S,
146°31.33’E, 4 Feb 1980, 68 m, mud with bryozoa and
sponges, M. Gomon, G.C.B, Poore and C.C. Lu (NMV
140492). d, o, 1 km E of Bicheno, Muirs Rock,
4153.0'S, 148°19.0°E, 21 Apr 1985, 15 m, sponge and
bryozoan epifauna, airlift, R.S. Wilson (NMV
J26144). d, same data as previous, except from red
algal turf, infauna (NMV J 26145). Many, D'Entrecas-
teaux Channel, 2.5 km E of Birches Bay, 43°11.0’S,
147°16.0°E, 16 Apr 1985, 10 m, R.S. Wilson (NMV
126140).

Australian Museum, Old Collections. 99, 9 km E of
Coogee, NSW, 33°57’S, 151*21'E, 15 Mar 1898, 89 m,
fine sand, E.R. Waite on HMCS Thetis (AM G2274).
d, 2 km E of Orient Point, NSW, 34°13’S, 150°48’E 20
Mar 1898, 23 m, sand and rock, E.R. Waite on HMCS
Thetis (AM G2273). 59, Thetis stn 48, E.R. Waite (AM
3902).

Description of male. Body about 2.2 times as
long as greatest width; lateral margins subparal-
lel, maximum width at pereonites 5; dorsal sur-
faces not polished, generally finely pitted anter-
iorly, otherwise granular, with scattered setae.
Cephalon dorsal surface finely nodulose;
anterior margin with series of 5-7 discrete acute
tubercles on either side of rostrum, lateralmost
being most prominent; with prominent bifur-
cate spike on rostrum. Pereonite | without
spikes or ornamentation. Pereonites 2 and 3
each with 2 fine transverse rows of low spikes,
pereonites 4-7 each with 2 transverse rows of
prominent acute spikes, anterior row larger than
posterior row; coxae 5-7 with posterior margins
evenly rounded. Pleon posterior margin with
posterior boss, posterolateral angles of boss each
with prominent posteriorly directed spikes.
Pleotelson with 2 prominent spikes opposing
those of pleon, pair smaller tubercles set
immediately posterior to these; posterolateral
flange with 1 submarginal and 2 marginal acute
tubercles; posterior margin with two prominent
submedian indentations either side of median
lobe.

Antennule peduncle article 1 with 6 (occasion-
ally 7) anterior spikes; with 1 long proximo-
posterior spike and 1 short posterior spike; dor-
sal surface of peduncular articles 1 and 2 pro-
vided with numerous roughened setae; flagellum
with 6 articles. Antenna peduncle articles 4 and
5 with long setae; flagellum of about 7
articles.

Epistome with 2 prominent widely separated
conical spikes set on transverse ridge. Mandible

spine row with 5-6 spines; molar process
toothed along margins, surface finely ridged,
distal margin not deeply indented; palp article 2
with 7 stout biserrate setae on medial margin,
distal 2 being longest, article 3 with 8 short and 4
long stout biserrate setae on medial margin.
Maxilla lateral lobe with 9 finely pectinate sctae,
middle lobe with 6, medial lobe with 5 CP spines
and distally a further 6 weakly plumose setae.

Pereopod 1 basis about 2.5 times as long as
wide, anterior margin with 2 sensory setae;
ischium 0.7 times as long as basis, 2.2 times as
long as wide, anterior margin with 2 short proxi-
mal spines and 1 distal longer and biserrate
spine; merus about half as long as ischium, 1.2
times as long as wide, anterolateral angle with 2
gently curving pectinate spines, posterior mar-
gin with 3 spines becoming progressively longer
distally and single long simple seta; carpus short,
0.7 times as long as merus, 1.2 times as long as
wide, posterior margin with 2 biserrate spines;
propodus about equal in length to ischium,
widest proximally, about 2.8 times as long as
wide, posterior margin with distinct scale spikes
and 3 large biserrate spines, row of 3 smaller
biserrate spines on medial margin; dactylus 0.6
length of propodus, unguis about 4496 length of
entire dactylus. Pereopods 2 and 3 similar to 1,
differing principally in having the carpus more
elongate and propodus more slender. Pereopod
2 basis 3.0 times as long as wide; ischium 2.5
times as long as wide; carpus about equal in
length to merus, 1.4 times as long as wide, with
further 5 biserrate on distal margin; propodus
0.8 times as long as ischium, 2.9 times as long as
wide, margins subparallel, posterior margin with
weakly biserrate spines. Pereopods 4-7 gener-
ally subsimilar except 6 and 7 provided with
more biserrate spines on carpus than 4 and 5.
Pereopod 6 similar to 7 except basis slightly
longer, carpus distal margin with 3 large trifid
spines Pereopod 7 basis 2.5 times as long as
wide, posterior margin with prominent scale
spikes, with several sensory setae; ischium
slightly shorter than basis, 3.7 times as long as
wide; merus 0.4 times as long as ischium, 3.7
times as long as wide, posterior margin with 3
biserrate spines, anterodistal angle with 1 large
spine; carpus 1.4 times as long as merus, 2.3
times as long as wide, posterior margin with 6
biserrate spines, distal margin with 2 large trifid
spines and 6 biserrate spines; propodus about
1.2 times as long as carpus and 0.6 time as long
as ischium, about 3.5 times as long as wide, pos-
terior margin with 3 biserrate spines, anterodis-
tal angle with 2 sensory setae.

Penial processes each about 3.5 times as long
as basal width, tapering slightly to an oblique

162 N. L. BRUCE

a m
«T coo 90900002557

a LE, VW I VIV m A

Figure 5. Oxinasphaera bisubula sp. nov. A-E, holotype, remainder paratype d 4.5 mm (NMV J40490). A, dorsal
view; B, lateral view; C, frons; D, cephalon and antennules, anterior view; E, pleon and pleotelson, ventral view;

F, antennule, f, seta from peduncular article 1; G, antenna; H, left mandible; I, left mandible, distal detail; J, right
mandible, distal detail. Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 163

Figure 6. Oxinasphaera bisubula sp. nov. All figs paratype d 4.5 mm (NMV J40490). A, mandible palp; B,
maxilla; C, maxillule; D, maxillule exopod apex; E, maxillule endopod apex; F, maxilliped, f, seta from palp
article 5; G, maxilliped endite distal margin, g, cuticular setule from endite.

164 N. L. BRUCE

Figure 7. Oxinasphaera bisubula sp. nov. All figs paratype 9 4.5 mm (NMV J40490). A, pereopod 1; B, pereopod
l, dactylus; C, pereopod 2; D, pereopod 3; E, pereopod 6, distal articles; F, pereopod 7, f, spine from anterodistal
angle merus, ff, spines from distal margin carpus, fff, spine from posterior margin of merus; G, pereopod 7,
dactylus medial view (a, anterior margin; b, posterior margin).

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 165

TED pu
UMEN t

h aue

DES M^; Mf,

CN ty l
Na Zi,
ps 7
y ld,

DU) “Uu

NN

Ne

Figure 8. Oxinasphaera bisubula sp. nov. All figs paratype ¢ 4.5 mm (NMV J40490). A-E, pleopods 1-5
respectively, a, scales from endopod, aa, scales from exopod; F, uropod; G, penes, g, scales from base of penes; H,
coupling hooks, pleopod 1.

166 N. L. BRUCE

Figure 9. Oxinasphaera bisubula sp. nov. Ovigerous 9 4.5 mm (NMV J26292). A, lateral view; B, dorsal view of
pleon and pleotelson; C, frons; D, mandible; E, maxillule; F, maxilla; G, maxilliped; H, pereopod 7; I, oostegite 1;
J, oostegite 4; K, uropod. Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 167

y i» vi
M P,

i S
UA

Figure 10. Oxinasphaera bisubula sp. nov. Non-ovigerous 9 4.3 mm (NMV J40490). A, dorsal view; B, lateral
view: C, frons; D, pleon and pleotelson, ventral view; E, antennule; F, uropod. Scale 1.0 mm.

168 N. L. BRUCE

Figure 11. Oxinasphaera bisubula sp. nov., SEMs. A-F d 4.8 mm, G, H d 4.5 mm (NMV 740490). A, cephalon,
lateral view (X 100); B, cephalon anterior margin and frons (X 110); C, cephalon, anterior margin (X110); D,
pleon and pleotelson ( X 40); E, pleotelson tubercles (X 270); F, spike rows, pereonite 5 (X 370); G, left mandible,
incisor, lacinia mobilis and spine row (X 750); H, molar process (X 700).

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

Figure 12. Oxinasphaera bisubula sp. nov., SEMs. All figs 9 4.5 mm (NMV J26267). A, maxillule, lateral lobe
(X950); B, seta, maxilliped palp article 4 (X 4000); C, pereopod 1, dactylus (7 450); D, pereopod 2, propodus
(X450); E, serrate spine, pereopod 1 propodus (X 1200); F, seta from uropod (X 2700); G, uropod rami, apices

(X 150); H, detail, pleotelson cuticle (X 2000).

170

subtruncate apex; proximolateral margin with
scale spikes.

Pleopod 1 exopod with c. 28 PMS, endopod
with c. 19 PMS. Pleopod 2 exopod and endopod
with c. 16 and 28 PMS respectively; appendix
masculina straight, 6.0 times as long as maxi-
mum width, narrowing shortly before apex to
sub acute point, extending beyond endopod
slightly (by about 0.15 of its length). Pleopod 3
exopod and endopod with c. 14 and 26 PMS
respectively. Pleopod 4 exopod with single seta
at distomedial angle, endopod with 6 thickened
fleshy ridges. Pleopod 5 exopod with 2 apical
lateral scale lobes 1 medial apical scale lobe and
one lateral proximal scale lobe, endopod with 7
thickened fleshy ridges. Uropod dorsally nodu-
lar, covered with roughened setae; exopod about
4.7 times as long as proximal width, about half as
long as endopod, apex deeply bifid with lateral
process prominent; endopod about 3.4 times as
long as wide, apex with prominent terminal and
3 downwardly directed prominent spikes.

Ovigerous Female. Body dorsally without
tubercles, with sparse setae; posterior margin of
pleotelson with thickened rim, with shallow
median indentation. Uropod rami thickened,
covered with fine setae and longer thickened
setae.

Non-ovigerous Female. Body dorsal surfaces
with weak transverse tubercles on pereonites 2-
7, pleon and pleotelson; pleon with 2 obvious
low submedian mounds; pleotelson with 4 indis-
tinct ridges, variable in development between
individuals, never prominent.

Colour. Pale yellow ground colour, often with
chromatophores which are more evident in
females. Cuticle of preserved specimens may be
clear or opaque.

Size. Males 3.3-5.1 mm, females 3.5-4.5 mm,
ovigerous females 3.8-4.5 mm, mancas 0.9-2.8
mm.

Variation. In some localities, notably southern
Tasmania, the males have the posterior pair of
pleotelsonic tubercles less well developed than
those of the type series.

Remarks. This species, restricted in its distri-
bution to south-eastern Australia, is the most
frequently collected member of the genus. Males
are readily recognized by the prominent pleonal
spikes, the presence of a second pair of pleotel-
sonic tubercles, a bifurcate spike on the rostrum,
and the antennule peduncle article 1 with 6 or 7
spikes. Females of Oxinasphaera bisubula can be

N. L. BRUCE

recognized by the submedian pair of domed
mounds on the pleon, a feature shared with few
other species either in the group or in the
genus.

There are several other species similar to Oxi-
nasphaera bisubula, and these are best separated
using the key provided.

Distribution. Here recorded from off Moreton
Bay, south-eastern Queensland (27778 ), along
the New South Wales coast to the Bass Strait and
Tasmania (43°11’S ) and westwards to Victoria
(143°24’E), at depths from the intertidal to 130
m, with only one record at a depth greater than
77 m; of the samples where depth was recorded,
about 71% are from less than 50 m. Twelve
samples are recorded from sponges or have
sponges mentioned in the habitat data.

Hosts. Sponges, including some identified as
Echinoclathria sp., Teichonella labrinthica and
Polymastea craticia.

Etymology. The epithet is derived from the
Latin subula (= awl-shaped or pointed), and
alludes to the prominent spikes on the epis-
tome.

Oxinasphaera parodia sp. nov.
Figures 13, 14

Material examined. Holotype. & (5.0 mm), S of point
Hicks, Vic, 38°17.7’S, 149*1 1.3'E, 24 Jul 1986, 400 m,
coarse sand, gravel, mud, many sponges, M.F. Gomon
(NMV J40486).

Paratypes. 124 (3.5-4.8 mm, mean = 4.1 mm), 139
(ovig 4.0, 4.5 mm; 11 non-ovig 2.8-4.0 mm, mean =
3.5 mm), 31 mancas (1.6-2.3 mm), same data as holo-
type (NMV 719159 [9], 119160 [9 and mancas]).

Description of male. Body about twice as long as
greatest width; dorsal surfaces not polished, gen-
erally granular, with scattered setae. Cephalon
dorsal surface laterally finely nodulose; anterior
margin with 3-4 small lateral tubercles on either
side of rostrum; with prominent bifurcate spike
on rostrum. Pereonite | without ornamentation,
Pereonite 2 with 2 fine transverse rows of low
spikes, pereonites 3-7 each with 2 transverse
rows of prominent acute spikes, anterior row
larger than posterior row; coxae 5 and 7 with
posterior margins evenly rounded, coxa 6 pos-
teriorly subtruncate. Pleon with posterior boss,
with posterolateral angles of boss each with
prominent posteriorly directed spikes. Pleotel-
son with 2 prominent spikes opposing those of
pleon, dorsal surface heavily granular, without
distinct tubercles; posterolateral flange with 1
submarginal and 2 marginal tubercles; posterior

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 171

COUVY v 09

K vi

U Yves

Figure 13. Oxinasphaera parodia sp. nov. A-E holotype, F-I paratype d 4.8 mm (NMV JJ19159). A, lateral view;
B, cephalon, dorsal view; C, pleon and pleotelson, dorsal view; D, frons; E, antennules, anterior view; F,
antennule; G, pereopod 1 propodus; H, pleopod 2; I, penes; J, non-ovigerous 9 3.5 mm, pleon and pleotelson.
Scale 1.0 mm.

Figure 14. Oxinasphaera parodia sp. nov. SEMs. d 4.2 mm (NMV 719159). A, cephalon, anterior view (X95); B.
frons (X90); C, pleon and pleotelson (X 77); D, pleotelson apex (X 120); E, detail, pleotelson cuticle (X 160); F,
pereopod | dactylus, medial aspect (X400); G, uropod (X110).

| margin; proximolateral

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

margin with 2 submedian indentations on either
side of median lobe.

Antennule peduncle article 1 with 8 anterior
spikes; with 1 long proximo-posterior spike and
l short posterior spike; article 2 with small spike
at anterodistal margin; dorsal surface of pedun-
cular articles 1 and 2 provided with numerous
roughened setae; flagellum with 6 articles.

Epistome with 2 prominent widely separated
conical spikes set on transverse ridge.

Pereopod l propodus about equal in length to
ischium, widest proximally, about 2.7 times as
longas wide, posterior margin with distinct scale
Spikes and 4 large biserrate spines, row of 4
smaller biserrate spines on medial margin; dac-
tylus 0.5 length of propodus, unguis about 8096
length of entire dactylus.

Penial processes each process about 3.0 times
as long as basal width, tapering slightly, distola-
teral margin curving smoothly to meet medial
margin with scale
spikes.

Pleopod 2 appendix masculina straight, about
as long as endopod, 5.2 times as long as maxi-
mum width, extending slightly beyond endopod
(by about 0.18 of its length). Uropod exopod
about 4.0 times as long as proximal width, about
0.4 times as long as endopod, apex deeply bifid
with lateral process prominent; endopod about
4.0 times as long as wide, apex 3 prominent ven-
trally directed spikes.

Female. Ovigerous and non-ovigerous females
could not reliably be distinguished from females
of O. bisubula other than by lack of chromato-
phores, and lack of pleonal mounds.

Colour. White, chromatophores not apparent.

Size. Males 3.5-5.0 mm, females 2.8-4.4 mm,
ovigerous females 3.8-4.5 mm, mancas 1.6-2.3
mm.

Remarks. This species is very similar to Oxinas-
phaera bisubula, but differs consistently in sev-
eral male characters: antennule with 8-9
anterior spikes (versus 6-7); anterior cephalic
margin only laterally nodulose, weakly so (ver-
sus entire anterior margin); pleotelson without
posterior pair of prominent tubercles (versus
with) pleotelson with prominent rough
tubercles (versus tubercles not as prominent);

| appendix masculina blunt (versus with apically

narrowed and obliquely truncate).

Females are virtually indistinguishable from
O. bisubula, and without males probably cannot
be identified with certainty. Both males and

la

females are, in contrast to O. bisubula, without
chromatophores.

Distribution. Known only from the type locality,
off the eastern Victoria coast at a depth of 400
metres,

Hosts. Host identity not known.

Oxinasphaera lobivia sp. nov.
Figures 15, 16

Material examined. Holotype. ¢ (3.7 mm), S of Saint
Helena Is., Moreton Bay, south-eastern Qld, 2 Sep
1979, 6 m, from trawled sponge, N.L. Bruce (QM
W20034).

Paratypes. 129 (2.6-3.4 mm, 3.3 [dissected], mean =
3.1 mm), 139 (4 ovig 2.9-3.8 mm, mean = 3.3 mm; 7
non-ovig 2.1-2,9 mm, mean = 2.5 mm), same data as
holotype (QM W20035, W20036, W20037, W20045,
28, 20 ZMUC CRU1387). 45 (2.8, 2.8, 2.9, 2.9, mm),
179 (4 ovig 3.0-3.3 mm, mean = 3.1 mm; 13 non-ovig
2.1-3.1 mm, mean - 2.6 mm), manca (1.8 mm), Shag
Rock, Stradbroke Is., south-eastern Qld, 22 Aug 1979,
in sponge, M. Ready and Niels Svennivig (QM
W20041).

Non-paratypic material. NSW. 49 (3 damaged or
poor), 159, W of Spit Bridge, Middle Harbour Port
Jackson, 33°8.2’S, 151°14.6’E, 19 Jun 1981, 8 m, tele-
stacean bed with mussels, J.K. Lowry (AM P41177);
26, non-ovig 9, 2 mancas, Jibbon Point, Port Hacking,
34°05’S, 151°10’E, 13 Aug 1981, 15 m, sponge, J.K.
Lowry and R.T. Springthorpe (AM P44208).

Description of male. Body about 2.2 times as
long as greatest width; dorsal surfaces polished,
finely pitted on cephalon and pereonite 1, other-
wise generally granular, with scattered setae.
Cephalon dorsal surface not nodulose; anterior
margin with 7 small close set nodules on either
side of rostrum; with prominent broad based
bifurcate spike on rostrum. Pereonite | without
tubercles or ornamentation. Pereonite 2 with 2
fine transverse rows of low spikes, pereonites 3-
Teach with 2 transverse rows of prominent acute
spikes, anterior row larger than posterior row;
coxae 5 and 7 with posterior margins evenly
rounded, coxa 6 ventro-posteriorly subtruncate.
Pleon with posterior boss, with posterolateral
angles of boss each with prominent posteriorly
directed spikes, each with 2 tubercles anteriorly.
Pleotelson with 2 prominent spikes opposing
those of pleon, dorsal surface granular, with 2
tubercles set posterolaterally to each primary
spike; posterolateral flange without prominent
spikes; posterior margin with 2 submedian
indentations on either side of median lobe.
Antennule peduncle article 1 with 7 anterior
spikes, distalmost spike being distinctly smaller
than remainder; with 1 long proximo-posterior

174 N. L. BRUCE

ccoo 9
co 0000

Figure 15. Oxinasphaera lobivia sp. nov. A-E holotype, F- I paratype 3 3.3 mm (QM W20045). A, dorsal view; B,
lateral view; C, cephalon; D, frons; E, antennules, anterior view; F, antennule; G, pleopod 2: H, penes; I, uropod.
Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 175

Figure 16. Oxinasphaera lobivia sp. nov. SEMs. ¢ 3.5 mm (QM W20037). A, cephalon, anterior view (X 100); B,
cephalon, anterior margin (X 110); C, pleon and pleotelson (X45); D, pleotelson apex (X270).

spike and 1 short posterior spike; dorsal surface
of peduncular articles 1 and 2 provided with few
roughened setae; flagellum with 6 articles.

Epistome with 2 prominent widely separated
conical spikes.

Pereopods essentially the same as O. bisu-
bula.

Penial processes each process about 3.2 times
as long as basal width, tapering slightly, distola-
teral margin curving smoothly to meet medial
margin; proximolateral margin with scale
spikes.

Pleopod 2 appendix masculina straight, 4.8
times as long as maximum width, about as long
(0.94) as endopod, extending slightly beyond
endopod (by about 0.12 its length), apex nar-
rowly rounded. Uropod exopod about 2.8 times
as long as proximal width, about 0.4 times as
long as endopod, apex deeply bifid with lateral
process prominent; endopod about 3.5 times as
long as wide, apex with 3 prominent ventrally
directed spikes.

Female. Females are not distinguishable from
those of O. bisubula.

Colour. Abundant dark brown and black chro-
matophores giving an overall dark brown
appearance.

Size. Males 2.6-3.7 mm, females 2.1-3.1 mm,
ovigerous females 2.9-3.8 mm, manca 1.8
mm.

Remarks. Oxinasphaera lobivia is distinguished
from others of the O. bisubula group of species
by the cephalic ornamentation, additional pair
of pleonal tubercles and the two pairs of second-
ary pleotelsonic tubercles. The characters which
distinguish this species are: cephalon anterior
margin with 7+7 low spikes; antennule peduncle
article 1 with 7 anterior spikes; pleon with 2
prominent nodules anterior to the posterior
spikes; pleotelson with 2 pairs of tubercles set
posterolaterally to the pleotelson spikes; ros-
trum with short bifurcate spike, basally wide;
uropod wide distally with 3 prominent terminal
spikes.

The presence of the posterior pairs of
tubercles is not always easy to observe, but this
and the additional pleonal spikes reliably separ-
ates this species from the type species.

176 N. L. BRUCE

Distribution. South-eastern Queensland (More-
ton Bay and Stradbroke Is.), Sydney and Port
Hacking, central NSW, all locations shallow
inshore habitats.

Hosts. Collected from unidentified sponges.
Oxinasphaera frailea sp. nov
Figures 17-19

Material examined. Holotype. ¢ (3.9 mm), Table
Head, Port Essington, Cobourg Peninsula, NT,
11°14.4’S, 132°10.8’E, 13 May 1983, 3 m, rock wash-
ings, N.L. Bruce (NTM Cr0011333).

Paratypes. NT. l (3.8 mm), 29 (non-ovig 3.3, 3.5
mm), 3 mancas (2.7, 2.8, 3.0 mm), same data as holo-
type (NTM Cr0011334). d (3.1 mm), same data as
holotype but 14 May 1983, 3 m, algal covered rocks,
N.L. Bruce and D. Staples (ZMUC CRUI384). 39
(3.2, 3.3 broken, 3.4 mm), 9 (non-ovig 3.3 mm), Wal-
ford Point, Coral Bay, Port Essington, Cobourg Pen-

c6 009754 SS
ADF

VU veg
= So

00006

3355

insula, 11°12.0’S, 132°03.0’E, 18 May 1983, 3-4 m,
coral reef, N.L. Bruce (NTM Cr01335). € (2.9 mm),
Table Head, Port Essington, Cobourg Peninsula,
11°14.4’S, 132°10.8’E, 13 May 1983, 4 m, Jaspis sp.,
AJB stn CP/51/8-10, N.L. Bruce (QM W20038).

Non-paratypic material. NT. 3d (3.2, 3.2 broken, 3.4
mm), 9 (non/ovig 2.9 mm), Fannie Bay, Darwin,
11°24.0’S, 130*48.0'E, 26 Oct 1982, 8 m, hydroids,
J.K. Lowry (NMV J42643). d (3.0 mm), Arafura Sea,
NE of Cobourg Peninsula, NT, 11°09’S, 134°27’E, 21
Oct 1989, 30.2 m, BRR (QM W20043).

WA. d (3.9 mm), North West Shelf, 19°29.9’S,
118°52.0’E, 24 Oct 1983, 37 m, bottom 85% sand, T.
Ward (ZMUC CRU1385). l (3.6 mm), near Point
Murat, Bundegi Reef, Exmouth Gulf, 21*49'S,
113°] 1/E, 4 Jan 1984, 9 m, pink sponge on dead coral,
J.K. Lowry (AM P44198).

Description of male. Body about 2.1 times as
long as greatest width; dorsal surfaces not pol-
ished, generally granular, with scattered setae.

Figure 17. Oxinasphaera frailea sp. nov. F paratyped 3.8 mm (NTM Cr001 1334), remainder holotype. A, dorsal
view; B, lateral view; C, cephalon; D, frons; E, antennules, anterior view; F, antennule. Scale 1.0 mm.

Figure 18. Oxinasphaera frailea sp. nov. F paratype & 3.8 mm (NTM Cr001 1334), remainder holotype. A,
pleopod 1; B, pleopod 2: C, setae from distomedial margin of pleopod 1 endopod; D, apex, appendix masculina:
E, penes; F, uropod.

Cephalon dorsal surface not nodulose; anterior
margin with 4 prominent widely separated nod-
ules on either side of rostrum; with prominent
broad based bifurcate spike on rostrum. Pereon-
ite 1 without tubercles or ornamentation. Per-
eonites 2 and 3 with 2 rows of distinct low
rounded spikes, pereonites 4-7 each with 2
transverse rows of prominent acute spikes,
anterior row larger than posterior row; coxae 5
and 7 with posterior margins evenly rounded,
coxa 6 ventro-posteriorly weakly concave. Pleon
with posterior boss, with posterolateral angles of
boss each with prominent posteriorly directed
spikes. Pleotelson with 2 prominent spikes
opposing those of pleon, dorsal surface granular,
with 1 tubercle set posterolaterally to each pri-
mary spike; posterolateral flange with 1 promi-
nent tubercle; posterior margin with 2 sub-
median indentations on either side of median
lobe.

Antennule peduncle article 1 with 7 anterior
spikes; with 1 long proximo-posterior spike and
I short posterior spike; dorsal surface of pedun-
cular articles 1 and 2 provided with few
roughened setae.

Epistome with 2 prominent narrowly separ-
ated conical spikes, basally somewhat flat-
tened.

Pereopods essentially the same as O. bisu-
bula.

Penial processes each process about 2.7 times
as long as basal width, tapering slightly, distola-
teral margin curving smoothly to meet medial
margin, mediodistally margin weakly oblique;
proximolateral margin with scale spikes.

