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RECORDS OF THE 
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8 June 2005 


RECORDS OF THE AUSTRALIAN MUSEUM 


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© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 153-164. ISSN 0067-1975 


Algal-tube Dwelling Amphipods in the Genus Cerapus 
from Australia and Papua New Guinea 
(Crustacea: Amphipoda: Ischyroceridae) 


J.K. Lowry* and P.B. Berents 


Australian Museum, 6 College Street, Sydney NSW 2010, Australia 
jimlowry@crustacea.net • pennyb@austmus.gov.au 


Abstract. Three new species of ischyrocerid amphipods in the genus Cerapus (C. bundegi, C. murrayae, 
and C. volucola ) are described from Australia and Papua New Guinea. Although these species have all 
of the morphological characteristics of Cerapus, they differ from other species in the genus in the 
construction of their tubes which are wrapped and parchment-like and sometimes elaborately covered 
with pieces of cut algae and seagrasses. 


Lowry, J.K., & RB. Berents, 2005. Algal-tube dwelling amphipods in the genus Cerapus from Australia and 
Papua New Guinea (Crustacea: Amphipoda: Ischyroceridae). Records of the Australian Museum 57(2): 153-164. 


Within Australia about 20 species (described and 
undescribed) occur in the Cerapus clade of Lowry & 
Berents, 1996 ( Bathypoma, Cerapus, Notopoma, Para- 
cerapus and Runanga). These species can be grouped by 
the type of tube they build. The most common tube type is 
made of minute sand grains and detritus held together with 
amphipod silk, known as detrital-tubes. The ends may be 
straight or fluted. In other species the females attach coarse 
sand grains to one end of the tube to form a holdfast that is 
buried in the substrate, known as anchor-tubes. The species 
described here make their tubes by wrapping algae into 
tubes, that may be simple or elaborately decorated with 
pieces of algae or seagrasses and these are known as algal- 
tubes. These species live on algae and the tubes are assumed 
to form an effective camouflage. Although the tubes are 
bizarre compared with those of other species of Cerapus, 
the morphology of the species in this group is similar to the 
type species C. tubularis Say, 1817 (Lowry & Berents, 
1989), which build their tubes of minute sand grains. In 
this paper we describe three new species (C. bundegi, C. 
murrayae, and C. volucola ) that build algal-tubes. 

* author for correspondence 


J.L. Barnard (1973) placed Ericthonius and related 
genera and Cerapus and related genera in the Ischyro¬ 
ceridae, but not Siphonoecetes and related genera, which 
he placed in the Corophiidae. Bousfield (1979; 1982) and 
Just (1983) maintained the classification of J.L. Barnard 
(1973). Lowry & Berents (1996) were able to demonstrate 
the monophyly of the combined Ericthonius, Cerapus and 
Siphonoecetes clades, but they were not able to link the 
group with a known family level taxon. Myers & Lowry 
(2003), in their revision of the corophiidean amphipods, 
co nfir med that the siphonoecetin clade (including the above 
three clades) is a sister taxon to the ischyrocerin clade which 
makes up the subfamily Ischyrocerinae. 

Diagnosis are generated with the aid of Intkey (Dallwitz 
et al., 1993 onwards; Dallwitz et al., 1998). Characters in 
bold face distinguish each taxon in at least two respects 
from every other taxon in the genus Cerapus. Material used 
in this study is lodged in the Australian Museum, Sydney 
(AM). The following abbreviations are used on the plates: 
A, antenna; D, dactylus; G, gnathopod; P, pereopod; PL, 
pleopod; UR, urosome. 

www.amonline.net.au/pdf/publications/1439_complete.pdf 


154 


Records of the Australian Museum (2005) Vol. 57 


Superfamily Photoidea Boeck, 1871 

(Myers & Lowry, 2003) 

Family Ischyroceridae Stebbing, 1899 

Subfamily Ischyrocerinae Stebbing, 1899 

(Myers & Lowry, 2003) 

Tribe Siphonoecetini Just, 1983 

(Myers & Lowry, 2003) 

Cerapus bundegi n.sp. 

Figs. 1-2 

Type material. Holotype, S, 4.77 mm, AM P62291; paratype, 2, 
3.85 mm, AM P62385; 41 paratypes, AM P62381; paratype, S, 3.54 
mm, AM P62383; PARATYPE, S, 2.74 mm, AM P62384; north of jetty, 
Cottesloe Beach, Perth, Western Australia, Australia, 31°59'S 115°45'E, 
just outside limestone reef, sand with scattered seagrass, 4 m, J. Just, 3 
April 1984, AU 28. 21 PARATYPES, AM P62382, off end of South Mole, 
Arthur Head, Fremantle, Western Australia, Australia, 32°3'S 115°44'E, 
brown & red algae, 6 m, R.T. Springthorpe, 25 December 1983, WA 274. 

Additional material. 2 specimens, AM P62292, Sargassum sp. and sand, 
2.3 m; 6 specimens, AM P62332, broad-leaf Sargassum sp., sand & algal 
bottom, 2.8 m; 3 juveniles, AM P62334, fine Dictyota sp. and sand, 2.6 
m; 2 specimens, AM P62338, broad-leaf Sargassum sp. and sand, 2.9 
m; Id, AM P62339, fine Sargassum sp. and sand, 2.9 m; Id, AM 
P62340, Padina sp., Lobophora sp. and sand, 2.7 m; 1 2, AM P62341, 
fine Dictyota sp. and sand, 2.3 m; 3 juveniles, AM P62387, Padina sp. 
and sand, 2.3 m; Id, AM P62392, Dictyotales and sand, 2.8 m; 4 
specimens, AM P62395, Sargassum sp. and sand, 2.7 m; 3 specimens, 
AM P62397, Dictyotales and sand, 2.8 m; 2 specimens, AM P62404, 
Sargassum sp. and sand, 2.9 m, west side of Malus Island, Dampier 
Archipelago, Western Australia, Australia, 26°30.61’S 116°38.92’E, R.A. 
Peart, 27 September 1999, WA 661. lcl, AM P62388, south side of 
Kendrew Island, Dampier Archipelago, Western Australia, Australia, 
20°20.63'S 116°31.99E, Sargassum sp. and sand, 3.8 m, R.A. Peart, 30 
August 1999, WA 694. 4 specimens, AM P62389, mixed brown algae, 
17 m, M. Hewitt; 3 juveniles, AM P62391, Dictyopteris sp., 17 m, P. 
Morrison, Nelson Rocks, Dampier Archipelago, Western Australia, 
Australia, 20°26.51'S 116°40.23'E, 7 September 1999, WA 715. ASS, 
AM P62386, Dictyopteris sp., intertidal zone, 0.5 m; 1 2, AM P62394, 
Padina sp., intertidal, 0.5 m, Tish Point, Rosemary Island, Dampier 
Archipelago, Western Australia, Australia, 20°29.67'S 116°35.89'E, R.A. 
Peart, 30 August 1999, WA 685. 11 specimens, AM P62335, beach at 
north end of Bundegi Reef, Exmouth Gulf, Western Australia, Australia, 
21°49’S 114°11’E, rocky rubble, coralline algae with green epiphyte, 2 
m, H.E. Stoddart, 4 January 1984. Many specimens, AM P62336, 
seagrass with bases and sand, 2 m; 12 specimens, AM P62398, mixed 
algae, 2 m, J.K. Lowry; many specimens, AM P62393, south Ned’s Camp, 
Cape Range National Park, Western Australia, Australia, 22°00'S 
113°55'E, seagrass, 2 m, R.T. Springthorpe, 31 December 1983. many 
specimens, AM P62337, Ned’s Camp, Cape Range National Park, 
Western Australia, Australia, 21°59'S 113°55E, green algae, 1.5 m, R.T. 
Springthorpe, 2 January 1984, WA 380. 6 specimens, AM P62290, 4 m, 
R.T. Springthorpe; 3 specimens, AM P62333, mixed coralline algae, 4 
m, J.K. Lowry, Red Bluff, Kalbarri, Western Australia, Australia, 27°42'S 
114°9'E, 10 January 1984. 4 specimens, AM P62286, 16 specimens, 
AM P62285, reef close to shore, Champion Bay, Geraldton, Western 
Australia, Australia, 28°45.88'S 114°36.83'E, Ecklonia radiata on 
shallow rocky reef at low tide, 0.5 m, R.A. Peart, 28 November 2000. 4 
specimens, AM P62396, Seven Mile Beach, Western Australia, Australia, 
29°11'S 114°53E, Amphibolis sp. (seagrass), 1 m, G.J. Edgar, 6 June 
1987. 2 specimens, AM P62402,10 specimens, AM P62406,5 specimens, 
AM P62403,22 2, AM P62405, Cottesloe Beach, Perth, Western Australia, 
Australia, 31°59'S 115°45E, sand patch on limestone reef with algae & 
seagrass ( Amphibolis sp.), 2 m, J. Just, 2-3 March 1984. 1 S, AM P62390, 
reef west of groyne, 2 km south of Cape Peron, Western Australia, Australia, 
32°16'S 115 0 4TE, Caulerpa sp. in deep channels in limestone reef, 6 m, 
J.K. Lowry, 26 December 1983, WA 302. 12, AM P62399, sponges, 6 m, 
R.T. Springthorpe; 5 specimens, AM P62400, Caulerpa sp., 6 m, J.K. Lowry; 
2 juveniles, AM P62401, orange gorgonacean, 6 m, R.T. Springthorpe, end 
of South Mole, Arthur Head, Fremantle, Western Australia, Australia, 32°3'S 
115°44E, 25 December 1983, WA 286. 


Diagnosis. Head, rostrum long, apically acute; anteroventral 
comer subquadrate. Antenna 1 peduncular article 1 longer 
than article 3, swollen along posterior margin. Pereonite 
2 with sternal keel. Pereopod 5 merus posterior lobe with 
4 plumose setae. 

Description. Based on holotype male, 4.77 mm. Head, 
rostrum long, apically acute, length 0.4x head; lateral 
cephalic lobe with ventral corner rounded, subocular margin 
weakly recessed, anteroventral corner subquadrate, ventral 
margin horizontal, posterior margin vertical. Antenna 1 long, 
length 0.5x body length; peduncular article 1 longer than 
(1.2x) peduncular article 3, not produced anterodistally and 
anteromedially, swollen along posterior margin, postero- 
distal corner not produced; flagellum 6-articulate; article 1 
short. Antenna 2 length 0.9x antenna 1; flagellum 5- 
articulate; article 1 short. Epistome and upper lip fused, 
straight. Mandible with palp article 2 long and slender, 
length 3.lx breadth, 1.2x article 3; palp article 3 slender, 
blade-like, long, 3.3x breadth. 

Pereon. Pereonite 1 without lateral keel; without sternal 
keel. Pereonite 2 with sternal keel. Pereonite 3 without 
sternal keel. Pereonite 5 length 2.lx depth. 

Gnathopod 1 coxa not fused to pereonite 1, length 1.4x 
depth, without anteroventral lobe; basis length 1.6x depth; 
carpus, length 1.2x depth with setose posterior lobe, broad; 
palm extremely acute, with barbed robust setae. Gnathopod 
2 carpochelate; coxa not fused to pereonite 2, length 1.7x 
depth, without anteroventral lobe or cusp; basis short, broad, 
length 1.3x breadth; carpus long, broad, length 1.2x breadth; 
palm shallowly excavate, anterodistal tooth small, located 
near articulation with propodus, posterodistal tooth well 
defined, medium, length 1.2x width; propodus broad, 
slightly curved, length 3.2x width, without proximal tooth 
on posterior margin, posterodistal corner smooth, without 
tooth; dactylus, length 0.7x propodus. Male gnathopod 2 
changes significantly through growth stages. Figure 1 
illustrates these changes for specimens ranging in size from 
2.74 to 4.77 mm. 

Pereopod 3 coxa not fused to pereonite 3, length 2.2x 
depth, without anteroventral lobe; basis, length 1.7x breadth, 
evenly rounded, with simple setae along anterior margin, 
without denticles along anterior margin; ischium long, 
length 1.6x breadth; merus short, length 0.8x breadth, 
without ridges. Pereopod 4 coxa not fused to pereonite 4, 
length 2x depth, without anteroventral lobe; basis length 
1.3x breadth, with simple setal group midway along anterior 
margin; ischium long, length 2x breadth; merus short, length 
lx breadth. Pereopod 5 coxa length 1.3x depth, without 
patches of small setae, without setae along ventral margin; 
merus with anterior lobe not extending beyond anterior 
margin of carpus, posterior lobe with 4 plumose setae; 
propodus with 1 seta along posterior margin; dactylus short, 
uncinate with one accessory hook. Pereopod 6 coxa without 
setal fringe ventrally, without patch of small setae near 
anterior margin; basis without patch of small setae near 
anterior margin; merus, length 1.2x breadth; dactylus short, 
uncinate, with two accessory hooks. Pereopod 7 coxa 
without posterodorsal lobe, without patch of small setae; 
merus, length 1.7x breadth; dactylus, short, uncinate, with 
two accessory hooks. 


Lowry & Berents: Algal-tube dwelling amphipods 155 


Fig. 1. Cerapus bundegi n.sp., holotype male, 4.77 mm (P62291), paratype female, 3.85 mm (P62385), paratype 
male “a”, 2.74 mm (P62384), paratype male “b”, 3.54 mm (P62383). South Mole, Arthur Head, Fremantle, Western 
Australia. Scales represent 0.1 mm. 


156 Records of the Australian Museum (2005) Vol. 57 


Fig. 2. Cerapus bundegi n.sp., holotype male, 4.77 mm (P62291), paratype female, 3.85 mm (P62385). South 
Mole, Arthur Head, Fremantle, Western Australia. Scales represent 0.1 mm. 


Pleon. Pleopods 1 to 3 decreasing in size. Pleopod 1 inner 
ramus 8-articulate; outer ramus 8-articulate, article 1 evenly 
swollen. Pleopod 2 biramous; inner ramus reduced, 1- 
articulate; outer ramus, broad, 1-articulate. Pleopod 3 present; 
inner ramus present, reduced; 1-articulate. Uropod 1 biramous, 
peduncle with distoventral corona of cuticular teeth, length 


1.4x outer ramus; rami with distoventral corona of cuticular 
teeth; outer ramus with lateral row of denticles, without medial 
setae, with 9 lateral setae, with large apical robust seta without 
smaller slender setae; inner ramus, length 0.5x outer ramus, 
without medial or lateral setae, with large apical robust seta 
without smaller slender setae. Uropod 2 uniramous, length of 


Lowry & Berents: Algal-tube dwelling amphipods 157 


peduncle 2.4x breadth, 3.8x ramus; ramus small, with 4 
denticles and 1 apical seta. Uropod 3 uniramous, peduncle 
length 1.5x breadth; ramus with 3 curved hooks. Telson length 
0.5x breadth, moderately cleft, 0.6x length, each lobe with 
17-18 anteriorly directed hooks, in 2 rows. 

Female (sexually dimorphic characters). Based on paratype 
female, 3.85 mm (P62385). Antenna 1 flagellum 4- 
articulate. Pereonite 2 without sternal keel. Pereonite 5 
length 1.5x depth. Gnathopod 1 coxa length 1.7x depth; 
basis length 2.3x depth; carpus, length l.lx depth with 
setose posterior lobe. Gnathopod 2 subchelate; coxa length 
1.9x depth; basis slender, length 1.8x breadth; carpus short, 
compressed, length 1.3x breadth. Pereopod 5 coxa length 
1.5x depth, with setae along ventral margin. Oostegites from 
gnathopod 2 to pereopod 5. 

Etymology. The species is named after Bundegi Reef, in 
the northern part of its range. 

Tube. Wrapped, parchment-like; seagrass and algae 
“wrapped” to form a tube. 

Habitat. Littoral, 0.5 to 6 m depth. 

Life-style. Epifaunal sediment, algal or seagrass dwellers. 

Remarks. Cerapus bundegi is a common, widespread 
species along the western Australian coast. The tubes of C. 
bundegi and C. volucola are almost identical, but 
morphologically the species differ in many ways. The most 
obvious differences are the shape of male gnathopod 2, and 
the number of dorsal recurved hooks on the telson (17-18 
in C. bundegi and 14-15 in C. volucola). 

Distribution. Western Australia: Malus Island, Kendrew 
Island, Nelson Rocks, Tish Point, Rosemary Island, all in 
the Dampier Archipelago; Bundegi Reef, Exmouth Gulf, 
Ned’s Camp, Cape Range National Park; Red Bluff, 
Kalbarri; Champion Bay, Geraldton; Seven Mile Beach; 
Cottesloe Beach, Perth; Cape Peron; South Mole, Arthur 
Head, Fremantle (20°S to 32°S). 

Cerapus murrayae n.sp. 

Figs. 3-6 

Type material. Holotype, 3, 7.04 mm, AM P31037; paratype, $, 
4.64 mm, AM P51220; PARATYPE, S, 4.32 mm, AM P51218; PARATYPE, 
6 , 3.04 mm, AM P51219; paratype, 6 , 6.08 mm, AM P61568; 
paratype, 6 , 8.9 mm, AM P62508; 36 paratypes, AM P61567; near 
bridge, Queenscliff Lagoon, Queenscliff, New South Wales, Australia, 
33°47.1'S 151°16.8'E, washed in at high tide on drift algae Lobophora 
variegatus ovZonaria, intertidal, A. Murray, 26 October 1980. PARATYPE, 
6, 5.08 mm, AM P61569, Balmoral Beach, New South Wales, Australia, 
32°49.6'S 151°15.2'E, C. Short, 3 m, sand, 7 September 1978. Paratype, 
6, AM P61570,150 mN of Horseshoe Bay, Trial Bay, New South Wales, 
Australia, 30°53’S 153°03'E, sediment and detritus from around reef 
edge, hand dredge, 7 m, R.T. Springthorpe, 15 June 1986. PARATYPE, 
S', AM P61571, Split Solitary Island, New South Wales, Australia, 
30°14.61'S 153°10.73'E, Zonaria sp., soft corals, bryozoans and sponges, 
18 m, P.B. Berents, J.K. Lowry and R. Peart, 12 February 2000. 

Diagnosis. Head, rostrum short, apically acute; 

anteroventral corner subquadrate. Antenna 1 peduncular 
article 1 longer than article 3, slightly swollen along 
posterior margin. Pereonite 2 without sternal keel. Pereopod 
5 merus posterior lobe with 7 plumose setae. 

Description. Based on holotype male, 7.04 mm (P31037). 
Head, rostrum short, apically acute, length 0.2x head; lateral 
cephalic lobe with ventral comer rounded, subocular margin 


deeply recessed, anteroventral corner subquadrate, ventral 
margin horizontal, posterior margin vertical. Antenna 1 long, 
length 0.5x body length; peduncular article 1 longer than 
(1.2x) peduncular article 3, not produced anterodistally and 
anteromedially, slightly swollen along posterior margin, 
posterodistal corner not produced; flagellum 6-articulate; 
article 1 short. Antenna 2 subequal in length to antenna 1; 
flagellum 6-articulate; article 1 long. Epistome and upper 
lip fused, straight. Mandible with palp article 2 long and 
slender, length 3.4x breadth, 1.4x article 3; palp article 3 
slender, blade-like, long, 3.lx breadth. 