Pleopod 1 endopod distally acute, distal half
of lateral margin with conspicuously stout cir-
cumplumose setae, subapically without MS,
with setulose patch. Pleopod 2 appendix mascu-
lina straight, 6.5 times as long as maximum

178

N. L. BRUCE

width, about as long as endopod, extending
slightly beyond endopod (by about 0.13 of its
length); apex subtruncate, distally irregular;
endopod margins subparallel, distal margin
broadly rounded. Uropod exopod about 3.1
times as long as proximal width, about 0.4 times
as long as endopod, apex deeply bifid with lat-
eral process prominent and additional lateral
spike; endopod about 2.9 times as long as wide,
apex 3 prominent spikes.

Female. Females are not distinguishable from
others of this group.

Colour. Abundant dark brown and black chro-
matophores giving an overall dark brown
appearance.

Size. Males 3.0-3.9 mm, females 2.9-3.5 mm,
ovigerous females not observed, manca 2.7-3.0
mm.

Remarks. While the overall appearance of Oxi-
nasphaera frailea is similar to that of others of
the O. bisubula group, particularly the details of
the pleon and pleotelson, there are several
characters by which the species can be immedi-
ately recognized. The most obvious of these
being the prominent acute tubercles on the

anterior margin of the cephalon. The setation of
pleopod 1, shape of the endopods of pleopods 1
and 2, and shape of the appendix masculina are
unique within the genus.

The characters by which the species can be
separated from others of the group are: cephalon
anterior margin with 4-4 prominent widely
spaced acute tubercles; antennule peduncle
article | with 7 anterior spikes; pleon without
tubercles anterior setae; pleopod 2 appendix
masculina apically truncate, endopod broadly
rounded.

Distribution. Tropical western and northern
Australia from Exmouth Gulf, WA to Darwin
and Cobourg Peninsula, NT, at depths between
3 and 37 m. All records are from inshore waters
except that from the North West Shelf; appar-
ently absent from coral reefs.

Hosts. Jaspis sp., one other record directly from
an unidentified sponge.

Oxinasphaera denmoza sp. nov.
Figures 20, 21

Material examined. Holotype. d (4.2 mm), Blue
Lagoon, Lizard Is., Great Barrier Reef, Qld, 7 Jun
1987, c. 10 m, in red sponge, isolated patch reef near

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 179

500
ere CC O VO 5
È conc A - 7 "a :
= ki. € 2 vwa
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Figure 20. Oxinasphaera denmoza sp. nov. A-E holotype, F-H paratype d 3.8 mm (QM W14949). A, dorsal view;
B, lateral view; C, antennules, anterior view; D, cephalon; E, frons; F, antennule; G, pleopod 2; H, penes; I,
uropod. Scale 1.0 mm.

180 N. L. BRUCE

Figure 21. Oxinasphaera denmoza sp. nov. SEMs. à 3.8 mm (QM W14949). A, cephalon, frontal view (X 110); B,
pleon and pleotelson (7X 50); C, pleotelson apex (7X 300); D, pereopod dactylus unguis, showing single distal seta

(X 16C9); E, uropod. lateral view (X 180).

lagoon entrance, P.J.F. Davie and J. Short (QM
W 20039).

Paratypes. Qld. 4¢ (3.7, 3.8, 3.8, 4.0 mm), same data
as holotype (OM W14949), d (3.3 mm), 9 (ovig 3.2
mm), Watsons Bay, Lizards Is., 5 Jun 1987, 10-13 m,
in sponge on sand/mud, P.J.F. Davie and J. Short (QM
W 14953). 68 (3.2, 3.3, .34, 3.5, 3.6, 3.6 mm), Watsons
Bay, Lizard Is., 14°40’S, 145°28’E, 5 Dec 1975, 12 m,
sandy bottom, W. Ponder, P. Colman and I. Loch (AM
P41134).

Non-paratypic material. Qld. 49, 29, 70-80 imma-
ture, in poor condition, Mrs Watsons Beach, Lizard
Is., 14*40'S, 145°28’E, 24 Nov 1978, 18.3 m, in sponge
on sand, D.F. Hoese and H. Larson (AM P28849). 66,
Wistari Reef, Capricorn Group, southern Great

Barrier Reef, 16 Dec 1978, in sponge, 24 m, D. Fisk
(OM W8060, 1 d ZMUC CRU1381).

Description of male. Body about 2.3 times as
long as greatest width; dorsal surfaces not pol-
ished, generally granular, with scattered setae.
Cephalon dorsal surface not nodulose; anterior
margin without nodules; with prominent broad
based short bifurcate spike on rostrum. Pereon-
ite 1 without tubercles or ornamentation. Per-
eonite 2 with 2 fine transverse rows of low
spikes, pereonites 3-6 each with 2 transverse
rows of prominent acute spikes, anterior row
larger than posterior row; pereonite 7 without
posterior row of spikes, covered with small pos-

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 181

teriorly directed acute tubercles; coxae 5 and 7
with posterior margins evenly rounded, coxa 6
ventro-posteriorly concave. Pleon with pos-
terior boss, with posterolateral angles of boss
each with prominent posteriorly directed spikes.
Pleotelson with 2 prominent spikes opposing
those of pleon, dorsal surface granular, without
additional; posterolateral flange without | acute
tubercle; posterior margin with 2 submedian
indentations on either side of median lobe.

Antennule peduncle article 1 with 9 anterior
spikes, distalmost spike being distinctly smaller
than remainder; with 1 long proximo-posterior
spike and 1 very short posterior spike; dorsal
surface of peduncular articles 1 and 2 provided
with few roughened setae; flagellum with 6
articles.

Epistome with 2 prominent conical spikes,
basally somewhat flattened, on basal transverse
ridge.

Pereopods essentially the same as O. bisu-
bula.

Penial processes each process about 3.5 times
as long as basal width, tapering slightly, distola-
teral margin curving smoothly apex, medial
margin straight; proximolateral margin with
scale spikes.

Pleopod 2 appendix masculina straight, 6.2
times as long as maximum width, about as long
(0.92) as endopod, extending slightly beyond
endopod (by about 0.14 of its length), apex nar-
rowly rounded. Uropod exopod about 3.4 times
as long as proximal width, about 0.4 times as
long as endopod, apex deeply bifid with lateral
process prominent; endopod about 3.0 times as
long as wide, apex with 3 prominent ventrally
directed spikes.

Female. Females are not distinguishable from
those of O. lobivia or O. frailea .

Colour. Dark brown and black chromatophores
giving an overall brown appearance.

Size. Males 3.2-4.2 mm, ovigerous female 3.2
mm.

Remarks. The lack of any tubercles on the
anterior margin of the cephalon and the number
of antennular spikes (9) distinguishes O. den-
moza from others of the O. bisubula group.

The characters by which this species can be
identified are: cephalon anterior margin without
tubercles; antennule peduncle article 1 with 9
anterior spikes; pleon without anterior
tubercles; pleotelson without tubercles; rostrum
with bifurcate basally flattened spike; uropod
exopod distally narrow.

Distribution. Great Barrier Reef, Queensland,
Wistari Reef, Capricorn Group, in the south and
Lizard Island in the north; from the coral reef
itself, at depths from 10 to 24 m.

Hosts. From unidentified sponges; only one
sample was not recorded as having been col-
lected directly from a sponge.

Oxinasphaera copiapoa sp. nov.
Figures 22, 23

Material examined. Holotype. ¢ (4.4 mm), off Moona
Moona Creek, Jervis Bay, NSW, 35°03’S, 150°41’E, 19
Jun 1982, 3 m, from sponges, J.K. Lowry (AM
P44207).

Paratypes. NSW. 11g (3.2-4.5 [dissected] mm,
mean = 3.6 mm), 29 (non-ovig 4.1, 5.2 mm), 2 imma-
ture (3.2, 3.5 mm), same data as holotype (AM
P41165, slide P44213; 29 and e ZMUC CRUI1380). 8
(3.4, squashed 4.6 mm), 9 (non-ovig 6.2 mm), off
Moona Moona Creek, Jervis Bay, 35°03’S, 150741" E,
15 Aug 1981, 5 m, from kelp holdfasts, P.B. Berents
(AM P41182).

Non-paratypic material. NSW. d (5.2 mm), 9 (non-
ovig 4.2 mm), inside Box Head, Broken Bay, 337338,
151°21°E, 22 Nov 1982, sponge Echinoclathria, J.K.
Lowry and R. T. Springthorpe (AM P44195). d (3.5,
squashed), 2 imm/o (1.9, 2.0 mm), off Moona Moona
Creek, Jervis Bay, 35°03.5’S, 150°41.0’E, 15 Aug 1981,
4.5 m, on test of solitary ascidian Herdmania momus,
P.B. Berents (AM P41200).

Description of male. Body about 2.4 times as
long as greatest width; lateral margins subparal-
lel. Cephalon anterior margin without tubercles;
rostrum with weak trifurcate spike. Pereonite 1
unornamented. Pereonites 2-7 each with 2
transverse rows of spikes, anterior row distinctly
larger than posterior row; coxae 5 posteriorly
subtruncate, coxae 6 indented, coxae 7 rounded.
Pleon with weak posterior boss, with 2 rounded
tubercles at lateral angles of boss with 2 promi-
nent tubercles set anterior to these, posterola-
teral margin of pleon with 3-4 with tubercles.
Pleotelson granular, with spikes opposing
pleonal spikes, 2 prominent tubercles set lat-
erally and posteriorly on each side; posterola-
teral flange with 3 marginal rounded tubercles;
posterior margin flattened, somewhat produced,
with two submedian triangular excisions.

Antennule peduncle article 1 with 7 anterior
spikes; with 1 long posteroproximal and 1 pos-
teromedial spike; dorsal surface of peduncular
articles 1 and 2 with few setae; flagellum with 5
articles.

Epistome with 2 basally separated distinct
spikes; posterolateral lobes each with 2 distinct
spikes.

182 N. L. BRUCE

“repos, vi APOD IS This

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Figure 22. Oxinasphaera copiapoa sp. nov. A-F holotype, F-I paratype 4.5 mm (AM P41165). A, dorsal view;
B, lateral view; C, cephalon, anterior margin, ; D, antennules, anterior view and frons; E, pleotelson posterior
margin; F, pleotelson posterior margin, ventral view; G, antennule; H, pereopod 1; I, pereopod 2; J, pereopod 7;
K, penes. Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 183

by

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sna o: e “evo ki
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Figure 23. Oxinasphaera copiapoa sp. nov. A, B paratype 3 4.5 mm, C-E non-ovigerous 9 5.2 mm (AM P41165).
A, pleopod 2; B, uropod; C, female, dorsal view; D, female, pleon and pleotelson, dorsal view; E, female, pleon

and pleotelson, lateral view. Scale 1.0 mm.

Pereopod 1 basis about 2.4 times as long as
wide, anterior margin with 2 sensory setae, dis-
tally scaled; ischium 0.7 times as long as basis,
twice times as long as wide, anterior margin with
1 proximal spine and 1 distal longer feebly biser-
rate spine; merus 0.4 as long as ischium, 0.8
times as long as wide, anterolateral angle with 1
long and 1 short gently curving weakly pectinate
spines, posterior margin with 1 biserrate spine
and single long simple seta; carpus 1.2 times as
long as long as merus, 1.3 times as long as wide,
posterior margin with 2 biserrate spines third
submarginal small biserrate spine and 2 simple
setae; propodus slightly shorter (0.89) than
ischium, widest proximally, about twice as long
as wide, posterior margin with distinct scale
spikes and 3 large biserrate spines, row of 4
smaller biserrate spines on medial margin; dac-
tylus 0.6 length of propodus, unguis about 5596
length of entire dactylus. Pereopod 2 similar
others of genus. Pereopod 7 basis 2.7 times as

long as wide, anterior margin with 7 sensory
setae, and elongate scale setules; ischium slightly
shorter (0.88) than basis, 3.0 times as long as
wide; merus 0.4 times as long as ischium, 1.4
times as long as wide, posterior margin with 3
biserrate spines, anterodistal angle with 1 large
and 1 small spine; carpus about as long as merus,
1.8 times as long as wide, posterior margin with
3 biserrate, 2 simple and 1 trifid spines, distal
margin with 2 large trifid spines and 5 biserrate
spines; anterodistal angle with 2 slender weakly
biserrate spines; propodus 1.2 times as long as
carpus and 0.6 time as long as ischium, about
three times as long as wide, posterior margin
with 3 biserrate spines, anterodistal angle with 2
sensory setae.

Penial processes about 3 times as long as basal
width, apex narrowly rounded; proximolateral
margin with scale spikes.

Pleopod 2 appendix masculina slightly shorter
(0.96) than endopod, 6.9 times as long as

184

maximum width, extending beyond endopod by
0.14 of its length, very slightly curved laterally,
apex bluntly rounded. Uropod dorsally nodular,
covered with roughened setae; exopod about 4
times as long as proximal width, about half as
long as endopod, apex deeply bifid with pro-
cesses about equally prominent, ventral margin
with 2 prominent acute serrations; endopod
about 4 times as long as wide, apex with 3 promi-
nent bluntly rounded spikes, ventral margin
with 4-5 prominent bluntly rounded spikes.

Female. Pereonites 2-7 granular, with small
tubercles. Rostrum trifid. Pleotelson with 2
indistinct low submedian bumps anteriorly; pos-
terior margin with weak median indentation.

Colour. Pale tan in alcohol; chromatophores not
observed.

Size. Males 3.2-4.5 mm, females 4.1-6.2 mm.

Remarks. Oxinasphaera vopiapoa can be recog-
nized by the posterior margin of the pleotelson
being somewhat flattened and produced, the
pleon having a weakly produced medial portion,
the epistome with 2 discrete spikes, prominently
bifid uropod apices and the anterior margin of
the cephalon without submedian tubercles. O.
australis can be separated by the three promi-
nent tubercles on the anterior margin of the
cephalon and the acute apex to the appendix
masculina. O. matucana has two prominent
tubercles on the anterior margin of pereonite l,
and the posterior antennule spikes are set wide
apart with one at the distal extremity of ped-
uncle article 1.

The male specimen from Broken Bay, poss-
ibly senescent, differs from the type material in
lacking distinct tubercles on the median pleonal
process. The specimens, other than being larger,
agrees well with the type material.

Distribution. Sydney region to Jervis Bay, New
South Wales, depths between 3 and 5 m.

Hosts. Ecinoclathria sp., and the ascidian Herd-
mania momus, possibly an accidental associ-
ation

Oxinasphaera multidens
(Richardson, 1910) comb. nov.

Figure 24
Cymodoce multidens Richardson, 1910: 27, fig.
26.
Non Cymodoce multidens, — Kensley, 1984: 216 (=
O. kensleyi sp. nov.)

N. L. BRUCE

Material examined. Lectotype. d' (5.6 mm), off Jolo
Light, Philippine Islands, 15 Feb 1908, c. 53 m, on
surface of brown sponge, Albatross stn. 5141 (USNM
273516).

Paralectotypes. d (5.2 mm), 359 and immature
specimens, same data as holotype (USNM 40918).

Description of male. Body about 2.1 times as
long as greatest width; dorsal surfaces not pol-
ished, generally granular, with scattered setae.
Cephalon dorsal surface not nodulose; anterior
margin without nodules; without spike on ros-
trum. Pereonite | without tubercles or ornamen-
tation. Pereonites 2-3 with 2 transverse rows of
low spikes, pereonites 4-6 each with 2 trans-
verse rows of prominent acute spikes, anterior
and posterior rows about subequal in size; per-
eonite 7 posterior margin weakly produced, with
weak median indentation, without posterior row
of spikes, covered with small posteriorly
directed acute tubercies; coxae 5 and 7 with pos-
terior margins evenly rounded, coxa 6 ventro-
posteriorly subtruncate. Pleon with posterior
boss, with posterolateral angles of boss each with
prominent posteriorly directed spikes and 2
additional tubercles; pleonite 3 with sublateral
tubercle. Pleotelson with 2 prominent spikes
opposing those of pleon, dorsal surface moder-
ately granular, with lateral row of 5 prominent
sublateral tubercles running longitudinally, 3
prominent median tubercles placed longitudi-
nally anterior to pleotelson apex; posterolateral
flange with 2 acute tubercles; posterior margin
with 2 submedian indentations on either side of
median lobe, each point with prominent dorsal
tubercle.

Antennule peduncle article 1 with 8 anterior
spikes, with 1 long proximo-posterior spike, 1
very short posterior spike, and 1 posterodistal
spike; peduncle article 2 with prominent antero-
proximal tubercle and small medial tubercle;
dorsal surface of peduncular articles 1 and 2 pro-
vided with few roughened setae; flagellum with 7
articles.

Epistome with 2 prominent conical spikes,
basally somewhat flattened, on basal transverse
ridge, lateral lobes each with small tubercle.

Pereopods essentially the same as O. obre-
gonia.

Penial processes not examined in detail, simi-
lar to those of O. obregonia.

Pleopod 1 endopod medial margin with PMS
along proximal two-thirds. Pleopod 2 appendix
masculina straight, 5.3 times as long as maxi-
mum width, shorter than (0.74) as endopod, not

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 185

Figure 24. Oxinasphaera multidens (Richardson). A-E lectotype, F-G paralectotype d. A, dorsal view; B, lateral
view; C, cephalon; D, frons; E, pleotelson apex; F, antennule; G, pleopod 1; H, pleopod 2; Scale 1.0 mm.

186 N. L. BRUCE

extending beyond endopod, apex narrowly
rounded. Uropod not examined in detail, gener-
ally similar to O. obregonia; exopod about half as
long as endopod, apex with 3 prominent and 1
smaller ventrally directed spikes.

Female. Not differing from others of the

group.

Colour. Faded in alcohol to a pale yellow
colour.

Size. Males 5.2-5.6 mm.

Remarks. Richardson's (1910) description was
based on specimens from two stations, both
from near Jolo Light, Philippines. Implicit in her
remarks is that material from station 514, the
type locality, contained numerous males. Simi-
larly implicit is that the material from station
4145 contained only a single male. In her
description Richardson (1910: 27) stated that
there was “one small median point instead of
two.” In this she was not quite accurate as O.
multidens does, as do all species of the genus,
have a single rostrum, but lacks any trace of a
rostral spike. Given that there is some ambiguity
over the status of all the material that Richard-
son reported on, and also as a formal redescrip-
tion is given here, the larger male from the tpe
locality is designated as the lectotype.

The species is readily identified by the unique
pattern of pleonal tubercles, and the antennular
morphology of peduncle article 1 having 8
anterior and 3 posterior spikes and peduncular
article 2 having a prominent tubercle on the
anterior margin.

This species is one of a group of central Indo-
Pacific species, generally similar to O. bisubula,
but is principally characterized by having
elongate penial processes and a single rostral
spike when present. The other species are Oxi-
nasphaera obregonia sp. nov. and Oxinasphaera
tual sp. nov.

Distribution. Known only from the type locality,
at a depth of about 53 m.

Hosts. Unidentified sponge.

Oxinasphaera corypantha sp. nov.
Figure 25

Material examined. Holotype. d (4.3 mm), Lot Maitre,
Nouméa, New Caledonia, Oct 1978, 20 m. from
Siphonochalina, Yves Magnier (QM W20040).

Paratypes. 69 (non-ovig 3.0, 3.1, 3.3, 3.5, 3.6. 3.8
mm), 4 imm (2.5, 2.5, 2.7, 2.7 mm), 2 broken, 4 man-
cas (1.8-2.0 mm), same data as holotype (QM
W8083).

Description of male. Body about 2.2 times as
long as greatest width; dorsal surfaces polished,
generally granular, with scattered setae. Cepha-
lon dorsal surface not nodulose; anterior margin
without nodules on either side of rostrum; ros-
trum without spike. Pereonite l with indistinct
posterior ridge, surface of which is weakly
tuberculate. Pereonite 2 weakly tuberculate; per-
eonites 3-6 each with 2 transverse rows of
prominent acute spikes, anterior row slightly
larger than posterior row; pereonite 7 with only
anterior spike row distinct, posterior margin
bisinuate; coxae 4 and 5 posterior margins
weakly concave, coxa 7 posteriorly rounded.
Pleon with posterior boss, with posterolateral
angles strongly produced, produced portions
each with prominent posterior spike and | small
anterior spike; pleonite 3 with 1 lateral tubercle.
Pleotelson with 2 spikes opposing those of pleon,
dorsal surface weakly granular and setose, with 2
additional lateral tubercles on each side; poste-
rolateral flange with 2 acute tubercles; posterior
margin with 2 submedian indentations on either
side of median lobe; median lobe slender, dis-
tinctly shorter than sinus.

Antennule peduncle article 1 with 9 anterior
spikes; with 1 long proximo-posterior spike and
1 shorter posterior spike; peduncle article 2 with
I indistinct anterior spike; dorsal surface of ped-
uncular articles 1 and 2 provided with few
roughened setae.

Epistome with 2 prominent conical spikes,
basally somewhat flattened.

Pereopods essentially the same as O. bisu-
bula.

Penial processes essentially the same as O.
bisubula (examined in situ).

Pleopod 1 medial margin with subproximal
ventral groove. Pleopod 2 appendix masculina
straight, 7.6 times as long as maximum width,
distolateral margin curving at apex, longer (1.1)
than endopod, extending slightly beyond endo-
pod by about 0.16 of its length), apex bluntly
rounded. Uropod not examined in detail, simi-
lar to that of O. bisubula or O. multidens.

Female. No ovigerous females observed, and the
maturity of the “non-ovigerous” females in the
sample is unclear. Generally indistinguishable
from others of the group.

Colour. In alcohol, pale cream yellow colour,
females with scattered brown chromatophores
over dorsal surfaces.

Size. Males 4.3 mm, non-ovigerous females 3.0-
3.8 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 187

Figure 25. Oxinasphaera corypantha sp. nov. All figs of holotype. A, dorsal view; B, lateral view; C, frons; D.
antennules, anterior view; E, pleotelson apex; F, pleopod 1: G, pleopod 2. Scale 1.0 mm.

Remarks. 'The details given here for the anten-
nule and uropod are brief as these appendages
were not dissected off on order to preserve the
integrity of the single male (holotype) speci-
men.

The prominence of the pleonal posterior lobe
almost approaches the form of processes, and
that character along with the posterior ridge on
pereonite 1, the generally weakly developed per-
eonal spike rows and the short and slender
median telsonic process all serve to identify this
species. Oxinasphaera multidens is the only

other species that lacks a rostral spike, and that
species has a far more prominent pereonal
spikes, numerous secondary tubercles on the
pleon, a robust telsonic median lobe and the
appendix masculina not longer than the endo-
pod of pleopod 2.

Distribution. Known only from the type locality,
Ilot Maitre, Nouméa, New Caledonia.

Hosts. Recorded from the sponge Siphonochal-
ina sp.

188 N. L. BRUCE

Oxinasphaera obregonia sp. nov.
Figure 26
Material examined. Holotype. d (3.6 mm), N of Straits
of Sunda, Java, Indonesia, 05*40'S, 106°08’E, 28 Aug
1922, 54 m, numerous sponges [Sigsbee trawl]. stn. 71,
Th. Mortensen's Kei Is. Expedition 1922 (ZMUC
CRUI391).

Paratypes. 69 (3.5, 3.5, 3.6, 3.6, 3.7, 3.8 mm), imm
(25 mm). same data as holotype (ZMUC
CRU1390).

Non-paratvpic material. & (3.7 mm), 9 (3.0, 3.2, 3.3
mm), imm (2.7 mm), same data as holotype. but
station number 11 (ZMUC CRUI389).

Description of male. Body about 2.3 times as
long as greatest width: dorsal surfaces not pol-
ished, generally granular, with scattered setae.
Cephalon dorsal surface not nodulose; anterior
margin with 3-4 small lateral nodules on either
side of rostrum: with prominent single acute
spike on rostrum. Pereonite | without tubercles
or ornamentation. Pereonites 2 and 3 with 2
transverse rows of spikes, pereonites 4-6 each
with 2 transverse rows of prominent acute
spikes, anterior row slightly larger than posterior
row; pereonite 7 with posterior row of low
spikes; coxae 5 and 7 with posterior margins
evenly rounded, coxa 6 ventro-posteriorly con-
cave. Pleon with weakly developed posterior
boss, with posterolateral angles of boss each with
small spikes. Pleotelson with 2 spikes opposing
those of pleon, dorsal surface weakly granular,
without additional tubercles; posterolateral
flange with | acute tubercle; posterior margin
with 2 submedian indentations on either side of
median lobe, each point with distinct dorsal
tubercle.

Antennule peduncle article 1 with 7 anterior
spikes; with | long proximo-posterior spike and
| shorter posteromedial spike; dorsal surface of
peduncular articles 1 and 2 provided with few
roughened setae; flagellum with 6 articles.

Epistome with 2 prominent conical spikes,
basally somewhat flattened.

Pereopods essentially the same as O. hisubula,
but propodus of pereopod 7 with very promi-
nent spines, the longest of which are as long as
the propodus.

Penial processes each about 4.1 times as long
as basal width, both margins tapering towards
slender apex.

Pleopod 2 appendix masculina straight, 5.6
times as long as maximum width, about as long
(0.97) as endopod, extending slightly beyond
endopod (by about 0.10 of its length), apex nar-
rowly rounded. Uropod exopod about 4.7 times

as long as proximal width, about 0.5 times as
long as endopod, apex deeply bifid with lateral
process prominent; endopod about 3.0 times as
long as wide, apex with 3 prominent ventrally
directed spikes.

Female. Similar to other species of the O. bisu-
bula group.

Colour. Faded in alcohol to a pale yellow
colour.

Size. Males 3.5-3.8 mm, ovigerous female 3.0-
3.3 mm.

Remarks. The sample from station number 11 is
excluded from the type series as the label data is
incompatible with the station number, and
therefore the locality cannot be regarded as cer-
tain. All females from this sample are in poor
condition, and are therefore not described.

The single rostral spike immediately separates
O. obregonia from all other species of the genus
except O. tual. Both of these species are further
characterized by having slender elongate penial
processes. O. obregonia is distinguished from O.
tual by having close set epistome spikes without
an additional pair of smaller spikes between,
very weakly developed pleonal boss, antennule
peduncle article 1 with 7 spikes and article 2
without a spike, pleopod 1 medial margin simple
and the appendix masculina being of even
width.

Distribution. Known only from the type locality,
at a depth of about 54 m..
Hosts. Unidentified sponges.
Oxinasphaera tual sp. nov.