Pereon. Pereonite 1 with lateral keel; without sternal keel. 
Pereonite 2 without sternal keel. Pereonite 3 without sternal 
keel. Pereonite 5 length 2x depth. 

Gnathopod 1 coxa not fused to pereonite 1, length 1.4x 
depth, without anteroventral lobe; basis length 1.8x depth; 
carpus length 1.3x depth with setose posterior lobe, broad; 
palm extremely acute, with simple robust setae. Gnathopod 
2 carpochelate; coxa not fused to pereonite 2, length 1.6x 
depth, without anteroventral lobe or cusp; basis short, broad, 
length 1.3x breadth; carpus long, broad, length 1.2x breadth; 
palm shallowly excavate, anterodistal tooth small, located 
near articulation with propodus, posterodistal tooth well 
defined, medium in size, length 1.4x width; propodus very 
broad, slightly curved, length 2.2x width, without tooth on 
posterior margin, posterodistal corner smooth, without 
tooth; dactylus, length 0.7x propodus. 

Pereopod 3 coxa fused to pereonite 3, with broad 
anteroventral lobe; basis, length 1.7x breadth, evenly 
rounded, with simple setae along anterior margin, without 
denticles along anterior margin; ischium long, length 2.8x 
breadth; merus short, length lx breadth, without ridges. 
Pereopod 4 coxa fused to pereonite 4, with anterior lobe; 
basis length 1.3x breadth, with simple setal group midway 
along anterior margin; ischium long, length 2. lx breadth; 
merus short, subequal in length 1 to breadth. Pereopod 5 
coxa length 1.2x depth, without patches of small setae, with 
setae along ventral margin; merus with anterior lobe 
extending beyond anterior margin of carpus, posterior lobe 
with 7 plumose setae; propodus with 3 setae along posterior 
margin; dactylus short, uncinate with one accessory hook. 
Pereopod 6 coxa without setal fringe ventrally, without patch 
of small setae near anterior margin; basis without patch of 
small setae near anterior margin; merus, length 2x breadth; 
dactylus short, uncinate, with two accessory hooks. 
Pereopod 7 coxa without posterodorsal lobe, without patch 
of small setae; merus, length 1.6x breadth; dactylus, short, 
uncinate, with two accessory hooks. 

Pleon. Pleopods 1 to 3 decreasing in size. Pleopod 1 inner 
ramus 9-articulate; outer ramus 3-articulate, article 1 with 
medial lobe. Pleopod 2 biramous; inner ramus reduced, 1- 
articulate; outer ramus, broad, 1-articulate. Pleopod 3 
present; inner ramus present, reduced; 1-articulate; outer 
ramus broad, 1-articulate. Uropod 1 biramous, peduncle 
with distoventral corona of cuticular teeth, length 1.5x outer 
ramus; rami with distoventral corona of cuticular teeth; outer 
ramus with lateral row of denticles, without medial setae, 
and with 5 lateral setae, with large apical robust seta without 
smaller slender setae; inner ramus, length 0.7x outer ramus, 
without medial or lateral setae, with large apical robust seta 
without smaller slender setae. Uropod 2 uniramous, length 
of peduncle 2.8x breadth, 5.8x ramus; ramus small, with 4 
denticles and 1 apical seta. Uropod 3 uniramous, peduncle 


158 Records of the Australian Museum (2005) Vol. 57 


B 


Lowry & Berents: Algal-tube dwelling amphipods 159 


Fig. 4. Cerapus murrayae n.sp., Queenscliff 
Lagoon, New South Wales: (A), dorsal view 
of head and pereonites 1-3; ( B ), tube. 


length 1.8x breadth; ramus with 3 curved hooks. Telson 
length 0.4x breadth, moderately cleft, 0.3x length, each lobe 
with 12-13 anteriorly directed hooks in 2 rows. 

Adult male. Based on paratype male, 8.9 mm (P62580). 
Antenna 1, flagellum 8-articulate. Antenna 2, flagellum 8- 
articulate. Pereonite 1 with sternal keel. Gnathopod 2 basis 
length 2.3x breadth; carpus length 1.6x breadth; palm with 
posterodistal tooth well defined, length l.lx width; 
propodus broad, curved, length 4x width. Male gnathopod 
2 changes significantly through growth stages. Figure 5 
illustrates these changes for specimens ranging in size from 
3.04 to 8.9 mm. 

Female (sexually dimorphic characters). Based on paratype 
female, 4.64 mm (P51220). Antenna 1, flagellum 5- 
articulate. Pereonite 1 without lateral keel. Pereonite 5 length 
1.4x depth. Gnathopod 1 coxa length 1.5x depth. 
Gnathopod 2 subchelate; coxa length 1.8x depth; basis 
length 1.6x breadth; carpus short, compressed, length 1.3x 
breadth; propodus palm extremely acute. Pereopod 3 coxa 
not fused to pereonite 3. Pereopod 4 coxa not fused to 
pereonite 4. Pereopod 5 coxa length 1.4x depth. 

Etymology. Named for Anna Murray (Australian Museum), 
who first discovered the species and brought it to our 
attention. 

I\ibe. Wrapped algae and seagrass; decorated with small 
pieces of several kinds of seagrass, algae and wood. 

Habitat. Littoral, 3 to 18 m depth. 

Life-style. Algal dwellers or epifaunal sediment dwellers. 


160 Records of the Australian Museum (2005) Vol. 57 


Fig. 5. Cerapus murrayae n.sp., holotype male, 7.04 mm (P31037), paratype female, 4.64 mm (P51220), paratype male “a”, 3.04 mm 
(P51219), paratype male “b”, 4.32 mm (P51218), paratype male “c”, 6.08 mm (P61568) and paratype male “d”, 8.9 mm (P62508), 
Queenscliff Lagoon, New South Wales. Scales for MD represent 0.05 mm; remainder represent 0.1 mm. 


Remarks. Cerapus murrayae is currently known only from 
the New South Wales central coast, the most restricted 
distribution of the three known species that make parchment¬ 
like tubes. These spectacular tubes, decorated with pieces 
of algae, distinguish C. murrayae from C. bundegi and C. 
volucola. Morphologically C. murrayae can be distinguish¬ 
ed from both C. bundegi and C. volucola by the shape of 
the male gnathopod 2 and the number of dorsal recurved 
hooks on the telson (17-18 in C. bundegi and 14-15 in C. 
volucola ) is different. 

Distribution. New South Wales : Horseshoe Bay, Trial Bay; 


Split Solitary Island; Queenscliff Lagoon, Queenscliff; 
Balmoral Beach, Middle Harbour. 

Cerapus volucola n.sp. 

Figs. 7-9 

Type material. Holotype, S, 3.20 mm, AM P62408; 12 
PARATYPES, AM P62407; paratype, S, 2.84 mm, AM 
P62409; paratype, $, 3.92 mm, AM P62410; False Orford 
Ness, northeast of Cape York, Queensland, Australia, 
11°23'S 142°52'E, rock, brown algae and sand, 4 m, D. 
Blake, 18 February 1979. 


Lowry & Berents: Algal-tube dwelling amphipods 161 


Additional material. 1 6 , AM P62419; 2 specimens, AM P62413, Padoz 
Natun reef, Madang Lagoon, Papua New Guinea, 5°09.60'S 145°48.77'E, 
Halimeda & epiphytic algae on rubble consolidated by sponges, 8 m, 
D. Gochfeld, 15 December 1993; 1$, AM P62414, Padoz Natun reef, 
Madang Lagoon, Papua New Guinea, 5°09.60'S 145°48.77'E, Halimeda 
& epiphytic algae on rubble consolidated by sponges, 8 m, D. Gochfeld, 
2 January 1994. 3 specimens, AM P62411, half way between Lizard 
Island and Carter Reef, Queensland, Australia, 14°37’S 145°33'E, 38 m, 
J.Leis, 26 November 1981. 5 specimens, AM P62417, Magnetic Island, 
Queensland, Australia, 19°8'S 146°50E, among Sargassum, 3 m, G. Edgar 
13 August 1981. 1 ovigerous female, AM P62415, north Wistari Reef, 
Capricorn Group, Great Barrier Reef, Queensland, Australia, 23°29'S 
151°53E, coral sand, 24 m, D. Fisk, 11 November 1978; 1 $, AM P62416, 
northwest Wistari Reef, Great Barrier Reef, Queensland, Australia, 23°29'S 
151°53'E, sand, 24 m,D. Fisk, 16 December 1978.1 $, AM P62418, Middle 
Reef, North Stradbroke Island, Queensland, Australia, 27°24.4'S 153°32E, 
Zonaria, 20 m, K.B. Attwood and E.L.A. Ho, 4 June 1993. 


Diagnosis. Head, rostrum long, apically acute; antero- 
ventral corner rounded. Antenna 1 peduncular article 1 
longer than article 3, swollen along posterior margin. 

Pereonite 2 without sternal keel. Pereopod 5 merus posterior 
lobe with 2 plumose setae. 

Description. Based on holotype male, 3.20 mm (P62408) 
and paratype female, 3.92 mm (P62410). With chocolate- 
coloured stripe around margin of head and dorsal surface 
of peduncle of antenna 1. Head, rostrum long, apically 
acute, length 0.3x head; lateral cephalic lobe with ventral 
corner rounded, subocular margin weakly recessed, 
anteroventral corner rounded, ventral margin horizontal, 
posterior margin vertical. Antenna 1 long, length 0.5x body 


162 Records of the Australian Museum (2005) Vol. 57 


Fig. 7. Cerapus volucola n.sp., male, False Orford Ness, northeast of Cape 
York, Queensland: (A), whole animal; ( B ), tube; (C), dorsal view of head. 


length; peduncular article 1 longer than (1.2x) peduncular 
article 3, not produced anterodistally and anteromedially, 
swollen along posterior margin, posterodistal corner not 
produced; flagellum 6-articulate; article 1 short. Antenna 2 
length lx antenna 1; flagellum 6-articulate; article 1 short. 
Epistome and upper lip fused, straight. Mandible with palp 
article 2 long and slender, length 2.7x breadth, l.lx article 
3; palp article 3 slender, blade-like, long, 3.5x breadth. 

Pereon. Pereonites 1 to 3 without lateral or sternal keels. 
Pereonite 5 length 1.6x depth. 

Gnathopod 1 coxa fused to pereonite 1, without 
anteroventral lobe; basis length 1.75x depth; carpus, length 
1.25x depth with setose posterior lobe, broad; palm 
extremely acute, with barbed robust setae. Gnathopod 2 
carpochelate; coxa fused to pereonite 2, without antero¬ 
ventral lobe or cusp; basis short, broad, length 1.35x 
breadth; carpus long, broad, length l.lx breadth; palm 
deeply excavate, anterodistal tooth large, located distal to 
articulation with propodus, posterodistal tooth well defined, 
medium, length 1.3x width; propodus very broad, slightly 
curved, length 2.5x width, without tooth on posterior 
margin, posterodistal corner smooth, without tooth; 
dactylus, length 0.7x propodus. 

Pereopod 3 coxa fused to pereonite 3, with narrow 
anteroventral lobe; basis, length 1.5x breadth, evenly 
rounded, with plumose setal group and simple setae along 
anterior margin, without denticles along anterior margin; 
ischium long, length 1.6x breadth; merus short, length lx 
breadth, without ridges. Pereopod 4 coxa fused to pereonite, 
with anterior lobe; basis length 1.3x breadth, with plumose 
setal group midway along anterior margin; ischium long, 
length 1.8x breadth; merus short, length lx breadth. 
Pereopod 5 coxa length 1.3x depth, without patches of small 
setae, without setae along ventral margin; merus with 
anterior lobe extending beyond anterior margin of carpus, 
posterior lobe with 2 plumose setae; propodus with 1 seta 


along posterior margin; dactylus short, uncinate with one 
accessory hook. Pereopod 6 coxa without setal fringe 
ventrally, without patch of small setae near anterior margin; 
basis without patch of small setae near anterior margin; 
merus, length 1.6x breadth; dactylus short, uncinate, with 
two accessory hooks. Pereopod 7 coxa without postero- 
dorsal lobe, without patch of small setae; merus, length 2.2x 
breadth; dactylus, short, uncinate, with two accessory hooks. 

Pleon. Pleopods 1 to 3 decreasing in size. Pleopod 1 inner 
ramus 7-articulate; outer ramus 5-articulate, article 1 evenly 
swollen. Pleopod 2 biramous; inner ramus reduced, 1- 
articulate; outer ramus, broad, 2-articulate. Pleopod 3 
present; inner ramus present, reduced; 1-articulate. Uropod 

1 biramous, peduncle with distoventral corona of cuticular 
teeth, length 1.3x outer ramus; rami with distoventral corona 
of cuticular teeth; outer ramus with lateral row of denticles, 
without medial setae, with 5 lateral setae, with large apical 
robust seta without smaller slender setae; inner ramus, length 
0.6x outer ramus, without medial or lateral setae, with large 
apical robust seta without smaller slender setae. Uropod 2 
uniramous, length of peduncle 2.5x breadth, 4.6x ramus; ramus 
small, with 4 denticles and 1 apical seta. Uropod 3 uniramous, 
peduncle length 1.5x breadth; ramus with 3 curved hooks. 
Telson length 0.4x breadth, moderately cleft, 0.6x length, each 
lobe with 14-15 anteriorly directed hooks in 2 rows. 

Female (sexually dimorphic characters). Based on paratype 
female, 3.92 mm (P62410). Gnathopod 1 coxa not fused to 
pereonite 1, length 1.5x depth; basis length 2.3x depth; 
carpus, length l.lx depth with setose posterior lobe, narrow. 
Gnathopod 2 subchelate; basis length 1.6x breadth; carpus 
length 1.3x breadth; propodus palm extremely acute; 
dactylus length 0.8x propodus. Oostegites from gnathopod 

2 to pereopod 5. 

Etymology. The species name is derived from the Latin 
words volumen for scroll and cola for inhabit. 


Lowry & Berents: Algal-tube dwelling amphipods 163 


Tube. Wrapped, parchment-like; seagrass and algae 
“wrapped” to form a tube. 

Habitat. Littoral, 3 to 38 m depth. 

Life-style. Algal or seagrass dwellers. 

Remarks. Cerapus volucola is a widespread species known 
from Madang Lagoon in northern Papua New Guinea, all 
along the Great Barrier Reef in northeastern Australia, to 
North Stradbroke Island in southern Queensland. The tubes 
of C. bundegi and C. volucola are almost identical, but 
morphologically the species differ in many ways. The most 
obvious differences are the shape of male gnathopod 2, and 
the number of dorsal recurved hooks on the telson (17-18 
in C. bundegi and 14-15 in C. volucola ). 


Distribution. Papua New Guinea : Padoz Natun reef, 
Madang Lagoon. Australia, Queensland: False Orford Ness, 
northeast of Cape York; Lizard Island and Carter Reef; 
Magnetic Island; Wistari Reef, Capricorn Group; Middle 
Reef, North Stradbroke Island (5°S to 27 °S). 


Acknowledgments. We thank the late Sharne Wiedland for her 
beautiful illustrations of the whole animals and tubes; Roger 
Springthorpe for assistance with illustrations. This research was 
supported by a grant from the Australian Biological Resources 
Study. 


164 Records of the Australian Museum (2005) Vol. 57 


Fig. 9. Cerapus volucola n.sp., holotype male, 3.20 mm (P62408). False Orford Ness, northeast of Cape York, 
Queensland. Scales represent 0.1 mm. 


References 

Barnard, J.L., 1973. Revision of Corophiidae and related families 
(Amphipoda). Smithsonian Contributions to Zoology 151: 1-27. 
Boeck, A., 1871. Crustacea Amphipoda borealia et arctica. 
Forhandlinger iVidenskabs-Selskabet i Christiania 1870: 81- 
280, i-viii [index]. 

Bousfield, E.L., 1979. A revised classification and phylogeny of 
amphipod crustaceans. Transactions of the Royal Society of 
Canada , series 4, 16: 343-390. 

Bousfield, E.L., 1982. Amphipoda. In McGraw-Hill Yearbook of 
Science & Technology, pp. 96-100. New York: McGraw-Hill. 
Dallwitz, M.J., T.A. Paine & E.J. Zurcher, 1993 onwards. User’s 
Guide to the DELTA System: A General System for Processing 
Taxonomic Descriptions. 4th edition. 

http://biodiversity.uno.edu/delta/ 

Dallwitz, M.J., T.A. Paine & E.J. Zurcher, 1998. Interactive keys. 
In Information Technology, Plant Pathology and Biodiversity, 
ed. P. Bridge, P. Jeffries, D.R. Morse & PR. Scott, pp. 201- 
212. Wallingford: CAB International. 

Just, J., 1983. Siphonoecetinae subfam. n. (Crustacea, Amphipoda, 
Corophiidae) 1: Classification. Steenstrupia 9(6): 117-135. 


Lowry, J.K., & PB. Berents, 1989. A redescription of Cerapus 
tubularis Say, 1817, based on material of the first reviewer, 
S.I. Smith, 1880, (Crustacea: Amphipoda: Corophioidea). 
Journal of Natural History 23: 1341-1352. 

Lowry, J.K., & P.B. Berents, 1996. The Ericthonius group, a new 
perspective on an old problem (Crustacea: Amphipoda: 
Corophioidea). Records of the Australian Museum 48(1): 75-109. 

www.amonline.net.au/pdf/publications/281_complete.pdf 

Myers, A.A., & J.K. Lowry, 2003. A phylogeny and a new 
classification of the Corophiidea. Journal of Crustacean 
Biology 23(2): 443-485. 

Say, T., 1817. On a new genus of the Crustacea, and the species 
on which it was established. Journal of the Academy of Natural 
Sciences of Philadelphia 1: 49-52. 

Stebbing, T.R.R., 1899. Revision of Amphipoda (continued). 
Annals and Magazine of Natural History, series 7, 4: 205- 
211 . 


Manuscript received 14 June 2002, revised 1 August 2004 and accepted 
15 September 2004. 

Associate Editor: G.D.F. Wilson. 


© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 165-178. ISSN 0067-1975 


A Review of the Australian Fossil Storks 
of the Genus Ciconia (Aves: Ciconiidae), 
With the Description of a New Species 


Walter E. Boles 


Terrestrial Zoology, Australian Museum, 

6 College Street, Sydney NSW 2010, Australia, and 

School of Biological, Earth and Environmental Sciences, 
University of New South Wales NSW 2052, Australia 

walterb@austmus.gov.au 


Abstract. Only a single species of stork, the Black-necked Stork Ephippiorhynchus (= Xenorhynchus) 
asiaticus, occurs in Australia today, and is known from several fossil localities from the Early Pliocene. 
Two species of smaller fossil storks are also known, one previously named and one described here. The 
former, found in the Darling Downs, southeastern Queensland, was named Xenorhynchus nanus De Vis, 
1888. Some later authors suggested that this species should be transferred to the living genus Ciconia; 
this decision is confirmed here, the name for this species becoming Ciconia nana. The second species of 
small stork comes from several Late Oligocene and Early Miocene sites at Riversleigh, northwestern 
Queensland. This taxon is referred to the genus Ciconia and distinguished as a new species, C. 
louisebolesae. It constitutes the earliest record of the Ciconiidae from Australia. 