Figure 27
Material examined. Holotype. d ( 4.5 mm), off Tual,
Kei Is., (now Pulau Kai, Dulah), Indonesia, c. 05°37’S,
132743 E, 21 Mar 1922, 2 m, from sponges, Th. Mor-
tensen’s Kei Is. Expedition 1922 (ZMUC
CRUI394).

Paratypes. 40 (3.7, 3.9, 4.0, 4.2 mm), 59 (ovig 3.5,
non-ovig 3.0, 3.0, 3.2, 3.3 mm), same data as holotype
(ZMUC CRUI395). Note: No station number is
associated with this sample, presumably hand col-
lected.

Description of male. Body about 2.1 times as
long as greatest width; dorsal surfaces not pol-
ished, generally granular, with scattered setae.
Cephalon dorsal surface not nodulose; anterior
margin with 4-5 small nodules on either side of
rostrum; with single acute spike on rostrum. Per-
eonite | without tubercles or ornamentation.
Pereonites 2-7 with 2 transverse rows of small

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 189

co)

^T

On) Mo gb vo A

Figure 26. Oxinasphaera obregonia sp. nov. A-F holotype, G-J paratype ¢ 3.6 mm. A, dorsal view; B, lateral
view; C, cephalon; D, antennules, anterior view; E, frons; F, pleotelson apex; G, antennule; H, pereopod 7, distal
articles; I, pleopod 2; J, penes; K, uropod. Scale 1.0 mm.

N. L. BRUCE

190

MN
AZ ANN
/ Dani W

lj pi y \

E holotype, F-J paratype 9 4.0 mm. A, dorsal view; B, lateral view; C,
erior view: E, frons; F, antennule; G, pleopod 1; H, pleopod 2; I,

Figure 27. Oxinasphaera tual sp. nov. A-
cephalon, anterior margin; D, antennules, ant
uropod; J, penes. Scale 1.0 mm.

^ NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 191

spikes, pereonites 3-7 each with anterior spike
row more prominent acute; pereonite 7 with
posterior row of low spikes; coxae 5 and 6 with
posterior margins subtruncate, coxa 7 ventro-
posteriorly rounded. Pleon with prominent pos-
terior boss, each posterolateral angle with 2 dis-
tinct spikes positioned anteriorly to posterior
spike; pleonites 2 and 3 each with lateral acute
tubercle. Pleotelson with 2 spikes opposing
those of pleon, dorsal surface weakly granular,
without additional tubercles; posterolateral
flange with 2-3 acute tubercles; posterior mar-
gin with 2 submedian indentations on either side
of median lobe.

Antennule peduncle article 1 with 9 anterior
spikes; with 1 long proximo-posterior spike and
1 shorter posterior spike; peduncle article 2 with
l anteroproximal spike; dorsal surface of pedun-
cular articles 1 and 2 provided with few
roughened setae; flagellum with 4 articles.

Epistome with 2 prominent conical widely
separated spikes, between which lie 2 small
spikes; lateral lobes each with 1 low tubercle.

Pereopods essentially the same as O. bisu-
bula.

Penial processes basally distinctly separate,
each process about 4.5 times as long as basal
width, both margins tapering towards slender
apex.

Pleopod 1 medial margin with stiff setae set
proximally to prominent ventral lobe, medial
margin of which is densely setulose. Pleopod 2
appendix masculina straight, 9.0 times as long as
maximum width, about as long (0.99) as endo-
pod, extending slightly beyond endopod (by
about 0.08 of its length), distal 0.4 abruptly nar-
rowed, apex subtruncate. Uropod exopod about
3.7 times as long as proximal width, about 0.5
times as long as endopod, apex deeply bifid with
lateral process prominent; endopod about 4.0
times as long as wide, apex with 4 prominent
ventrally directed spikes.

Female. Similar to other females of the group;
ovigerous females not observed

Colour. Faded in alcohol to a pale yellow
colour.

Size. Males 3.7-4.5 mm, ovigerous females 3.0-
3.5 mm.

Remarks. This species belongs the group which
the pleon has a posterior boss and a distinct
spike at the posterolateral angles of the boss. Of
that group there are several Indo-Pacific species
that either lack the rostral spike or have a single
spike. These species are Oxinasphaera multidens

and Oxinasphaera corypantha sp. nov. (without
a rostral spike), Oxinasphaera obregonia sp. nov
and the present species (with a rostral spike).
Oxinasphaera obregonia and O. tual are readily
separated by differences in pleonal morphology,
the posterior pleonal process in O. twal being
both prominent and prominently armed, and by
O. tual having a medial lobe on pleopod | endo-
pod and the appendix masculina being flask
shaped.

Distribution. Known from the

locality.

only type

Hosts. Unidentified sponge.

Etymology. The epithet is taken from the type
locality (noun in apposition).

Oxinasphaera rebutia sp. nov.
Figure 28

Material examined. Holotype. ¢ (4.4 mm), E of Mal-
abar, 33°58’S, 151°17’E, NSW, 3 Dec 1973, 66 m,
AMSBS (AM P22196).

Paratypes. 39 (5.2, 5.0, 4.6 mm), 9?(3.8 mm), 9 km E
of Coogee, NSW, 33°57’S, 151°21’E, 89 m, no date,
about 1898 to 19147, fine sand, E.R. Waite on HMCS
Thetis (AM P44206).

Non-paratypic material. 39 (3.5, 3.7, 3.7 mm), E of
North Head, Port Jackson, NSW, 33°49’S, 151°18’E, 2
Feb 1973, 25.9 m, from unidentified sponge, AMSBS
(AM P22983).

Description of male. Body about 2.2 times as
long as greatest width; lateral margins subparal-
lel. Cephalon granular, anterior margin without
tubercles; rostrum without spike. Pereonite |
granular, with posteromedian cluster of 5 promi-
nent rounded tubercles, median tubercle being
twice as large as others. Pereonites 2-7 each with
2 transverse rows of rounded spikes, posterior
row weak; pereonites 2 and 3 with median 3
spikes conspicuously more prominent than
remainder; coxae 5 and 7 rounded, 6 subtrun-
cate. Pleon without posterior boss, with numer-
ous large rounded tubercles. Pleotelson granu-
lar, without spikes, posteriorly with distinct low
rounded tubercles; posterolateral flange without
prominent tubercles; posterior margin deep, lat-
erally depressed, posterior margin deeply exca-
vate, with short rounded median process dor-
sally with single distinct low rounded tubercle,
and longitudinal row of low rounded
tubercles.

Antennule peduncle article 1 with 9 elongate
subacute anterior spikes, medial spike being
smaller than remainder; with 1 posteroproximal

192 N. L. BRUCE

NON a ay nn
00005555;
FELIS
UV v 2,

Figure 28. Oxinasphaera rebutia sp. nov. All figs of holotype. A, dorsal view; B, lateral view; C, antennules,
anterior view; D, frons; E, pleotelson apex; F, pleopod 2. Scale 1.0 mm.

and | posteromedial spike subequal in size; dor-
sal surface of peduncular articles 1 and 2 with
few setae; flagellum with 7 articles.

Epistome with 2 long basally united distinct
spikes; lateral posterolobes with weak
tubercle.

Pereopods not examined in detail.

Penial processes examined in situ, about three
times as long as basal width, apex subacute.

Pleopod 2 appendix masculina shorter (0.85)
than endopod, 7.7 times as long as maximum
width, extending slightly (0.07) beyond endo-
pod, apex slender and acute. Uropod exopod
about 4 times as long as proximal width, about
half as long as endopod, apex deeply bifid with
processes about equally prominent, ventral mar-
gin with prominent acute serrations; exopod
about 4 times as long as wide, apex with 2

prominent downwardly projecting spikes, ven-
tral margin with 4-5 prominent serrations.

Female. Unknown.

Colour. Pale tan in alcohol; chromatophores not
observed.

Size. Males 3.5-5.2 mm.

Remarks. The characteristic tubercles on per-
eonite 1, granular surface of the cephalon and
pereonite 1, lack of a rostral spike and telson
morphology readily identifies this species. Only
O. aylostera has a similar telson morphology and
these two species can easily distinguished by O.
rebutia lacking a rostral spike, and having a
prominent cluster of tubercles on the posterior
of pereonite l.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

The three specimens AM P22983 differ
slightly from the others in that the pereonal and
antennular spikes are poorly developed, and are
excluded from the type series. Two of the speci-
mens have a poorly developed median spike on
the posterior of pereonite 1, and all agree with O.
rebutia in lacking rostral spikes, and in the mor-
phology of the antennule, uropods, pleon and
pleotelson.

The old specimens were too fragile to dissect,
and therefore the description given here is brief
and taken only from the holotype.

Distribution. All specimens taken in the vicinity
of Sydney, New South Wales, at depths from 26
to 89 m.

Hosts.
sponge.

One sample from an unidentified

Oxinasphaera aylostera sp. nov.
Figures 29, 30

Material examined. Holotype. & (4.2 mm), N of Fly
Point, Port Stephens, NSW, 32°43’S, 152°09’E, 8 Nov
1981, 20 m, orange sponge on dead mussel, R.T.
Springthorpe and D. Stracey (AM P44204).

Paratypes. NSW. 39 (ovig 3.8, non-ovig 3.8, 3.6
mm), same data as holotype (AM P41153). (4.3 mm),
Nelson Head, Port Stephens, 32°43’S, 152°09’E, 27
Oct 1980, 24 m, rubble on stones in channel, J. Hall
(AM P44193). 8 (4.2 mm), Barrenjoey Head, Broken
Bay, 33°35’S, 151°20’E, 22 Apr 1983, 5 m, sponge, J.K.
Lowry (AM P41157).

Vic. 23 (4.0, 4.0 mm), 49 (ovig 3.6, non-ovig 3.0, 3.4,
parasitized 3.8 mm), SW shore, Gabo Is, 34°58’S,
149*55.7'E, 19 Feb 1973, 28 m, sponge community
between lighthouse and jetty, J.E. Watson and S.A.
Shepherd (NMV J40480).

Non-paratypic material. NSW. About 300 non-ovi-
gerous females and mancas, same data as holotype
(AM P44203). 4 (4.5 mm), Barrenjoey Head, Broken
Bay, 33°35’S, 151°20’E, 22 Apr 1983, 2 m, yellow
sponge, J.K. Lowry (AM P41158).

Description of male. Body about 2.1 times as
long as greatest width; lateral margins subparal-
lel. Cephalon anterior margin without tubercles;
rostrum with prominent bifurcate acute spike.
Pereonite | unornamented. Pereonites 2-7 each
with 2 transverse rows of spikes, anterior row
distinctly larger than posterior on pereonites 2, 3
and 7, pereonite 2 with prominent median spike;
coxae 5-7 rounded. Pleon without posterior
boss, with 2 rounded tubercles at the usual pos-
ition of boss, posterolateral margin of pleonite 3
with prominent tubercles. Pleotelson granular,
without spikes, posteriorly with distinct low
rounded tubercles; posterolateral flange with 1
marginal rounded tubercle; posterior margin

193

deep, laterally depressed, posterior margin
deeply excavate, with short rounded median
process dorsally with single distinct low rounded
tubercle.

Antennule peduncle article | with 8 elongate
subacute anterior spikes; with | long postero-
proximal and 1 posteromedial spike; dorsal sur-
face of peduncular articles 1 and 2 with few
setae; flagellum with 7 articles.

Epistome with 2 long basally united distinct
spikes; lateral lobes unornamented.

Pereopod 7 carpus 1.4 times as long as merus,
about twice as long as wide, posterior margin
with 3 biserrate, 2 simple spines, distal margin
with 3 large trifid spines, largest nearly as long as
carpus, and 4 biserrate spines, anterodistal angle
with 2 slender weakly biserrate spines; propodus
1.1 times as long as carpus, 3.2 times as long as
wide, posterior margin with 3 biserrate spines,
anterodistal angle with 2 sensory setae.

Penial processes 3.3 times as long as basal
width; medial margin straight, lateral margin
curving to subacute apex; proximolateral mar-
gin with scale spikes.

Pleopod 2 appendix masculina shorter (0.8)
than endopod, 6.7 times as long as maximum
width, not extending beyond endopod, apex
bluntly rounded. Uropod dorsally nodular; exo-
pod about 4 times as long as proximal width,
about half as long as endopod, apex deeply bifid
with processes about equally prominent, ventral
margin with prominent acute serrations; exopod
about 4 times as long as wide, apex with 2 promi-
nent downwardly projecting spikes, ventral mar-
gin with 4-5 prominent serrations.

Female. Pleotelson smoothly rounded; posterior
margin with weak indistinct median inden-
tation.

Colour. Pale tan in alcohol; chromatophores not
observed.

Size. Males 3.6-4.5 mm, females (ovigerous and
non-ovigerous) 3.6-3.8 mm.

Remarks. This species and Oxinasphaera rebu-
tia can be separated from all others of the genus
by the posterior margin of the pleon not being
produced and pleotelson posterior margin
having a deep profile with a wide and deep
median excavation which houses a short median
process. This process is ornamented by a distinct
rounded tubercle, and the lateral lobes on either
side have a longitudinal row of similar tubercles.
The two species also have prominent and acute
antennule spikes and an elongate bifurcate

194 N. L. BRUCE

viv V V. VV JW QS S.

Figure 29. Oxinasphaera aylostera sp. nov. A-G, J holotype, remainder paratype 9 4.8 mm NMV J26423. A,
dorsal view; B, lateral view; C, cephalon; D, frons; E, antennules, anterior view; F, pleotelson posterior margin;
G, pleotelson posterior margin, ventral view; H, antennule; I, pereopod 7, distal articles; J, pleopod 2; K, penes.
Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 195

Figure 30. Oxinasphaera aylostera sp. nov., AM P41153. A, non-ovigerous 9, pleon and pleotelson; B, ovigerous
9, pleon and pleotelson; C, ovigerous female, pleon and pleotelson, lateral view.

epistome spike, Oxinasphaera aylostera is recog-
nized by pereonite 2 havinga prominent median
spike, and the anterior spike rows on pereonites
3 and 4 being larger than is usual in the genus. It
is readily distinguished from O. rebutia by
havinga rostral spike, and by lacking prominent
tubercles on pereonite |.

Distribution. Port Stephens, NSW to Gabo Is.,
eastern Victoria, at depths from 2 to 28 m.

Hosts. Unidentified sponges.

Oxinasphaera tuberculosa
(Stebbing, 1873) comb. nov,

Figures 31-33

Cymodocea tuberculosa Stebbing, 1873: 96, pl. 3 figs
I-Ib.

Cymodoce tuberculosa. — Baker, 1910: 76, pl. 21
figs 1-20.-Nierstrasz, 1931: 200.

Cymodoce tuberculosa tuberculosa. — Harrison and
Holdich, 1984: 392 (remarks).

Non Cymodoce tuberculosa. — Whitelegge, 1902:
258, fig. 28 (= Oxinasphaera thetisae sp. nov.).

Non Cymodoce tuberculosa. — Baker, 1908: 140, pl.
3 figs 12-15 [= O. bispinosa (Baker, 1910)].

Non Cymodoce tuberculosa bispinosa Baker, 1910:
78, pl. 21 figs 21-23, pl 22 figs 1-7 [= O. bispinosa
(Baker, 1910)].

Non Cymodoce tuberculosa tripartita. — Barnard,
1920: 363, pl. 15 fig. 28. — Barnard, 1940: 493 (key).
— Kensley, 1978: 100, Fig. 42C (= O. kensleyi sp.
nov).

Material examined. WA. d (6.0 mm), 7o (4.8, 4.8, 4.9,
5.0, 5.5, 5.5, 5.6 mm), Bundegi Reef, near Point
Murat, Exmouth Gulf, 21°49’S, 1 131 IE. 4 Jan 1984,
9 m, large grey cup sponge, R.T. Springthorpe (AM
P41351). 3 (6.2 mm), Arthur Head, Fremantle,
32°03’S, 115°44’E, 25 Dec 1983, 6 m, from Caulerpa,
J.K. Lowry (AM P41 116). 40 7 and o (previously partly
desiccated), western end, Lucky Bay. 33°59.0’S,

118°02.5°E, 12 Apr 1984, 20 m, gorgonians and soft
corals, G.C.B. Poore and H.M. Lew Ton (NMV
J26181). 28 (5.4, 5.8 mm), 9 (ovig 4.8 mm), N end of
Little Beach, Two Peoples Bay, 34°58.2’S, 1 1810.8 E.
18 Apr 1986, 5 m. yellow sponge, anemones, G.C.B.
Poore and H.M. Lew Ton (NMV 726166). 38 (5.5, 5.6.
5.8 mm), ? (damaged, 5.0 mm), off Possession Point,
King George Sound. 35°02.57S, 117955.0'E, 14 Dec
1983. 7 m, from purple finger sponges, R.T. Spring-
thorpe (AM P41099). 9 (5.8 mm), 9 (4.8 mm), 17
mancas and juveniles (in poor condition), off SE cor-
ner of Michaelmas Is., King George Sound. 35°03’S.
118°00’E, 17 Dec 1983, 24 m, branching sponge, R.T.
Springthorpe (AM P41108, slides PA4218). d (4.8 mm,
senescent), NE end, Vancouver Peninsula, 35°03.4’S,
117°56.2’E, 8 Apr 1986, 6 m, dictyotaleans, G.C.B.
Poore and H.M. Lew Ton (NMV J26163).

SA. 4d (4.5, 5.0, 5.3. 5.5 mm), 9 (4.5 mm), Coal Reef,
Tiparra Reef, Tiparra Bay, 34°04.0°S, 137923.0/E, 15
Mar 1985, 5 m, sponges with hydroids and algae.
G.C.B. Poore and H.M. Lew Ton (NMV J40481). £,
Penneshaw jetty, 31 Jan 1989, sand and weed on
pylons, 5-8 m, K.L. Gowlett-Holmes (SAM C 5605).
38, Cape D'Estaing, 27 Jan 1989, 10-13 m, rubble,
reef crevice, W. Zeidler and K.L. Gowlett-Holmes
(SAM C5606). 9, Pearson Is., Investigator Group, 10
Jan 1969, 33 m, algae on slope, S.A. Shepherd (SAM
C5607). 10 specimens (in poor condition), Wedge Is.,
Gambier Group, 29 Dec 1963, 26 m, from sponge,
SAORI (SAM C5608). o, West Is., Encounter Bay, 12
Feb 1966, ex sponge, S.A. Shepherd (SAM C 5609). 9,
Whyalla, Upper Spencer Gulf, 33°02.4’S, 137^37.6'E,
Sept 1987, 10 m, SA Fisheries Survey (C5610). g,
Upper Spencer Gulf, Commissariat Point, 35°35.00’S,
137°46.08’E, Feb 1987, 6 m, beacon 20, S.A. Fisheries
(C5611). 4 d, ‘South Australian Coast’, no other data,
probably Baker's 1910 specimens (SAM C368).

Vic. 7 (5.6 mm), SW shore midway between light-
house and jetty, Gabo Is., 37°34’S, 149°55’E, 19 Feb
1973, 28 m, sponge community, J.E. Watson and S.A.
Sheperd (NMV J40493).

Tas. d, 35 km N of Cape Wickham, King Is.,
39°13.6’S, 143°55.6’E, 23 Nov 1981, 85 m, fine sand,
R. Wilson (NMV 140495).

196 N. L. BRUCE

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Figure 31. Oxinasphaera tuberculosa (Stebbing). A-E ¢ 6.2 mm, Fremantle (AM P41116), remainder 5.8 mm,
King George Sound (AM P41108). A, dorsal view; B, lateral view; C, cephalon; D, frons; E, pleotelson posterior
margin; F, antennule; G, antenna; H, penes. Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 197

Figure 32. Oxinasphaera tuberculosa (Stebbing). All figs & 5.8 mm, King George Sound (AM P41108). A, per-
eopod 1; B. pereopod 2: C, pereopod 7; D, pleopod 1; E, pleopod Da

198

N. L. BRUCE

Figure 33. Oxinasphaera tuberculosa (Stebbing). A, D, F, G non-ovigerous 9 4.8mm, King George Sound (AM
P41108); B, C, Eovigerous o 4.8 mm, Two Peoples Bay (NMV J26166). A, pleon and pleotelson, dorsal view; B,
pleon pleotelson, dorsal view; C, pleon pleotelson, lateral view; D, frons; E, frons; F, antennule; G, uropod.

Types. l have been unable to locate the
material from the Swan River, WA on which
Stebbing based his description. It 1s not held at
The Natural History Museum, London, nor any
other institution that I have contacted.

Description of male. Body about 2.5 times as
long as greatest width; dorsal surfaces not pol-
ished, generally granular, with scattered setae.
Cephalon dorsal surface not nodulose; anterior
margin unornamented; rostrum without spike,
with pair of very weak nodules. Pereonite 1 pos-
terior margin finely granular. Pereonite 2 finely
granular, and 3-7 each with 2 transverse rows of
prominent spikes, anterior and posterior rows
subequal in size; coxae 5-7 with posterior mar-
gins evenly rounded. Pleon without posterior
boss, without prominent spikes. Pleotelson
without prominent spikes or acute tubercles,
prominent rounded tubercles on posterior lobe;
posterolateral flange without acute tubercles;

posterior margin with deep groove on either side
of distinctly dorsal median lobe, telson on either
side of apical notch forming distinct lobe, each
with prominent spike.

Antennule peduncle article 1 with 4 large and
1 small medial flat anterior spikes; without pos-
terior spikes; dorsal surface of peduncular
articles 1 and 2 provided with few roughened
setae; flagellum with 8 articles.

Epistome with 2 prominent elongate flat
truncate spikes, without subsidiary spikes.

Pereopod 1 basis about 2.8 times as long as
wide, anterior margin with 2 sensory setae;
ischium about 0.7 times as long as basis, about
2.4 times as long as wide, anterior margin with 1
short proximal spines and 1 distal longer and
biserrate spine; merus about half as long as
ischium, about 1.2 times as long as wide, ante-
rolateral angle with 1 gently curving pectinate
spine, posterior margin with 1 spine and single

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

long simple seta; carpus short, about 0.7 times as
long as merus, about 1.1 times as long as wide,
posterior margin with 2 biserrate spines; propo-
dus about 0.8 times as long as ischium, widest
proximally, about 2.5 times as long as wide, pos-
terior margin with distinct scale spikes and 4
large biserrate spines, row of 4 smaller biserrate
spines on medial margin; dactylus about 0.6
length of propodus, unguis about 5596 length of
entire dactylus. Pereopods 2 and 3 similar to 1,
differing principally in carpus being more
elongate and propodus more slender. Pereopod
2 basis about 3.0 times as long as wide; ischium
about 2.5 times as long as wide; carpus about
equal in length to merus, about 1.6 times as long
as wide, with 3 biserrate and 3 trifid spines on
posterior of distal margin; propodus about 0.7
times as long as ischium, about 3 times as long as
wide, margins subparallel, posterior margin with
3 weakly biserrate spines. Pereopod 7 basis
about 3.3 times as long as wide, posterior margin
with prominent scale spikes, with several sen-
sory setae; ischium slightly shorter than basis,
about 2.7 times as long as wide; merus about 0.5
times as long as ischium, about 1.6 times as long
as wide, posterior margin with 3 biserrate
spines, anterodistal angle with | large and 1
small spines; carpus about as long as merus,
about 2.0 times as long as wide, posterior margin
with 4 biserrate spines, distal margin with 4 large
trifid spines and 5 biserrate spines, anterodistal
margin with 1 weakly serrate spine; propodus
about as long as carpus and 0.6 time as long as
ischium, about 3.4 times as long as wide, pos-
terior margin with 3 biserrate spines, anterodis-
tal angle with | sensory seta.

Penial processes each about 1.8 times as long
as basal width, both margins converging evenly
to an acute point; proximolateral margins
densely scaled.

Pleopod 1 endopod without lobed or grooved
medial margin. Pleopod 2 appendix masculina
curving weakly laterally, 7.6 times as long as
maximum width, about as long (0.97) as endo-
pod, extending slightly beyond endopod (by
about 0.16 of its length), apex bluntly rounded.
Uropod exopod about 3.4 times as long as proxi-
mal width, about 0.45 times as long as endopod,
apex deeply bifid with lateral process promi-
nent; endopod about 3.4 times as long as wide,
apex with prominent terminal spike and 2
smaller ventrally directed spikes.

Female. Ovigerous females with unornamented
somites. Posterior margin of pleotelson
upturned, with wide shallow indentation; with

199

conspicuous marginal setae. Antennule ped-
uncle article 2 0.4 as long as article 1, article
3 1.7 times as long as article 2; flagellum 0.8
times as long as peduncle, with 7 articles. Uro-
pod endopod flat, distally with medial margin
evenly rounded; exopod flat, about half as long
as endopod, apex bifid. Non-ovigerous female
with posterior margin of pleotelson not
upturned, smoothly rounded, without wide shal-
low indentation. Antennule peduncle article 2
proportionally shorter than in ovigerous female,
0.24 as long as article 1; flagellum about three-
quarters as long as peduncle. Uropod as for ovi-
gerous female but endopod with rounded apical
point,

Colour. Pale brown to cream in preserved speci-
mens.

Size. Males 4.5-6.2 mm, females 4.5-5.6 mm.

Remarks. Oxinasphaera tuberculosa was one of
the first described of the Australian marine iso-
pods, but has since remained poorly known. The
species is immediately characterized by the deep
grooves extending anterolaterally from the pos-
terior margin of the telson, and the very large flat
truncate antennule and epistome spikes, charac-
ters that it shares with Oxinasphaera epostoa
and Oxinasphaera lowryi. The latter is easily
separated by having a single epistome spike, and
the pleon with a posterior boss with opposing
spikes on the pleotelson. Oxinasphaera epostoa
is very similar but males of that species have a
far more setose pleon and pleotelson, pereonite
2 and 3 are only weakly nodulose and the appen-
dix masculina is straight and distally nar-
rowed.

Distribution. From North West Cape, WA,
southwards and eastward along the coast of
southern Australia to Gabo Is., Victoria and
Tasmania.

Hosts. From sponges, none of which have been
identified.

Oxinasphaera epostoa sp. nov,
Figures 34, 35

Material examined. Holotype. ¢ (5.5 mm), Table
Point, Cobourg Peninsula, Port Essington, NT,
11°14.8’S, 132°10.5’E, 12 May 1983, 6 m, rock wash-
ings, N.L. Bruce (NTM Cr001 1329).

Paratypes. NT. 2o (4.0, 4.8 mm), same data as holo-
type (NTM Cr0011330). ¢ (4.3 mm), 139 (non-ovig
mm). 17 mancas (mm), same data as holotype, but 11
May 1983, from Callyspongia ?diffusa (NTM
Cr0011331), 32 (5.3, 6.0 dissected, 6.0 [SEM] mm), 69

200 N. L. BRUCE

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Figure 34. Oxinasphaera epostoa sp. nov. A-E, J holotype, G, Fd 6.0 mm, H, I non-ovigerous 9 5.3 mm (both
ZMUC CRU 1383). A, dorsal view; B, lateral view; C, frons; D, antennules, anterior view; E. pleotelson, posterior
margin; F, pleopod 2; G, penes; H. coxae 2-7; I, pleotelson, dorsal view. Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 201

Figure 35,Oxinasphaera epostoa sp. nov., SEMs. d 5.3 mm (ZMUC CRUI383). A, cephalon, anterior margin
(x75); B, pleon and pleotelson (x30); C, pleotelson apex (x100); D, uropod (x100).

(non-ovig 4.5, 4.7, 4.8, 5.0, 5.3 [drawn] mm), 2 mancas
( 3.5, 4.3 mm), same data as holotype except, from
mixed sponges (ZM UC CRU1383). 218 (3.8-6.4 mm,
mean = 5.5. mm), 129 (9 ovig 3.7-5.3 mm, mean = 4.5
mm, 3 non-ovig ), manca (2.5 mm), same data as holo-
type, except: 7 Aug 1986, 5-7 m, burrowing into
sponge, P.J.F. Davie (QM W20033, ZMUC
CRUI382).

Non-paratypic material. NT. ¢ (6.0 mm), Arafura
Sea, NE of Cobourg Peninsula, 11°09’S, 134°27’E, 21
Oct 1989, 30.2 m, BRR (QM W20044). 179 ( 4.4-6.4
mm, mean = 5.5 mm), 49 (ovig, all broken, non-ovig
4.8 mm), Arafura Sea, NE of Cobourg Peninsula,
11°27’S, 133°34’E, 18 Oct 1989, 20.1 m, BRR (QM
W20042). 3, 59, Table Head, Port Essington, Cobourg
Peninsula, 11?14.4/S, 132°10.8’E, 13 May 1983, 4 m,
Jaspis sp., N.L. Bruce (NTM Cr0011332).

Description of male. Body about twice as long as
greatest width; dorsal surfaces not polished, gen-
erally granular, with scattered setae, particularly
dense on pleon and pleotelson. Cephalon dorsal
surface not nodulose; anterior margin unorna-
mented; rostrum without spike. Pereonite |
anterior margin with row of fine nodules. Per-
eonite 2-6 each with 2 transverse rows of promi-
nent spikes, pereonite with additional middle
row of low nodules; anterior row largest on per-
eonite 4, both rows subequal in size on pereon-

ites 3, 4 and 6; pereonite 7 with posterior spikes
only; coxae 5 and 6 with posterior margins
straight, coxae 7 rounded. Pleon without pos-
terior boss, without prominent spikes. Pleotel-
son without prominent spikes or acute tubercles;
posterolateral flange without acute tubercles;
posterior margin with deep groove on either side
of distinctly dorsal median lobe; median lobe
with prominent rounded median tubercles and
adjacent rounded tubercles; telson on either side
of apical notch forming distinct lobe, each with
prominent spike.

Antennule peduncle article 1 with 4 large and
1 small medial flat anterior spikes; without pos-
terior spikes; dorsal surface of peduncular
articles 1 and 2 provided with few roughened
setae; flagellum with 8 articles.

Epistome with 2 prominent elongate flat
truncate spikes, without subsidiary spikes.

Pereopods as for O. tuberculosa.

Penial processes each twice (2.0) as long as
basal width, both margins converging evenly to
an acute point; proximolateral margins weakly
scaled.

Pleopod 1 endopod without lobed or grooved
medial margin. Pleopod 2 appendix masculina
straight, 7.6 times as long as maximum width,

202

tapering from about mid-length, longer (1.2)
than endopod, extending beyond endopod by
about 0.15 of its length, apex bluntly rounded.
Uropods as for O. tuberculosa.

Female. Not distinguishable from O. tubercu-
losa.

Colour. Pale tan to white in acohol, with scat-
tered chromatophores.

Size. Males 4.5-6.2 mm, females 4.5-5.6 mm.

Remarks. In most characters Oxinasphaera
epostoa differs little from O. tuberculosa. The
characters that do distinguish the two species
are, in Oxinasphaera epostoa the pleon having
fewer tubercles and being densely setose, a
shorter and rounder median process on the pleo-
telson, the lateral grooves on posterior of pleon
are less clearly defined. coxae 5 and 6 are pos-
teriorly straight (rounded in O. tuberculosa), the
anterior pereonites are less nodulose, pereonite
7 with only a single row of spikes, the appendix
masculina being longer and apically more slen-
der and the penes slightly longer (1.95-2,0 vs
1.87 in O. tuberculosa) and less densely set with
scale spikes.

Distribution. Cobourg Peninsula, Northern Ter-
ritory and adjacent Arafura Sea; shallow sub-
tidal to 30 m.

Hosts. Callyspongia ?diffusa.

Etymology. Epostoa is a genus of woolly cac-
tus.

Oxinasphaera lowryi sp. nov.
Figures 36, 37

Material examined. Holotype. 9 (8.0 mm), W side
Bowen ls., Jervis Bay, NSW, 35°07’S, 150°46’E, 28 Jun
1981, 6 m, from large grey sponge, J.K. Lowry and
R.T. Springthorpe (AM P44212).

Paratypes. NSW. c (7.9 mm), Wommin reef, S of
Cook ls., 28°12.0’S, 153°34.8’E, 4 Feb 1993, 21 m,
symbiotic with sponge, J.N.A. Hooper and S. Cook
(QM W18423).9(9.5 mm), same data as previous (QM
W 18427). & (8.5 mm), Julian Rocks, Byron Bay,
28°36.8’S, 15337. 7E, 2 Feb 1993, 18 m, “black coral
garden’, symbiotic with sponge, J. N.A. Hooper and S.
Cook (QM W 18352). 8 (8.5 mm, dissected), inside Box
Head, Broken Bay, 33°33’S, 151°21’E, 22 Nov 1982,
from sponge, J.K. Lowry and R.T. Springthorpe (AM
P41154, slides P44217). d (9.2 mm, intermoult), o
(non-ovig 6.3 mm), same data as holotype (AM
P41159). & (10.0 mm, damaged), Nielsen Park, Syd-
ney, 3 Oct 1979, in Galeolaria, N. Svennivig (ZMUC
CRUI388).

N. L. BRUCE

Description of male. Body about twice as long as
greatest width; dorsal surfaces not polished, gen-
erally granular, with abundant scattered setae,
particularly dense on pleon and pleotelson.
Cephalon dorsal surface not nodulose; anterior
margin unornamented; rostrum without spike.
Pereonite 1 granular, posterior margin with
transverse row of fine nodules; pereonite 2
densely granular, with indistinct anterior and
posterior nodule rows; pereonites 5 with 3 trans-
verse rows of prominent spikes, pereonite 6 and
7 with single transverse row of widely spaced
prominent spikes; coxae 5-7 with posterior mar-
gins evenly rounded. Pleon with prominent pos-
terior boss, with prominent posterolateral spikes
and 2 spikes set anteriorly to these. Pleotelson
densely granular; posterolateral flange with |
tubercle; posterior margin with deep groove on
either side of distinctly dorsal median lobe; tel-
son on either side of apical notch forming dis-
tinct lobe; lobes without prominent rounded
median tubercles or spikes.

Antennule peduncle article 1 with 4 large flat
anterior spikes; without posterior spikes; dorsal
surface of peduncular articles 1 and 2 provided
with few roughened setae; flagellum with 10
articles.

Epistome with single prominent elongate flat
truncate spike.

Pereopod 1 basis about 3 times as long as
wide, anterior margin with 2 sensory setae;
ischium about 0.7 times as long as basis, about
2.3 times as long as wide, anterior margin with 1
short proximal spines and 1 distal longer and
biserrate spine; merus about one-third as long as
ischium, about as long as wide, anterolateral
angle with 1 long and 1 short spine, posterior
margin with | spine and single long simple seta;
carpus short, about 0.9 times as long as merus,
about 0.8 times as long as wide, posterior margin
with 2 biserrate spines; propodus about 0.7
times as long as ischium, widest proximally,
about 2.7 times as long as wide, posterior margin
with 4 large biserrate spines, row of 3 smaller
biserrate spines on medial margin; dactylus
about half length of propodus. Pereopods 2 simi-
lar to 1, differing principally in having the car-
pus more elongate and propodus more slender
and the ischium anterior margin with abundant
setules. Pereopod 7 basis about 2.8 times as long
as wide, posterior margin with prominent scale
spikes, with several sensory setae, anterior mar-
gin with abundant long setules; ischium slightly
shorter than basis, about 2.7 times as long as
wide; merus about half as long as ischium, about
1.8 times as long as wide, posterior margin with

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 203

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Figure 36. Oxinasphaera lowryi sp. nov. A-E holotype, remainder d 8.5 mm (AM P41154). A, dorsal view; B,
lateral view; C, antennules, anterior view; D, frons; E, pleotelson posterior margin; F, antennule, f — setae from
dorsal surface of pedunular article 1; G, right mandible; H, pleopod 2; I, penes, in situ; J, penes. Scale 2.0

mm.

204 N. L. BRUCE

Figure 37. Oxinasphaera lowryi sp. nov. A-D, d 8.5 mm (AM P41154), E-G non-ovigerous 9 6.3 mm (AM
P41159). A, pereopod 1; B, dactylus apex, pereopod 1; C, pereopod 2; D, pereopod 7; E, lateral view; F, frons; G,
dorsal view.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

3 biserrate and 1 trifid spines, anterodistal angle
with 1 large and | small spines; carpus about 0.8
times as long as merus, about 1.8 times as long as
wide, posterior margin with 5 biserrate spines,
distal margin with 2 large trifid spines and 6
biserrate spines, anterodistal margin with 2
weakly serrate spine; ischium to carpus with
both anterior and posterior margins with abun-
dant long setules; propodus about 1.3 times as
long as carpus and half as long as ischium, about
3 times as long as wide, posterior margin with 6
spines, 2 of which are obviously biserrate, ante-
rodistal angle with | sensory seta and 3 simple
setae,

Penial processes each about 3 times as long as
basal width, medial margin straight, distal lat-
eral margin converging evenly to narrowly
rounded point; proximal margins weakly
scaled.

Pleopods as for O. tuberculosa. Pleopod 2

appendix masculina straight, articulating sub-
basally, tapering slightly from about mid-length,
9.2 times as long as maximum width, shorter
(0.8) than endopod, just extending beyond endo-
pod (by about 0.05 of its length), apex bluntly
rounded. Uropod exopod about half as long as
endopod, apex deeply bifid with lateral process
prominent; endopod about 2.6 times as long as
wide, apex with 2 prominent ventrally directed
spikes.
Female. Ovigerous females not observed. Non-
ovigerous females with distinct pleonal dome,
and strongly domed pleotelson; posterior mar-
gin of pleotelson with minute median notch.

Colour. In freshly collected preserved, dorsal
surfaces densely covered by chromatophores
giving an overall dark grey appearance.

Size. Males 7.9-9.5 mm, one non-ovigerous
female 6.3 mm.

Remarks. The species, one of the largest of the
genus, is immediately recognized by having a
single flat epistomal spike, the dorsal surfaces
are densely hirsute, on the pereonites the pos-
terior spike rows are prominent and on pereon-
ites 6 and 7 are in a single transverse row. This
species is the only species with heavily setulose
posterior pereopods.

Distribution. Byron Bay, northern NSW to
Jervis Bay, southern NSW (35'S), intertidal to
20 m.

Hosts. Unidentified sponges; also from the
massed worm tubes of the polychaete Galeo-
laria, although these could have been from
sponges in the tube masses.

205

Etymology. Named in recognition of Dr Jim K.
Lowry's contribution to knowledge of the Aus-
tralian amphipod fauna.

Oxinasphaera thetisae sp. nov.
Figures 38, 39

Cymodoce iuberculosa. — Whitelegge, 1902: 260,
fig. 28, part [non O. tuberculosa (Stebbing)].

Material examined. Holotype. d (11.5 mm), 9 km E of
Coogee, NSW, 33°57’S, 151°21’E, 15 Mar 1898, 89 m,
fine sand. E.R. Waite on HMCS Thetis (AM
G2270).

Paratypes. NSW. d (1 no head, 9.0, 9.5, 9.7, 10.5
previously dissected, 10.6 mm), o (non-ovig 8.5 mm), l
km S of Cape Bailey, 34*02.5'S, 151712.0'E, 11 Mar
1898, 39 m, sand to rock, E.R. Waite on HMCS Thetis
(AM G2194).

Whitelegge listed the stations from which his speci-
mens were taken as: “8 off Cape Hawke, 10 off Coogee
Bay in 25 and 50 fathoms: stns 31 and 44." Material
examined here is G2270 from stn 44 and G2194 from
stn 36. Other samples identified as C. tuberculata in
the AM collections are: G2274 (= O. rebutia sp. nov.
now AM P44206 and O. bisubula sp. nov.) from stn 48;
G2273 (= O. bisubula sp. nov.) from stn 55; G3902 (=
O. bisubula sp. nov.) from stn 48. Material from stn 31
has not been located.

Description of male. Body about twice as long as
greatest width; dorsal surfaces not polished, gen-
erally strongly granular, with scattered setae,
particularly dense on pleon and pleotelson.
Cephalon dorsal surface granular, with anterom-
edial nodule; anterior margin unornamented;
rostrum without spike. Pereonite 1 strongly
granular; pereonites 2-7 each with 2 rows of
transverse spikes, anterior row prominent,
spikes becoming robust on pereonites 5-7 ;
coxae 5-7 with posterior margins evenly
rounded. Pleon without posterior boss, with 2
prominent submedian spikes on posterior mar-
gin, further 5-6 spikes set laterally to these,
forming irregular row. Pleotelson densely gran-
ular; with 4 submedial clusters of 4 and 3 large
spikes; posterolateral flange with 1 acute
tubercle; posterior margin with deep groove on
either side of distinctly dorsal median lobe; tel-
son on either side of apical notch forming dis-
tinct lobe; median lobe with prominent acute or
spikes.

Antennule peduncle article 1 with 4-5 irregu-
lar flat anterior spikes, lateralmost being promi-
nent; without posterior spikes; dorsal surface of
peduncular articles 1 and 2 granular, provided
with few roughened setae; flagellum with 14
articles.

206 N. L. BRUCE

Figure 38. Oxinasphaera thetisae sp. nov. A-I holotype, remainder € 9.5 mm (AM G2194). A, dorsal view; B,
lateral view: C, frons ; D, antennules, anterior view; E, pleotelson posterior margin; F, pleotelson, ventral view of
apex; G, antennule: H, penes; I, uropod, in situ. Scale 3.0 mm.

^ NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

Figure 39. Oxinasphaera thetisae sp. nov. A. B, 9 9.5 mm, C, D, non-ovigerous 9 8.5 (AM G2194). A, pleopod 1;
B, Pleopod 2; C, female, pleon and pleotelson; D, female, cephalon, anterior view.

Epistome with single prominent short flat
truncate spike.

Pereopods similar to those of others of the
group (heavily encrusted, not observed in
detail).

Penial processes each about 2.5-3 times as
long as basal width, medial margin angling lat-
erally a little beyond midpoint, lateral margin
straight, curving medially just before apex; prox-
imolateral margins weakly scaled.

Pleopod 1 endopod two-thirds as long as exo-
pod, proximal margin densely setulose, distal
part angle abruptly to medial. Pleopod 2 appen-
dix masculina lateral margin smoothly curved,
medial margin straight, articulating subbasally,
5.5 times as long as maximum width, tapering
from about mid-length, longer (1.06) than endo-
pod, extending beyond endopod by 0.2 of its
length, apex narrow, truncate. Uropod exopod
about half as long as endopod, apex deeply bifid
with lateral process prominent; endopod about
2.6 times as long as wide, densely granular, apex
with 1 prominent and 2 smaller spikes.
Female. The single non-ovigerous female is dis-
tinctive in having the dorsal surfaces granular.
The posterior margin of the pleotelson is
entire.

Colour. Colour has faded from these nearly 100
year old specimens.

Size. Males 9.0-11.5 mm.

Remarks. The identity of Whitelegge's (1902)
specimens has always been uncertain, and the
material examined here is not identified in the
Australian Museum collection as having been
identified by Whitelegge, but by Baker as Cyrno-
doce tuberculata Stebbing, 1873, presumably a
lapsus. Nonetheless, the description given by
Whitelegge, particularly of the first antennules
and of the size of his specimens, suggests that the
present material and Whitelegge's are the one
species.

The species is distinctive within the genus in
its large size, strongly developed granulosity of
the cephalon and pereonite 1, single flat truncate
epistomal spike, the robustness of the large
pleonal and pleotelson spikes and the irregular
antennular spikes. Only Oxinasphaera lowryi
has a single epistomal spike, but the two species
are otherwise dissimilar in appearance, the
antennular, pleonal and pleotelson characteris-
tics being widely different.

Distribution. Two stations to the south of Syd-
ney, NSW, circa 34°S, at depths between 39 and
89 m.

208

Hosts. No host recorded for this species.

Etymology. Named after the research vessel
from which the specimens were collected, the
HMCS Thetis.

Oxinasphaera bispinosa (Baker, 1910) comb.
nov.
Figures 40-44

Cymodoce tuberculosa. — Baker, 1908: 140. pl. 3 figs
12-15 (non Cymodoce tubereulosa Stebbing, 1873).

Cymodoce tuberculosa n. var. bispinosa Baker, 1910:
78, pl. 21 figs 21-23, pl 22 figs 1-7 (part, see comments
under “types”) [= O. bispinosa (Baker, 1910)].

Material examined. Syntypes (presumed). 34 (7.5, 7.0
[head off], 7.0 [in 2 pieces] mm), 9 (ovig 7.5 mm).
"South Australia" (SAM C372).

Non-type material. Tas. 400+ g and 9, subsampled
169 (7.0-9.5 mm, mean = 8.2 mm [8.0 SEM, 7.8 SEM
dissected). 39 (imm 6.0, 6.5, 7.5 mm), 14 o (ovig 6.8,
7.5. 7.7. 8.5 mm, 10 non-ovig 6.0-8.0 mm, mean = 6.9
mm), 28 km E of Cape Farewell. King Is., 39°32.8’S,
144°16.0’E. | Nov 1980, 18 m, fine sand, M.F. Gomon
and G.C.B. Poore on FV Sarda (NMV 126414, 109,
102 ZMUC CRUI392). 3¢ (6.5. 8.0, 8.5 mm), 47 km E
of Cape Rochon, Three Hummock Is., 40723.8'S,
145*32.0'E, 3 Nov 1980, 66 m, muddy sand, M.F.
Gomon and G.C.B. Poore (NMV 126300). 24 (7.0, 7.5
mm) 32 km NW of Devonport, 40°56.04’S,
146°39.00’E, 4 Feb 1980, 66 m, muddy sand, M.
Gomon and G.C.B. Poore (NMV J26298). 4d (6.9,
6.6., 6.3, 5.4 mm), 7o ( ovig 6.6, 6.3, 6.0. non-ovig 6.6,
6.3, 6.1, 5.7 mm), Isthmus Bay, Bruny Is.. 13 Aug
1984, from sponge, J.R. Penprase (TM G2815). 20
(7.8, 6.6 mm), (non-ovig 5.4 mm), near Partridge Is.,
D'Entrecasteaux Channel, 22 Jul 1957, off sponge on
Mimachlamys asperrimus, vessel Te Rapunga (TM
16682/8445).

Vic. 38 (7.0, 7.8, imm 6.2 mm), 82 km SW of Cape
Liptrap, 39*06.83'S, 144°49.64’E, 11 Feb 1981, 65 m,
medium sand, M.F. Gomon and G.C.B. Poore (NMV
126286).

Additional material. Tas. 200+ d and 9, 35 km E of
Cape Farewell, King Is., 39°39.2’S, 144°21.0’E, 1 Nov
1980, 27 m, finesand, M. Gomon and G.C.B. Poore on
FV Sarda (NMV J26415). 26, 229 and mancas (poss-
ibly mixed species), 35 km N of Cape Wickham, King
Is.. 39*13.6'S, 143'55.6'E, 23 Nov 1981, 85 m, fine
sand, R. Wilson (NMV 126305). 3d, 2 ovigo, 59 non
ovigg and imm g , 5 km N of North Point. 40*40.3'S,
145*15.0'E, 4 Nov 1980, 33 m, medium shell, M.F.
Gomon and G.C.B. Poore (NMV J26255). 3d (l imm),
39 km NNE of Devenport, 40°49.75’S, 146°31.33’E, 4
Feb 1980. 68 m, mud with bryozoa and sponges, M.
Gomon, G.C.B. Poore and C.C. Lu (NMV J40492). 60°
(3 imm), lle (2 ovig), 2.5 km SE of Birches Bay,
D'Entrecasteaux Channel, 43°11.00’S, 147*16.00'E,
16 Apr 1985. 10 m, R.S. Wilson (NMV J40479).

Australian Museum Old Collections. 34, NSW, from
a sponge [det. as C. tuberculosa by Baker, 1926] (AM
P9554). 73, 89, between Merimbula and Tathra, NSW,

N. L. BRUCE

18 Jul 1925, 73 m, taken from sponge, W. Boardman
on trawler Bar-Ea-Mul [det. as C. tuberculosa by
Baker, 1926] (AM P9564). d, Oyster Bay, Tasmania,
42°40’S, 148°03’E, sorted from a bottle of weed wash-
ings, FIS Endeavour Expedition 1909-14 (AM
E6610). d, eastern slope Bass Strait, 6 Sep 1930,
Endeavour (AM E6753). 6, 16 km N of Circular Head,
Tasmania, FIS Endeavour Expedition 1909-14 (AM
E6739). d, Spencer Gulf, South Australia, 29 m, FIS
Endeavour Expedition 1909-14 (AM E6770).

Types. The locality was given by Baker (1910)
only as "South Australian coast", and I have not
been able to identify with absolute certainty the
specimens mentioned by Baker, which would be
the syntypes of this species. The South Aus-
tralian Museum has two samples determined by
Baker as Cymodoce tuberculosa var bispinosa.
One of these (SAM C372, labelled “syntypes”)
contains three males and an ovigerous female of
O. bispinosa. The other (SAM C371, also
labelled “syntypes”) contains two specimens of
Cymodoce tuberculosa Stebbing, 1873 and two
specimens of a large species of uncertain generic
disposition. Another sample exists (SAM C369)
which is identified as Cymodoce tuberculosa, but
these five females do not appear to be Cymodoce
tuberculosa, and furthermore are not a species of
Oxinasphaera, and appear most similar to the
genus Neosphaeroma Baker, 1926. Baker (1910)
unfortunately failed to make clear on which
specimens he was basing his description, and the
sample SAM C372 is here taken to be the pre-
sumed syntypes. Owing to the uncertain type
status of SAM C372, a lectotype has not been
selected.

Description of male (based on material from
Cape Farewell, Tasmania, NMV J26414).
Cephalon with dorsal median nodule; anterior
margin without nodules; with prominent bifur-
cate rostral point. Pereonite | with obscure
longitudinal grooves on lateral surfaces, other-
wise without tubercles or ornamentation. Per-
eonites 2 and 3 each with 2 transverse rows of
distinct low spikes, pereonites 4-7 each with 2
transverse rows of prominent acute spikes,
anterior row larger than posterior row on 4 and
7, subequal on 5 and 6; coxae 5 and 7 with pos-
terior margins evenly rounded, coxae of pereon-
ite 6 posteriorly straight. Pleon posterior boss
with 2 submedian prominent processes extend-
ing posteriorly over pleotelson, proximally well
separated, pleon otherwise without prominent
spikes. Pleotelson with 2 prominent spikes
opposing those of pleon, posterior margin with
two prominent submedian indentations either
side of median lobe .

209
A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

BESATE
CRT

osa (Baker). d 8.5 mm (NMV 414 iew; B, lateral view; C, frons; D,
Figure 40. Oxinasphaera bisp J26414). A, dorsal view; B, sure
: vx eat y^ .F N a d 7, distal articles; I,
IL ; Ge keta VIRO sciare e , antennule; G, pereopod 1; H, pereopo
antennule terior view; E, (
(YE ; p od apices, lateral view. Scale 2.0 mm.
u n2, pi p

210

N. L. BRUCE

Figure 41. Ovinasphaera bispinosa (Baker). & 8.5 mm (NMV 126414). A, maxilliped; B, pleopod 1; C, pleopod 2;

D, appendix masculina apex; E, penes.

Antennule peduncle article 1 with 5 broadly
rounded anterior spikes; without posterior
spikes; flagellum with 12 articles.

Epistome with 2 prominent narrowly separ-
ated flat distally truncate spikes. Mandible
molar process distal margin deeply indented.

Pereopod | basis about 2.6 times as long as
wide, anterior margin with l sensory setae;
ischium about 0.9 times as long as basis, about
2.7 times as long as wide, anterior margin with 1
short proximal spines and | distal longer and
biserrate spine; merus about 0.4 times as long as
ischium, about 1.2 times as long as wide, ante-

rolateral angle with 2 gently curving pectinate
spines, posterior margin with 4 long setae; car-
pus short, about 0.4 times as long as merus,
about 1.3 times as long as wide, posterior margin
with 2 biserrate spines; propodus about 0.8
times as long as ischium, widest proximally,
about 3.2 times as long as wide, posterior margin
with distinct scale spikes and 5 large biserrate
spines, row of 9 smaller biserrate spines on
medial margin; dactylus about 0.4 length of pro-
podus, unguis about 4796 length of entire dacty-
lus. Pereopod 7 carpus about 2.3 times as long as
wide, posterior margin with 6 biserrate spines, 3

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 211

Figure 42. Oxinasphaera bispinosa (Baker), A-C. non-ovigerous 9, 7.0 mm, D, E ovigerous 9, 8.5 mm. A, dorsal
view: B, lateral view; C, frons; D, dorsal view; E, pleon and pleotelson, lateral view. Scale 2.0 mm.

biserrate spines on posteromedial margin, distal
margin with 2 large trifid spines and 6 biserrate
spines; propodus about 1.2 times as long as car-
pus and about 2.6 times as long as wide, pos-
terior margin with 7 biserrate spines, anterodis-
tal angle with 4 simple and | sensory setae.

Penial processes each about 6.7 times as long
as basal width, tapering to an elongate narrowly
rounded apex, extending to pleopod rami; distal
one third of penial process with plane twisted 90
to proximal part; proximolateral margin with
scale spikes.