Boles, Walter E., 2005. A review of the Australian fossil storks of the genus Ciconia (Aves: Ciconiidae), with 
the description of a new species. Records of the Australian Museum 57(2): 165-178. 


The classification of living storks (Ciconiidae) by Kahl 
(1979) admitted 17 species in six genera in three tribes, 
whereas that of Hancock etal. (1992) recognized 19 species 
in six genera in two tribes. The family is represented in 
Australia by a single living species, the Black-necked Stork, 
or Jabiru, Ephippiorhynchus (= Xenorhynchus auct.) 
asiaticus (Latham, 1790). 

Storks are rather well represented in the world fossil 
record, although no comprehensive review of them has been 
attempted. The earliest records come from the Late Eocene 
of Egypt (Ciconiidae gen. and sp. indet. and Leptoptilos sp. 
indet.) (Miller et al., 1997). After taxa incorrectly referred 
to this family were removed (Olson, 1985), the earliest 
named species became Palaeoephippiorhynchus dietrichi 
Lambrecht, 1930 (Late Oligocene; Egypt). The identity of 


the older Eociconia sangequanensis Hou, 1989 (Middle 
Eocene; China) as a stork needs to be confirmed (Unwin, 
1993). Other Tertiary-aged storks are known from North 
America, Europe and Asia (references in Olson, 1985; 
Bickart, 1990). Quaternary-aged palaeospecies are known 
for several extant genera. 

The fossil record of this family in Australia has not been 
studied in detail. Much of the Australian fossil stork material 
is comparable in size and morphology to E. asiaticus. 
Specimens assigned to this species are known from Pliocene 
and Pleistocene localities in northeastern and southeastern 
Queensland and northeastern South Australia (Archer, 1976; 
Baird, 1991a; Boles & Mackness, 1994; Molnar & Kurz, 
1997; Vickers-Rich, 1991). 

www.amonline.net.au/pdf/publications/1440_complete.pdf 


166 Records of the Australian Museum (2005) Vol. 57 


The first stork reported from Australia was described by 
C.W. De Vis, who named several species (De Vis, 1888, 
1892,1905); however, all but Xenorhynchus nanus are now 
known to have been misidentified to family (amended family 
identifications summarized by van Tets & Rich, 1990). 
Material of a new species of stork from Oligo-Miocene 
deposits at Riversleigh, northwestern Queensland, was 
mentioned briefly in the literature with little elaboration 
(Boles, 1991, 1997; Vickers-Rich, 1991). It is the purpose 
of this study to review X. nanus and the undescribed 
Riversleigh stork. Both are here considered to belong to 
the extant genus Ciconia. This genus has an extensive fossil 
record. Three of the living species of Ciconia have been 
recorded from Quaternary deposits (Brodkorb, 1963). 
Several fossil taxa have been assigned to Ciconia, but many 
are based on single specimens. A large, but unnamed species 
of Ciconia from the Late Miocene-Early Pliocene of Arizona 
is known from numerous skeletal elements (Bickart, 1990), 
as is another large form, C. maltha L. Miller, 1910, from 
the Quaternary of North America and Cuba (Miller, 1910; 
Howard, 1942; Feduccia, 1967). Other palaeospecies 
include C. stehlini Janossy, 1992 (Early Pleistocene, 
Hungary, tarsometatarsi, tibiotarsi, ulna, phalanges), C. 
gaudryi Lambrecht, 1933 (Late Pliocene of Greece, 
humerus), C. minor Harrison, 1980 (Late Miocene, Kenya, 
distal femur) and C. sarmatica Grigorescu & Kessler, 1977 
(Late Miocene, Romania, proximal carpometacarpus). 
Lambrecht (1933) cited records of indeterminate species 
of Ciconia from the Pleistocene of California and Late 
Pliocene of France, and Olson & Rasmussen (2001) 
recorded two indeterminate species from North Carolina, 
one Middle Miocene in age, the other Early Pliocene. Late 
Pleistocene or Quaternary reports of this genus include those 
by Ono (1984; Honshu, Japan; Ciconia sp.), Steadman et 
al, (1994; northeast Mexico; Ciconia sp. or Mycteria sp.) 
and Suarez & Olson (2003; Cuba; Ciconia sp.). 

Species of Ciconia and Mycteria are rather generalized 
in their morphology compared to the large, long-legged 
Ephippiorhynchus and Jabiru, the heavy-bodied Leptoptilos 
and somewhat aberrant Anastomus. Any fossil stork remains 
not exhibiting characters of these more distinctive genera 
were frequently allocated to one of the more “typical” ones. 
The problem of deciding whether a fossil form based on 
single or fragmentary elements has been correctly assigned 
to genus is compounded by the heavy reliance by the current 
taxonomy on behavioural (Kahl, 1972, 1979; Slikas, 1998) 
or molecular characters (Slikas, 1997). 

Materials and methods 

Taxonomic nomenclature follows Kahl (1979). Osteological 
terminology follows Baumel & Witmer (1993), except that 
as terms of position and direction anterior is used rather 
than rostral or cranial and posterior rather than caudal. Most 
of the measurements follow the methods of Steadman 
(1980) or van den Driesch (1976), and were made with 
digital calipers and rounded to the nearest 0.1 mm. 

Several factors hamper the ease of using the fossil record 
of storks from elsewhere for evaluating that of Australia. 
Generic-level taxonomy of the Ciconiidae has changed 
substantially, with several formerly monotypic genera now 
merged with others. New palaeogenera were often based 
on material that exhibited some morphological intermediacy 


between two nominal genera that have since been 
synonymised; this is particularly so in the expanded concept 
of Ciconia. The more inclusive generic concepts result in a 
broader morphological range across the constituent species, 
into which the palaeospecies may fit comfortably. Published 
diagnoses of such fossil forms must be assessed with caution 
because some of the characters may no longer apply to the 
genus sensu lato. 

Another difficulty is that many of the species of fossil 
storks have been based on isolated fragments, confounding 
comparison between nominal taxa for which common 
osteological elements are not known. Moreover, many extant 
taxa are poorly represented in skeletal collections and of 
those specimens that do exist, individuals from zoos form a 
high proportion. In addition to any developmental 
abnormalities the latter may have, most likewise lack 
provenance and are frequently unsexed. 

Osteological diagnosis of Ciconiidae 

The skeletal elements can be recognized as belonging to 
this family on the basis of the following suites of characters. 
Diagnoses are restricted to those portions of the elements 
represented by the fossils, both here for the family and 
subsequently for generic level taxa in the respective species 
accounts. 

Cranium. The lateral indentations at the orbits are shallow 
(in dorsal view); fossae glandulae nasalis are absent. The 
processus postorbitalis is long, and the temporal fossae well 
defined and rather extensive posteriorly. There is a single 
small circular fontanelle orbitocranialis situated at the 
posterior border of septum interorbitalis where it joins the 
braincase. 

Quadrate. The anterior and posterior borders of the blade¬ 
like processus orbitalis are straight or slightly tapering 
through most of its length. The process is more or less 
straight (in posterior view) but not strongly flattened, with 
the distal end somewhat inflated; it is not incised posteriorly, 
twisted nor inflected medially or ventrally. The processus 
oticus is broad and not compressed laterally; the processus 
mandibularis is deep mediolaterally (in ventral view). The 
condylus medialis and combined condyli lateralis and 
caudalis are long and thin, and converge laterally at an acute 
angle; the sulcus intercondylaris is moderately large, 
particularly on its medial half. The short, broad projection 
of the condylus lateralis extends anteriorly along the lateral 
side, at its anterior end supporting the cotyla quadratojugalis, 
which is located just above the posteroventral border of the 
element; the part of the projection between the cotyla and 
the posterior end of the quadrate comprises about half of 
its length. 

Humerus. The element has a pronounced sigmoid 
curvature, with a particularly marked anterior bend in the 
distal end (in dorsal view). Proximal end. In anterior view, 
the long axis and distal border of the caput humeri are 
oriented dorsodistally-ventroproximally; the caput humeri 
is moderately short. The sulcus ligamentum transversus and 
incisura capitis are deep. The tuberculum dorsale is distinct 
and triangular. The fossa pneumotricipitalis is large. The 
distal margin of the crista bicipitalis forms a nearly right 


Boles: fossil birds of the stork genus Ciconia 167 


angle with the shaft. The intumescentia humeri is inflated, 
particularly distally. The crista deltopectoralis is prominent, 
with its apex more or less level with the distal end of the 
crista bicipitalis. Distal end. The fossa musculus brachialis 
is large and deep, particularly ventrodistally, and is angled 
sharply dorsoproximally-ventrodistally relative to the shaft. 
The tuberculum supracondylare ventrale is elongate and 
situated along a prominent ridge. The epicondylus ventralis 
is strongly produced as a triangular projection. The 
epicondylus dorsalis and processus flexorius are rudiment¬ 
ary. The ventral side of the distal end is flat (in anterior 
view) with the processus supracondylare dorsalis prominent, 
angling moderately to very abruptly to shaft. The fossa 
olecrani is broad and shallow, and extends proximally from, 
and dorsoventrally across, the condylus ventralis humeri. 

Ulna. Proximal end. The proximal end is straight in relation 
to the shaft, i.e. there is no inflection from the midline of 
the shaft. The margins of the impressio m. brachialis are 
pronounced, with the anterior margin the more extensive 
distally. The tuberculum lig. collateralis ventralis is slightly 
bulbous but does not overhang the impressio m. brachialis 
and has a relatively short distal extension along its border. 
The incisura radialis is more proximodistally oval (narrower, 
longer) than circular and the impressio m. scapulotricipitalis 
is small with little distal extension. 

Tibiotarsus. Proximal end. The proximal end is deeper than 
wide because the region level with the incisura tibialis 
between the cristae cnemialis and the facies articularis is 
elongated. The surface is mostly level, with a small to at 
most moderate rise towards the cristae cnemialis. The cristae 
cnemialis are not strongly developed proximally, but are 
rather broad (in proximal view). They form a more or less 
90° angle, and from this junction, the crista cnemialis 
lateralis is about twice the length of the crista cnemialis 
cranialis. The crista cnemialis cranialis is situated towards, 
but not at, the medial edge, with only a slight indentation 
separating them; it is long distally, angling smoothly into 
the shaft (in medial view). There is an expanded articular 
surface at the end of the crista cnemialis lateralis with a 
flattened anterolateral face, which projects both antero- 
medially and posterolaterally (in proximal view). Distal end. 
The shaft is long, thin and straight, with the posterior surface 
rounded and the anterior surface flattened for most of its 
length, taken up by a very broad and shallow sulcus 
extensorius, which deepens for a short extent just proximal 
of the pons supratendineus. There is a large, prominent 
papilla for M. tibialis cranialis centred directly proximal to 
the area intercondylaris and level with the distal border of 
the pons supratendineus. The pons supratendineus is 
restricted to the medial half of shaft, with its distal border 
strongly developed into a ridge. The scar on the lateral face 
of the shaft is large and proximodistally elongated. The 
sulcus m. fibularis is moderately deep. The distal end of the 
element has little mediolateral expansion, and the medial 
border of the shaft does not flare strongly outwards proximal 
to the condylus medialis. The condyli lateralis and medialis 
are more or less parallel and directly distal to the respective 
borders of the shaft, are longer anteroposteriorly than 
proximodistally, and have about the same distal extension; 
the condylus lateralis extends further proximally. The 
condylus medialis is notched distally. The area inter¬ 


condylaris is a deep circular pit centred on the midline of 
shaft, extending between the pons supratendineus and the 
condylus medialis. The sulcus intercondylaris is deep (in 
distal view) and the trochlea cartilaginis tibialis is shallow 
with prominent borders. 

Tarsometatarsus. Proximal end. The eminentia inter- 
cotylaris is narrow, with the lateral border abrupt and the 
medial one sloping (in dorsal view). In proximal view, the 
rims of the cotylae are rounded and (in dorsal view) the 
medial rim of cotyla medialis is blunt or rounded. The 
hypotarsi comprises two parallel cristae hypotarsi separated 
by a single large sulcus hypotarsi, which is deep throughout 
its length; it is centred mediolaterally on the plantar face. 
There is no small secondary groove within the sulcus 
hypotarsi. Distal end. The sulcus extensorius occupies the 
greater part of the length of the anterior surface, making 
the distal third of the shaft relatively flat, and then angling 
from the midline of the shaft to the lateral side at the distal 
end, extending into the foramen vasculare distale but not 
beyond that into the incisura intertrochlearis lateralis. The 
fossa metatarsi I is a long proximodistally elongated oval, 
terminating distally on a ridge extending towards the 
trochlea metatarsi II. The fossa supratrochlearis plantaris is 
markedly excavated lateral to this ridge. The trochleae are 
not inflated proximally nor do they join the shaft abruptly; 
the shaft bulges laterally just proximal to the trochlea 
metatarsi IV, meeting it with relatively little demarcation. 
The trochleae form a shallow but obvious curve (in distal 
view). The trochleae metatarsi II and IV are more or less 
equal in length and shorter than trochlea metatarsi III. 

Genus Ciconia Brisson, 1760 

Ciconia Brisson (1760). Ornithologia sive Synopsis Methodica, 
1: 48, 361—type species: Ciconia =Ardea ciconia Linnaeus, 
1758. 

In the original concept of Ciconia, the genus comprised 
two species, C. ciconia Linnaeus, 1758 (Eurasia, Africa) 
and C. nigra Linnaeus, 1758 (Eurasia, Africa). The generic 
limits were expanded by Kahl (1979) and Wood (1983, 
1984) to incorporate three species that were long kept in 
monotypic genera: (Sphenorhynchus) abdimii Lichtenstein, 
1823 (Africa), (Dissoura) episcopus Boddaert, 1783 
(Africa, southern Asia), and (Euxenura) maguari Gmelin, 
1789 (South America). This has considerably expanded the 
size range of the species in both directions and added 
variability in the morphology. 

The represented elements can be diagnosed as Ciconia 
and separated from those of other genera of storks by the 
following suites of characters: 

Cranium. Most of the characters on which a generic 
diagnosis might be based are missing in the fossil. It does 
permit separation from Ephippiorhynchus by having the 
fossae temporalis moderately shallow and moderately 
concealed by the cristae temporalis, rather than deep and 
unconcealed (in dorsal view); the nuchal area (supra- 
occipital) is slightly convex around the prominentia 
cerebellum, rather than somewhat concave; and the crista 
nuchalis transversus is low and does not project posteriorly 
beyond the extent of the prominentia cerebellum. In these 
characters, the fossil agrees with Ciconia. 


168 Records of the Australian Museum (2005) Vol. 57 


Quadrate. The processus oticus is thin (in lateral view). The 
processus orbitus is thin. The sulcus between the processus 
mandibularis and condylus pterygoideus is moderately deep. 
The processus mandibularis is markedly longer mediolaterally 
than anteroposteriorly, the condyli form an acute angle and 
the sulcus intercondylaris is relatively narrow. 

Humerus. The tuberculum ventrale is situated distal to the 
caput humeri. The fossa pneumotricipitalis does not extend 
proximally well beyond the attachment for M. scapulo- 
humeralis caudalis nor as far distally past the midpoint of 
the crista bicipitalis. The intumescentia humeri is moderately 
inflated distally. There is a slight notch where the distal end 
of the crista bicipitalis joins the shaft, but the sulcus nervus 
coracobrachialis is obsolete. The impressio m. coraco- 
brachialis is flat, not depressed. The dorsal edge of the crista 
deltopectoralis is generally straight, not concave. The 
attachment for M. scapulohumeralis cranialis is situated at 
the proximal end of the linea m. latissimus dorsi rather than 
ventral to it and directly distal to the fossa pneumo¬ 
tricipitalis. In dorsal view, the dorsal side of the shaft 
posterior to the condylus dorsalis is shallow, with the 
anterior and posterior sides straight and roughly parallel, 
forming a rectangular surface; the anterior face of the shaft 
meets anteroproximal corner of the processus supracon- 
dylaris dorsalis gradually; and the tuberculum supra- 
condylare dorsale is not strongly developed. The epicon- 
dylus ventralis is moderately produced (in anterior view). 
The scar for M. pronator profundus is moderately short and 
shallow and that for M. flexor carpi ulnaris is moderately 
small; thus the area of the ventral side distal to the 
epicondylus ventralis is not markedly excavated (in ventral 
view) and the epicondylus ventralis is less undercut (in 
anterior view). The sulcus humerotricipitalis is moderate 
in width. The condylus ventralis humeri extends further 
distally relative to the condylus dorsalis humeri (in anterior 
view); in distal view, its posterior surface faces more 
posteriorly and less distally. The ventrodistal corner, distal 
to the epicondylus ventralis, is only slightly to moderately 
excavated. 

Ulna. The condition of the ulnar fragment considered in 
this study is not suitable for useful comparisons between 
taxa. This element is not diagnosed further. 

Tibiotarsus (taken in part from Howard, 1942 and Olson, 
1991). There are limited characters of the proximal end that 
are useful in separating the genera of storks, and most of 
these are related to the angles and extent of the cristae and 
articular surfaces. On the distal end, the tuberculum 
retinaculi m. fibularis proximal to the condylus lateralis 
forms a prominent triangular ridge, which is pointed 
proximally and broadens distally (prominent papilla in 
Ephippiorhynchus ); the proximomedial corner of the 
condylus lateralis is not incised by expansion of the area 
intercondylaris; the proximomedial border of the condylus 
medialis lacks a prominent round fossa; the posterior sides 
of the condyli extend prominently and are more oval than 
circular (in lateral view); the distal border of pons 
supratendineus is horizontal (tilted or arched in Ephippio¬ 
rhynchus ); the distal opening of the canalis extensorius is 
moderately to strongly horizontally elongate (rounded in 
Ephippiorhynchus ); and the incisura intercondylaris is broad 
and relatively flat at its base (in distal view). 


Tarsometatarsus (taken in part from Howard, 1942). The 
hypotarsus is slender relative to the proximal width of the 
cotylae and to the length of crista hypotarsi lateralis, the 
longer of the cristae; the cristae hypotarsi are slender. The 
eminentia intercotylaris is situated on the proximodistal 
midline rather than medial to it, and the lateral side of its 
base is only slightly excavated, if at all. The area between 
the cotylae and the hypotarsus consists of a gradual drop 
with a pit of moderate depth proximal to the cristae 
hypotarsi. The ridge leading to the distal end of the 
hypotarsus is generally low and broad. The cotyla lateralis 
is elongate; the cotyla medialis much more circular (in 
proximal view). The trochlea metatarsi II is situated dorsally 
and is little rotated laterodorsally-medioplantarly. The fossa 
metatarsi I is flush with the surface of the bone or only 
slightly elevated. 

Ciconia nana (De Vis, 1888) 

Fig. 1 

Xenorhynchus nanus De Vis, 1888. Proc. Linn. Soc. N.S.W. 3: 

1287, Qld: Darling Downs: Condamine River: Chinchilla. 
Ciconia nana (De Vis, 1888). Rich & van Tets, 1982: 306A; van 

Tets, 1984: 470; van Tets & Rich, 1990: 166; Vickers-Rich, 

1991: 752. 