Pleopod | exopod with ventral lateral submar-
ginal groove. Pleopod 2 appendix masculina
basal two thirds swollen, distal third narrow, 4.3
times as long as maximum width, apex bluntly

truncated. Uropod dorsally nodular, covered
with roughened setae; exopod about 4.6 times as
long as proximal width, about 0.6 times as long
as endopod, apex deeply bifid with lateral pro-
cess prominent; endopod about 3.7 times as long
as wide, apex with prominent point and, in lat-
eral view, 2 prominent downwardly projecting
spikes.

Ovigerous Female. Cephalon with median dor-
sal tubercle; pereonites 2-7 each with single row
of indistinct low tubercles; pleon smooth, with-
out nodules or domes; pleotelson posterior mar-
gin with wide median excavation.

Non-ovigerous Female. Cephalon with median
dorsal tubercle. Body smooth, finely granular,

212 N. L. BRUCE

Figure 43. Oxinasphaera bispinosa (Baker), SEMs. A-E d 7.8 mm, G, H, d 7.5 mm (NMV J26414). A, frons
(X65): B. spike rows, pereonites 5 and 6 (X 110); C, pleon and pleotelson (X 22); D, pleotelson, tubercles and pits
(7X 400): E. pleotelson apex (X 130); F, molar process (X 300); G, left mandible. incisor, lacinia mobilis and spine
row (7 430); H. right mandible, incisor and spine row (X 500).

$ LI DI
E d \ \ 4 / 4 =
Tw | ^ j Po L
1 i ti VS E o ;
G È 1 i " | B Uu L
, o 1 3 B "(il y fin

era bispinosa (Baker), SEMs. ¢ 7.5 mm (NMV J26414). A, maxillule, lateral lobe (X 500); B,
dite, dorsal surface showing serrate spines (X 430); D, pereopod
d 7, trifid spines on carpus (X 450); F, pleopod 5 (X 150); G,

d 5 (X 3000); uropod, distal ends of rami (X 170).

Figure 44. Oxinaspha
maxillule, medial lobe (X 430); C, maxilliped en
1, unguis, distal spine and setae (X650); E, pereopo

scale from scale lobe, pleopo

214

unornamented. Epistome with rudimentary
undivided flat spike,

Colour. Pale yellow ground colour, usually with-
out chromatophores. Cuticle usually opaque.

Size, Males 5.4-9.5 mm, immature males 6.0-
7.5 mm, females 5.7-8.0 mm, ovigerous females
6.0-8.5 mm.

Remarks. The lack of material from further west
than the east Victorian coast, and the evident
south-eastern distribution of the recent records,
suggests that Baker's data may be incorrect. As
there is no fresh or recent material from the
South Australia coast, the redescription 1s here
based on the large sample of specimens from off
Tasmania's northern coast (NMV J26414).

Oxinasphaera bispinosa is one of the most eas-
ily recognised species of the genus, and is the
only Australian species with prominent pleonal
processes. It is further distinguished by the
elongate penial processes and bottle-shaped
appendix masculina. It shares all of these charac-
ters with Oxinasphaera tripartita from Papua
New Guinea and the Philippines and Oxinas-
phaera kensleyi sp. nov, from South Africa. It
can be separated from those two species by the
different shape of the processes, the regular
antennular spikes and by lacking a large promi-
nent pleotelsonic lobe overriding the pleotel-
sonic apex.

Distribution. All recent material is from around
Tasmania and the Bass Strait, from about 39° to
43'S and westwards to 147°E, at depths between
18 and 85 metres. Substrata recorded are fine
sand, and medium shell. Old material includes
one lot from Spencer Gulf, S.A, collected by
F.I.S. Endevour in 1909-1914, and one sample
taken off the coast between Tathra and Merim-
bula, NSW, in 1925,

Hosts. Some records are from sponges, none
identified.

Etymology. The epithet given by Baker presum-
ably alludes to the prominent pleonal pro-
cesses.

Oxinasphaera tripartita
(Richardson, 1910) comb nov.

Figures 45-47

Cymodoce tripartita Richardson, 1910: 29, fig. 27.

Non Cymodoce tuberculosa tripartita. — Barnard,
1920: 363, pl. 15 fig. 28. — Barnard, 1940: 504, —
Nierstrasz, 1931: 200. — Kensley, 1978: 100, fig. 42C
(= Oxinasphaera kenslevi sp. nov.).

N. L. BRUCE

Material examined. Syntype. d (3.4), Jolo Jolo, Phil-
ippines, 11 Feb 1908, from interior of a pearl oyster
(USNM 40919) [See “Remarks'].

Non-type material. 4d (3.4, 3.3, 3.3, 3.2 mm).
So ( 2.3, 2.6, 2.6, 2.7, 2.8, 2.8 mm), between Kranket
Is and Paenwai Is, Madang, Papua New Guinea,
5°11.2°S, 145^50.9'E, 22 May 1989. 20 m,
from sponge, N.L. Bruce and M. Jebb (ZMUC
CRUI393).

Description of male (based on Madang speci-
mens). Body about twice as long as wide, cuticle
clear. Cephalon anterior half irregularly and
finely nodulose; anterior margin without nod-
ules; with very weak obscure bifurcate rostral
spike. Pereonite 1 unornamented. Pereonites 2
and 3 each with 2 transverse rows of distinct low
spikes, pereonites 4-7 each with 2 transverse
rows of prominent acute spikes, anterior row
larger than posterior row on 4 and 5, posterior
row largest on pereonite 7; coxae 5 and 6 pos-
terior margins straight, those of pereonite 7 pos-
teriorly rounded. Pleon posterior boss, with 2
submedian prominent processes extending pos-
teriorly over pleotelson, medial separating mar-
gin concave. Pleotelson dorsal surfaces con-
spicuously granular; with 2 prominent spikes
opposing those of pleon; lateral flange with 1
prominent spike; posterior margin with promi-
nent posteriorly rounded lobe overlying telsonic
excision and extending posterior to submedian
indentations.

Antennule peduncle article 1 with 7-8 irregu-
lar anterior spikes and posteromedial blade; fla-
gellum with 4 articles.

Epistome without distinct spikes, with indis-
tinct transverse blade armed with irregular small
tubercles.

Pereopod 1 basis about 3 times as long as
wide, anterior margin with 2 sensory setae;
ischium 0.7 times as long as basis, 2.6 times as
long as wide, anterior margin with | short proxi-
mal spines and 1 distal longer and biserrate
spine; merus about 0.4 times as long as ischium,
about 1.3 times as long as wide, anterolateral
angle with 1 gently curving pectinate spine, pos-
terior margin with l long seta; carpus short,
about 0.8 timesas long as merus, about 1.3 times
as long as wide, posterior margin with 2 biserrate
spines; propodus about 0.8 times as long as
ischium, widest proximally, about 2.8 times as
long as wide, posterior margin with distinct scale
spikes and 4 large biserrate spines, row of 3
smaller biserrate spines on medial margin; dac-
tylus about 0.5 length of propodus, unguis about

Figure 45. Oxinasphaera tripartita (Richardson). A-E d 3.3 mm, remainder ¢ 3.2 mm (ZMUC CRU1393). A,
dorsal view: B, lateral view; C, frons; D, antennules, anterior view, in situ; E, pleon, ventral view; F, antennule: G,
antenna; H. maxilliped; I, pereopod 1; J, pereopod 2, distal articles: K. pereopod 7, distal articles; L. penes. Scale

1.0 mm.

216

N. L. BRUCE

Figure 46. Oxinasphaera tripartita (Richardson). A-C 7 3.2 mm, D, E non-ovigerous 9 2.8 mm (ZMUC
CRU1393). A, pleopod 1; B. pleopod 2; C, uropod; D, lateral view; E, dorsal view.

5296 length of entire dactylus. Pereopod 7 carpus
about 2.3 times as long as wide, posterior margin
with 4 biserrate spines and 2 weakly biserrate
spines, distal margin with 2 large trifid spines
and 5 simple or very weakly biserrate spines;
propodus 1.3 times as long as carpus and 3.1
times as long as wide, posterior margin with 3
spines, anterodistal angle with 2 simple and 1
sensory setae.

Penial processes each about 7.5 times as long
as basal width, tapering to an elongate slender
apex, extending to pleopod rami; proximolateral
margin with scale spikes, distal margin with
scale setules.

Pleopod l exopod with ventral lateral submar-
ginal groove. Pleopod 2 appendix masculina
basal four-fifths swollen, distal fifth narrow,
apex obliquely truncated, 7.3 times as long as
maximum width. Uropod not distinctly nodu-
lar, setae smooth; exopod 4.2 times as long as
proximal width, about half as long as endopod,
apex deeply bifid with lateral process promi-
nent; endopod 3.2 times as long as wide, apex
with prominent point and 1 prominent down-
wardly projecting spike.

Female. Pleon and pleotelson without distinct
domes; posterior margin of pleotelson with api-
cal triangular protrusion.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 217

Figure 47. Oxinasphaera tripartita (Richardson). SEMs. 7 3.3 mm (ZMUC CRU 1393). A, frons (X 130); B, pleon
and pleotelson (X 55); C, pleonal and pleotelson spikes, lateral view (X 190); D, pleotelson apex (X100); E,

uropod rami (X 130).

Colour. Pale yellow ground colour, with abun-
dant dark brown chromatophores. Cuticle
usually clear.

Size. Males 3.2-3.4 mm, females 2.3-2.8 mm.

Remarks. Richardson (1910) recorded two male
and two female syntypes. The present type
material consists only of one male. This male is
labelled lectotype, but I am unaware of any pub-
lication ofa lectotype designation, and the speci-
men is here regarded as a syntype. Richardson
recorded this specimen as having been collected
from the interior of a pearl oyster, but the vial in
which the specimen is contained was found, on

examination, to be full of sponge spicules,
suggesting that the actual habitat is in
sponges.

Two species of Oxinasphaera, O. tripartita
and O. kensleyi have a posterodorsal lobe on the
posterior margin of the pleotelson in conjunc-
tion with antennular peduncle article 1 having
an irregular row of anterior spikes and a poste-
romedial blade. Both species have in addition
prominent posteriorly directed processes on the
pleon, a character shared with the otherwise dis-
similar O. bispinosa. O. tripartita can be dis-
tinguished from O. kensleyi by having pereon-
ites 2 and 3 weakly ornamented, the pereonal

218

spikes of all pereonites being less strongly devel-
oped than those of O. kensleyi, by having weaker
antennular spikes, a wider posterior antennular
blade, and the penes and appendix masculina
being straight rather than sinuate.

Distribution. Jolo, Philippines and Madang,
Papua New Guinea.

Hosts. The specimens from Madang, were col-
lected directly from a sponge, probably a species
of Cribrochalina.

Etymology. The epithet given by Richardson
presumably alludes to the prominent pleonal
processes.

Oxinasphaera kensleyi sp. nov.
Figures 48, 49

Cymodoce tuberculosa tripartita. — Barnard, 1920:
363. pl. 15 fig. 28. — Barnard, 1940: 504, — Nier-
strasz, 1931: 200. — Kensley, 1978:100, fig. 42C.

Cymodoce tubereulosa-Kensley, 1984: 216.

Non Cymodoce tuberculosa Stebbing, 1873: 96, pl. 3
figs 1, Ib.

Non Cymodoce tripartita Richardson, 1910:29, fig,
27.

Material examined. Holotype. d (4.3 mm), off East
London, South Africa, 3304.6'S, 28°06.6’E, 26 May
1978. 90 m, stn 163, R.V. Meiring Naude (SAfM
A41309),

Paratypes. 56 (3.9, 4.0 [dissected], 4.1, 4.2, 4.5 mm),
same data as holotype (SAfM A19308).

Description of male. Cephalon anterior half
irregularly and finely nodulose, with low median
tubercle; anterior margin with row of small
close-set nodules; with weak obscure bifurcate
rostral spike. Pereonite 1 with anterior submar-
ginal transverse row of small tubercles. Pereon-
ites 2 -7 each with 2 transverse rows of acute
spikes, those of pereonites 2-4 being somewhat
rounded, anterior row larger than posterior row;
coxae 5-7 posterior margins rounded. Pleon
posterior boss with 2 submedian prominent pro-
cesses extending posteriorly over pleotelson,
each process with prominent proximal ventral
spike, medial separating margin straight. Pleo-
telson dorsal surfaces conspicuously granular;
with 2 prominent bluntly rounded spikes oppos-
ing those of pleon; lateral flange weakly devel-
oped, with 1 prominent spike; posterior margin
with prominent posteriorly acute lobe overlying
telsonic excision and extending posterior to sub-
median indentations.

Antennule peduncle article | with 5 irregular
anterior spikes, distal most being most promi-

N. L. BRUCE

nent, with distally truncate posteromedial blade;
flagellum with 6 articles.

Epistome without distinct spikes, with indis-
tinct transverse blade armed with irregular small
tubercles.

Pereopod generally similar to those of O. tri-
partita.

Penial processes 7.2 times as long as basal
width, tapering to an elongate slender acute
apex, extending to pleopod rami; proximolateral
margin with scale spikes, distal medial margin
with scale setules.

Pleopod | exopod with ventral lateral submar-
ginal groove. Pleopod 2 appendix masculina
sinuate, tapering to narrow bluntly rounded
apex, 6.8 times as long as maximum width. Uro-
pod nodular, with abundant setae; exopod 4.2
times as long as proximal width, about half as
long as endopod, apex deeply bifid with lateral
process prominent; endopod 3.5 times as long as
wide, apex with prominent terminal spike.

Female. No females examined.

Colour. Pale yellow ground colour, chromato-
phores not apparent in preserved specimens.

Size. Males 3.2-3.4 mm, females 2.3-2.8 mm.

Remarks. This is currently the only species ofthe
genus known from the Western Indian Ocean,
and cannot be confused with any other sphaero-
matids in the region (but see undescribed
species). The only similar species is Oxinas-
phaera tripartita, a species known from Papua
New Guinea and the Philippines, and O. kens-
leyi can be separated from that species by the far
more prominent pereonal spikes and the sinuate
penes and appendix masculina. Other differ-
ences are given in the ‘Remarks’ for Oxinas-
phaera tripartita.

The material examined here is that of Kensley
(1984) which was incorrectly attributed to
Richardson's species.

Distribution. Indian Ocean, off Durban [c.
30°S](Barnard 1920) and East London, South
Africa; from c. 65 to 90 m.

Hosts. Barnard (1920) reported on the speci-
mens that he examined: “in each case from
siliceous sponges."

Etymology. The name is in recognition of Dr
Brian Kensley's contributions to knowledge of
the isopods and Crustacea of South Africa.

Oxinasphaera australis Baker, comb. nov.
Figures 50, 51
Cymodoce multidens var. australis Baker, 1929: 52,

pl. 1 figs 4-6 [Non Oxinasphaera multidens (Richard-
son, 1910)].

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 219

SP SCOR BO DOG Swen
o
o

E
ZN)

U RAB

EM — 7 > E

CO Togo OS USAS,

e EL BI e So GIA
IU

i
HA
wih

Figure 48. Oxinasphaera kensleyi sp. nov. A-D, holotype, E, d 4.2 mm, F-G, d 4.0 mm (SAfM A19308). A, lateral
view: B, dorsal view (in two parts); C, frons; D, antennules, anterior view; E, pleotelson apex; F, antennule; G
maxilliped; H, penes. Scale 1.0 mm. d

N. L. BRUCE

Figure 49. Oxinasphaera kensleyi sp. nov. d 4.0 mm (SAfM A19308). A, pleopod 1; B, pleopod 2; C, uro-
pod.

Material examined. Syntypes. 69 (5.7-6.6 mm, mean
= 6.1 mm), 15 9 (ovig 8.4 [in 2 pieces], 10.2, non-ovig
7.5-9.3 mm, mean = 8.4 mm), Cottesloe, Perth, WA,
no other data, L.G. Glauert (WAM 10385, 10484).

Description of male. Body about 2.2 times as
long as wide. Cephalon anterior margin with 8-9
nodules, anterior median and 2 submedian nod-
ules being prominent and elongate giving a tri-
cornate appearance; with very weak obscure
simple rostral spike. Pereonite l unornamented.
Pereonites 2 -7 granular, each with 2 transverse
rows of small low spikes, pereonite 1 with dis-
tinct anteromedial tubercle; coxae 5 and 6 pos-
terior margins convex, with distinct posterior
join to ventral margin, those of pereonite 7
entirely rounded posteriorly. Pleon without pos-
terior boss, with 2 submedian clusters of
tubercles; lateral flange with row of 3-5 spikes.
Pleotelson dorsal surfaces granular; without
prominent spikes; posterior margin somewhat
flattened and produced, with 2 submedian tri-
angular excisions.

Antennule peduncle article 1 with 8-9 irregu-
lar anterior spikes and 2 posteromedial spikes.

Epistome with 3 small distinct spikes, surface
with numerous small granular tubercles.

Pereopod 1 propodus about 2.6 times as long
as wide, posterior margin with distinct scale

spikes and 6 large biserrate spines, row of 5
smaller biserrate spines on medial margin; dac-
tylus about 0.4 length of propodus, unguis about
50% length of entire dactylus.

Penial processes examined in situ, generally
similar to O. bisubula and O. copiapoa.

Pleopod 2 appendix masculina about as long
as endopod, 6.4 times as long as maximum
width, extending beyond endopod by 0.2 of its
length, distally acuminate, recurved apex acute.
Uropod not distinctly nodular, ventral margins
of both rami with acute prominent downwardly
projecting spikes; uropod exopod apex with two
spikes, appearing bifid.

Female. Retains male pattern of ornamentation
in reduced form; epistome spike more distinctly
bifid. Pereonite 1 with low rounded submedian
subanterior tubercles. Pleotelson posterior mar-
gin with shallow submedian excisions.

Colour. Pale yellow in the faded preserved speci-
mens.

Size. Males 5.7-6.6 mm, ovigerous females 8.4-
10.2 mm, non-ovigerous females 7.5-9.3 mm.

Remarks. There is no new material available for
O. australis. The male syntypic specimens
appear to have dried out at some time, and are in
a fragile and brittle condition, with the pleopods

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

An
DEAS X
"E Cet, 006809

221

Figure 50. Oxinasphaera australis (Baker). A-E syntype d 6.3 mm (WAM 10385/10484), remainder as indicated.
A, dorsal view; B. lateral view; C, antennules, anterior view, in situ; D, frons; E, cephalon, dorsal view; F, pleon,
ventral view; G, penes, in situ, € 6.3 mm; H, pleopod 2, drawn from Baker's slide. Scale 1.0 mm.

and pereopods immovable. In none of the males
were the penes clearly visible, and all specimens
were too fragile to dissect. Oxinasphaera aus-
tralis belongs to a group of species characterised
by having the posterior margin of the pleotelson
somewhat produced and flattened, and dis-
tinctly trilobate. The somatic morphology
allows males and females of the species to easily
distinguished from others of the genus. In par-
ticular the male of O. australis has prominent
single rostral point flanked by two prominent
tubercles on either side of the anterior margin of
the cephalon, giving it a tricornate appearance;
additionally there is a prominent median
tubercle on the anterior of pereonite 2; the epis-
tome anterior process is trilobate; antennule

peduncle article | has 8-10 teeth on the anterior
margin and 2 teeth on the posteromedial mar-
gin. All syntypic males showed this character,
non-ovigerous females have smaller and more
numerous (13) teeth on the antennule peduncle
article 1 anterior margin and 3 on the posterom-
edial margin.

This species (and the other related species)
also differ from the remainder ofthe genus in the
females being manifestly larger than the
males.

Distribution. Known only from the type locality,
Cottesloe, Perth, WA.

Hosts. Not known

N. L. BRUCE

Figure 51. Oxinasphaera australis (Baker). A, E, F syntype, ovigerous 9 10.2 mm; B-D syntype, non-ovigerous 9
9.0 mm. A, dorsal view; B, lateral view; C, cephalon, dorsal view; D, frons; E, pleotelson posterior margin; F,
pleon and pleotelson, lateral view; G, pereopod 1, non-ovigerous 9 drawn from Baker's slide. Scale 1.0 mm.

Oxinasphaera matucana sp. nov.
Figures 52, 53

Material examined. Holotype. 8 (5.6 mm), Pearson Is.,
Investigator Group, SA, 9 Jan 1969, 50 m, coarse
gravel, S.A. Shepherd (SAM C5604).

Paratypes. 9 (4.6 mm), NE side of Topgallant Is.,
Investigator Group, SA, 33°43.0’S, 134936.6/E,21 Apr
1965, 12 m, algae, bryozoa, sponges, S.A. Shepherd
and G.C.B. Poore (NMV J40487). o (5.0 mm), manca
(2.8 mm), Bastion Point, Mallacoota, Vic., 37°34.3’S,
149°46.2’E, 6 Apr 1989, reef 300 m offshore, 5 m,

sponges, G.C.B.

140483).

Description of male. Body about 2.2 times as
long as greatest width; lateral margins subparal-
lel. Cephalon dorsal surface finely pitted;
anterior margin without tubercles; rostrum with
bifurcate spike. Pereonite 1 with 2 distinct sub-
median rounded spikes posterior to anterior
margin. Pereonites 2-7 each with 2 transverse
rows of spikes, anterior row on pereonite 2
prominent with medial 5 being distinctly larger

Poore and R. Wilson (NMV

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 223

Figure 52. Oxinasphaera matucana sp. nov. A-D, F holotype, remainder 7 4.6 mm (NMV 740487). A, dorsal
view; B, lateral view; C, cephalon, dorsal view; D, antennules, anterior view; E, antennules, anterior view; F,
frons; G, antennule; H, pereopod 1; I, pereopod 7. Scale 1.0 mm.

[SS]
to
pay

N. L. BRUCE

Figure 53. Oxinasphaera matucana sp. nov. d 4.6 mm (NMV J40487). A, pleopod 1; B, uropod.

than all other pereonal spikes; pereonite 3 with
anterior row larger than posterior but not as
prominent as that of pereonite 2; pereonites 4-7
rows of about equal prominence; coxae 5-7 with
posterior margins rounded. Pleon without pos-
terior boss, with 2 longitudinally oriented sub-
median tubercles, pleonite 3 with 2 sublateral
tubercles. Pleotelson without spikes or promi-
nent tubercles; posterolateral flange with 3 mar-
ginal acute tubercles; posterior margin flattened,
somewhat produced, with two submedian tri-
angular excisions.

Antennule peduncle article 1 with 10 anterior
spikes; with 1 long posteroproximal and 1 pos-
terodistal spike; dorsal surface of peduncular
articles 1 and 2 provided with roughened setae;
flagellum with 9 articles.

Epistome without distinct spikes, with 4—5
teeth set on transverse ridge; lateral lobes each
with 2 distinct spikes.

Pereopod 1 basis about 2.5 times as long as
wide, anterior margin with 1 sensory seta;
ischium 0.7 times as long as basis, 2.1 times as
long as wide, anterior margin with | short proxi-
mal spine and 1 distal longer feebly biserrate
spine; merus about one third as long as ischium,
0.7 times as long as wide, anterolateral angle
with 1 gently curving weakly pectinate spine,
posterior margin with 1 spine and single long
simple seta; carpus about as long as merus, about
as long as wide, posterior margin with 2 biserrate
spines; propodus about equal in length (0.92) to
ischium, widest proximally, about 2.4 times as
long as wide, posterior margin with distinct scale
spikes and 4 large biserrate spines, row of 5
smaller biserrate spines on medial margin; dac-
tylus 0.5 length of propodus, unguis about 53%
length of entire dactylus. Pereopod 7 basis 3.9
times as long as wide, anterior margin with 7
sensory setae; ischium slightly shorter (0.8) than

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

basis, 3.6 times as long as wide; merus about half
as long as ischium, 1.8 times as long as wide,
posterior margin with 3 simple spines, anterod-
istal angle with 1 large and 2 small spine; carpus
1.3 times as long as merus, 2.6 times as long as
wide, posterior margin with 5 biserrate and 3
simple spines, distal margin with 3 large trifid
spines, longest equal in length to propodus and 5
spines at anterodistal angle; propodus about 0.9
times as long as carpus and 0.5 time as long as
ischium, about 3.0 times as long as wide, pos-
terior margin with 3 biserrate spines, anterodis-
tal angle with 2 sensory setae.

Pleopods conforming to the genus; appendix
masculina not observed. Uropod dorsally nodu-
lar, covered with roughened setae; exopod about
4 times as long as proximal width, about half as
long as endopod, apex deeply bifid with pro-
cesses about equally prominent, ventral margin
with 3-4 prominent acute serrations; endopod
about 3 times as long as wide, apex prominently
bifid, ventral margin with 4-5 prominent acute
serrations.

Female. Unknown.
Colour. Pale tan in alcohol.
Size. Males 4.6-5.6 mm, females 3.5-4.5 mm.

Remarks. Of the males examined here only the
holotype had partly developed penial processes
and a developing appendix masculina, not
detached from the endopod of pleopod 2, and
therefore all three specimens are regarded as
immature. However, all specimens had clearly
developed dorsal sculpting, and the pleotelson
and uropods are of the adult male form. The
ornamentation of the antennule, pereonite 1 and
2, and pleotelson and uropods clearly separates
Oxinasphaera matucana from other species of
the O. australis group and the remaining species
of the genus, and a new species is therefore estab-
lished.

Distribution. Mallacoota, Victoria, westwards to
the Investigator Group, South Australia, at
depths between 5 and 50 m.

Hosts. One specimen recorded from an
unidentified sponge.

Oxinasphaera poorei sp. nov.
Figures 54, 55

Material examined. Holotype. ¢ (2.9 mm), Bastion
Point, Mallacoota, Vic., 37°34.3’S, 149°46.2’E, 6 Apr
1989. reef 300 m offshore, 5 m, sponges, G.C.B. Poore
and R.S. Wilson (NMV J40488).

225

Paratypes. 2¢ (2.7, 3.3 mm), same data as holotype
(NMV J40498).

Description of male. Cephalon irregularly pitted;
with prominent flattened indistinctly bifurcate
rostral point; anterior margin without spikes,
with single prominent anteriorly projecting
flange. Pereonite 1 without tubercles or orna-
mentation. Pereonites 2-6 each with anterior
transverse row of distinctly flattened rounded
spikes, pereonite 7 spikes not markedly flat-
tened; posterior rows of pereonal spikes all
weakly developed; coxae 5-7 with posterior
margins nearly straight, ventral margin subtrun-
cate, that of coxa 6 weakly concave. Pleon with-
out posterior boss, although posterior margin
weakly produced; pleonites 3 and 4 each with
large sublateral tubercle, otherwise without
prominent spikes. Pleotelson generally nodular,
with 4 prominent rounded tubercles across
anterior, posterior margin with two prominent
submedian indentations either side of median
lobe .

Antennule peduncle article 1 with 3 wide
rounded anterior spikes, and large proximal
length set just posterior to front row; without
posterior spikes; flagellum with 6 articles.

Epistome with 2 basally united, distally acute,
flat spikes.

Pereopod | basis about 2.6 times as long as
wide, anterior margin with 2 sensory setae, dis-
tally with scale spikes; ischium about 0.6 times
as long as basis, about 2.3 times as long as wide,
anterior margin with | short proximal and 1 dis-
tal longer and biserrate spines; merus 0.5 times
as long as ischium, about as long as wide, ante-
rolateral angle with 1 gently curving pectinate
spine, posterior margin 1 biserrate spine and 1
long seta; carpus as long as merus, about 1.5
times as long as wide, posterior margin with 2
biserrate spines; propodus about (1.1) as long as
ischium, widest proximally, about 2.5 times as
long as wide, posterior margin with 2 large biser-
rate spines; dactylus about 0.6 length of propo-
dus, unguis about 5296 length of entire dactylus.
Pereopod 2 similar to others, merus distal mar-
gin with 4 biserrate spines. Pereopod 7 basis 3.6
times as long as wide, anterior margin with scale
spikes, with prominent seta at anterodistal
angle; ischium 0.68 as long as basis, 2.8 times as
long as wide; merus about half as long as
ischium, 1.6 times as long as wide, posterior
margin with 3 feebly biserrate spines, 2 biserrate
spines at anterodistal angle; carpus slightly
longer (1.1) than merus, 22 times as long as
wide, posterior margin with 4 large biserrate

226 N. L. BRUCE

Figure 54. Oxinasphaera poorei sp. nov. A-D, holotype, remainder d 2.7 mm (NMV J40487). A, dorsal view (two
parts); B. lateral view; C, antennules, anterior view, in situ; D, frons; E, antennule; F, pereopod 1; G, pereopod 2;
H, pereopod 7; I, penes. Scale 0.5 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

227

Figure 55. Oxinasphaera poorei sp. nov. d 2.7 mm (NMV J40487). A, pleopod 1; B, pleopod 2; C, uropod.

spines, distal margin with 2 trifid and 3 biserrate
spines, anterodistal angle with single weakly
biserrate spine; propodus about 1.2 times aslong
as carpus and about 2.6 times as long as wide,
posterior margin with 3 biserrate spines, ante-
rodistal angle with 2 simple and 1 sensory
setae.

Penial processes each about 2.8 times as long
as basal width, medial margin straight, lateral
curving to subacute apex; proximolateral mar-
gin with scale spikes.

Pleopod l endopod medial PMS inserted dor-
sally. Pleopod 2 appendix masculina stout, 4.0
times as long as maximum width, about as long
as endopod, extending beyond endopod by 0.2
of its length, apex bluntly truncated. Uropod
dorsally nodular; exopod about 3.3 times as long
as proximal width, about 0.4 times as long as
endopod, apex deeply bifid with lateral process
prominent; endopod about 3.5 times as long as
wide, apex with 2 prominent spikes, additional
rounded nodules along ventral margin.

Female. Unknown.
Colour. Pale yellow in alcohol.
Size. Males 2.7-3.3 mm.

Remarks. This species is unique within the
genus in possessing flattened pereonal spikes,
and in having the anterior margin of the cepha-

lon with a distinct hardened ridge. The anten-
nule, with only 4 flattened spikes, the lack of
pleonal boss and simple posterior notches to the
posterior margin of the pleotelson all further
serve to identify the species.

Distribution. Known only from the type

locality.
Hosts. Unidentified sponge.

Etymology. Named in recognition of Gary C.B.
Poore's contribution to the knowledge of Aus-
tralian isopod and crustacean fauna.

Oxinasphaera islaya sp. nov.
Figures 56-58

Material examined. Holotype. d (2.8 mm), 2 km S of
Cape Peron, WA, 32°16’S, 115°41’E, 26 Dec 1983, 6
m, from sponges, cave in reef, J.K, Lowry (AM
P44197).

Paratypes. 160 (2.0-2.5 mm, mean= 2.3 mm), ll
mancas (2.1, 2.0, 1.9, 1.8, 1.7 mm and 6 1.2-1.8 mm),
same data as holotype (AM P41118, slides P44215,
P44216; ZMUC CRUI386).

Additional material. 53 (2.2 imm, 2.3, 3.0, 3.0, 3.5
mm), “The Hotspot" reef, 5 nm W of N end of Flinders
Is., SA, 33°40.5’S, 134°22.0’E, 19 Apr 1985, 12 m,
assorted algae, S. Shepherd (NMV J40484).

Description of male. Body about 2.3 times as
long as greatest width; lateral margins subparal-
lel; dorsal surfaces polished, anteriorly with

228 N. L. BRUCE

uj a Yo;
i
j

La,
SU

Figure 56. Oxinasphaera islaya sp. nov. A-D, F holotype, remainder d 2.1 mm (AM P41118). A, dorsal view; B.
lateral view; C, frons; D, antennules, anterior view, in situ; E, posterior margin of pleon; F, pleon, ventral view; G,
antennule; H, antenna; I, maxilliped; J, penes. Scale 1.0 mm.

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE) 229

Figure 57. Oxinasphaera islaya sp. nov. H, F, ¢ 2.0 mm, remainder d 2.1 mm (AM P41118). A, pereopod 1; B,
pereopod 7; C-G, pleopods 1-5 respectively; H, appendix masculina apex; I, uropod.

N. L. BRUCE

Figure 58. Oxinasphaera islaya sp. nov. SEMs. d 2.7 mm (AM P41118). A, cephalon, anterior view (X 100); B,
cephalon anterior margin (X 150); C, pleon and pleotelson (X 75); D, detail, pleotelson tubercles (X550); E,

pereopod dactylus (X 1100); F, uropods (X 190).

large pits, otherwise sparsely setose. Cephalon
anterior margin with 3-4 small lateral tubercles;
rostrum spike simple. Pereonite 1 with smooth
low nodules, large shallow pits. Perconites 2-4
with only anterior spike row distinct, pereonites
5-7 each with 2 transverse rows of spikes,
anterior row large, posterior row indistinct; all
spikes rounded; coxae 5 with posterior margins
evenly rounded, coxae 6 ventrally subacute,
coxae 7 rounded. Pleon with weak posterior
boss, posterolateral angles of boss each with 2
rounded spikes; lateral margin of pleonite 4 with

2 prominent rounded tubercles. Pleotelson with
2 rounded spikes opposing those of pleon, other-
wise surface irregularly nodulose; posterolateral
flange with 1 submarginal and 3-4 rounded
tubercles; posterior margin appearing serrate,
with 2 weak submedian indentations either side
of median lobe .

Antennule peduncle article 1 with 7 anterior
spikes; with 1 long proximo-posterior spike and
l short posterior spike; dorsal surface of pedun-
cular articles | and 2 provided with few setae;
flagellum with 4 articles. Antenna peduncle

A NEW GENUS OF MARINE ISOPOD (SPHAEROMATIDAE)

articles 4 and 5 with few long setae; flagellum of
about 6 articles.

Epistome with 2 prominent widely separated
conical spikes, lateral lobes each with 1 small
tubercle.

Pereopod 1 basis 2.9 times as long as wide,
anterior margin with 2 sensory setae, few scale
spikes, anterodistal angle with prominent single
seta; ischium 0.7 times as long as basis, 2.5 times
as long as wide, anterior margin with 1 short
proximal spine; merus about half as long as
ischium, about as long as wide, anterodistal
angle with 2 stout curving pectinate spines, pos-
terior margin with single long simple seta; car-
pus about as long as merus, 1.4 times as long as
wide, posterior margin with l biserrate spine
and short simple seta; propodus about equal in
length to ischium, widest proximally, about 2.6
times as long as wide, posterior margin irregu-
larly notched and 3 large biserrate spines, 2
smaller biserrate spines on medial margin; dac-
tylus about half length of propodus, unguis
about 51% length of entire dactylus. Pereopods 2
and 3 similarto 1, differing principally in having
the carpus more elongate and propodus more
slender, and having additional biserrate seta on
distal margin of carpus. Pereopod 7 ischium 3.1
times as long as wide; merus 0.5 times as long as
ischium, 1.6 times as long as wide, anterodistal
angle with 1 large spine, posterodistal angle with
2 simple setae; carpus 1.2 times as long as merus,
2.5 times as long as wide, posterodistal angle
with 2 biserrate and 1 large trifid spines, ante-
rodistal angle with 2 biserrate and 2 simple
spines; propodus about 1.2 times as long as car-
pus and 0.8 time as long as ischium, about 3.9
times as long as wide, posterodistal angle with 1
biserrate spine, anterodistal angle with 1 sensory
and 1 simple setae.

Penial processes each about 3.7 times as long
as basal width, tapering slightly to a subacute
apex; proximolateral margin with scale spikes.

Pleopod 2 endopod appendix masculina
straight, 5.8 times as long as maximum width,
apex rounded and narrowed, longer (1.3) than
endopod, extending beyond endopod by 0.35 of
its length. Pleopod 4 exopod and exopod with
single seta at distomedial angle, endopod with-
out thickened fleshy ridges. Pleopod 5 exopod
with 1 apical lateral scale lobe 1 medial apical
scale lobe, endopod without thickened fleshy
ridges. Uropod dorsally finely nodular; exopod
about 3.8 times as long as proximal width, about
two-thirds as long as endopod, apex not dis-
tinctly bifid, with lateral process being distal
part of series of prominent ventral serrations;

231

endopod about 3.6 times as long as wide, apex
with row of 4 terminal prominent spikes.

Female. Unknown.
Colour. Pale brown to dark brown.
Size. Males 2.0-3.5 mm, mancas 1.2-2.1 mm.

Variation. The posterior margin of the pleotel-
son in some specimens shows the typical trilo-
bate or bi-excavate appearance that is common
to most species of the genus. In some specimens
this character was very unclearly expressed.

Remarks. This species is the smallest of the
genus, the nearly entire pleotelson posterior
margin enabling easy recognition. The anterior
dorsal surfaces are pitted but, with the polished
cuticle, this does not give a roughened appear-
ance. Other distinguishing characters are the
relatively long appendix masculina, the lack of
ridges on pleopods 4 and 5 and the distinctly saw
edged appearance of the uropodal exopod.

Distribution. Cape Peron, southern Western
Australia and Flinders Is., South Australia, 6 to
12 metres depth.

Hosts. Type series taken from an unidentified
sponge.

Undescribed Species
Oxinasphaera sp. 1. (Fig. 59)

Material examined. 3 d/o (4.4, 4.5, 4.6 mm), 4 o (ovig
6.0, non-ovig 4.6, 4.7, 5.0 mm), 2 km S of Cape Peron,
WA, 32°16’S, 115941'E, 6 m, from sponges, gorgona-
ceans from cave in reef, J.K. Lowry (AM P44198).

Remarks. Among the specimens from Western
Australia is a species of Oxinapshaera that could
not be assigned to any of the species in the pres-
ent work. It is most similar to O. australis, from
which it differs in lacking a prominent median
tubercle on pereonite 2, having the posterior
margin of the pleotelson entire in intersex and
female specimens, and in the deeply serrated
uropods. It is included here in order to draw
attention to this species and avoid its confusion
with O. australis, also known only from Western
Australia.

There are no male specimens in this series.
Those marked as "c/o" lack penes, appendix
masculina and show a degree of ornamentation
that is not normally found in females. Those
identified as females are unornamented and, as
in O. australis, are larger than the males. The
cephalic, antennular and epistome ornamen-
tation of the intersex specimens and the orna-
mentation of the uropods and posterior margin

Figure 59. Ovinasphacra sp. | (AM P41118).

of the pleotelson precludes assignment to any of
the species treated here. Specimens that are
clearly recognizable as mature are needed before
the species can be named and fully described.

Oxinasphaera sp. 2. 18, western Bass Strait,
39*16.7'S, 143°06.7’E. 95 m (NMV J40478).
Senescent male, without clearly developed
antennule spikes; differs from other species in
having a transverse row of 4 spikes across the
pleon.

Oxinasphaera sp. 3. 19, Burrewarra Point,
NSW, 35°50’S, 150°14’S, 17 m (NMV J26234).
Heavily granular pleon, one of the O. bisubula
group.

Oxinasphaera sp. 4. 19, Spiky Bridge Coastal
Reserve, Tasmania, 42°08’S, 148°08’E (NMV
140485). Antennule with 5 spikes, dorsal per-
conal tubercles prominent and rounded, cepha-
lon anterior margin with 2-3 flattened teeth;
pleotelson similar to the O. bisubula group.
Although distinctive, more males are needed to
confirm that the cephalic spike development is
typical.

Oxinasphaera spp. 5-7. Northern Indian
Ocean, USNM. These sublittoral species, cur-
rently under study by Brian Kensley and Mari-
lyn Schotte, all have prominent pleonal pro-
cesses, and show a diverse pereonal spike,
pleotelson, antennule, epistome and uropod
morphology. They are noted here as these speci-
mens clearly demonstrate the presence of the
genus in the western and northern Indian Ocean
(Fig. 2).

Acknowledgements

This contribution was funded by a grant from
the Australian Biological Resources Study
(ABRS 89/1844)

N. L. BRUCE

I thank all those who assisted when visiting
their institutions and who lent the specimens
used in this study: Dr Penny B. Berents (AM), Dr
A.J. Bruce and Ms Karen Coombes (NTM), Mr
Peter J.F. Davie and Mr. John Short (QM), Dr
Gary C. B. Poore and Dr Robin S. Wilson
(NMV), Mr Wolfgang Zeidler (SAM), Dr Brian
Kensley and Marilyn Schotte (USNM, Smith-
sonian Institution), Ms Melissa Hewitt (WAM),
and Ms Liz Hoensen (SAfM). I also thank Mr
Jorgen Olesen (ZMUC) for his careful inking
and assistance with running the PAUP pro-
gramme, and Dr John N. A. Hooper (QM) for
confirming and correcting the nomenclature of
the sponge identifications.

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N. L. BRUCE
Appendix 1

Character Matrix. Character state and distribution of the 31 character used
in the analysis (? = character not present; — = state unknown).

0 0 l l 2 2
Characters 12345 67890 12345 67890 12345 67890
Cymodoce 00000 00010 00010 00000 00000 01000
Sphaeroma 00000 00000 0000? 22000 00000 00000
bisubula 12000 11011 11010 00001 00000 00000
parodia 02000 11011 11010 00001 00100 00001
lobivia 12000 11011 01010 00001 00000 00000
frailea 12001 11011 01010 00001 00000 00001
denmoza 02000 11111 11010 00001 00000 00000
copiapoa 02001 11011 11010 — 00001 00000 00001
multidens 00000 — 00111 11010 00001 00100 01000
corypantha | 00000 — 00111 11010 00001 00100 00100
obregonia 11000 11001 01010 00001 00000 01000

tual 11000 — OIIII 11010 00001 00100 OI1II
aylostera 02010 11000 10010 11101 00000 00000
rebutia 00111 11000 — 00010 11101 00000 00000

tuberculosa 02000 10000 00020 "71010 00000 10001
epostoa 00000 11000 00020 "71010 00000 10001
lowryi 00000 10011 11020 71010 00001 10000
thetisae 00001 11000 10020 — ?0010 10001 10000
bispinosa 02000 11020 01010 00010 00000 T1111
tripartita 02000 10020 110?1 22000 1101? 21111
kensleyi 02001 11020 11021 79000 1101? ?1100
australis 10000 | 00000 10110 00000 0001? ?0000
matucana 00101 10000 10110 00000 0001? NON

poorei 12001 11000 00010 00010 00000 00000
islaya 11101 11000 10010 00001 00000 00000

Memoirs of the Museum of Victoria 56(1):235-244 (1997)

SPERMATOLOGICAL EVIDENCE SUPPORTS THE TAXONOMIC PLACEMENT

OF THE AUSTRALIAN ENDEMIC HAIRY STONE CRAB, LOMIS HIRTA
(DECAPODA: ANOMURA: LOMIDAE)

CHRISTOPHER C. TUDGE

Zoology Department, University of Queensland, Brisbane, Qld 4072, Australia

Present address: Crustacea, Museum of Victoria, 71 Victoria Crescent. Abbotsford, Vic. 3067, Australia

Abstract

Tudge, C.C., 1997. Spermatological evidence supports the taxonomic placement of the
Australian endemic hairy stone crab, Lomis hirta (Decapoda: Anomura: Lomidac).
Memoirs of the Museum of Victoria 56(1y; 235-244.

An ultrastructural investigation of the spermatozoal morphology of the Australian
endemic hairy stone crab, Lomis hirta, has revealed a unique sperm morphology supporting
placement within its own family and superfamily. The spermatophore and spermatozoal
morphology agree with morphological evidence that place this crab within the infraorder
Anomura. the relatively unmodified spermatozoon suggests a basal origin for this taxon
(with respect to other anomurans) and may be similar to the ancestral sperm type for many
families in this infraorder. An independent lineage from the base of the Anomura has been
previously suggested for Loris on the basis of many morphological synapomorphies being

shared with representatives from disparate families of the Anomura.

Introduction
Previous classifications of the decapod
infraorder Anomura (Borradaile, — 1907;

Glaessner, 1969), recognised the four superfam-
ilies, Thalassinoidea, Paguroidea, Galatheoidea
and Hippoidea. More recently the thalassinoids
have been excluded from the Anomura and the
constituent anomuran superfamilies redefined
asthe Paguroidea, Lomoidea. Galatheoidea and
Hippoidea (McLaughlin, 1983b; McLaughlin
and Holthuis, 1985; Poore, 1994). McLaughlin
and Holthuis stated that the term “Anomala' was
the more appropriate name for these constituent
superfamilies (minus the thalassinoids), but the
term ‘Anomura’ is more commonly used and has
been adopted herein.

The superfamily Lomoidea contains the
monospecific genus Lomis in the family Lomi-
dae. This hairy stone crab is endemic to the
rocky southern coasts of Australia and like its
porcellanid relatives is very crab-like in appear-
ance. Lomis hirta was first described as a hairy
porcellanid by Lamarck (1818), then Bouvier
(1894, 1895) considered it a symmetrical hermit
crab, but this enigmatic crab has since been elev-
ated to its own family and superfamily
(McLaughlin, 19832). Pilgrim (1965) reassessed
the literature and examined fresh specimens and
came to the conclusion that Lomis was a basal
offshoot of the paguroids (along with the Pylo-
chelidae) and that this basal position was indica-

tive of its sharing many characters with the
paguroids, galatheoids and thalassinoids.
McLaughlin (1983a) similarly placed Lomis ina
basal position, with respect to the Anomura, but
had it evolving independently as a distinct evol-
utionary line from a common ancestral stock.
Lomis continues to be problematic in regard to
its relationship to the remainder of the Anomura
(McLaughlin, 1983b; Martin and Abele, 1986;
Richter and Scholtz, 1994).

The use of spermatozoal morphology for sys-
tematic, taxonomic and phylogenetic studies,
initially pioneered by workers such as Koltzoff
(1906) and Retzius (1909), has been successfully
applied to the Crustacea (Jamieson, 1991) and
in the Anomura and Brachyura in particular
(Tudge, 1992, 1995a, b; Jamieson, 1994, 1995;
and references therein). In this paper, the ultra-
structure of the spermatozoa of Lomis hirta is
described for the first time and the phylogenetic
implications are discussed.

Methods

The single male specimen of Lomis hirta
(Lamarck, 1818) was collected and fixed by Dr
Gary Poore, Curator of Crustacea, Museum of
Victoria, Australia from Flinders Reef, Victoria
in September 1993. The male reproductive
material (both testes including the ducts of the
vasa deferentia) was removed from the crab and
immediately fixed in cold glutaraldehyde for a

235

minimum of 2 hours at 4°C then posted to Bris-
bane at ambient temperature where the remain-
der of the fixation and embedding process was
carried out.

Light microscopy

For light microscopy, glutaraldehyde-fixed
sperm were viewed and photographed under an
Olympus BH2 Nomarski interference contrast
microscope. Micrographs were taken with an
attached Olympus OM-2 camera.

Transmission electron microscopy

The standard fixation procedure (outlined
below) for transmission electron microscopy
was carried out in a Lynx -el. Microscopy Tissue
Processor (Australian Biomedical Corporation,
Ltd, Mount Waverley, Victoria, Australia), after
the initial glutaraldehyde fixation and first phos-
phate buffer wash.

Portions of the testis (approximately | mm?)
were fixed in 396 glutaraldehyde in 0.2 M phos-
phate buffer (pH 7.2), with 396 sucrose added,
for a minimum of | h at 4°C. They were washed
in phosphate buffer (3 washes in 15 min), post-
fixed in phosphate buffered 1% osmium tetrox-
ide for 80 min; similarly washed in buffer and

dehydrated through ascending concentrations of

ethanol (40-10090). After being infiltrated and
embedded in Spurr's epoxy resin (Spurr, 1969),
thin sections (500-800 A thick) were cut on a
LKB 2128 UM IV microtome with a diamond
knife. Sections were placed on carbon-stabilised
colloidin-coated 200 um mesh copper grids and
stained (according to Daddow, 1986) in Rey-
nold's lead citrate (Reynolds, 1963) for 30 s,
rinsed in distilled water, then 6% aqueous uranyl
acetate for 1 min, Reynold's lead citrate again
for 30 s and a final rinse in distilled water.
Micrographs were taken on an Hitachi H-300
transmission electron microscope at 80 kV.

Results

Spermatophore morphology

The spermatophores of Lomis hirta were only
viewed in thick sections for transmission elec-
tron microscopy and the dimensions and mor-
phology can only be estimated. The spermato-
phores are pedunculate, having the sperm-filled
ampulla attached to a long thin stalk which in
turn is connected to a basal plate or pedestal.
The entire spermatophores have been estimated
at approximately 300 um in length and the
ampullar dimensions at 100 um long X 50 pm
wide (Fig. 1A). The spermatophore wall (Fig.
LE), which constitutes the ampullar region of the

C. C. TUDGE

spermatophore, is shown at the electron micro-
scope level to be composed of a single homo-
geneously granular electron-dense layer (Fig.
2A) and to be of variable thickness.

Spermatozoal morphology

At the light microscope level the spermatozoa
of Lomis hirta are irregular in outline, but basi-
cally globular in form with one to three small
vertices which may be extended into microtubu-
lar arms (Figs IB-E). Up to two microtubular
arms have been observed in a single spermato-
zoon but sometimes three vertices are apparent
(Fig. IB). A ring-shaped acrosome vesicle is
obvious on one surface of the sperm cell. The
entire sperm cell can be up to 6um in diameter
(refer to Figs 2C and 4 throughout). In all figures
the acrosomal or apical end ofthe spermatozoon
is considered the anterior pole while the
opposite, basal or nuclear end is posterior.

The acrosome vesicle is an irregular, inverted
cup-shaped structure, approximately 2.5 um
wide, and is broadly penetrated posteriorly by
the perforatorial chamber (Figs 2B, C; 3A, B, D).
The acrosome vesicle wall is relatively thin and
1s composed of three distinctive zones. The most
external region of the vesicle is made up of an
interrupted layer of extremely electron-dense
material embedded in a granular zone of lesser
electron density. The extremely electron-dense
material is interpreted to be the equivalent of the
dense operculum, which normally caps the acro-
some vesicle in anomurans, but its extension
around the entire acrosome vesicle is unusual
(Figs 2B, C; 3A, B, D, F). As mentioned this
opercular layer is not a continuous, evenly dis-
tributed structure but forms an interrupted
dense layer. The electron-dense, granular zone,
here interpreted as the outer acrosome zone, sur-
rounds the operculum and composes the
majority of the thin acrosome vesicle. Occurring
between this combined dense outer acrosome
layer and the perforatorial chamber wall is a
zone of similar granularity, the inner acrosome
zone (Figs 2C; 3A, B, D-F). This latter zone
is more electron-pale than the outer acrosome
zone.

The perforatorial chamber invaginates into
the acrosome vesicle from the posterior end and
extends almost to the anterior most tip. It has a
broad posterior region which tapers anteriorly to
a blunt point (Figs 2C; 3A). The open, posterior
end is slightly constricted when compared to the
greatest width of the perforatorial chamber. The
perforatorial chamber contents can be divided
into two distinct regions. Internally there is a

TAXONOMIC POSITION OF THE HAIRY STONE CRAB 237

Figure 1. A-E Lomis hirta (Lomidae). A. Toluidine stained thick resin section showing partially sectioned
spermatophore and contained spermatozoa. B-D. Light micrographs of spermatozoa showing one or two micro-
tubular arms. E. Light micrograph of spermatozoa extruded from the vas deferens. Note: the thick spermato-
phore wall. Scale bars as indicated (original).

a = ampulla; av = acrosome vesicle; ma = microtubular arm; ped = pedestal; s = spermatozoon; st = stalk; sw =

spermatophore wall.

lw

38 C. C. TUDGE

Figure 2. A-C Lomis hirta (Lomidae). Transmission electron micrographs of spermatozoa. A. Detail of sper-
matophore wall surrounding spermatozoa. B. Low power shot of three spermatozoa. C. Longitudinal section (LS)
of a spermatozoon. Arrowhead indicates probable actin filaments. Scale bar = lum, except where indicated
(original).

av = acrosome vesicle; cy = cytoplasm; ia = inner acrosome zone; m = mitochondrion; ms = membrane system; mt
= microtubules; n = nucleus; npm = nucleo-plasma membrane; o = operculum; oa = outer acrosome zone; p =
perforatorial chamber; sw = spermatophore wall.

TAXONOMIC POSITION OF THE HAIRY STONE CRAB

core of longitudinally arranged striations, simi-
lar in appearance to actin filaments, while exter-
nal to this is a homogeneous, finely granular
region of moderate electron density (Figs 2C;
3A, B; 4). The posterior contents of the perfor-
atorial chamber appear continuous with the
cytoplasm beneath the acrosome vesicle.

The cytoplasm is very extensive and com-
pletely surrounds the acrosome vesicle, with the
widest areas lateral to it (Figs 2C; 3D, F; 4).
Anterior and posterior to the acrosome vesicle
the cytoplasm is reduced to a thin layer with no
obvious organelles present, except for a pair of
centrioles which occur beneath the perforatorial
chamber (Fig. 3C). Lateral to the acrosome ves-
icle the cytoplasm contains many organelles
including an anterior region of loosely aggre-
gated membranes and associated with these,
many small spherical mitochondria, of which
some appear cristate and others more degener-
ate (acristate) (Figs 2C; 3D, F). Scattered
amongst these organelles are short and long
bundles of microtubules which represent
internal sections of the microtubular arms.
Some bundles of microtubules are seen to extend
into the nuclear material (Figs 3D, E).