De Vis (1888) based Xenorhynchus nanus on material from 
the Condamine River, near Chinchilla, in the Darling Downs 
of Queensland. The material comprised a distal tibiotarsus, 
collected by J. Daniels, and a proximal ulna, a later 
acquisition but described at the same time. Subsequently 
De Vis (1905) reported this species from Wurdulumankula, 
Cooper Creek, South Australia, based on a distal tibiotarsus, 
collected by Professor J. Gregory. The original tibiotarsal 
fragment was designated as the lectotype by Brodkorb 
(1963). Similarities in size and shape to species of Ciconia 
were noted by Rich & van Tets (1982), who provisionally 
transferred this form to this genus, where it has been 
listed in subsequent reviews (e.g., van Tets, 1984; van 
Tets & Rich, 1990; Vickers-Rich, 1991), although no 
detailed comparisons had been made. 

The Darling Downs, southeastern Queensland, feature 
deposits of two discrete periods. Pleistocene deposits occur 
on the east side of the Condamine River (26°48'S 150°41'E), 
producing the Darling Downs Local Fauna (Molnar & Kurz, 
1997). The older, Pliocene-aged assemblage, the Chinchilla 
Local Fauna, which yielded the holotype of X. nanus, comes 
from the fluviatile Chinchilla Sands along the western banks 
of the Condamine River, near Chinchilla. On the basis of 
closer faunal resemblances of this fauna to the Early 
Pliocene Bluff Downs Local Fauna than to the Pleistocene 
Darling Downs Local Fauna, its age has been put at Early 
to Middle Pliocene (T. Rich et al., 1991). 

Many important specimens from Gregory’s trip along 
Cooper Creek have their locality listed as Wurdulumankula, 
although no similar place name has been found on Gregory’s 
maps (Gregory, 1906), and the exact location of this site is 
uncertain (Tedford & Wells, 1990). It is considered to be 
located in the Piranna Soakage of Cooper Creek, in the 
eastern Lake Eyre basin, South Australia, and to be one of 
a number of sites from which fossils of the Malkuni Fauna 
have been recovered, one of two faunas in the fluviatile 
deposits of the Katipiri Formation (Tedford & Wells, 1990). 


Boles: fossil birds of the stork genus Ciconia 169 


Fig. 1. Specimens of fossil stork Ciconia nana. (A-C) lectotype (QM FI 131), distal right tibiotarsus; (A) anterior view; ( B ) lateral 
view; (C) medial view; ( D ) paralectotype (QM F5514), proximal right ulna, anterior view. Scale = 10 mm. 


These assemblages represent fluviatile/lacustrine faunas in 
the Great Artesian basin that predate the formation of inland 
dunes and are probably Late Pleistocene in age (Woodburne 
et al., 1985). The fossils are found as “float” or in place. 

Lectotype. Distal tibiotarsus (QM FI 131; Fig. la-c; 
Brodkorb, 1963). 

Type locality. North bank of the Condamine River, 5 km 
from Chinchilla, Darling Downs, Queensland. 

Paralectotype. Proximal ulna (QM F5514, Fig. Id). 

Etymology. Nanus (Latin, a dwarf), in reference to the small 
size of this bird in comparison with its putative congener, 
E. (X.) asiaticus. 

Diagnosis. Ciconia nana is diagnosed from other living 
species in the genus by the following combination of 
characters: the anterior extension of the condylus lateralis 
relative to the condylus medialis is greater; the ridge from 
the papilla for M. tibialis cranialis to the condylus medialis 
is only slightly incised by a furrow; and the tuberculum 
retinaculi m. fibularis is nearly confluent with the lateral 
border of the shaft (in anterior view). 

Of the fossil forms, few can be compared with C. nana 
for lack of common elements. Ciconia maltha was much 
larger (Table 1), as was the unnamed Mio-Pliocene species 
from Arizona, which Bickart (1990) characterized as a 
“giant, equalling in size large individuals of the extant Jabiru 
mycteria , \ The somewhat younger Ciconia stehlini had a 
tibiotarsus of comparable proximal width to C. nana. 
Janossy (1992) did not provide any characters that serve to 
separate these species morphologically. Of living species, 
C. ciconia and C. nigra are similar to C. nana in proximal 
width of the tibiotarsus but the condyli of C. nana are deeper. 


Referred material. Distal tibiotarsus (QM F5513), 
Wurdulumankula, Cooper Creek, South Australia. 

Measurements. Table 1. 

Description 

De Vis’ (1888) description of the original tibiotarsus (QM 
FI 131) was detailed, and identified diagnostic characters 
at family, generic and specific levels. Descriptions of the 
other specimens amounted to just a few adjectives. The 
second tibiotarsus “adds nothing to our information about 
the smaller Jabiru than that it attained a rather larger size 
than the tibia already described”. Its distal width was greater 
and “all parts of the bone are proportionately larger” (De 
Vis, 1905). About the ulna De Vis (1888) remarked only 
that it was of compatible size with the first tibiotarsus but 
“unfortunately its worn condition unfits it for description”. 

Tibiotarsus. The lectotypical tibiotarsal fragment QM 
FI 131 consists of the distal end with the shaft broken distal 
to the crista fibularis (length 118.6 mm as preserved). It is 
abraded on the proximal borders of both condyli, the cristae 
of trochlea cartilaginis tibialis, the epicondylus medialis and 
the papilla for M. tibialis cranialis. The anterior face of the 
shaft is flattened, with the linea extensorius developed into 
a low ridge along its distal end. The posterior face is strongly 
rounded. The tuberculum retinaculi m. fibularis is confluent 
with the lateral border of the anterior face of the shaft; a 
nutrient foramen is proximal to this. There is a large, broad, 
elongate scar on the lateral face of the shaft; the distal end 
is level with the tuberculum. The tuberositas retinaculi 
extensorius on the linea extensorius is small but obviously 
elongate. The sulcus extensorius is of uniform depth, not 
deepening markedly proximal to the pons supratendineus. 
The distal border of the pons supratendineus is developed 


170 Records of the Australian Museum (2005) Vol. 57 


Table 1. Measurements (mm) of the tibiotarsus of Xenorhynchus nanus and other fossil and living species of 
storks, giving mean, standard deviation, range and sample size (in parentheses), f indicates fossil taxa; values for 
Ciconia stehlini and C. maltha from Janossy (1992) and Howard (1942), respectively. QM, Queensland Museum, 
Brisbane. 


distal width 

depth, 

depth, 


across 

condylus 

condylus 


condyli 

lateralis 

medialis 

f Ciconia nana 


QMF1131 (lectotype) 

14.3 

19.5 

18.5 

QM F5513 

16.1 

20.6 

20.5 

Ciconia ciconia 

15.3; 0.7 

18.5; 0.8 

18.1; 1.0 


14.5-16.2 (4) 

17.4-19.2 (4) 

16.6-18.9 (4) 

Ciconia nigra 

14.8 (1) 

18.4 (1) 

17.5 (1) 

Ciconia maguari 

18.2; 1.3 

21.8; 1.0 

21.7; 0.7 


17.0-19.5 (3) 

20.9-22.9 (3) 

20.9-22.3 (3) 

Ciconia episcopus 

11.6; 0.5 

13.5; 0.8 

13.4; 0.7 


11.0-12.0 (3) 

12.3-14.2 (3) 

12.4-13.6 (3) 

Ciconia abdimii 

10.9; 0.8 

12.6; 0.6 

12.7; 0.7 


9.7-12.1 (6) 

11.8-13.7 (6) 

11.8-13.9 (6) 

f Ciconia stehlini 

14.5 

— 

— 

f Ciconia maltha 

18.0-21.5 

— 

— 

Anastomus lamelligerus 

11.3; 0.6 

13.0; 0.5 

13.0; 0.5 


10.6-11.8 (3) 

12.6-13.5 (3) 

12.7-13.6 (3) 

Anastomus oscitans 

10.8 (1) 

11.8(1) 

12.0(1) 

Mycteria ibis 

13.3; 0.5 

17.1; 0.6 

17.2; 1.0 


12.6-14.2 (6) 

16.1-18.3 (6) 

16.2-18.9 (6) 

Mycteria leucocephala 

13.1; 0.4 

16.6; 1.4 

16.9; 1.0 


12.7-13.4 (3) 

15.3-18.1 (3) 

16.0-18.0 (3) 

Mycteria cinerea 

13.2 (1) 

18.1 (1) 

18.0 (1) 

Mycteria americana 

13.5; 0.6 

17.0; 1.2 

16.9; 1.4 


12.6-13.8 (4) 

16.0-18.5 (4) 

15.6-18.6 (4) 

Leptoptilos dubius 

20.3; 0 

26.5; 0.4 

25.9; 0 


20.3 (2) 

26.2-26.8 (2) 

(2) 

Leptoptilos javanica 

16.4 (1) 

20.5 (1) 

19.6 (1) 

Leptoptilos crumeniferus 

19.1; 0.8 

23.4; 0.8 

23.8; 0.6 


18.1-20.1 (4) 

22.3-23.7 (4) 

22.9-24.3 (4) 

Jabiru mycteria 

21.1; 0.2 

27.4; 0.1 

28.4; 0.3 


20.9-21.3 (3) 

27.3-27.5 (3) 

28.1-28.7 (3) 

Ephippiorhynchus senegalensis 

18.1; 1.1 

25.0; 1.6 

25.4; 1.3 


17.1-19.5 (4) 

23.7-27.3 (4) 

24.5-27.3 (4) 

Ephippiorhynchus asiaticus 

17.1; 0.9 

23.0; 1.5 

22.9; 1.2 


15.6-18.4 (9) 

21.5-25.2 (9) 

21.6-24.8 (9) 


anteriorly into a strong ridge. The condyli are similar in 
shape and size, with the condylus medialis extending slightly 
further distally and more markedly anteriorly. The depressio 
epicondylus lateralis is deeper and more extensive than the 
depressio epicondylus medialis; both have prominent rims 
anteriorly. Despite abrasion, the cristae of the trochlea 
cartilaginis tibialis are prominent, extending well away from 
the shaft. Measurements, Table 1. 

The other tibiotarsal fragment, QM F5513, is slightly 
larger and has less abrasion of the distal end. It retains about 
half the length of the shaft, as does the lectotype (length 
62.6 mm as preserved) and the medial half is missing for 
much of this. It agrees closely with the lectotype in 
morphology other than that the sulcus extensorius is slightly 
deeper, the tuberositas retinaculi extensorius is more raised, 
the tuberculum retinaculi m. fibularis is a small distance 
from the lateral border rather than confluent with it, and 
the distal opening of the canalis extensorius is somewhat 
larger and rounder. 


Ulna. Specimen QM F5514 consists of the proximal end of 
a right ulna. It is rather damaged, with the olecranon missing, 
and moderate to heavy abrasion on the cotyla dorsalis, facies 
articularis radialis and crista intercotylaris. It is broken distal 
to the proximalmost papilla. The impressio m. brachialis is 
long, moderately deep proximally and shallow distally. It 
is bounded posteriorly by a broad, rounded tuberculum lig. 
collateralis ventralis and anteriorly by a heavy ridge, which 
separates it from a prominent incisura radialis. Although, 
because of the abrasion, measurements of this element 
cannot be compared directly with those of other taxa, overall 
the specimen is slightly smaller than the ulna of Ciconia 
ciconia. The measurements of the specimen as preserved 
are, length 46.0 mm; proximal width 16.6 mm; proximal 
depth 11.8 mm. 

De Vis (1888) placed this species in the same genus as 
the living Ephippiorhynchus asiaticus, “noting further its 
strong resemblance to the Jabiru’s tibia in the massiveness, 
direction, and sculpture of the bridge traversing the 


Boles: fossil birds of the stork genus Ciconia 


171 


intercondylar space, we cannot but admit congeneric affinity 
between the two”. A comparison of the distal tibiotarsal 
fragments with other living species of the Ciconiidae, and 
E. asiaticus in particular, demonstrates that De Vis’ generic 
allocation for these specimens is not supported. The 
comparative material available to De Vis was limited to 
selected taxa, almost all of Australian origin. Thus, his 
placement of the fossil specimens in the same genus as the 
only Australian species is not surprising; it is doubtful that 
he had access to osteological representatives of any other 
genera of storks. 

De Vis (1888) noted size differences between the fossil 
tibiotarsus and that of E. asiaticus: “the fossil tibia ... is in 
the mean two-ninths less in its dimensions than the recent 
bone, indicating a bird but little more than half the bulk of 
the jabiru of the present day”. The morphological differences 
he mentioned were that “the rotular channel is shallower; 
there is considerably less intercondylar space behind the 
posterior edge of the bridge, the canal under the bridge is 
relatively much wider, the ectocondylar tubercle is not 
prominent, and the double flexure inwards and forwards 
apparent in the living jabiru between the shaft and the 
articular end is scarcely appreciable”. Most of these are 
either actually differences between Ciconia and Ephippio- 
rhynchus or have no generic significance. 

The condition of the ulnar fragment is not suitable to permit 
a useful comparison. The shallow, round impressio m. 
scapulotricipitalis may be of generic significance but it is also 
likely that the possible slight morphological differences 
between the fossil and recent specimens are due to abrasion. 

Riversleigh stork 

The presence of a new species of stork from Oligo-Miocene 
deposits at Riversleigh, was briefly mentioned by Vickers- 
Rich (1991). Boles (1991) noted that it “was not close to 
the living ... Xenorhynchus” without further elaboration, 
and subsequently (Boles, 1997) stated that the material was 
“probably referable to Ciconia , \ This taxon is described 
below. 

The Riversleigh deposits are located 5 km west of the 
Riversleigh homestead (19°02'S 138°45'E), 200 km north 
of Mt Isa, northwestern Queensland, where they occur as 
an outcrop of Tertiary limestone overlying the Cambrian 
Thorntonia Limestone. There are now over 200 named 
Oligo-Miocene deposits at Riversleigh. An informal system 
of grouping has been used (Systems A-C). These systems 
are “regionally clustered sites that appear to be super- 
positionally-related (differing in age but not significantly 
in position) and/or space-related (spatially isolated but 
approximately contemporaneous)” (Archer etal., 1989). The 
principal accumulations are thought to have occurred in several 
episodes involving large lakes, shallow pools and cave 
deposits. Undoubted stork material has been recovered from 
three sites and a referred specimen comes from a fourth. 

White Hunter Site, Hal’s Hill Sequence, D-Site Plateau, 
is considered to be part of System A, of Late Oligocene/ 
Early Miocene age (Creaser, 1997; Myers & Archer, 1997). 
The White Hunter Local fauna also contains other birds, 
including the small casuariid Emuarius gidju (Patterson & 
Rich, 1987) (Boles, 1992), the dromornithid Barawertoris 
tedfordi Rich, 1979, a flightless rail (Boles, 2005) and 
several passerines. Wayne’s Wok Site is in the central section 


of the D-Site Plateau. Its age is still unclear, but may be 
System A or B (?Early to Middle Miocene) (Black, 1997; 
Cooke, 1997a; Creaser, 1997). Birds found here also include 
Emuarius, dromornithids and passerines. Bitesantennary 
Site is a cave deposit in the Verdon Creek Sequence, on the 
northern section of the D-Site Plateau, where it intrudes 
into the widespread D-Site layer. It is possibly a System B 
site (Cooke, 1997a). 

Specimens of five skeletal elements were obtained at 
Bitesanntennary Site in close proximity and are assumed to 
have been associated. Proximal tarsometatarsal fragments 
from White Hunter Site and Wayne’s Wok Site allow direct 
comparison with each other but not with a distal tarsometa¬ 
tarsal fragment from Bitesantennary Site. Because the 
tarsometatarsal fragments all come from storks of 
comparable size and morphology, they are referred to the 
same taxon. A cervical vertebra from Neville’s Garden Site 
(Early Miocene) is tentatively referred to this species 
because of its size and morphological similarity to that of 
living storks. 

Ciconia louisebolesae n.sp. 

Fig. 2 

Holotype. QM F30290, right distal humeral fragment with 
surface damage to the anterior face of the condylus dorsalis, 
tuberculum supracondylare ventrale and dorsal border of 
sulcus humerotricipitalis. 

Type locality. Bitesantennary Site, Riversleigh, north¬ 
western Queensland, currently considered to be Early 
Miocene; Bitesantennary Local Fauna. 

Paratypes. All from Bitesantennary Site. Cranium —QM 
F20910, neurocranium, lacking skull roof; quadrate —QM 
F20893, complete right element; humerus —QM F20911, 
proximal right element broken through distal to the midpoint 
of the crista bicipitalis and to the fossa pneumotricipitalis, 
and missing the tuberculum dorsalis and processus 
deltopectoralis; tibiotarsus —QM F31350, extreme proximal 
left element broken through the shaft through proximal end 
of the foramen interosseum proximale; damage to most 
projecting features, including both the cristae cnemialis, 
particularly the crista cnemialis caudalis, the medial edge 
of the facies articularis medialis, and extensively on the 
posterior edge along the area of contact between the facies 
articularis medialis and lateralis; Tarsometatarsus —QM 
F36446, right distal fragment broken through the shaft 
proximal to the fossa metatarsi I. 

Etymology. Dedicated with love and respect to my mother, 
Louise Boles, for her guidance in my development as a 
person and her tolerance of my many transgressions. 

Diagnosis. Similar in size to C. ciconia and C. nigra, but 
sufficiently different to recognize as a new species. It differs 
by the following suite of characters of the distal humerus: 
the condylus ventralis humeri is proportionally smaller, not 
extending as far ventrally; the epicondylus ventralis is closer 
to the distal end; the muscle scars distal to the epicondylus 
ventralis are more extensive (in ventral view) and more 
excavated (in anterior view), although less than in genera 
such as Ephippiorhynchus; and the posterodorsal corner is 


172 Records of the Australian Museum (2005) Vol. 57 


Fig. 2. Specimens of the fossil stork Ciconia louisebolesae. (A) partial skull (QM F20910, Bitesantennary Site), lateral view; (B) right 
quadrate (QM F20893, Bitesantennary Site), lateral view; (C) proximal right humerus (QM F20911, Bitesantennary Site), posterior 
view; ( D-E ) distal right humerus (holotype: QM F30290, Bitesantennary Site), (D) anterior view, (E) posterior view; ( F-G ) proximal 
left tarsometatarsus (QM F36447, Wayne’s Wok Site), (F) anterior view, (G) lateral view; (//-/) distal right tarsometatarsus (QM 
F36446, Bitesantennary Site), ( H) dorsal view, (/) plantar view. Scale = 10 mm. 


more rounded (in dorsal view). The crista bicipitalis is not 
short (as in C. maguari ) and is prominently extended. In its 
size and comparative narrowness of the incisura tibialis, 
the tibiotarsus more closely resembles that of the small 
species of Ciconia (e.g., episcopus and abdimii ) rather than 
C. ciconia or the larger taxa. 

Referred material. QM F50428 (Neville’s Garden Site) 
cervical vertebra with only minor damage; QM F36445 
(Wayne’s Wok Site) left proximal tarsometatarsus with 
damage to the crista hypotarsi medialis and plantar face of 
cotyla lateralis; QM F36447 (White Hunter Site) proximal 
left tarsometatarsus missing most of hypotarsus. 

Measurements. Tables 2-4. 

Description 

Cranium. Measurements as preserved: 31.7 mm width of 
parietal; 40.1 mm width at midline of orbits (front of 
specimen as preserved); 40.7 mm width at processes 
temporalis. The specimen comprises the rear of the skull 
missing the dome of skull dorsal to the fossa temporalis 
and in a line just across top of the orbitae, retaining most of 
the parietal and frontal on a level with the orbital rims; the 
lamina parasphenoidalis is present but palatal elements are 
missing; the interorbital septum is present to in front of the 
fontaculi interorbitales and about half way through the 
orbits. Many of the external structures are damaged, with 
the processus postorbitalis lost and the posteroventral border 
of the processus paraoccipitalis and edges of the lamina 
parasphenoidalis slightly to strongly abraded; a large 
opening in the centre of the septum interosseus is a post¬ 


mortem artefact. The orbitae are broad and rounded. The 
fossa temporalis is broad, with the crista temporalis nuchalis 
strongly developed. The processus zygomaticus is short and 
blunt. The foramen magnum is squarish oval, with the dorsal 
border broken. The condylus occipitalis is hemispherical, 
with the dorsal border abraded. The recessus tympanicus 
dorsalis is moderately small and round; the fenestra 
vestibule, foramen pneumaticum caudale and fenestra 
cochleae are tightly grouped in a small recess. The cotylae 
quadratica otici and squamosi are of similar circular shape 
and size. The interior of the cranial cavity is largely intact. 
The fossa cranii caudalis and fossa tecti mesencephale are 
broad and circular, with the crista tentorialis prominent. 

Quadrate. Viewed anterodorsally, the element is very 
transversely expanded across the processus mandibularis 
and less so across the processus oticus; the midbody is 
comparatively thin between these processes, with the lateral 
side markedly concave. The crista tympanicum is low. The 
processus orbitalis is straight on its anterior border; its 
posterior border is straight until dorsal end, which curves 
to meet the anterior border at the apex. The capitula 
squamosum and oticum are elliptical, their long axes 
converging anteriorly (in posterodorsal view). The incisura 
intercapitularis is broad but shallow; it extends to a large 
round foramen pneumaticum located centrally on the 
posterodorsal surface. The region from the base of the 
capitulum squamosum, through this foramen, diagonally 
towards the base of the condylus lateralis is anteroposteriorly 
compressed compared to the rest of the element. The 
posterior surface between the processus mandibularis and 
processus oticus is very slightly concave (in lateral view). 


Boles: fossil birds of the stork genus Ciconia 


173 


Table 2. Measurements (mm) of the quadrate of Ciconia 
louisebolesae and living species of storks, giving mean, standard 
deviation, range and sample size (in parentheses). QM, Queensland 
Museum, Brisbane. 


greatest length, 

greatest depth, 

processus oticus 

processus orbitalis 

through processus 

through processus 

mandibularis 

mandibularis 


Ciconia louisebolesae 


QM F20893 

20.2 

20.2 

Ciconia ciconia 

22.3; 0.8 

20.5; 1.0 


21.7-23.2 (3) 

19.3-21.7 (4) 

Ciconia niger 

20.6; 0.2 

19.3; 0.1 


20.4-20.7 (2) 

19.2-19.4 (2) 

Ciconia maguari 

24.2; 0.5 

23.7; 2.7 


23.7-24.6 (3) 

21.8-26.8 (3) 

Ciconia episcopus 

19.2; 1.0 

17.4; 1.0 


18.8-20.8 (5) 

15.8-18.6 (6) 

Ciconia abdimii 

17.5; 0.5 

15.1; 0.6 


17.0-18.0 (4) 

14.5-15.8 (4) 

Anastomus lamelligerus 

20.1 (1) 

17.5; 0.4 


17.2-17.8 (2) 


Mycteria ibis 

23.5; 0.3 

24.7; 0.7 


23.3-23.9 (3) 

24.5-25.3 (4) 

Mycteria leucocephala 

23.8; 0.7 

24.9; 0.6 


23.3-24.3 (2) 

24.5-25.3 (2) 

Mycteria americana 

23.4; 1.3 

25.2; 2.0 


22.5-24.3 (2) 

23.8-26.8 (2) 

Leptoptilos crumeniferus 35.1 (1) 

34.0 (1) 

Jabiru mycteria 

35.6; 0.1 

39.1; 0.2 


36.5-36.6 (3) 

38.9-39.3 (3) 

Ephippiorhynchus 

29.6; 1.9 

28.6; 2.1 

senegalensis 

28.2-30.9 (2) 

26.7-31.1 (4) 

Ephippiorhynchus 

29.2; 2.1 

2.3; 1.5 

asiaticus 

26.9-31.5 (4) 

25.8-29.1 (4) 


The condylus medialis is the most medially projecting part 
of the element, while the projection of the condylus lateralis 
is the greatest laterally, exceeding that of the processus 
oticus. The condyli lateralis/caudalis lie perpendicular to 
the processus orbitalis. The small, oval condylus ptery- 
goideus stands discrete from the condylus medialis, 
separated by a small sulcus. 

Cervical vertebra. This is very elongate (greatest length 
39.7 mm, proximal width 26.2 mm, proximal depth 19.8 
mm, distal width 18.6 mm, distal depth 19.5) and agrees 
with modern Ciconia ciconia in morphology. It is not 
diagnosed further. 

Humerus. In anterior view, the caput humeri is rounded, 
sloping dorsally from the apex more steeply than on the 
ventral side; it is moderately broad, becoming expanded at 
its anteroposterior midpoint. Its distal border on the posterior 
surface is obscure. The sulcus lig. transversus is broad and 
deep but short, confined to the ventral third of the anterior 
face, barely reaching the ventral border of the caput; it ends 
abruptly at the edge of the anterior face of the element. Its 
depth is augmented by a bordering ridge on its proximal 
side and the intumescentia humeri on the distal. The 
proximal surface of the intumescentia is flat and smooth 
and (in proximal view) is little inflated; it is tilted dorsally 
to join smoothly with the shallow, indistinct impressio 
coracobrachialis. 


On the posterior surface, the broad incisura capitis 
separates the caput humeri from a long, deep and thickened 
area along the proximoventral border of the element, 
proximal to the fossa pneumotricipitalis, before merging 
with the proximal end of the posterior shaft surface without 
an obvious demarcation. Ventrally the prominent proximal 
section attenuates towards the crista bicipitalis, but protrudes 
substantially more posteriorly than the crista. On its ventral 
end, it supports the elongate scar for M. biceps brachii. 

The tuberculum ventrale is relatively large, triangular- 
trapezoidal in shape and oriented proximodorsally. In 
proximal view, it sits at the apex of a triangular block 
bounded along its dorsal base by the sulcus lig. transversus. 
This triangular block protrudes posteriorly well beyond the 
rest of the proximal end of the element. The deep but 
moderately thin crus dorsale fossae runs distally from this, 
forming the dorsal border of the fossa pneumotricipitalis. The 
fossa is elliptical and highly pneumatic. The area between it 
and the crus ventrale fossae is excavated as a broad concave 
basin. The thick crus has a well marked, elongate scar for the 
attachment of M. scapulohumeralis caudalis. The crista 
bicipitalis is not well-developed ventrally. 

The ridges bordering the sulcus scapulotricipitalis are 
short but moderately pronounced; they do not extend to the 
distal border. The ridge on the ventral side of the sulcus 
humerotricipitalis is broader and longer but about the same 
height as those defining the sulcus scapulotricipitalis. The 
sulcus humerotricipitalis is broad and shallow, with no 
obvious separation from the flat posterior surface of the 
shaft proximal to it; it is apparent only because of the 
bordering ridges. The fossa m. brachialis becomes confluent 
with the shaft on its dorsal and proximal sides and is poorly 
demarcated on its ventral side. The ridge along the ventral 
border supporting the tuberculum supracondylare ventrale 
is strongly developed and rather broad. In distal view, the 
condylus dorsalis humeri extends further anteriorly than 
does this ridge, which is turn projects only slightly more 
than the condylus ventralis humeri. The distal border of the 
condylus dorsalis humeri just reaches the dorsodistal corner. 
The condylus ventralis humeri is a rounded oval without 
obvious inflation. The incisura intercondylaris is moderately 
broad but shallow. The scars for M. flexor carpi ulnaris and 
M. pronator profundus face distally; they are adjacent, 
separated by only a thin ridge. The scars for M. ectepicon- 
dylo-ulnaris and M. extensor digitorum communis are 
shallower and less extensive. 

Tibiotarsus. The specimen is too damaged to permit 
standard measurements for comparison with other taxa. The 
following measurements, taken from the specimen as 
preserved, give an indication of the general size: proximal 
width 15.5 mm; proximal depth (measured from the facies 
articularis medialis to the crista cnemialis lateralis in a direct 
anteroposterior line) 20.2 mm; in both cases, the actual value 
would be considerably greater, particularly for the latter 
measurement. 

The crista cnemialis cranialis is broken; the remaining 
base is straight and does not extend far distally, although 
some distance further than the crista cnemialis lateralis. 
Their junction, at about a right angle, is marked by a shallow 
sulcus intercnemialis. The crista patellaris is straight, in both 
proximal and lateral views. The incisura tibialis is deeply 
incised, concave and moderately narrow; at its proximal 


174 Records of the Australian Museum (2005) Vol. 57 


Table 3. Measurements (mm) of the humerus of Ciconia louisebolesae and other fossil and living species of 
storks, giving mean, standard deviation, range and sample size within parentheses, f indicates fossil taxa; values 
for Ciconia maltha from Howard (1942). QM, Queensland Museum, Brisbane. 


proximal width 

distal width 

depth, condylus 
dorsalis humeri 

Ciconia louisebolesae 

QM F20911 

32.9 


QM F30290 

— 

28.2 

14.9 

Ciconia ciconia 

39.6; 1.7 

30.6; 0.8 

16.1; 1.0 


37.2-40.9 (6) 

28.8-31.5 (10) 

15.2-17.3 (10) 

Ciconia boyciana 

— 

33.8 (1) 

18.9 (1) 

Ciconia nigra 

40.1; 1.1 

30.6; 0.6 

16.4; 0.8 


38.9^10.9 (3) 

30.0-31.2 (2) 

15.6-17.2 (2) 

Ciconia maguari 

43.7; 1.0 

33.5; 1.8 

17.9; 0.8 


42.7^14.7 (4) 

31.6-36.1 (5) 

16.6-18.5 (5) 

Ciconia episcopus 

32.2; 2.8 

23.9; 1.2 

12.8; 0.8 


28.2-35.6 (6) 

22.1-24.6 (4) 

11.7-13.6 (4) 

Ciconia stormi 

— 

21.9 (1) 

11.4(1) 

Ciconia abdimii 

28.3; 1.4 

22.9; 0.5 

12.3; 0.1 


26.5-30.0 (5) 

22.4-23.2 (3) 

12.2-12.4 (3) 

Ciconia maltha t 

46.2-53.3 

38.7-40.7 

— 

Anastomus lamelligerus 

26.5; 0.0 

21.6; 0.9 

11.5; 0.2 


26.3-26.6 (2) 

20.6-22.3 (3) 

11.3-11.7 (3) 

Anastomus oscitans 

— 

20.0; 1.3 

11.4; 0.6 

Mycteria ibis 

34.6; 2.1 

18.6-21.1 (3) 

26.3; 1.2 

10.9-12.0 (3) 

14.3; 0.7 


33.1-38.2 (5) 

24.5-27.5 (4) 

13.8-15.3 (4) 

Mycteria cinerea 

— 

27.2; 2.2 

15.0; 1.1 

Mycteria leucocephala 

31.9; 2.1 

25.6-28.7 (2) 

28.9; 0.9 

14.2-15.8 (2) 

14.9; 0.7 


31.7-32.0 (2) 

28.2-29.5 (2) 

15.4-16.4 (2) 

Mycteria americana 

34.1; 2.9 

27.6; 1.6 

14.9; 0.8 


31.0-36.7 (3) 

25.9-29.7 (18) 

13.0-16.5 (18) 

Leptoptilos dubius 

53.0 (1) 

46.8 (1) 

24.7 (1) 

Leptoptilos crumeniferus 

48.8; 0.7 

47.7; 3.3 

25.4; 1.6 


48.4^19.6 (3) 

43.5-51.9 (7) 

23.5-28.1 (7) 

Leptoptilos javanicus 

45.0; 3.9 

36.1; 2.3 

19.7; 0.9 


42.6-49.5 (3) 

34.4-38.7 (3) 

19.0-20.8 (3) 

Jabiru mycteria 

54.1; 1.4 

40.8; 3.2 

22.4; 1.9 


52.9-55.7 (3) 

34.6-45.7 (10) 

18.2-25.5 (10) 

Ephippiorhynchus senegalensis 

46.6; 2.2 

36.1; 1.3 

18.7; 0.5 


44.0-48.9 (4) 

35.1-37.0 (2) 

18.3-19.0 (2) 

Ephippiorhynchus asiaticus 

44.1; 1.8 

35.9; 2.7 

19.2; 1.6 


42.1-41.0 (9) 

32.9-37.6 (9) 

18.0-20.8 (9) 


end, it merges with a shallow excavation on the proximal 
surface. Its posterior border flares abruptly medially as the 
anterior side of the rather narrow and strongly rounded facies 
articularis lateralis. Much of the proximal surface of the 
facies is missing but what remains indicates that this was 
markedly raised. The circular fossa retropatellaris is deeper 
and smaller than the flattened and shallowly concave facies 
articular medialis. The circular attachment for M. 
femorotibialis medialis is on the end of the crista medialis, 
facing posteromedially. The facies gastrocnemialis is 
relatively flat from the medial side of the crista cnemialis 
cranialis along the medial side of the shaft, before curving 
smoothly into the posterior face. 

General agreement in morphology and the association 
of the Riversleigh tibiotarsal fragment with other stork 
material from Bitesantennary Site are used as the basis for 
allocating it to this taxon. 


Tarsometatarsus. The eminentia intercotylaris is directed 
anteroproximally at about 50° from the horizontal, 
projecting further anteriorly than the cotylae. The rounded 
anterior side is elliptical (in anterior view) with its long 
axis directed proximodistally; from the apex, it slopes 
posterodistally on its posterior side. The lateral side drops 
directly to the cotyla lateralis; the medial side extends as a 
long sloping ridge, confluent with the anterior rim of the 
cotyla medialis to past its mediolateral midpoint. The apex 
is slightly broader than the base (in medial view). 

The cotylae are roughly equal in both their anterior and 
posterior extents and thus in overall depth. In proximal view, 
the cotyla medialis is roughly circular over most of its 
border, with the anterior side flattened. Posterior to the 
anterior rim, the cotyla meets medial side of the eminentia 
intercotylaris in a smooth, gradual slope. The medial rim 
projects strongly proximally to a narrow edge. In anterior 


Boles: fossil birds of the stork genus Ciconia 


175 


Table 4. Measurements (mm) of the tarsometatarsus of Ciconia louisebolesae and other fossil and living species of storks, giving 
mean, standard deviation, range and sample size (in parentheses), f indicates fossil taxa; values for Ciconia maltha from Howard 
(1942). QM, Queensland Museum, Brisbane. 


proximal 
width across 
cotylae 

depth, 

cotyla 

lateralis 

depth, 

cotyla 

medialis 

distal 

width 

medial depth, trochlea 
metatarsi metatarsi metatarsi 

II III IV 

greatest 

distal 

depth 

Ciconia louisebolesae 

QM F36445 c. 18.2 


9.2 


QM F36447 

16.3 

7.9 

9.4 

— 

— 

— 

— 

— 

QM F36446 

— 

— 

— 

19.6 

9.6 

9.0 

8.5 

14.3 

Ciconia ciconia 

17.4; 1.5 

9.0; 0.9 

9.1; 0.6 

19.9; 1.3 

8.8; 0.4 

9.3; 0.6 

8.5; 0.5 

15.3; 1.0 


14.2-19.3 (9) 

7.4-10.1 (9) 

8.4-9.8 (9) 

17.7-21.7 (9) 

8.3-9.6 (9) 

8.3-10.2 (9) 

7.8-9.3 (9) 

13.5-16.6 (9) 

Ciconia nigra 

16.2; 0.4 

8.6; 0.4 

8.4; 0 

18.9; 0.5 

7.0; 0.3 

8.9; 0.3 

7.6; 0.2 

15.1; 0.2 


15.9-16.7 (3) 

8.2-8.9 (3) 

8.4 (3) 

18.4-19.3 (3) 

6.6-7.3 (3) 

8.5-9.1 (3) 

7.5-7.8 (3) 

14.9-15.2 (3) 

Ciconia maguari 

21.3; 0.4 

11.2; 0.5 

10.1; 0.7 

23.2; 0.9 

10.0; 0.8 

10.9; 0.3 

10.0; 0.3 

18.0; 1.2 


20.7-21.6 (4) 

10.7-11.7 (4) 

9.3-10.8 (4) 

22.2-24.2 (4) 

9.4-11.2 (3) 

10.5-11.2 (3) 

9.5-10.3 (3) 

17.0-19.7 (3) 

Ciconia episcopus 

14.4; 1.3 

7.8; 0.8 

7.8; 0.7 

16.7; 1.1 

7.2; 0.3 

8.0; 0.6 

7.2; 0.6 

12.2; 0.7 


12.8-16.2 (6) 

7.1-8.9 (6) 

6.7-8.9 (6) 

15.7-18.5 (6) 

6.4-7.5 (6) 

7.3-8.9 (6) 

6.5-8.0 (6) 

11.5-13.0 (6) 

Ciconia abdimii 

12.5; 0.9 

7.0; 0.4 

6.6; 0.4 

14.9; 1.1 

6.8; 0.6 

7.3; 0.4 

6.3; 0.4 

10.7; 0.8 


11.1-13.5 (5) 

6.6-7.4 (5) 

6.2-7.1 (5) 

13.9-15.6 (5) 

6.2-1.1 (5) 

6.1-1.5 (5) 

5.9-7.0 (5) 

9.6-11.8 (5) 

f Ciconia maltha 

20-24.5 

— 

— 

23-28 

— 

— 

— 

— 

Anastomus 

13.4; 0.2 

7.0; 0.6 

7.4; 0.1 

14.9; 0.6 

7.1; 1.3 

7.0; 0.5 

5.7; 1.3 

10.6; 1.3 

lamelligerus 

13.2-13.5 (2) 

6.5-7.4 (2) 

7.3-7.5 (2) 

14.4-15.3 (2) 

6.2-8.0 (2) 

6.6-7.3 (2) 

4.8-6.6 (2) 

9.7-11.5 (2) 

Mycteria ibis 

15.4; 0.8 

8.8; 0.9 

8.9; 1.0 

18.9; 0.8 

8.4; 0.9 

9.1; 0.6 

7.8; 0.5 

16.1; 1.5 


14.6-16.2 (4) 

7.7-9.6 (4) 

7.8-9.9 (4) 

18.2-19.9 (4) 

7.4-9.3 (4) 

8.4-9.7 (4) 

7.1-8.4 (4) 

14.1-17.4 (4) 