The nucleus of the spermatozoa of Lomis hirta
is irregular in form but retains a basically globu-
lar appearance (Figs 1B-D). It is composed of
homogeneous, coarsely granular material of
moderate electron density and is surrounded
externally by a thickened nucleo-plasma mem-
brane (Figs 2B, C; 3D; 4). No distinct membrane
separates the cytoplasm from the nucleus. As
previously mentioned small sections of the
internal microtubular bundles are apparent in
the nucleus (Fig. 2C).

Discussion

The paucity of information concerning the mor-
phology of the spermatophores of Lomis hirta
make it difficult to do more than speculate about
their structure. It can be ascertained that the
spermatophores are pedunculate (Fig. 1), and
therefore typical of the Anomura studied to date
(Tudge, 1991, 19952). Until further morphologi-
cal and ultrastructural studies are carried out on
the spermatophore of this crab it will not be
known if it has a spermatophore morphology
that substantiates its morphological and sper-
matozoal uniqueness. The homogeneous, granu-
lar appearance of the spermatophore wall in
Lomis (Fig. 2A) is similar to that recorded in the
spermatophores of several pagurid hermit crabs,
the parapagurid, Sympagurus, the galatheid,

239

Allogalathea, the hippid, Hippa, and several
porcellanid species (Tudge, 19952).

The spermatozoon of Lomis hirta (Figs 2, 3, 4)
possesses spermatozoal characters, such as
microtubular arms (possible three?) and a con-
centrically arranged acrosome vesicle poster-
iorly penetrated by a perforatorial chamber,
which justify its position in the Anomura but its
spermatozoal morphology is distinct enough to
warrant its own family and superfamily. Some of
the characteristics of the spermatozoon of
Lomis are considered unique. The variation in
sperm cell shape shown within an individual is
unusual and the amorphous form of the sperma-
tozoa (Figs I B-E; 2B), and to a lesser extent, the
acrosome vesicle, distinguishes Lomis from
other investigated anomurans (Hinsch, 1980,
1991; Tudge, 1992, 1995a, b). The small acro-
some vesicle completely embedded in the cyto-
plasm has not been recorded for any other
anomuran spermatozoon and is only approxi-
mated by that seen in the thalassinoids, Axius
and Callianassa (Tudge, 1995a, b). The micro-
tubular bundles, which appear to be concen-
trated in the cytoplasm, can also extend into the
nucleus (Figs 2C; 3D-F). It is not known if these
microtubular bundles, which are the bases of the
microtubular arms, are cytoplasmic in origin (as
in other investigated anomurans) or nuclear in
origin (as seen in the spermatozoa of astaci-
deans, palinurans and some thalassinoids and
brachyurans). Although the sperm cells appear
to have three vertices, only one or two microtu-
bular arms have been seen on a single sperma-
tozoon (Figs I B-D) and it is not known if three
microtubular arms is standard for Loris. Three
microtubular arms have been recorded for all
investigated paguroids and galatheoids, with the
exception of the Porcellanidae; which, with the
hippids and thalassinoids, have four or more
microtubular arms (Tudge, 1995a). Another
unique feature of the spermatozoa of Lomis is
the discontinuous electron-dense operculum,
which surrounds the entire acrosome vesicle
(Figs 2C; 2A, B, D-F) and not just the apical pole,
as in all other investigated anomurans. An
alternative theory concerning this structure is
that it is not an operculum at all, but simply
another dense zone of the acrosome vesicle. This
possibility assumes that the operculum is there-
fore lacking which would similarly be an auta-
pomorphy.

The Lomidae have been assigned to a basal
position in the Anomura, based on adult somatic
characters (Pilgrim, 1965; McLaughlin, 1983a,
b; Martin and Abele, 1986; Richter and Scholtz,

240 C. C. TUDGE

Figure 3, A-F. Lomis hirta (Lomidae). Transmission electron micrographs of spermatozoa. A. Detail of longi-
tudinal section (LS) of acrosome vesicle, Arrowhead indicates probable actin filaments. B, Transverse section
(TS) through acrosome vesicle. C. Detail of pair of centrioles. D. Oblique section through a spermatozoon
showing long internal bundle of microtubules. E. Detail of microtubular bundle. F. Detail of cytoplasmic orga-
nelles, Scale bars = lum, except where indicated (original).

ce = centriole: cy = cytoplasm; ia = inner acrosome zone; m = mitochondrion; ms = membrane system; mt =
microtubules: n = nucleus: o = operculum; oa = outer acrosome zone; p = perforatorial chamber.

TAXONOMIC POSITION OF THE HAIRY STONE CRAB 24]

operculum

inner acrosome zone

LIN
SA

AVE
MIS Z:
AZI AN
isa DA
e uo ur]

Nil
"PETI =
WARM Mr

Figure 4. Lomis hirta (Lomidae). Semidiagrammatic longitudinal section of a spermatozoon, based on a tracing
of a micrograph. Scale bar = Ium (original).

242

1994), but there are differences between their
conclusions. Not all authors dealt with all ano-
muran taxa. Pilgrim (1965) thought the family to
be an early offshoot of the Anomura (within
which he included the Thalassinidea).
McLaughlin (1983a: 435) stated “the Lomidae
represents a distinct evolutionary lineage within
the Anomala, sharing the basic characters of the
infraorder, but having evolved independently
from the ancestral stock common to all of its
major taxa", and decided it was a specialised
and highly evolved taxon within the infraorder.
Martin and Abele (1986) thought the Lomidae
to be the sister family to the Lithodidae, together
sister taxa to the Paguroidea. Richter and
Scholtz (1994) placed the Lomidae as the sister
group of the Paguroidea (including Lithodidae
in the latter superfamily) based on uropod mor-
phology.

The fact that the phylogenetic relationship of
Lomis has sparked such debate and still remains
unclear is reflected in its lack of recognisable
morphological apomorphies and its sharing of
numerous synapomorphic characters with more
remotely related anomuran taxa. Like many
other families in the Anomura (the Hippidae,
Lithodidae, Pylochelidae and Aeglidae, for
example) the Lomidae is only represented as fos-
sils from the Recent (Glaessner, 1969) and there-
fore palaeontological evidence of their evolution
is limited.

The spermatozoal data supports the views
presented by Pilgrim (1965) and McLaughlin
(1983a) that Lomis is basal to, or a separate lin-
eage from, the ancestor of the Anomura + Thal-
assinidea (Pilgrim, 1965) or Anomala (anomu-
rans excluding thalassinideans) (McLaughlin,
1983a). This same spermatozoal data is less sup-
portive of the view expressed by Richter and
Scholtz (1994) that the Lomidae is the sister
taxon of the Paguroidea and does not support
the close relationship between Lomis and litho-
dids suggested by Martin and Abele (1986).
Although the spermatozoal morphology in the
Lithodidae is only described from light micro-
scope drawings of Lithodes maja (see Retzius,
1909), it can be clearly ascertained that the over-
all spermatozoal morphology is unlike that of
Lomis and more similar to investigated pagu-
roids (Tudge, 1992, 1995a, b). A close link
between Lomis and lithodids seems unlikely on
the present spermatological evidence. Poore
(1994) clearly showed a sister group relationship
between the Thalassinidea and the Anomura
based on morphological characters but the pos-
ition of Lomis in this analysis is not known as it

C. C. TUDGE

was not one of the anomuran taxa used. The
spermatozoa of Lomis appear to share several
characters in common with two previously
investigated thalassinoids (Axius and Callia-
nassa) but a detailed comparison of spermato-
zoal morphology between the Lomoidea and
Thalassinoidea will have to wait pending a more
thorough investigation into the spermatozoal
form and diversity within the latter super-
family.

The ancestor of some or all of the anomuran
families may have had a spermatozoal mor-
phology very similar to that of Lomis and the
diversity (admittedly restricted by a conserva-
tive ground-plan) of sperm morphology encoun-
tered in extant families of the Anomura could
have emerged through modification of this
ancestral sperm form. It is conceivable that the
anomuran spermatozoal type (in the Paguroidea
in particular) developed from a lomid-like mor-
phology by 1) increasing the size and complexity
of the acrosome vesicle, 2) restricting the oper-
culum to the apical pole, 3) shifting the acro-
some vesicle to a superior position on the
cytoplasm and 4) establishing three, equidistant
microtubular arms. The lomid lineage may have
independently evolved from this ancestor but
retained the spermatozoal morphology which
was ancestral to the remainder of the Anomura.
The lack of distinct morphological apomor-
phies, the sharing of morphological characters
with as many as nine other anomuran families
and the newly presented spermatological evi-
dence, indicating a unique and particularly
unmodified sperm morphology, would appear
to vindicate a basal or ancestral position of
Lomis in relation to the remainder of the Ano-
mura.

Acknowledgments

The author wishes to thank Dr Gary Poore
(Museum of Victoria) for collecting and under-
taking the initial fixation of the specimen of
Lomis hirta. Mrs Lina Daddow (Zoology
Department, The University of Queensland) is
thanked for technical assistance with electron
microscopy. The support and careful reading of
the manuscript by Prof. Barrie Jamieson
(Zoology Department, The University of
Queensland) is gratefully acknowledged.

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397-409.

Tudge, C.C., 1995a. The ultrastructure and phylogeny
of anomuran crab spermatozoa. Ph.D. Thesis,
Zoology Department, University of Queensland,
Brisbane, Australia. 346 pp.

Tudge, C.C., 1995b. Ultrastructure and phylogeny of
the spermatozoa of the infraorders Thalassinidea
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toire naturelle (Paris) 166: 251—263.

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Memoirs of the Museum of Victoria 56(1):245-253 (1997)

A REMARKABLE NEW PYGMY SEAHORSE (SYNGNATHIDAE: HIPPOCAMPUS) FROM
SOUTH-EASTERN AUSTRALIA, WITH A REDESCRIPTION OF H. BARGIBANTI
WHITLEY FROM NEW CALEDONIA

MARTIN F. GOMON

Department of Natural Sciences, Museum of Victoria, GPO Box 666. Melbourne, Vic. 3000, Australia

Abstract

Gomon. M.F., 1997. A remarkable new pygmy seahorse (Syngnathidae: Hippocampus) from
south-eastern Australia, with a redescription of H. bargibanti Whitley from New Caledonia.
Memoirs of the Museum of Victoria 56(1): 245-254.

A new syngnathid fish, Hippocampus minotaur, is described from southeastern Australia.
It can be distinguished from congeners by its prominent head and distinctive thick ‘neck’,
tiny dorsal fin with only 7 rays, low number of pectoral fin rays (11), low number of trunk
segments (8), lack of definition of trunk and caudal segments and lack of inferior and median
ventral trunk ridges. It appears to be restricted to mid-continental shelf depths. A complete
description of the New Caledonian H. bargibanti Whitley, 1970 is also presented,

Introduction

Species of Hippocampus (family Syngnathidae),
widely referred to as seahorses, are extremely
variable (e.g., Ginsburg, 1937). Their well pub-
licised male brooding habits have been respon-
sible for a low rate of dispersal and consequent
clinal variation in at least some populations. In
such cases, subpopulations have occasionally
been recognised as separate species. Although a
comprehensive generic revision of Hippocam-
pusis still to come, the perception that the genus
is less diverse and species are more widely dis-
tributed than long thought is obvious from treat-
ments that have appeared in regional studies
over the last 15 years (e.g., Vari, 1982, Dawson,
1986a, 1986b, 1994). Nevertheless, previously
unrecognised taxa do occur. One example is an
extraordinary, diminutive, southeastern Aus-
tralian species which occurs at depths that are
generally greater than those inhabited by its
sympatric congeners. A description of the
species is presented here, bringing the number of
currently recognised Australian representatives
ofthe genus to seven. The new species is similar
in many regards to the New Caledonian H. bar-
gibanti Whitley, 1970, described in a brief note
in the abstracts of an ordinary meeting of the
Linnaean Society of New South Wales. A
detailed description of that species is also
provided.

Methodology follows that of Vari (1982),
except that a camera lucida on a binocular
microscope was used to make tracings from
which proportional measurements were

recorded. Because of the fleshy nature of the two
species treated and the reduction in ossification
of their trunk and tail segments, it is not possible
to report the numbers of segments based on
external features, such as tubercles on the lateral
trunk ridge. À comparison of body segments,
determined as described by Vari (1982), with
vertebrae in H. abdominalis and H. breviceps
revealed that the first two precaudal vertebrae
are not represented by tubercles on the lateral
trunk ridge. The numbers of trunk segments as
reported by previous workers is therefore
assumed to be two fewer than the numbers of
precaudal vertebrae. The numbers of tail seg-
ments in the two species are equal to the num-
bers of caudal vertebrae. In the following
descriptions, the numbers of trunk and tail seg-
ments reported were calculated on the basis of
the numbers of precaudal and caudal vertebrae
observed on radiographs. The descriptions of
the two species are of the holotype and lectotype;
variations observed in paratypes, paralectotypes
and other material are noted in parentheses. All
body lengths are reported as millimeters total
length (TL) as recorded by Vari (1982). The
snout, orbit and post orbital lengths are com-
pared with the head length (HL).

Specimens are from the collections of the Aus-
trallan Museum, Sydney (AMS) and the
Museum of Victoria, Melbourne (NMV). In
addition to specimens of H. bargibanti and H.
minotaur mentioned in the descriptions below,
the following material was examined for this
study: H. abdominalis: NMV A2899 (42.6,
immature male?) Victoria, Port Phillip Bay,

245

246

Rye; NMV A8945 (60.5, female) South Aus-
tralia, Point Sinclair; NMV A8975 (56.1, male)
Victoria, off Lakes Entrance; NMV A14219(12:
13.2-14.5, juveniles) Victoria, Port Phillip Bay,
aquarium bred; /7. angustus: AMS 1.35463-001
(18.2, juvenile) Western Australia, entrance to
Swan Esturary; H. breviceps: NMV A2898 (3:
23.2-28.6, juveniles) Victoria, Port Phillip Bay,
Rye; NMV AI1099 (48.4, male) Victoria,
southern Port Phillip; NMV A11469 (2: 45.8-
57.2, male and female) Victoria, Port Welshpool
Jetty; NMV A14180 (2: 15.4-22.9, juveniles)
Victoria, Western Port; M. whitei AMS
1.28289-002 (16.6, juvenile) New South Wales,
Lake Macquarie.
Hippocampus minotaur sp. nov.
bullneck seahorse
Figures l, 2a, 3a

Material examined. Holotype. NMV A192 (48.7,
male), Australia, New South Wales, off Eden, 35-40
fm (64-73 m), 30 Dec 1960, Danish seine trawl, R.
Slack-Smith.

Paratypes. NSW. AMSTA.3509 (42.2, female), coast
of NSW, trawled; AMS IA.3560 (52.6, female), 8 mi E
of Eden, 50-60 fm (91.5-109.8 m).

Vic. NMV A14161 (19.2, juvenile), Bass Strait, 38
km SW of Cape Paterson, 38°56.4’S, 145°16.6’E, 70 m,
fine sandy bottom, 12 Nov 1981, RV Tangaroa, R.
Wilson.

Diagnosis. Dorsal fin rays 7; pectoral fin rays 11;
anal fin rays 4; no exaggerated constriction div-
iding head and body; head and body extremely
fleshy without recognisable bony segments,
spines or other ornamentation; ventral trunk
ridges undeveloped.

Description. Head prominent, rather immobile,
maintained at an angle of about 90° to axis of
trunk; head length 16.4 (16.7-21.3)% TL; snout
very short, 29.3 (28.0-31.1)% HL; orbital diam-
eter 25.6 (20.3-24.4)% HL; postorbital length
46.3 (47.7-48.8)% HL; dermal flaps absent; cor-
onet low, appearing as a shallow mound; “neck”
extremely deep (= broad), only slightly shallower
than head depth; head and body very fleshy, seg-
ments indistinct in fresh material, devoid of
prominent spines or tubercles; fleshy filaments
absent; ventral portion of trunk segments
incomplete; inferior trunk ridge and median
ventral trunk ridge absent; other trunk and tail
ridges very poorly developed, remnants of lat-
eral trunk and inferior tail ridges not perceptibly
confluent, remnants of superior trunk and tail
ridges overlapping on ultimate trunk segment;
trunk segments 8; caudal segments 41; first

M. F. GOMON

caudal segment quadrangular; last trunk seg-
ment incomplete; trunk short, 31.0 (31.3-
36.496 TL; abdomen very broad, 19.7 (14.6-
15.9)% TL; pouch in brooding males occupying
most of abdominal cavity, positioned opposite
last two trunk segments and first five caudal seg-
ments; dorsal fin tiny, with 7 rays, base
extremely short, 2.0 (2.6-3.8)% TL, situated at

juncture of last trunk segment and first caudal

segment, totally confined to posterior and
anterior portions of these segments respectively
(subdorsal rings 2); pectoral fin rays 11; anal fin
rays 4, vestigial or absent in adults. Colour in
preservative, mostly cream, speckled with tiny
brown dots with cream centres (remaining speci-
mens mostly brownish). Life colors unavaila-
ble.

Etymology. Minotaur (noun in apposition), a
mythical creature with the head of a bull and the
body of a human, referring to the contrasting
massive bull-like head and graceful trunk and
tail of this species.

Distribution and ecology. Specimens for which
detailed collection information is available
came from off Eden, New South Wales, and
southwest of Cape Paterson, Victoria. A. Steffe
(personal communication) has indicated that
the species has also been taken in environmental
monitoring surveys off Wollongong, NSW. One
of the paratypes was recorded as having been
collected on a bottom with “fine sand’.

Remarks. The prominent head with very short
snout, relatively thick ‘neck’ region and tiny dor-
sal fin supported by only seven rays make this
diminutive species unmistakable. Its very low
numbers of dorsal and pectoral fin rays in com-
parison with other recognised species of the
genus (Fig. 4) approach those of several other
“pygmy” species, such as the Western Atlantic H.
zosterae Jordan and Gilbert, 1882, the Japanese
H. coronatus Temminck and Schlegel, 1850, and
the New Caledonian H. bargibanti Whitley,
1970. Similarities between some of these species
may be paedomorphic. Like two of its three sym-
patric congeners, H. minotaur has a high num-
ber of caudal segments, its value overlapping
those of H. breviceps and being just less than in
H. abodominalis. Hippocampus abdominalis has
the highest numbers of caudal segments in the
genus. The number in M. minotaur, however,
does overlap with those of the Chinese H. tata-
kurae and the Indo-Pacific H. trimaculatus.

Hippocampus minotaur shares with the unusual
H. bargibanti the lack of any obvious demar-
cation of body segments and a great reduction of

A REMARKABLE NEW PYGMY SEAHORSE 247

9 Es.

Figure 1. Hippocampus minotaur sp. nov. a, holotype, NMV A192, male, 48.7 mm TL, and b, paratype, AMS
IA.3509, female, 42.2 mm TL. Scale = 5 mm.

Figure 2. Radiographs: a, Hippocampus minotaur sp. nov., holotype, NMV A192, male, 48.7 mm TL; b, Hip-
pocampus bargibanti lectotype, AMS 1.15418-002, 20.9 mm TL; c, Hippocampus breviceps NMV A2898, 27.4

mm TL.

248 M. F. GOMON

Figure 3. Juveniles of temperate Australian species of Hippocampus: a, Hippocampus minotaur sp. nov., para-
type, NMV A14161, 19.2 mm TL; b, Hippocampus abdominalis, NMV A14219, 14.0 mm TL; c, Hippocampus
breviceps, NMV A14180,22.9 mm TL; d, Hippocampus whitei, AMSI.28289-002, 16.6 mm TL; e, Hippocampus
angustus, AMS 1.35463-001, 18.2 mm TL. Scale = 5 mm.

A REMARKABLE NEW PYGMY SEAHORSE

both ventral trunk ridges and the lower portions
of trunk segments, as well as an abbreviated
snout, a reduction in the constriction separating
the head from the body, and small size. Hippo-
campus minotaur and H. bargibanti differ from
other previously described members of the
genus in having the ventral portions of the trunk
segments sufficiently reduced that they no
longer have the segmented appearance typical of
syngnathids (Fig. 2c). In H. bargibanti dermal
ossifications on the belly are reduced to uncon-
nected star-like plates (Fig. 2b). In H. minotaur,
dermal ossifications are absent in this region, the
only visible remnant of the posterior trunk seg-
ments ventrally in radiographs of females being
a crescentic process curving posteriorly and ven-
trally from the position of the lateral trunk ridge
of the last trunk segment. This may provide
structural support to the posterior margin of the
‘belly’. In the holotype of this species, a brooding
male, the ossified process curves anteriorly and
ventrally and separates the anterior margin of
the brood pouch from the thoracic cavity. The
level of the process in this specimen is near the
centre of the second penultimate trunk segment
(Fig. 2a). The space remaining for abdominal
organs in the holotype is in a narrow strip of the
trunk anteriorly.

Despite the disparity in their number of cau-
dal segments, the modifications shared by H.
minotaur and H. bargibanti support the hypoth-
esis that they are sister species. Other superficial
features may provide further evidence for such a
relationship. In the paratypes of H. minotaur,
noticeable bud-like processes somewhat remi-
niscent of those on the body of H. bargibanti are
present on the dorsal surface of the tail. In the
latter, the bumps closely resemble the polyps of
the gorgonian coral (Muricella sp.) to which it
has been observed to cling and apparently which
it mimics in life (Whitley, 1970). The structures
in H. minotaur may provide a similar form of
camouflage, especially if the species is found in
association with southern soft corals. Unfortu-
nately, collection information for type speci-
mens of H. minotaur is insufficiently detailed to
be able to assess this possibility.

Even at a very small size, H. minotaur 18
readily separable from its sympatric congeners
by its very short snout and absence of readily
recognisable body segments or associated
spinous ornamentation (Fig 3a). The shortsnout
seahorse H. breviceps (Fig. 3c) at this size has an
extremely elongate snout, comparable with
those in H. abdominalis (Fig. 3b), H. whitei (Fig.
3d)and the warm temperate Western Australian

249

H. angustus (Fig. 3e), even though the last three
species have a comparatively longer snout than
H. breviceps in adults. Hippocampus abdomi-
nalis, H. breviceps and H. whitei, which co-occur
in southeastern Australian are easily separated
at all sizes by their dorsal fin ray and caudal seg-
ment counts. Hippocampus angustus is separ-
able from H. abdominalis and H. breviceps by
caudal segment counts (Fig. 4).

Hippocampus bargibanti Whitley, 1970
Figures 2b, 5, 6

Hippocampus bargibanti Whitley, 1970: 294. —
Burgess et al., 1988: pl.79, bottom.

Material examined. Lectotype: AMS I.15418-002
(20.9, female), New Caledonia, Nouméa, on gorgonian
coral Muricella sp., 30 m, July 1969, collected by
Georges Bargibant.

Paralectotype: AMS 1.15418-001 (19.5 mm TL,
female), same collection information as lectotype.

Other specimens: New Caledonia. AMS 1.15997-
001 (2: 21.4-22.0, male and female), off Nouméa.
Canal Woodin, 20-25 m, 26 Sep 1971, S. Catala; AMS
1.19834-001 (24.2, female), Nouméa lagoon, Jul
1969.

Diagnosis. Dorsal fin rays 14; pectoral fin rays
10; anal fin absent in adults; very weak constric-
tion separating head and body; head and body
extremely fleshy mostly without recognisable
bony segments, body ornamentation in the form
of prominent bulbous tubercles; ventral trunk
ridges poorly developed.

Description. Head moderately large, with restric-
ted movement, held at an angle of about 90° to
axis of trunk; head length 19.6 (16.9-19.2)% TL;
snout extremely short 34.1 (24.4-28.6)% HL.
with dorsal constriction midway between tip
and eye giving tip a bulbous appearance; orbital
diameter 26.8 (19.5-24.4)% HL; postorbital
length 61,0 (51.4-61.0)% HL; prominent fleshy
bulbous spine above each eye, smaller crest-like
spine on dorsal midline of head in front of eye;
small blunt spine centrally on cheek below pos-
terior half of eye; slightly larger blunt spine on
ventral midline of head below posterior margin
of eye; corona prominent with broad base and
broadly flattened apex; prominent fleshy bul-
bous spine on each side of head below forward
half of corona, level with upper edge of eye;
slightly smaller spine immediately preceding
lower half of pectoral fin base; extremely promi-
nent fleshy bulbous spines based on dorsal trunk
ridge at about first, fifth and twelfth segments,
the last pair on either side of dorsal fin base
between seventh and tenth rays; fleshy bulbous

250 M. F. GOMON

Dorsal Fin Pectoral Fin Trunk Rings Caudal Rings
Ñ 10 is 20 z w as [w 3 mis.» sls » ES 40 A toj
ingens! 1921 1517 mg
erectus ? 1625 17 fen] 3338
reidi? 1619 1537 m 3337
zosterae ? 1013 1012 su [ aw]
hippocampus ? 1619 1315 [212] La]
ramulosus 3 1821 1518 1012 3640
camelopardalis * 1921 [ris m 3538
capensis * 1618 1517 [xm 313
histrix * 1719 1748] mi pss
kuda * 1518 1517 mi 3637
trimaculatus * 1921 17 E 3842
coronatus $ EZ m io [3840]
japonicus 5 an E m EJ
sindonis 5 BI n DI pr
tatakurae 5 E] mi m a
spinosissimus © 7] B a] E]
angustus ? 1521 n m 3335
zebra? 7] s m E
abdominalis ? EX LU [E “a
breviceps 9 1922 [3 m 3842
whitei 9 1547 Tu Mi 33-36
bargibanti 13-14] fo] (TEDI E
minotaur 7 MI [s] [s]
L

Figure 4. Selected meristic values for species of Hippocampus. Meristic values shown came from: 1 Ginsburg,
1937, 2 Vari, 1982, 3 Dawson, 1986b, 4 Dawson, 1986a, 5 Masuda et al., 1984, 6 Weber and de Beaufort, 1922, 7
Günther, 1870, 8 Whitley, 1964 and 9 Dawson, 1994.

Figure 5. Hippocampus bargibanti Whitley, 1970, lectotype, AMS I.15418-002, 20.9 mm TL, female. Scale = 5
mm.