Mycteria 

14.8; 0.7 

9.3; 0.8 

9.3; 0.3 

17.8; 0 

7.8; 1.3 

8.8; 0 

8.0; 0.2 

15.4; 0.4 

leucocephala 

14.3-15.3 (2) 

8.7-9.9 (2) 

9.1-9.5 (2) 

17.8 (2) 

6.9-8.7 (2) 

8.8 (2) 

7.8-8.1 (2) 

15.1-15.7 (2) 

Mycteria 

15.5; 0.3 

9.3; 0.2 

9.2; 0.1 

18.7; 1.5 

8.4; 0.7 

8.8; 0.9 

8.1; 0.4 

16.4; 0.8 

americana 

15.2-15.9 (2) 

9.1-9.4 (2) 

8.7-9.2 (2) 

17.4-19.7 (2) 

7.2-8.9 (2) 

7.9-9.1 (2) 

7.6-8.8 (2) 

14.5-18.2 (2) 

Leptoptilos dubius 

23.8 (1) 

12.8 (1) 

12.9 (1) 

28.1 (1) 

12.4 (1) 

14.0 (1) 

11.7 (1) 

22.9 (1) 

Leptoptilos 

22.0; 0.9 

13.1; 0.3 

12.1; 0.9 

26.6; 1.0 

12.1; 1.0 

13.5; 0.4 

11.3; 0.9 

20.7; 1.4 

crumeniferus 

20.8-23.2 (5) 

12.8-13.6 (5) 

11.1-13.0 (5) 

25.0-27.6 (5) 

10.9-13.2 (5) 

13.1-14.0 (5) 

10.3-12.2 (5) 

18.2-21.4 (5) 

Jabiru mycteria 

24.8; 0.1 

13.3; 0.5 

13.5; 0.4 

30.0; 0.1 

12.2; 0.1 

13.2; 1.3 

12.6; 0.1 

21.9; 0.9 


24.7-24.8 (3) 

12.8-13.7 (3) 

13.1-13.8 (3) 

29.5-29.6 (3) 

10.7-13.1 (3) 

13.1-13.3 (3) 

11.8-13.5 (3) 

20.7-23.0 (3) 

Ephippiorhynchus 

21.1; 1.6 

12.4; 1.6 

11.4; 0.7 

23.6; 1.6 

10.3; 1.1 

11.4; 0.1 

10.5; 1.1 

18.1; 1.6 

senegalensis 

19.6-23.3 (4) 

11.0-14.6 (4) 

10.9-12.4 (4) 

22.3-22.6 (4) 

9.8-11.8 (4) 

10.5-12.7 (4) 

9.8-12.0 (4) 

17.0-20.5 (4) 

Ephippiorhynchus 

19.4; 1.3 

11.5; 1.0 

10.7; 0.7 

22.4; 1.1 

10.5; 1.1 

10.8; 0.6 

10.2; 0.6 

19.4; 1.4 

asiaticus 

18.1-21.6 (9) 

10.4-12.8 (9) 

10.0-12.0 (9) 

21.3-24.0 (9) 

9.3-11.2 (8) 

9.7-11.4 (9) 

9.1-10.7 (9) 

17.3-21.7 (9) 


view, the cotyla medialis is situated more proximally than 
the cotyla lateralis. 

The area intercotylaris immediately posterior to the 
eminentia intercotylaris has a shallow depression between 
the eminentia intercotylaris and the borders of the cotylae. 
It then slopes posteriorly to another, much more extensive 
depression just anterior to the hypotarsus. The medial side 
of this section is higher than the lateral side. 

The hypotarsus has two cristae hypotarsi separated by a 
broad, open, moderately deep sulcus hypotarsi. There is a 
very thin, low ridge running through the sulcus parallel to 
the crista lateralis hypotarsi. On the lateral side of this low 
ridge, the sulcus slightly excavates the base of the crista 
lateralis hypotarsi such that this is narrower than the 
posterior margin. The configuration of the crista medialis 
hypotarsi cannot be assessed. The main axis of the 
hypotarsus runs through the eminentia intercotylaris about 
20° medial to the anteroposterior midline. The hypotarsus 
is separated from the ridge of the lateral shaft surface by a 
moderately deep furrow and from the medial shaft surface 
by broader and shallower furrow; both furrows contain a 
small nutrient foramen. 

The anterior face is excavated at the proximal end by a 
deep fossa infracotylaris dorsalis, which continues distally 
as a wide sulcus extensorius. The deepest part of this basin 


contains two foramina vasculare proximale at the same level, 
immediately proximal to the round tuberositas m. tibialis 
cranialis. On their medial side, the sulcus extensorius is 
prominent, bordered both medially and laterally by narrow 
ridges. 

Viewed anteriorly, the lateral and medial margins of the 
proximal fragment of shaft converge distally. Distal to the 
rim on the medial side, the shaft curves medially to a 
projection level with the cotyla, before angling proximally 
into the medial rim of the cotyla. The lateral side also swings 
outwards to meet the rim of cotyla lateralis, but not as 
strongly. The curvature of the anterior shaft face to the 
eminentia intercotylaris is greater on the lateral side because 
of an anterior projection distal to the cotyla lateralis. A 
rounded projection lateral to the cotyla lateralis originates 
just distal to the rim, then collapses to a low, flat, broad 
ridge forming the lateral shaft surface. There is a thin, low 
ridge on the lateral shaft surface that starts near the midpoint 
of the cotyla, trending posterodistally towards the midline 
of the shaft, then more distally. On the medial side, the 
projection on the proximal end of the medial shaft face 
continues distally as the broad, rounded shaft margin. 

On the distal end, the small portion of shaft that is retained 
is narrow before flaring to meet the trochleae. Its lateral 
margin is straight, the medial more curved to join trochlea 


176 Records of the Australian Museum (2005) Vol. 57 


metatarsi II (in dorsal view). There is a deep groove running 
proximodistally proximal to, and level with, the lateral 
border of the trochlea metatarsi III. It meets the foramen 
vasculare distale, which is obscured on the dorsal surface. 
The medial side of the dorsal face slopes to join the medial 
surface; the junction of the dorsal and lateral faces is rather 
abrupt. 

On the plantar surface, the sulcus supratrochlearis 
plantaris is moderately shallow. It is not strongly demarcated 
laterally, but medially is confined by a moderately high, 
narrow ridge connecting the base of the trochlea metatarsi 
II and the fossa metatarsi I. This long, elliptical fossa is 
situated on the proximal end of the ridge. There is a low, very 
thin ridge running lateral to the midline and meeting the 
foramen vasculare distale, which is prominent on this side. 

This fragment is markedly compressed dorsoplantarly. 
The dorsal surface is more or less straight (in medial view), 
with the dorsal side of the trochlea metatarsi III projecting 
beyond it. The ridge supporting the fossa metatarsi I 
protrudes further plantarly than does the rest of the plantar 
shaft surface. The sulcus intertrochlearis lateralis is broader 
and deeper than the sulcus intertrochlearis medialis. 

The trochleae are arranged in a curve, viewed distally. 
The trochlea metatarsi II lies at an angle of about 10° medial 
to the dorsoplantar midline; the trochleae metatarsi III and 
IV are tilted laterally about 10° and 20° from this midline, 
respectively. The trochlea metatarsi IV projects further 
dorsally and distally than does the trochlea metatarsi II, but 
neither projects as far as the trochlea metatarsi III. The 
trochlea metatarsi II extends further plantarly than the 
trochlea metatarsi IV. The trochlea metatarsi II is rounded 
dorsally and distally; its plantar border comprises a 
triangular projection, with a wide fovea lig. collateralis 
occupying much of the medial surface. The trochlea 
metatarsi III is grooved on its dorsal surface, with the lateral 
side projecting slightly further distally. The other trochleae 
lack grooves on their dorsal surfaces, although the trochlea 
metatarsi IV has a shallow groove on its distal surface. The 
dorsal surface of the trochlea metatarsi IV is tilted laterally. 
The distoplantar corner forms a triangular projection, 
projecting beyond the rest of the trochlea, but it and the 
fovea lig. collateralis are less prominent than on the trochlea 
metatarsi II. 

The two proximal fragments differ in size, but share the 
same morphology. The magnitude of the morphometric 
differences are within that exhibited by modern taxa (Table 
4; see also Hancock et al., 1992 for measurements of living 
species). 

Taphonomy 

The occurrence of a stork in Wayne’s Wok and White Hunter 
Sites is not surprising. Both support rich local faunas 
comprising species of a range of vertebrate groups, 
including many indicative of aquatic environments. In 
contrast, Bitesanntennary Site is an initially unlikely source 
of such a bird. It has also yielded many species (almost all 
bats with 11 species recorded; Hand, 1997), but on both 
faunistic and geological bases it is considered to be a cave- 
fill deposit. The other somewhat anomalous occurrence 
found here is that of the bulungamayine kangaroo Ganguroo 
bilamina (Cooke, 1997b). 


There are several explanations for the presence of the 
non-bats Ciconia louisebolesae and G. bilamina in this cave 
deposit (see Baird, 1991b). It is possible that they used the 
cave as a shelter ( Ganguroo ) or there may have been a pool 
or other suitable foraging habitat within its immediate 
entrance. Otherwise the cave may have served as a predator’s 
lair and these species represent prey items, or their carcasses 
may have been washed into the cave from outside after 
death. An entrance of the cave may have opened upward 
through the roof rather than to the side and thus served as a 
pitfall trap for unwary animals. The skeleton of a Yellow¬ 
billed Spoonbill Plataleaflavipes, a bird of roughly similar 
size, shape and habits as Ciconia storks, has been recovered 
from Weekes Cave, South Australia; van Tets (1974) 
considered that it “may have become trapped in the sink 
hole after blundering into it” in search of residual water 
after a wet period in the usually dry environment. 

Discussion 

Despite the number of putative palaeospecies assigned to 
Ciconia, it is difficult to interpret the fossil history of this 
genus. The fragmentary nature of many of the remains 
makes comparisons troublesome; these do not lend 
themselves to clarifying any trends. The only species that 
are well represented are the large C. maltha and unnamed 
Ciconia of North America. 

As currently construed, the genus Ephippiorhynchus 
occurs in equatorial Africa, represented by E. senegalensis, 
and in southern and southeastern Asia (but not Malaysia or 
Indonesia), southern New Guinea and northern and eastern 
Australia, represented by E. asiaticus. Its entry into 
Australasia from the north is unlikely to have been possible 
until Australia approached Asia and the New Guinean 
landmass was formed during the Late Miocene. Its current 
fossil record in Australia is compatible with this timetable 
and suggests that congeneric storks should not be recovered 
from earlier deposits. The Australasian and Asian 
populations are considered only subspecifically different 
and are separated by a substantial distributional gap. It seems 
unlikely that E. asiaticus was preceded into Australia by an 
earlier and now extinct congener. As there is no evidence 
of other large storks, such as marabous, all ephippio- 
rhynchine-like storks in Australia should be expected to 
belong to this species. 

The distribution of neospecies of Ciconia is throughout 
Eurasia, Africa and South America. During the Pleistocene, 
this extended to North America (C. maltha ) and Australia 
(C. nana). It is not clear why the latter landmasses lost their 
representatives of this genus. In Australia, Ciconia had a 
rather long history (late Oligocene-Pleistocene). Species of 
Ciconia and Ephippiorhynchus are sympatric across much 
of their ranges in Africa and Asia, so there is no reason to 
believe that the arrival of E. asiaticus in Australia was a 
contributing factor to the extinction of C. nana. There are 
also marked differences in size and presumably in ecology, 
which would have precluded any direct competition. 
Possible competition might be more likely between C. nana 
and ibises of the genera Threskiornis and Plegadis. 
Threskiomis cf. T molucca has been recorded from the Early 
Pliocene Bluff Downs Local Fauna, so it and C. nana would 
have co-existed, at least in time, if not in space. 


Boles: fossil birds of the stork genus Ciconia 


111 


Acknowledgments. The specimens described here form part 
of the collection of the Queensland Museum, Brisbane; the 
Riversleigh material was made available through the Vertebrate 
Palaeontology Laboratory, School of Biological Sciences, 
University of New South Wales. I thank the curators and collection 
managers of the following institutions permitted me to work with 
comparative specimens in their care: American Museum of Natural 
History, New York; Australian Museum, Sydney; Australian 
National Wildlife Collection, CSIRO Sustainable Ecosystems, 
Canberra; Field Museum of Natural History, Chicago; University 
of Kansas Museum of Natural History, Lawrence; United States 
National Museum of Natural History, Smithsonian Institution, 
Washington D.C.; Museum Victoria, Melbourne; Queensland 
Museum, Brisbane. My special gratitude goes to S. Olson for 
providing measurements of a number of specimens in the USNM. 
He and P. Vickers-Rich provided useful criticisms of the 
manuscript. The pictures were taken by the Photographic 
Department of the Australian Museum. The Australian Museum 
provided a venue to carry out this work. The Riversleigh project 
has been supported by the Australian Research Council, 
Department of the Environment, Sport and Territories, National 
Estate Programme Grants (Queensland), Queensland National 
Parks and Wildlife Service, Australian Geographic Society, 
Linnean Society of New South Wales, ICI, Australian Museum, 
Queensland Museum, University of New South Wales and 
Pasminco Pty Ltd. 


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Manuscript received 28 March 2003, revised 26 February 2004 and 
accepted 3 March 2004. 

Associate Editor: G.D. Edgecombe. 


© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 179-190. ISSN 0067-1975 


A New Flightless Gallinule (Aves: Rallidae: Gallinula) 
from the Oligo-Miocene of Riversleigh, 
Northwestern Queensland, Australia 


Walter E. Boles 


Terrestrial Zoology, Australian Museum, 

6 College Street, Sydney NSW 2010, Australia, and 

School of Biological, Earth and Environmental Sciences, 
University of New South Wales NSW 2052, Australia 

walterb@austmus.gov.au 


Abstract. Flightlessness in birds occurs in a taxonomically diverse array of families, but is best 
exemplified in the rails (Rallidae). Most flightless species of rails live on islands, where the absence of 
native mammalian predators may make flight superfluous. Fossil rails from Oligo-Miocene sites at 
Riversleigh, northwestern Queensland, Australia, are considered to represent a single species of gallinule 
Gallinula, described here as new. Compared with four Quaternary species of Gallinula from Australasia 
(two volant, two non-volant), it shows similarities with the flightless species in the development of the 
fore- and hindlimb elements and in other characteristics of limb bone morphology associated with 
flightlessness. These indicate that the Riversleigh species was non-volant. Its relationships with the 
Quaternary species, including the flightless Gallinula mortierii, now restricted to Tasmania, but known 
from Plio-Pleistocene deposits in eastern mainland Australia, are considered. 


Boles, Walter E., 2005. A new flightless gallinule (Aves: Rallidae: Gallinula ) from the Oligo-Miocene of 
Riversleigh, northwestern Queensland, Australia. Records of the Australian Museum 57(2): 179-190. 


Flightlessness in birds occurs in a taxonomically diverse 
array of families. Flightlessness per se probably conveys 
no adaptive value; instead, it is an consequence of 
morphological changes that convey other selective 
advantages to the bird (Livezey & Humphrey, 1986). The 
general consensus is that the loss of volancy results as the 
flight muscles and pectoral assemblage become reduced as 
energy-saving processes during ontogeny (Olson, 1973a). 
Such modifications are advantageous when they do not 
render the birds more susceptible to predation. The presence 
of flightless birds on islands is strongly correlated with the 
absence of predators, particularly mammalian ones. Other 
factors can moderate this relationship, such as the 
availability of sufficient cover to avoid or reduce predation, 


allowing birds to exist in the presence of predators, and the 
stability of resources, removing the need for far-ranging 
mobility (dispersal) (Worthy, 1988). 

The developmental mechanisms involved in the loss of 
flight have been explored in greatest depth in the rails 
(Rallidae) (Olson, 1973a), the family that best exemplifies 
the phenomenon. About a fourth of the world’s 125 or so 
living or recently extinct species have lost the power of 
flight. Most, but not all, of these are populations on islands, 
where the absence of native mammalian predators has 
reduced the benefit of the ability to fly. Those species that 
have become extinct usually did so at least in part from the 
inability to cope with the introduction of exotic predators. 
Prominent among the few instances of flightlessness in rails 
www.amonline.net.au/pdf/publications/1441_complete.pdf 


180 Records of the Australian Museum (2005) Vol. 57 


on larger landmasses are the three species of native-hens of 
Australia and New Zealand ( Gallinula , subgenus Tribonyx ) 
(Fig. 1). The Black-tailed Native-hen G. ventralis, a volant 
species, is widespread through mainland Australia except 
for the east coast and far tropical north, but does not occur 
in Tasmania, to where the much larger and flightless 
Tasmanian Native-hen G. mortierii is restricted. The extinct 
G. hodgenorum of New Zealand, well represented in 
subfossil deposits, was also flightless (Olson, 1975a). The 
living volant Dusky Moorhen G.(G.) tenebrosa is common 
in both Australia and New Zealand. 

The remains of rails have been recovered from several 
Oligo-Miocene sites at Riversleigh, northwestern Queens¬ 
land (Fig. 1). These are considered to represent a single 
species, a new gallinule of the genus Gallinula. Although 
only one skeletal element is represented by an intact 
specimen, there is adequate material of the wings, legs and 
coracoid to indicate that this form was flightless. 


Fig. 1. Distribution of Recent and fossil Australian species of 
Gallinula. Key: black shading, current distribution of G. mortierii ; 
grey shading, current distribution of G. ventralis; ■ Riversleigh 
(G. disneyi ); □ Quaternary mainland sites producing G. mortierii 
(taken from De Vis 1888, 1892; Baird 1984, 1985, 1986, 1991b, 
1992; McNamara & Baird 1991; Olson 1975b; and this work). 


Materials and methods 

Measurements were made with digital callipers and rounded 
to the nearest 0.1 mm. Length measurements of bones of 
G. hodgenorum were taken from specimens and from Olson 
(1975a); all other measurements were taken from 
specimens. Weight, wing chord and tarsus measurements 
of living species of Gallinula were taken from Marchant & 
Higgins (1993) and represent means for adult males. 
Osteological nomenclature follows Baumel & Witmer 
(1993), except that as terms of position and direction anterior 
is used rather than cranial and posterior rather than caudal. 
Olson (1973b) placed Tribonyx as a subgenus of the 
gallinules Gallinula; this was accepted by Condon (1975) 
and Christidis & Boles (1994), and is followed here. 


Ratio-diagrams of the log differences between measure¬ 
ments of compared taxa were constructed following the 
method of Simpson (1941), wherein measurements are 
converted to logarithms, and one taxon is arbitrarily chosen 
as a standard. The difference between its converted 
measurements and the corresponding ones for each taxon 
are calculated (the logarithms of the ratios). The standard 
taxon thus has all ratios of 0 (zero difference in logarithms), 
which when plotted along a vertical axis on arithmetic graph 
scale, form a straight line. The logarithmic ratios for each 
taxon are plotted such that the points on a single horizontal 
line represent different values of the same variable across 
the taxa. Those values larger than the standard fall to the 
right of the standard line, the smaller ones to the left of it. 
The points of each taxon are connected with a line. Taxa 
with proportions identical to those of the standard taxon 
will have lines parallel to that of the standard. Variations 
from a parallel line are indicative of variations in the 
proportions from that of the standard taxon. Because only 
one of the fossil elements is complete, measurements of 
other features of the bones were used in lieu of total lengths 
because these should also reflect the changes in overall sizes 
of the elements. For the purposes of these comparisons, the 
values used were the means in Table 1 unless otherwise 
indicated. 

Geology and geographical setting 

The fossils described in this study were collected from the 
Riversleigh deposits, which are located 5 km west of the 
Riversleigh homestead (19°02'S 138°45'E), 200 km north 
of Mt Isa, northwestern Queensland, where they occur as 
an outcrop of Tertiary limestone overlying the Cambrian 
Thorntonia Limestone. There are now over 200 named 
Oligo-Miocene sites at Riversleigh. An informal system of 
grouping has been used (Systems A-C). These systems are 
“regionally clustered sites that appear to be super- 
positionally-related (differing in age but not significantly 
in position) and/or space-related (spatially isolated but 
approximately contemporaneous)” (Archer et al., 1989). 
The principal accumulations are thought to have occurred 
in several episodes involving large lakes, shallow pools and 
cave deposits. 

Rail material has been recovered from six sites, ranging 
from Late Oligocene to Middle Miocene in age. Current 
understanding of the age of these sites is taken from Creaser 
(1997); other birds represented at these sites are taken from 
Boles (1995,1997). White Hunter Site, Hal’s Hill Sequence, 
D-Site Plateau, considered to be part of System A (Late 
Oligocene), has yielded the greatest diversity of rail 
specimens. The White Hunter Local Fauna also contains 
other birds, including the small casuariid Emuarius gidju 
(Patterson & Rich, 1987) (Boles, 1992), the dromornithid 
Barawertornis tedfordi Rich, 1979, a stork (Boles, 2005) 
and several passerines. LSO Site (LSO Local Fauna), from 
the Verdon Creek Sequence, in the northern section of the 
D-Site Plateau, is also regarded as System A. Another site 
from the D-Site Plateau, but considered part of System B 
(Early Miocene), is Camel Sputum Site, Godthelp Hill 
Sequence (Camel Sputum Local Fauna). In addition to rails, 
it has also provided Emuarius, Barawertornis, another new 
genus and species of dromornithid, a swift Collocalia buday 
(Boles, 2001) and several passerines. Creaser’s Ramparts 


Boles: Riversleigh flightless gallinule 181 


Site and Dirks Towers Site are in the central and northern 
sections of the D-Site Plateau, respectively. Both occur at 
about the same level and are possibly correlated. The ages 
are still unclear, but may be System A or B. Other birds 
recovered from Dirks Towers Site are Emuarius and 
passerines. Ringtail Site forms part of the Ray’s Amphi¬ 
theatre Sequence on Gag Plateau. This site is included in 
System C, considered to be of Middle Miocene age. The 
Ringtail Local Fauna includes waterfowl and a number of 
passerines. 

Systematic palaeontology 
Order Gruiformes Bonaparte, 1854 
Family Rallidae Rafinesque, 1815 

The fossils are referred to the Rallidae and subordinate taxa 
on the following suites of characters (adapted in part from 
Baird, 1992; Gilbert etal., 1981; McCoy, 1963; Olsen, 1979; 
and Worthy, 1997). 

Coracoid. The tuberculum brachiale is not undercut. The 
processus procoracoideus is pronounced, extending further 
medially than the processus acrocoracoideus. The cotyla 
scapularis is large and deep. The facies articularis humeralis is 
round, about as wide as long, and flares strongly laterally. The 
impressio m. stemocoracoidei is deep, extending far anteriorly. 

Humerus. The incisura capitalis is deep and forms a shallow 
angle with the main axis of the shaft. The fossa pneumotrici- 
pitalis is shallow. The long axis of the caput humeri is 
roughly parallel with that of the incisura capitis. The sulcus 
ligamentosus transversus is shallow. The crista delto- 
pectoralis is high, triangular and directed anteriorly. The 
processus supracondylaris dorsalis is small and blunt. The distal 
end of the element is narrow, not markedly produced laterally 
or medially. The processus flexorius extends further than the 
condylus ventralis. The fossa m. brachialis is shallow. 

Carpometacarpus. The processus extensorius tends slightly 
proximoventrally (in anterior view). The dorsal rim of the 
trochlea carpalis extends far proximally and is acute at its 
proximalmost point. The fossa infratrochlearis is distinct, 
deep and circular. The os metacarpale minus is slightly 
curved. The facies articularis digitalis major and minor 
extend distally to the same extent. 

Femur. The crista trochanteris is low but strongly developed 
proximolaterally and curves medially; there are strong 
ridges extending distally from its distal edge onto the 
anterolateral face of the shaft, where it joins the linea 
intermuscularis cranialis, and from near the anteriormost 
projection to the anterior border of the facies articularis 
antitrochanteris. The impressio m. iliotrochantericus 
caudalis is restricted to the extreme proximal end of the 
trochanter femoris. The collum trochanteris is distinctly 
narrowed both anteriorly and posteriorly. The shaft curves 
to meet the caput femoris in a broad, gentle curve (in anterior 
view). The proximal half of the shaft has a distinctive 
posterior inflection (in lateral view). The linea inter¬ 
muscularis cranialis extends far distally from the ventral 
side of the crista trochanteris. The sulcus intercondylaris is 
moderately shallow. The fossa poplitea is shallow. 


Tibiotarsus. The cristae cnemialis are strongly developed. 
The crista cnemialis lateralis is flattened proximodistally. 
The crista fibularis is well developed. The pons supra- 
tendineus is well developed. The condylus lateralis is much 
broader than the condylus medialis. The incisura intercon¬ 
dylaris is narrow and displaced medially by a broad condylus 
lateralis. The facies lateralis of condylus lateralis is rounded. 

Tarsometatarsus. The hypotarsus is an elongated triangle 
(in proximal view). The crista lateralis hypotarsi is 
prominent, extending distally. There are two sulci hypotarsi 
(usually one or both enclosed), and a proximal pons 
tendineus on the medial side of the dorsal face. The dorsal 
surface of the shaft is flat, not concave. The trochlea 
metatarsi IV is shorter than the trochlea metatarsi III. 
Another useful character, which cannot be assessed on the 
fossil, is that the trochlea metatarsi II is much shorter than 
the trochlea metatarsi IV and is recessed plantarly. 


Genus Gallinula Brisson, 1760 

Gallinula Brisson, 1760, Ornithologia sive Synopsis Methodica 
vol. 1, Paris: Ad Ripam Augustinorum [50], vol. 6[2]—type 
species: Gallinula Brisson = Fulica chloropus Linnaeus, 1758. 

Tribonyx has been distinguished from Gallinula by the 
shorter, heavier toes, longer tails, absence of white on the 
undertail coverts (Olson, 1973a) and short, wide bills with 
very short premaxilla (Olson, 1975a), all characters that 
cannot be assessed from the available fossil material. 
Osteological comparisons of a “typical” gallinule, G. 
tenebrosa, and two species of native-hens Gallinula 
(Tribonyx), G. ventralis and G. mortierii, found few useful 
characters that might permit separation of skeletal elements 
to subgeneric level. In the humerus of Gallinula (Gallinula ), 
the proximal end of the condylus dorsalis extends over the 
condylus ventralis. Brodkorb (1967) used this character to 
distinguish Gallinula (s.s.) from Fulica, in which the ventral 
extent of the condylus dorsalis is less, just reaching the 
dorsal border of the condylus ventralis. The species of 
Gallinula (Tribonyx) are somewhat intermediate between 
typical Gallinula and Fulica in this character, with a shorter, 
but still overlapping ventral extent of the condylus ventralis. 
The condylus dorsalis on the one distal humeral fossil 
fragment is abraded and the state of this character is 
equivocal. In the tarsometatarsus of Gallinula (s.s.), the 
lateral border of the shaft is about even with that of the 
trochlea metatarsi IV and they thus join smoothly with little 
lateral flaring. In contrast, species of Gallinula (Tribonyx) 
have the trochlea metatarsi IV curving outwards laterally, 
away from the border of the shaft. This character cannot be 
evaluated for the fossil material. Until skull material is 
recovered, it cannot be ascertained with certainty whether 
the Riversleigh flightless rail was indeed a native-hen. 

This taxon is here diagnosed only as Gallinula. In the 
following discussion, however, extensive reference and 
comparisons are made to native-hens Gallinula (Tribonyx) 
because these are the only gallinules in Australia and New 
Zealand in which flightlessness occurs and the only ones 
well represented in the fossil record. 

Gallinula (s.l.) can be diagnosed on the material available 
by the following suite of characters. 


182 Records of the Australian Museum (2005) Vol. 57 


Coracoid. The processus procoracoideus extends about half 
way along the shaft posteriorly, joining it gradually. The 
impressio m. sternocoracoidei is deeply excavated 
mediodistally, and has a rounded border to the facies 
articularis sternalis. 

Humerus. The crista bicipitalis extends distally only Vi -% 
as far as the crista deltopectoralis. The border of the 
epicondylus ventralis is concave (in anterior view). The 
proximal end of the condylus dorsalis extends over the 
proximal end of the condylus ventralis. 

Carpometacarpus. The processus alularis is broad (in 
proximal view). The os metacarpale minus is somewhat 
curved proximally, less so distally, but overall more than in 
Fulica. The distoposterior corner of the symphysis 
metacarpalis distalis is obliquely angular, rather than square. 

Femur. The junction of the impressiones obturatoriae and 
trochanter femoris forms an acute angle of c. 50°. The 
trochanter femoris is deep, flaring from the shaft both 
anteriorly and posteriorly (in lateral view). There is general 
agreement in the size and location of the impressiones 
iliotrochanteria. The condylus lateralis is well produced (in 
lateral view). The sulcus intercondylaris is situated near the 
midline of the element. The condylus medialis is moderately 
robust (in posterior view) and well produced posteriorly 
(in medial view). 

Tibiotarsus. The impressio lig. collateralis medialis is deep. 
The crista fibularis is short. There is a deep U-shaped notch 
on the posterolateral margin of the area interarticularis. The 


crista cnemialis lateralis extends to a point well proximal 
to the proximal end of the crista fibularis. The crista 
cnemialis cranialis continues as a crest along the anterior 
medial edge of the shaft. The fossa retropatellaris is 
moderately deep. The anterior surface of the proximal end 
is slightly convex. The condylus lateralis extends far 
proximally, overlapping the distal V 3 -V 2 of the pons 
supratendineus. 

Tarsometatarsus. The hypotarsus extends distally, rather 
than being truncate (in posterior and lateral views). The 
shaft has roughly parallel sides throughout its length. The 
foramen vasculare distale is situated far distally. Other useful 
characters, which cannot be assessed on the fossil, are that 
the dorsal margin of the trochlea metatarsi II is roughly 
even with the plantar margin of the trochlea metatarsi III; 
the area proximal to the trochlea metatarsi III, medial to 
the foramen vasculare distale and lateral to the trochlea 
metatarsi II, is broad with parallel sides; and the incisura 
intertrochlearis lateralis is wide. 


Gallinula disneyi n.sp. 

Fig. 2 

Holotype. Queensland Museum, QM F20906, right 
proximal humeral fragment, broken through the shaft, with 
minor damage to the tuberculum ventrale. 


Fig. 2. Specimens of the fossil gallinule Gallinula disneyi. (A) coracoid, shoulder end (QM F31470; White Hunter Site); ( B ) coracoid, 
sternal end (QM F31477; Camel Sputum Site); (C) humerus, proximal end (QM F20906: holotype; White Hunter Site); (D) humerus, 
proximal end (QM F31471; White Hunter Site); (E) humerus, distal end (QM F31472; White Hunter Site); (F) carpometacarpus (QM 
F31478; Camel Sputum Site); (G) femur, proximal end (QM F36452; LSO Site); ( H ) femur, distal end (QM F31479; Ringtail Site); (/) 
tibiotarsus, proximal end (QM F31473; White Hunter Site); (7) tibiotarsus, distal end (QM F31475; White Hunter Site); ( K) tibiotarsus, 
distal end (QM F31474; White Hunter Site); (L) tarsometatarsus, proximal end (QM F20799; Ringtail Site); (M) tarsometatarsus, distal 
end (QM F31476; White Hunter Site). Scale equals 10 mm. 


Boles: Riversleigh flightless gallinule 183 


Type locality. White Hunter Site, Hal’s Hill Sequence, 
Riversleigh, northwestern Queensland. 

Age and local fauna. Late Oligocene/Early Miocene 
(System A); White Hunter Local Fauna. 

Paratypes. Coracoid. QM F30692 (Dirks Towers), right 
shoulder fragment broken through the processus procora- 
coideus on the sternal side of the cotyla scapularis; QM F31469 
(White Hunter), left shoulder fragment, broken on the sternal 
side of the foramen n. supracoracoidei; QM F31470 (White 
Hunter), right shoulder fragment, broken on the sternal side of 
the foramen n. supracoracoidei; QM F31477 (Camel Sputum), 
left element, missing shoulder end beyond the sulcus m. 
supracoracoidei, with some damage to the medial margin. 
Humerus. QM F31471 (White Hunter), right proximal 
fragment, broken through the shaft, with damage to the caput 
humeri and the tuberculum ventrale, and abrasion to the crista 
deltopectoralis; QM F31472 (White Hunter), left distal 
fragment, broken through the shaft. Tibiotarsus. QM F31473 
(White Hunter), right proximal fragment, broken through the 
shaft distal to the proximal end of the crista fibularis, with 
abrasion to the proximal edge of the crista cnemialis 
medialis; QM F31474 (White Hunter), right distal fragment, 
broken through the shaft, with abrasion to the rims of the 
condyli; QM F31475 (White Hunter), right distal fragment, 
broken through the shaft. QM F24130 (Camel Sputum), 
left proximal fragment, broken through the shaft proximal 
to the distal end of the crista fibularis, with damage to the 
crista cnemialis medialis and edges of the facies articularis 
medialis; QM F31480 (Dirks Towers), right distal fragment, 
broken through the shaft proximal to the crista fibularis, 
missing the crista cnemialis cranialis and much of the facies 
articularis medialis. Tarsometatarsus. QM F23723 (White 
Hunter), proximal right fragment, broken through shaft; QM 
F20799 (Ringtail), proximal left fragment, broken through 
shaft; QM F30720 (Creaser’s Ramparts), proximal right 
fragment, broken through shaft. 

Referred specimens. Carpometacarpus. QM F30908 
(Dirks Towers), left element lacking the distal end and the 
distal half of the os metacarpale minus; QM F31478 (Camel 
Sputum), complete left element. Ulna. QM F30693 (Dirks 
Towers), proximal left fragment with some abrasion to the 
rims of the articular surfaces. The olecranon is low and the 
processus cotylaris dorsalis is hooked (Olsen, 1979). Its 
proximal width is 6.3 mm. Although this specimen agrees 
in configuration with ulnae of the Rallidae and of G. 
ventralis in particular, it is only tentatively assigned to this 
taxon. It is not considered further. Femur. QM F36542 (FSO), 
proximal left fragment, broken through the shaft about % of 
way to the distal end; complete except for slight damage to the 
proximal border of the crista trochanteris; QM F 31479 
(Ringtail), left distal fragment, broken through the shaft, with 
damage to the medial side of the condylus medialis. 
Tarsometatarsus. QM F31476 (White Hunter), distal right 
fragment, comprising the trochleae metatarsi III and IV. 

Etymology. Named for Henry John de Suffren Disney, 
formerly Curator of Birds at the Australian Museum, in 
honour of his contributions to the study of Australian birds, 
particularly another flightless rail, the Ford Howe Island 
Woodhen Gallirallus sylvestris. 


Diagnosis. Gallinula disneyi is distinguished from other 
species of the genus by the following suite of humeral 
characters. It agrees with G. mortierii, and differs from other 
species, by having the proximal end round rather than 
elongate proximodistally (in posterior view); it is broader 
than in G. hodgenorum. The tuberculum ventrale is situated 
distal to the tuberculum dorsale, and is more pronounced 
than in G. mortierii. The crista deltopectoralis is short 
distally; it is more produced anteriorly compared to G. 
hodgenorum. The crista bicipitalis is short distally, joining 
the shaft more abruptly; ventrally it is rounder than in G. 
mortierii and more produced anteriorly than in G. 
hodgenorum. The condylus lateralis is thinner, not bulbous 
distally, and does not extend to the lateral margin of the 
bone (in anterior view). The tuberculum supracondylare is 
narrow, not round. Compared to that of G. hodgenorum the 
condylus dorsalis does not extend as far either distally or 
dorsally, relative to the shaft, and the fossa olecrani is 
shallower with less sharply defined edges. 

Measurements. See Table 1. 

Description 

Coracoid. The processus acrocoracoideus is flattened, more 
so than in the other species, and is rotated such the tip is 
directed ventrally, more so than in G. ventralis ; it is not 
directed anteriorly but is directed more laterally than in G. 
hodgenorum. The processus procoracoideus is broader 
distally and more rectangular than in the other species with 
a blunter tip (this is pointed in G. hodgenorum ). The facies 
articularis humeralis and facies articularis clavicularis are 
smaller. Compared with the other species, the anterior end 
is more gracile while the sternal end is larger and more 
robust, with the area encompassing impressio m. stemocora- 
coidei broader, particularly anteriorly, than it is in G. 
ventralis. The sulcus m. supracoracoidei is more rounded 
(in ventral view) than in G. ventralis. 

Carpometacarpus. The element is rather stout; it is between 
those of G. mortierii and G. hodgenorum in size and 
robustness. The os metacarpale minus is curved posteriorly; 
in this it resembles G. mortierii and G. hodgenorum and 
differs from G. ventralis, in which it is straight. The 
symphysis metacarpalis proximalis is shortened, agreeing 
with G. mortierii. The dorsal surface is flattened, more so 
than in G. hodgenorum. Compared to G. ventralis, the 
symphysis metacarpalis distalis is broader and shorter and 
the spatium intermetacarpale shorter. The distal end is 
narrower than in G. mortierii. 

Femur. The collum trochanteris is rather short and deep (in 
anterior view). The distal end of the medial branch of the 
linea muscularis caudalis is prominent, more so than in G. 
ventralis and G. hodgenorum. The trochlea fibularis is broad 
and robust (in posterior view), with the distolateral corner 
square (in posterior view) and only moderately produced 
laterally compared to G. hodgenorum. The condylus 
medialis is well produced posteriorly. The ridge extending 
proximolaterally from the condylus lateralis is slightly 
pronounced, less so than in G. hodgenorum. 

Tibiotarsus. Both cristae cnemialis are proportionally 
deeper than in G. hodgenorum. The foramen interosseum 


184 Records of the Australian Museum (2005) Vol. 57 


Table 1. Measurements (mm) of fossil and Recent species of Australasian Gallinula. Values used for G. disneyi are indicated in bold 
type. For Recent species, the mean, (standard deviation) and range are given. Means for Recent species were used for constructing log- 
ratio diagrams (Figs. 4-5). 