A REMARKABLE NEW PYGMY SEAHORSE

Figure 6. Hippocampus bargibanti Whitley, 1970, on Muricella sp.

process on lateral trunk ridge at eighth trunk seg-
ment; similar process immediately ventrad;
lower, broader process on ventral profile of
“belly” flanking anus on last trunk segment; sev-
eral smaller, low, blunt, fleshy spines scattered
on belly; dermal filaments absent; “neck” broad,
depth at constriction only slightly less than head
depth; ventral portion of trunk segments incom-
plete; inferior trunk ridge and median ventral
trunk ridge reduced to star-shaped, embedded,
dermal ossifications anteriorly, ossifications
forming structural bases for low fleshy spines on
belly, posteriormost two trunk segments more
completely ossified; other trunk and tail ridges
very poorly developed, remnants of lateral trunk
and inferior tail ridges not perceptively conflu-
ent, remnants of superior trunk and tail ridges
apparently overlapping on ultimate trunk seg-
ment; trunk segments 12; caudal segments 31;
first caudal segment quadrangular; last trunk
segment incomplete; trunk of moderate length
43.1 (36.4-41.0)% TL; abdomen broad, 15.3
(12.4-20.6)% TL; pouch structure unknown;
dorsal fin small, with 14 rays, base distinctly
angled, the anterior end positioned much more
ventrally thàn posterior end, length 11.0 (9.1—
9.8)% TL, situated above last 2.5 trunk segments
and first caudal segment (subdorsal rings 3.5);
pectoral fin rays 10; anal fin rays absent in
adults. Colour in preservative mostly cream. In
life, variable depending on color of Muricella sp.
with which it isliving. Types "creamy white with
spaced orange-yellow streaks becoming ring-like
on the tail, and the bumps over its head (coronet
and nuchal plates) and along the body segments
imitate the yellow and orange clumps of spicules
of the gorgonian" (Whitley, 1970). Others
greyish white with scarlet markings (Fig. 6).

Distribution and ecology. So far, only known
from New Caledonia where it has been taken
solely on the gorgonian Muricella sp.

Remarks. Whitley (1970) did not select a holo-
type from the syntypic series. The larger, AMS
I.15418-002 (Fig. 5), is here designated lecto-
type. The specimen registered as AMS 1.19834-
001 was apparently collected at the same time as
the two types but was not sent to the AMS until
after Whitley had published his description (J.
Paxton, personal communication).

Although seahorses are known for their cryp-
tic appearance, few, if any, have developed the
extreme protective coloration and morphology
of H. bargibanti. The resemblance of this species
to the gorgonian with which it lives is so close

M. F. GOMON

that the type series was only noticed after a stalk
of gorgonian was collected and placed in an
aquarium (J. Paxton, personal communication).
Subsequent specimens of H. bargibanti have
been taken by targeting its more highly visible
gorgonian partner.

The five specimens examined are very similar
morphologically. Although none has the obvious
external pouch development found in other
species of the genus, one of the three non-type
specimens examined by A. Vincent and J. Prit-
chard (personal communication) is a male. It
differs slightly from the other four in the ossified
posterior trunk segments that are directed per-
pendicularly from the vertebral column (as
observed in radiographs) with one of the ventral
ossifications expanded basally. This difference
is considerably less pronounced than observed
in other species of the genus. The inflated
appearance of the trunk in most of the speci-
mens of H. bargibanti may be related to the
reduced ossification of the more anterior seg-
ments and thus the greater flexibility of the
region.

Discussion

Sufficient evidence appears to exist to recognise
H. bargibanti and H. minotaur as constituting a
monophyletic lineage (see H. minotaur
“Remarks'). The temptation to propose a new
generic name for the two is resisted because little
information is available on the overall inter-
relationships of the species currently referred to
Hippocampus.

Acknowledgments

The illustration of H. angustus was provided by
F. Neira (Victorian Fisheries Research Insti-
tute). All others were prepared by R. Campbell
(NMV). The colour photo of H. bargibanti was
kindly supplied by J. Rivaton (ORSTOM). l am
grateful to A. Steffe (New South Wales Fisheries)
for conveying his observations on the distri-
bution of H. minotaur and to A, Vincent and J.
Pritchard (University of Oxford) for their
opinions on the sexes of three of the five speci-
mens of H. bargibanti. M. McGrouther (AMS)
facilitated the loan of specimens and forwarded
information. J. Paxton (AMS) provided useful
information on specimens of H. bargibanti and
commented on the manuscript. L. Jonker
(NMV) assisted with radiographs. P. Gunn
(NMV) prepared Figure 4. R. Vari (US National
Museum of Natural History) made helpful
suggestions which improved the manuscript.

A REMARKABLE NEW PYGMY SEAHORSE

References

Burgess, W.E., Axelrod, H.R. and Hunziker, R.E.,
1988. Dr. Burgess's atlas of marine aquarium
fishes. T.F.H. Publications: Neptune City. 736
pp., 563 pls.

Dawson, C.E., 1986a. Syngnathidae. Pp. 445-458 in:
Smith, M.M. and Heemstra, P.C. (eds), Smiths'
sea fishes. MacMillan South Africa: Johannes-
burg. 1047 pp.

Dawson, C.E., 1986b. Syngnathidae. Pp. 628-639 in:
Whitehead, P.J.P., Bauchot, M.-L., Hureau, J.-C.,
Nielsen, J. and Tortonese, E. (eds), Fishes of the
north-eastern Atlantic and the Mediterranean.
UNESCO: Paris. Volume 2, pp. 511-1008.

Dawson, C.E., 1994. Syngnathidae. Pp. 440-475 in:
Gomon, M.F., Glover, C.J.M. and Kuiter, R.H.
(eds), The fishes of Australia's south coast. State
Print South Australia: Adelaide. 992 pp.

Ginsburg, I., 1937. Review of the sea horses (Hippo-
campus) found on the coast of American conti-
nents and of Europe. Proceedings of the United
States National Museum 83: 497—594.

Günther, A., 1870. Catalogue of the fishes in the British
Museum. Volume 8. Catalogue of the Physostomi,
containing the families Gymnotidae, Symbranchi-
dae, Muraenidae, Pegasidae, and of the Lopho-
branchii, Plectognathi, Dipnoi, Ganoidei, Chon-
dropterygii, Cyclostomata, Leptocardii, in the

253

British Museum. British Museum (Natural His-
tory): London. xxv + 549 pp.

Masuda, H., Amaoka, K., Araga, C., Uyeno, T. and
Yoshino, T. (Eds) 1984. The fishes of the
Japanese Archipelago. Tokai University Press:
Tokyo. 2 vols. 437 pp., 370 pls.

Vari, R., 1982. The seahorses (subfamily Hippocam-
pinae). Pp. 173-189 in: Bóhlke, J.E., Cohen,
D.M., Collette, B.B., Eschmeyer, W.N., Gibbs,
R.H., Jr., Pietsch, T.W., Richards, W.J., Smith,
C.L. and Thomson, K.S. (eds), Fishes of the west-
ern North Atlantic. Part Eight. Order Gasterostei-
formes, Suborder Syngnathoidei. Syngnathidae
(Doryrhamphinae, Syngnathinae, Hippocampi-
nae) Memoir Sears Foundation for Marine
Research 1(8). 198 pp.

Weber, M. and de Beaufort, L.F., 1922. The fishes of
the Indo-Australian Archipelago. Volume IV.
Heteromi, Solenichthyes, Synentognathi, Perce-
soces, Labyrinthici, Microcyprini. Brill: Leiden.
410 pp.

Whitley, G.P., 1964. Fishes from the Coral Sea and the
Swain Reefs. Records of the Australian Museum
26 (5): 145-195, pls VIII-X, figs 1-15.

Whitley, G.P., 1970. In: Abstract of proceedings. Ordi-
nary general meeting. 26th November, 1969. Pro-
ceedings of the Linnaean Society of New South
Wales 94 (3): 292-295.

dne
*

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POLOJ MALE

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Memoirs of the Museum of Victoria 56(1):255-259 (1997)

^ NEW GRENADIER, GENUS TRACHONURUS, FROM NEW ZEALAND
AND AUSTRALIA (PISCES: GADIFORMES: MACROURIDAE)

TOMIO IWAMOTO! AND PETER MCMILLAN?

"California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA
"MAF Fisheries, Greta Point, PO Box 297, Wellington, New Zealand

Abstract

Iwamoto, T. and McMillan, P., 1997, A new grenadier, genus Trachonurus, from New Zeal-
and and Australia (Pisces: Gadiformes: Macrouridae). Memoirs of the Museum of Victoria

56: 255-259.

A new grenadier fish, Trachonurus gagates, is described from depths of 436-1240 m off
New Zealand and Australia. The species is unusual in lacking a grooved lateral line, which
differentiates it from all other known species of the genus. An undescribed species from
Australia, however, also lacks a lateral line, but that species has much larger scales and
differs in a number of proportional measurements and its paler colour.

Introduction

The generic relationships of Trachonurus has
recently been considered by Sazonov and
Shcherbachev (1985), and its worldwide distri-
bution by Shcherbachev et al. (1979). The latter
paper followed Marshall (1973) in treating the
five nominal species as synonyms of 7. villosus
(Günther, 1877) Studies by TI, however,
revealed that more than one of those five species
are probably valid, and several other undes-
cribed species are present, Here, a new species
from New Zealand and Australia without a
grooved lateral line is described.

Abbreviations and methods for making
counts and measurements generally follow Iwa-
moto (1970) and Iwamoto and Sazonov (1988).
Because the new species lacks a grooved lateral
line and has only widely spaced free neuromasts,
counts of scales above or below the “lateral line”
were not possible. Total scale-row counts along a
diagonal to the anal fin were useful and are used
for this species. Institutional abbreviations fol-
low Leviton et al. (1985) and Leviton and Gibbs
(1988).

Trachonurus Günther

Trachonurus Günther, 1887: 142 (as subgenus of

Macrurus; type species, Coryphaenoides villosus

Günther, 1877).

See Marshall (1973: 619) and Sazonov and
Iwamoto (1992: 77) for descriptions of the
genus.

Trachonurus gagates Sp. nov.
Figure |

Trachonurus sp. B: Paulin et al., 1989: 125 (in
key).

Material examined. Holotype: AMS 1.24059-009
(70.2 HL, 430+ TL); Australia, NSW, off Norah Head:
33°32’S, 15209'E; 978 m; 1983.

Paratypes: Australia, Old. CSIRO H.1157-01 (73.4
mm HL, 430+ mm TL) WNW of Marion Reef:
1851.1'/S, 149*33 E; 599-591 m; FRV Soela stn SO6-
85-39, CSIRO H.1158-01 (70.0 HL. 412+ TL); NW of
Marion Reef; 18°46.0°S, 150°32,1°E: 1188-1200 m:
FRV Soela stn SO6-85-38: 25 Nov 1985.

NSW. AMS 1.24355-001 (53.6 HL, 3524 TL) and
CAS 82134 (2: 66.2-72.3 HL, 400+-400+ TL); off
Shoalhaven; 3454’S, 151178'E; 1150 m; 1983. AMS
1.24173-007 (3: 68.7-74.7 HL, 4274-4784 TL); off
Shoalhaven Bight; 34°56’S, 151^1 3E: 1115 m; 26 Oct
1983. AMS I.24356-007 (71.7 HL, 425+ TL); 34°51’S,
151715^E; 1043-1061 m; FRV Kapala stn K83-14-05;
26 Jan 1983.

Vic. AMS1.24157-002 (76.0 HL. 450+ TL); off Cape
Everard; 38°17’S. 149947'E; 1015 m; 1983. CSIRO
T538 (81.6 HL. 4354 TL); off Cape Martin: 3748'S,
139°33’E; 1007 m; 25 Apr 1983. CSIRO H.2638-01
(49.5 HL, 3054 TL) SE of Portland; 38'53.8'S,
142°00.3’E: FRV Soela stn SO2-89-11; 1989.

Tas. CSIRO H.2640-01 (68.7 HL, 410+ TL); FRV
Soela stn SO2-89-94; 1989. CSIRO H.2634-03 (45.5
HL, 305 TL); SW of Macquarie Harbour; 42°28.6’S,
144*44.0'E; 1000 m; FRV Soela stn SO2-89-85; 1989,
CSIRO H.886-03 (71.0 HL, 430+ TL) E coast:
41°24.3’S, 14843.7'E; 890-1052 m; FRV Soela stn
SO4-87-09; 1987. CSIRO T829 (69.3 HL, 400+ TL);
off Bicheno; 41°48’S, 14838'E; 1099 m; Challenger
stn 40/02; 21 Apr 1982. CSIRO T739 (60.6 HL, 3424
TL); off Cape Sorell; 4224’S, 144"40'E; 1130-1150 m;
FV Margaret Phillipa stn 03/07; 22 Oct 1983,

SA. NMV A.5880 (2: 54.9-59.4 HL, 334-381 TL);
38°31.67°S, 140*43.91'E: 1100 m. NMV A,5882 (56.8
AL, 350 TL); 38°36.1’S, 140°58.9’E; 1080-1110 m:
FRV Soela stn SOIO-88-83; 8 Feb 1988. NMV
A.8995 (75.3 HL, 3934 TL): 33°57.7’S, 131”26.4'E;
1000-1030 m; FV Comet stn RP26; 19 Feb
1990.

255

256 TOMIO IWAMOTO AND PETER McMILLAN

Figure L. Holotype of Zrachonurus gagates, AMS 1.24059-009, 70.2 mm HL, 430+ mm TL, from off New South
Wales, Australia, in 978 m. Photograph by Susan Middleton.

Figure 2. Paratype of Zrachonurus gagates, NMNZ P.15855, 70.6 mm HL, 4354 mm TL, from west coast of
North Island, New Zealand, in 800-848 m. Drawing by Peter McMillan.

A NEW GRENADIER 257

CSIRO T316 (61.9 HL, 390+ TL); Great Australian
Bight; 3425/8, 132705'E; 1175 m; FV Margaret Phil
lipa stn 6/19; 14 Feb 1984, CSIRO T747 (69.7 HL,
370+ TL); 33°49’S, 129°33/E; 1000-1052 m; Endeav-
our stn 04/09: 5 Jun 1983.

WA, CSIRO T289 (59.4 HL, 3704 TL); Great Aus-
tralian Bight; 332785, 128°36’E; 1027 m; FV Margaret
Phillipa stn 7/3; 24 Feb 1983. CSIRO H.3022-04 (69,1
HL, 405+ TL); 3317’S, 11413’E; 976 m; 1989. CSIRO
H.3023-08 (81.8 HL, 465« TL) W of Bunbury;
33920'5, 114°30’E; 435 m; 25 Dec 1989, CSIRO
H.3002-05 (51.2 HL, 3334 TL); SE of Albany; 35°23’S,
11827'E; 1030 m; 21 Dec 1989, CSIRO H.2617-07
(64.5 HL, 4204 TL); W of Bunbury; 33"15,8'S,
114°] |.1"E; 982 m; FRV Southern Surveyor stn SSI-
91-85; 1991,

New Zealand. NMNZ P. 11303 (72.3 HL, 4254 TL):
off Hawke Bay, 39°41.7’S, 177958.4/E; 1160-1240 m;
24 Nov 1981. NMNZ P.11415 (75.8 HL, 470 TL); W
of Cape Reinga; 34°57.4’S, 171°53.2’E; 1060-1080 m;

19 Nov 1981, NMNZ P.11561 (65.7 HL, 392+ TL); off

Wairarapa; 41704.5'S, 176°32.6’E; 960-1100 m; 18

Dec 1981, NMNZ P.12969 (62.0 HL, 355+ TL); NE of

Chatham lL; 42749,9'S, 176"08.2'W; 1050 m; 7 Aug
1982. NMNZ P,14763(53.0 HL, 327+ TL); Challenger
Plateau; 3830.3’S, 17041.9'E; 827-836 m; 25 Oct
1983. NMNZ 14770 (48.8 HL, 300+ TL); Challenger
Plateau; 39°31.8’S, 167°47.0°E; 804-817 m; | Sep
1983. NMNZ P. 15855 (70.6 HL, 435+ TL); west coast,
North Island; 37°41.3’S, 173"53.1'E; 801-848 m; 25

Apr 1981, NMNZ P.15856 (73.4 HL, 428+ TL); off

Wairarapa; 41^11.0'S, 176'40.0'E; 1070-1180 m; 4
Apr 1984. NMNZ P.15857 (72.4 HL, 432+ TL); north
Chatham Rise; 42°37.4’S, 176“19.0'E; 1065-1070 m;
18 Aug 1982. NMNZ P.15858 (70.0 HL, 404+ TL);
north Chatham Rise; 42°42,3°S, 175*32.6'E; 1060-
1070 m; 29 Aug 1982. NMNZ P.16105 (70,3 HL, 4234
TL); Challenger Plateau; 40°01.9’S, 168"10.9'E; 865-
870 m; 14 Jul 1984, NMNZ P.16106 (73.2 HL, 4234
TL) Challenger Plateau; 39°57.5°S, 167759.5'E; 886-
900 m; 11 Jul 1984.

Diagnosis. Trachonurus without a grooved lat-
eral line; chin barbel short, 4-8% of head length
(HL) suborbital width 13-1596; postorbital
length of head 50-55%; gill rakers on first (outer)
arch (mesial) 11—13 total, gill rakers on second
arch (lateral/mesial) 10-13 total/10-12 total;
scale rows below mid-base of first dorsal fin 25-
31; scale rows below origin of second dorsal fin
20-25; scale rows over distance equal to predor-
sal length usually 35-42; scale rows between
pelvic fin base and gill cover 11-14.

Measurements and counts. Total lengths 300—
4784 mm; head lengths 45.5-81.8 mm. The fol-
lowing in percent of head length (figures in par-
entheses are exceptions to the usual range):
postrostral length 75-80; snout length 24-27;
preoral length (height) (12)14-17; internasal
width 20-23; interorbital width 31-38 (40);

orbit diameter 25-30; suborbital width 13-15;
postorbital length 50-54; distance orbit to preo-
percle angle 36-42; length upper jaw 34-39;
length barbel 4-8; length outer gill slit 13-17;
preanal length 149-174; distance outer pelvic to
anal-fin origin 26-40; distance isthmus to anal-
fin origin 86-107; greatest body depth 74-102;
ID-2D interspace 14-35; height ID. 45-53;
length P. 38-54; length V, 30-42; length pos-
terior nostril 4-8.

ID. 11,7-9; P. (19, 111) 112-116; V. 6-7; total
gill-rakers outer arch (lateral/mesial) 3-8 / 11-
13, second arch 10-13 / 10-12; scales below 1D.
8-10, below mid-base of ID. (to A.) (25) 27-31,
below 2D. (20) 21-25, over distance equal to
predorsal length 35-42; scales from pelvic base
to gill cover 11-14; pyloric caeca 10-14.

Description of holotype (with. additions from
paratypes). Body long and gradually tapering
from trunk to end of tail. Head about sixth total
length; laterally compressed, completely lacking
strong sharp ridges. Snout bluntly rounded in
lateral profile, scarcely extending beyond jaws;
snout narrow in dorsal view, obtusely conical,
broadening notably above nostrils at anterodor-
sal corner of orbits. Interorbital space broad,
mostly flat, dorsal orbital margin forming shal-
low curved incursion into interorbital space.
Suborbital region vertical, flattened, and lacking
well-developed horizontal ridge. Mouth moder-
ately large, jaws extending to vertical slightly
behind midorbit; lips thick and fleshy. Preoper-
cle broadly rounded, without developed ridge.
Opercle and subopercle forming deep, narrow,
inverted triangle. Interopercle broadly exposed
laterally and posteriorly, densely scaled and con-
nected to similarly densely scaled lower jaw.
Chin barbel short, thick at base, tapering rapidly
to tip. Abdominal cavity long, extending poster-
iorly to over 8th-10th anal ray. Gill rakers on
outer side of first (outer) arch few small nubs;
mesial rakers more tubercular, with forwardly
directed tufts of long, recurved spinules.

Fins all small and typical of genus. Second
spinous ray of first dorsal fin slender, flexible,
tapering to thin tip and wholly lacking serrations
on leading edge. Second dorsal scarcely devel-
oped anteriorly and consisting of short uncon-
nected rays over most of posterior length, Pec-
toral fins small, short, tips fine and easily broken
(about half head length in better specimens). Pel-
vic fins small, outer ray with fine tips that barely
extend to anal fin origin; origin far posterior,
almost vertical to hind edge of first dorsal fin.

258

Teeth small but relatively stout, in narrow
bands in both jaws; about 3 or 4 teeth wide near
symphysis in upper jaw, narrowing to 2 rows and
then | row at extreme posterior end; outer series
slightly larger than inner series. Lower jaw in 3
irregular series at symphysis, narrowing rapidly
to 2 rows, continuing well posterior to end of
rictus; inner series slightly larger than outer
series.

Body scales relatively small, covered with
fine, slender, conical, erect spinules, some spin-
ules over dorsum with strongly curved tips.
Spinules generally arranged in a row along
anterior edge of exposed field, those posteriorly
in somewhat "V" arrangement. Spinules with
shallow buttresses or "roots," some intercon-
nected with adjacent spinules. Gular membrane
in holotype with dense, elongated patch of small
spiny scales; scales absent over bases of lower-
most branchiostegal rays. (In paratypes scale
cover on gular and branchiostegal membranes
variable, but generally heavily present on both
membranes.) Ventral scales along base of anal
fin somewhat stouter, larger, and with larger
spinules than other scales. Dorsal scales along
second dorsal fin scarcely larger than scales
more ventrad, but with some slightly enlarged
and strongly recurved spinules. Scales of head
highly variable in shape and size. Those along
posterior margin of orbits somewhat larger and
more elongate; series along dorsal edge of preo-
percle larger and slightly stouter. Scales over
opercle and subopercle somewhat larger than
others of head. Scales on snout, suborbital, man-
dible, interorbital, and interopercle small. Scales
abruptly absent under gill cover, as typical of
genus and characteristic of most genera related
to Cetonurus (as discussed by Sazonov and
Shcherbachev, 1985).

Periproct region broad, extensive, subtending
most of distance between pelvic and anal fins (as
typical of genus). Pyloric caeca short, thick, and
simple; length about equal to least suborbital
width or much shorter, length usually two to
three times caecum width.

Colour in alcohol overall dark brown to black-
ish. Fins, lips, lining of mouth and gill chambers
black. Gill membranes and barbel dark.
Abdominal region not noticeably darker exter-
nally than adjacent areas.

Size. To about 48 cm total length.

Distribution.New Zealand and Australia (Qld,
NSW, Vic., SA, Tas., WA) in 435-1240 m. In
New Zealand waters it is found between 32°24’S
and 43°30”S (1.e., in central and northern waters

TOMIO IWAMOTO AND PETER McMILLAN

on both coasts) at depths of 801—1240 m. It
occurs sympatrically with the rarer Trachonurus
sp. A of Paulin et al. (1989) in waters north of
latitude 40°S.

Etymology. From the Greek, gagates, velvety
black, with reference to the dark scale covering
of the body.

Remarks and comparisons. A second undes-
cribed species from Western Australia also lacks
alateral line, but is readily distinguished from 7.
gagates by its much larger scales, longer chin
barbel, narrower internasal and suborbital,
wider orbits, shorter postorbital and orbit-to-
preopercle distances, and generally paler
colour.

Aside from the absence of a lateral line, 7.
gagates appears most similar to 7. villosus from
Australian waters, the two agreeing in almost all
counts, proportional measurements, and squa-
mation and dentition features. The exceptions
arc few and slight with much overlap: the snout
length in T. gagates is slightly shorter (24-2796
HL vs 25-3196), the postorbital length is slightly
greater (50-5590 HL vs 46-52%), the orbit-to-
preopercle distance slightly longer (36-42% HL
vs 32-3896), and the chin barbel is shorter (4-8%
HL vs 8-14%). Additionally, the number of scale
rows along the lateral line over a distance equal
to the predorsal length amounts to about 35-42
for most individuals of 7. gagates, but 39-50 in
T. villosus.

Acknowledgments

We thank the many curators and assistants who
lent specimens and provided other assistance
and courtesies during the preparation of this
paper: J. Paxton, D. Hoese, M. McGrouther and
others (AMS) C. Paulin and A. Stewart
(NMNZ); M. Gomon (NMV); P. Last, A. Wil-
liams, A. Graham (CSIRO). Susan Middleton
(CAS) prepared the excellent photograph of the
holotype. Jon Fong and David Catania (CAS)
provided technical assistance. An extended visit
to Australia by TI was made possible by a visit-
ing fellowship from the Australian Museum. We
are grateful to Paxton, Hoese, and J. Leis for
assistance. with. that. fellowship, to Last for
wrangling funds from the Marine Laboratory,
CSIRO (Hobart) to extend TUS visit to that insti-
tution, to Gomon for providing housing and
hospitality, and to the CAS In-House Reseach
Fund for providing support for visits in 1989 to
Australia and New Zealand.

A NEW GRENADIER

References

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1873-76. Report on the Scientific Results of

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CONTENTS

Revision of the Tribe Archimantini (Mantodea: Mantidae: Mantinae)

G.A. Milledge neiden aiit edipi EI l
A new species of Progradungula Forster and Gray (Araneae: Gradungulidae) from Victoria
G.A. Milledges ili a ae RISO aa E DNA] 65

Supplement to a revision of Australian members of the bee genus Homalictus (Cockerell)
(Hymenoptera: Halictidae)
K. Walker... teve E aaa ea a I Tes ee IDEE OCT LIESS QT LT ete LUE 69

Descriptions of new Leptophlebiidae (Insecta: Ephemeroptera) from Australia. I. Tillyardophlebia

Phylogeny and biogeography of Australian genera of Chlorocystini (Insecta: Homoptera:

Tibicinidae)

AJ, de BOOP iis tete re edite ridado sa pos SCOTI Tene ae nee eae ee DI 91
The type species of A/eurodicus Douglas, a whitefly genus of economic importance (Homoptera:
Aleyrodidae)

JSH. PI OLTRE OR RORIS EE E PNEU EL 125

Additions to the taxonomy of the Limnoriidae (Crustacea: Isopoda)
LJ CDOKSONS. docce eee tenue RA AE CC SE CT AA DELE 129

A new genus of marine isopod (Crustacea: Flabellifera: Sphaeromatidae) from Australia and the
Indo-Pacific Ocean
NO l LON NN FE IE la tH CY V PCR ITO ELI a Ye pr demova neiro 145

Spermatological evidence supports the taxonomic placement of the Australian endemic hairy stone
crab, Lomis hirta (Decapoda: Anomura: Lomidae)
QU. Tudge uite RISORTO 235

A remarkable new pygmy seahorse (Syngnathidae: Hippocampus) from south-eastern Australia, with a
redescription of H. bargibanti Whitley from New Caledonia
ME GOMORRA IRE SN VENTI LL 245

A new grenadier, genus Trachonurus, from New Zealand and Australia (Pisces: Gadiformes:
Macrouridae)
T. Iwamoto and P, McMillan asas on RAR URINE 255