Gallinula 

disneyi 

Gallinula 

ventralis 

n=6 

Gallinula 

hodgenorum 

Gallinula 

mortierii 

n=3 

Gallinula 

tenebrosa 

n=3 

Coracoid 


anterior tip of processus 

QM F30692 

6.6 

7.9 (0.3) 

4.1 

9.1 (0.4) 

8.3 (0.1) 

acrocoracoideus to 

QM F31469 

6.0 

7.1-8.8 


8.4-9.8 

8.1-8.5 

posterior border of facies 

QM F31470 

6.0 


articularis humeralis 


Humerus 


proximal width 

QM F20906 

11.4 

12.2 (0.4) 

7.8 

14.2 (0.4) 

13.3 (0.2) 


QM F31471 

c.11.0 

11.3-13.0 


13.5-14.6 

13.0-13.7 

distal width 

QM F31472 

9.7 

8.8 (0.2) 

5.2 

9.9 (0.1) 

9.6 (0.2) 


8.0-9.5 


9.7-10.1 

9.3-9.9 

Carpometacarpus 


total length 

QM F31478 

28.6 

39.2 (1.0) 

19.5 

35.5 (0.4) 

38.7 (0.5) 


35.8-42.4 


35.1-36.4 

37.8-39.4 

proximal depth 

QM F30908 

8.3 

8.2 (0.2) 

4.9 

9.1 (0.2) 

8.2 (0.2) 


QM F31478 

7.3 

7.6-8.2 


8.9-9.5 

8.0-8.6 

Femur 


proximal width 

QM F36542 

10.5 

10.8 (0.3) 

10.3 

15.5 (1.2) 

11.6 (0.2) 


10.0-11.4 


13.2-16.9 

11.2-11.9 

proximal depth 

QM F36542 

7.9 

8.5 (0.3) 

8.1 


9.5 (0.4) 


7.8-9.3 


14.3-14.5 

9.1-10.2 

distal width 

QM F31479 

>13 

10.2 (0.3) 

9.7 

16.8 (0.4) 

11.2 (0.3) 


9.1-11.2 


16.2-17.5 

10.6-11.6 

depth, condylus lateralis 

QM F31479 

11.2 

8.8 (0.2) 

8.1 

14.7 (0.3) 

9.6 (0.2) 


8.3-9.3 


14.4-15.4 

9.4-9.9 

depth, condylus medialis 

QM F31479 

>9.5 

7.9 (0.1) 

7.3 

13.3 (0.2) 

8.9 (0.2) 


7.5-8.2 


13.1-13.7 

8.6-9.2 

Tibiotarsus 


proximal width 

QM F24130 

9.7 

9.0 (0.3) 

7.9 

15.6 (1.1) 

9.8 (0.2) 


QM F31473 

9.5 

8.1-9.8 


14.1-17.7 

9.6-10.2 


9.6 


distal width 

QM F31474 

8.0 

7.9 (0.3) 

6.8 

12.9 (0.1) 

8.8 (0.1) 


QM F31475 

8.2 

7.0-8.5 


12.6-13.0 

8.6-8.9 


8.1 


depth, condylus lateralis 

QM F31474 

c.7.8 

7.7 (0.2) 

6.7 

12.2 (0) 

8.9 (0.2) 


QM F31475 

8.0 

7.0-8.3 


12.2 

8.5-9.3 


7.9 


depth, condylus medialis 

QM F31474 

>7.0 

8.4 (0.3) 

7.1 

13.1 (0) 

9.4 (0.1) 


QM F31475 

8.6 

7.5-8.9 


13.0-13.1 

9.1-9.6 

Tarsometatarsus 


proximal width 

QM F20799 

10.4 

8.5 (0.3) 

7.4, 7.7 

13.5 (0.1) 

9.3 (0.1) 


QM F23723 

9.6 

7.5-9.4 


13.4-13.8 

9.1-9.5 


QM F30720 

10.3 


10.1 


proximal depth 

QM F20799 

9.5 

8.3 (0.2) 

7.6, 8.0 

13.1 (0.1) 

9.4 (0.2) 


QM F23723 

9.2 

7.5-9.0 

13.0-13.2 


9.1-9.7 


QM F30720 

10.6 


9.8 


depth, trochlea metatarsi III 

QM F31476 

c. 4.3 

4.6 (0.2) 

4.1, 4.5 

7.9 (0.2) 

5.5 (0.2) 


4.2-5.1 


7.6-8.1 

5.3-5.8 

depth, trochlea metatarsi IV 

QM F31476 

c. 4.4 

5.1 (0.1) 

4.7, 5.1 

7.7 (0.2) 

6.4 (0.2) 


4.6-5.8 


7.4-7.9 

6.1-6.7 


proximale is moderately long, extending proximally and 
cutting into the distal side of the facies articularis lateralis, 
which is consequently shortened. The impressio lig. 
collateralis mediate is situated more proximally than in the 
other species. The mediodistal portion of the shaft is not as 
curved medially as in the other species, while being more 
robust distally than in G. ventralis. The distal end of the 


sulcus extensorius is narrower and both it and the pons 
supratendineus are situated more medially than in the other 
species. The condylus lateralis is moderately flattened on 
the distal and anterodistal borders, and there is a greater 
difference between its anterior extension and that of the 
condylus medialis than in G. hodgenorum. 


Boles: Riversleigh flightless gallinule 185 


Tarsometatarsus. The proximal end is broader than in G. 
ventralis. The proximal end is broader relative to the shaft 
compared to G. mortierii. The hypotarsus is placed more 
medially than in the other species and the plantar apex is 
rounded. Both canales hypotarsi are enclosed. The shaft is 
more robust than that of G. ventralis. Compared to that of 
G. hodgenorum, the ridge extending distally from the 
hypotarsus is higher, longer and more centrally situated, 
thus making the plantar surface of the shaft more angular 
and less flattened. 

Discussion 

Comparison of Gallinula disneyi with other species 

Three species of Gallinula live in Australia today: the typical 
G. tenebrosa and the two native-hens ( Tribonyx ), the volant 
G. ventralis and flightless G. mortierii. Gallinula tenebrosa 
is a heavier bird (male weight: 570 gm) than G. ventralis 
(410 gm), yet the legs are only slightly longer (male tarsus: 
tenebrosa , 63 mm; ventralis, 61 mm) and the wings are shorter 
(male wing: tenebrosa, 208 mm; ventralis, 218 mm). The wings 
of the much larger G. mortierii (males: weight, 1334 gm; tarsus, 
84 mm) are both actually and proportionally small compared 
to those of the two volant species (wing: 202 mm). 

The differences evident in external measurements are for 
the most part mirrored in the long bones (Fig. 2). The 
elements of G. ventralis are shorter than those of G. 
tenebrosa, except for the carpometacarpus, which is about 
the same length. The leg elements are more gracile in G. 
ventralis than in G. tenebrosa. The coracoid and hindlimb 
elements in G. mortierii are longer and more robust than 
both these species, whereas the ulna and carpometacarpus 
are shorter; the humerus is somewhat intermediate, 
approaching G. tenebrosa most closely. Gallinula 
hodgenorum is consistently smaller in all length measure¬ 
ments except for the femur, which is midway between the 
values for G. ventralis and G. tenebrosa. The carpometa¬ 
carpus is the only complete element known for G. disneyi. 
It is larger than in G. hodgenorum but substantially smaller 
than in any of the living species (Table 1). 

Because no direct comparisons of element lengths can 
be made between G. disneyi and the other taxa (other than 
for carpometacarpus), a number of measurements were 
taken from the fragments available for G. disneyi, with 
comparable ones from the living species. The measurements 
are given in Table 1 and comparisons of the elements 
between the taxa are shown in Fig. 4. The wing elements of 
G. disneyi are smaller than in the volant species, somewhat 
approaching those of G. mortierii and being of similar 
robustness. The legs of G. mortierii are considerably more 
robust. The femur of G. disneyi is larger and the lower leg 
elements are roughly intermediate between those of G. 
tenebrosa and G. ventralis. Its coracoid is particularly 
reduced, both actually and proportionally, compared to all 
living species. Gallinula hodgenorum is smaller and more 
gracile overall than G. disneyi, most markedly in the 
coracoid and elements of the wing (Table 1). 

Using a log ratio diagram for the long bones lengths of 
Quaternary species of Gallinula (Fig. 2) shows that G. 
ventralis is similar to G. tenebrosa, with the major 
differences being the former’s proportionally shorter 
coracoid and longer distal wing elements and tarsometa¬ 


tarsus. The trend in the wing is not unexpected owing to the 
more pronounced mobility of this highly nomadic bird. 
Gallinula mortierii and G. hodgenorum resemble each other 
(except for size), but differ from the other taxa by having 
the forelimb elements much shorter relative to those of the 


Fig. 3. Log-ratio plot of long bone lengths (coracoid, fore- and 
hindlimbs) of living and recently extinct Australasian species of 
Gallinula. The standard species is Gallinula tenebrosa. 
Abbreviations: cor, coracoid; hum, humerus; uln, ulna; car, 
carpometacarpus ;fem, femur; tib, tibiotarsus; tar, tarsometatarsus. 

hindlimb; the femur, in particular, is proportionally long. 
There are slight differences between the flightless species 
in the relative lengths of some outer limb elements. 

Plots of the measurements from Table 1 produce overall 
patterns reminiscent of that in Fig. 3. Figure 5, based on the 
living species of Gallinula, is more indicative of relative 
robustness of elements than relative lengths, but the 
congruence in patterns supports the use of these measure¬ 
ments as indicators of major trends in the analysis. The 
relationship between proportions of G. ventralis and G. 
tenebrosa is similar to that in Fig. 3. Likewise, G. mortierii 
shows the less robust forelimbs and more robust hindlimbs. 

A comparison of G. disneyi with the three species of 
Gallinula (Tribonyx) (Fig. 6), using G. ventralis as the 
standard taxon, exhibits roughly parallel tendencies of the 
three flightless species, with similar trends in the pattern of 
reduction of the wings and coracoid and increase in the 
robustness of the hindlimb elements. Gallinula disneyi 
differs from the other flightless forms in proportionally 
narrower proximal end of the carpometacarpus, smaller 


Fig. 4. Skeletal elements of Gallinula disneyi n.sp. compared with 
those of living and recently extinct Australasian species. Left to 
right (top to bottom for B ), G. hodgenorum, G. ventralis, G. 
mortierii, G. disneyi. To facilitate comparisons, some figures have 
been reversed so that all appear from the same side of body. The 
registration number(s) of the specimen(s) of G. disneyi is given. 
(A) Coracoid, shoulder end, QM F31470; sternal end, QM F31477, 
QM F31477 reversed. ( B ) Carpometacarpus, QM F31478. (C) 
Humerus, proximal end, QM F20906; distal end, QM F31472; 
G. hodgenorum and QM F31472 reversed. ( D ) Femur, proximal 
end, QM F36542; distal end, QM F31479; proximal G. 
hodgenorum and G. ventralis reversed. (E) Tibiotarsus, proximal 
end, QM F24130; distal ends, QM F31475, QM F30696; G. 
hodgenorum, G. ventralis and QM F30696 reversed. (F) 
Tarsometatarsus, proximal end, QM F30720; distal end, QM 
F31476; QM F30720 reversed. Scale equals 10 mm. 


proximal end of the femur, several aspects of the tibiotarsus 
and shallower trochlear depths on the tarsometatarsus. This 
variation among the species may reflect real differences in 
the proportions of G. disneyi. Alternatively, in contrast to 
the chronologically constrained samples of G. hodgenorum 
and G. mortierii, that of G. disneyi comes from a broad 
period from the Late Oligocene to the Middle Miocene. 
Within that span there may have been changes in this rail’s 
body size in response to environmental shifts or other 
factors, such as Olson (1975b) proposed for Pleistocene G. 
mortierii relative to Recent animals. This pattern might also 
be due in part to sexual differences between specimens; 
males average larger than females in living Australian 
species of Gallinula, both volant and non-volant (Marchant 


Boles: Riversleigh flightless gallinule 187 


Fig. 5. Log-ratio plot of selected osteological measurements (see 
Table 62) of living and recently extinct Australasian species of 
Gallinula. The standard species is Gallinula tenebrosa. 
Abbreviations are co, coracoid; h, humerus; ca, carpometacarpus; 
f femur; tb, tibiotarsus; tm, tarsometatarsus; die, depth of condylus 
lateralis; dmc, depth of condylus medialis; dt3, depth of trochlea 
metatarsi III; dt4, depth of trochlea metatarsi IV; dw, distal width; 
paf, processus acrocoracoideus to facies articularis humeralis; pd, 
proximal depth; pw, proximal width; tl, total length. 

& Higgins, 1993). The most likely cause(s) cannot be 
identified until more material of common skeletal elements 
is available from the same sites. 

Worthy (1997) pointed out exceptions to the assertion 
that “in all flightless birds, flightlessness is associated with 
increased body size” (Livezey & Humphrey, 1986). This 
contradiction is also demonstrated by the gallinules. While 
G. mortierii is consistent with this statement relative to other 
members of the genus, the flightless G. hodgenorum is the 
smallest member of Gallinula (Tribonyx). 

Flightlessness in Gallinula disneyi 

The major morphological consequence of flightlessness is 
a reduction of the pectoral assemblage and forelimb. 
Concurrent with this is an increase in the size of the pelvic 
limb. The most obvious morphological change in the legs 
is an increase in the robustness of the elements. In the wing 
there are several structures that undergo obvious and 
characteristic modifications with the loss of flight. 

Olson (1975a) and Rich et al. (1985) presented a range 
of features that characterize the humerus of non-volant birds. 


These are evident in a comparison between the Riversleigh 
fossils and the similar-sized humerus of Gallinula ventralis. 
In the fossils, the incisura capitis is directed more 
proximodistally, bringing it more in line with the shaft; this 
is caused by the entire proximal end of the humerus being 
tilted laterally relative to the rest of the element. By virtue 
of this tilting, the tuberculum ventrale becomes on the same 
proximal level as the caput humeri. The caput itself is 
considerably flattened and elongate. The crista delto- 
pectoralis is thickened, reduced and rotated proximo- 
medially in respect to the shaft. The crista bicipitalis is 
reduced. The shaft is curved and stout; however, this has 
not been preserved in the fossils. Other changes are seen in 
the fossil carpometacarpus. Compared to this element in 
the volant species, it is reduced, becoming shorter as well 
as stouter, os metacarpale majus is bowed in anterior view, 
and os metacarpale minus is more curved. Changes in the 
coracoid related to flightlessness include a more robust 
processus acrocoracoideus and a broader and more medially 
directed processus procoracoideus lacking the ventrally 
directed twist of the tip. 

The proportional reduction in length is not uniform across 
the wing elements. There is a gradient in this proportion, 
with increased reduction from the proximal to the distal 
elements (Livezey, 1995). This is characteristic of flightless 
birds, not just rails (see, for example, Gadow, 1902; Livezey, 
1989, 1990, 1992; Livezey & Humphrey, 1986; Worthy, 
1988). Direct comparison of the fossils with comparable 
elements of the volant Gallinula ventralis demonstrates that 
the fossil rails also exhibit this trend (Fig. 6). The proximal 
end of the fossil humerus is slightly smaller than that of G. 
ventralis , primarily through the reduction of the caput 
humeri and cristae deltopectoralis and bicipitalis, while the 
distal end is larger. The carpometacarpus of the fossil is 
substantially shorter (78% of length) and more robust. In 
contrast, all fragments of the fossil’s hindlimb elements are 
considerably larger than the comparable sections of the bones 
of G. ventralis. Its larger legs and smaller wings compared to 
G. ventralis are a good indication that it was unable to fly. 

Although Gallinula hodgenorum had greater reduction 
of the wings and pectoral apparatus, and more pronounced 
morphological differences from G. ventralis than had G. 
mortierii (Olson, 1975b), it showed trends in the hindlimb 
that are also evident in the Riversleigh bird. The shafts of 
both the tibiotarsus and tarsometatarsus are heavier those 
of G. ventralis. The crista cnemialis lateralis of the 
tibiotarsus is thicker. The proximal end of the tarsometa¬ 
tarsus is more expanded, as are the trochleae, which are 
also heavier. It is difficult to compare these usefully with 
the trochleae of G. disneyi because abrasion to the latter 
gives an underestimate of their size (see Fig. 6). 

Distribution of Gallinula disneyi 

Gallinula disneyi , like G. mortierii, is unusual because of 
its continental distribution. Gallinula mortierii, now 
restricted to Tasmania, once extended well into eastern 
mainland Australia, where it has been recorded from 
Pleistocene and possible Pliocene deposits (Fig. 1), with 
the youngest record at 4670±90b.p. (Baird, 1991a). It is 
extensively represented in deposits in southeastern Australia 
in the Murray-Darling River system, with a northernmost 
record from Wyandotte Creek, northeastern Queensland 


188 Records of the Australian Museum (2005) Vol. 57 


Fig. 6. Log-ratio plot of selected osteological measurements (see 
Table 62) of Gallinula disneyi and living and recently extinct 
species of native-hens Tribonyx. The standard species is Gallinula 
ventralis. Abbreviations are co, coracoid; h, humerus; ca, 
carpometacarpus; /, femur; tb, tibiotarsus; tm, tarsometatarsus; 
die, depth of condylus lateralis; dmc, depth of condylus medialis; 
dt3, depth of trochlea metatarsi III; dt4, depth of trochlea metatarsi 
IV; dw, distal width; paf, processus acrocoracoideus to facies 
articularis humeralis; pd, proximal depth; pxv, proximal width; tl, 
total length. 

(Olson, 1975b; Baird, 1984, 1986; McNamara & Baird, 
1991). A new record from the Plio-Pleistocene Floraville 
Local Fauna (west of Leichhardt River, south of Floraville 
Homestead, northwestern Queensland; 18°17'S 139°52'E), 
represented by a tarsometatarsus (QM F24605), extends the 
western edge of the known distribution (unpublished data). 
During the Plio-Pleistocene, G. mortierii persisted despite 
the presence of native marsupial carnivores, even as it does 
today in Tasmania in the company of the Tasmanian Devil 
Sarcophilus harrisii and, previously, the Thylacine 
Thylacinus cynocephalus. Baird (1984, 1986, 1991a,b) 
hypothesized that its extinction on the mainland was 
probably due to a combination of changing environmental 
conditions and the introduction of the Dingo Canis 
familiaris dingo (earliest known occurrence 3450±95b.p.; 
Milham & Thomson, 1976). Ridpath (1972) noted that the 
native marsupial species were/are nocturnal, and the 
diurnally active G. mortierii has evolved several methods 
of avoiding ground predators. The Dingo, in contrast, 
forages extensively during the day when the native-hens 


would receive no benefit from darkness. Gallinula disneyi 
also co-existed with numerous native marsupial carnivores 
(large Dasyuridae, Thylacinidae, Thylacoleonidae), and it 
is doubtful that any of these could have caused its eventual 
extinction (see Baird, 1991a). As discussed below, it is 
possible that rather than becoming extinct, G. disneyi may 
have evolved into the living G. mortierii. 

Species relationships in Gallinula (Tribonyx) 

The pattern and sequence of speciation in the native-hens 
Gallinula (Tribonyx) are unclear. The progenitor of G. 
hodgenorum would have colonized New Zealand from 
Australia (Olson, 1975a,b). Olson (1975b) considered that 
G. hodgenorum “probably arose from an ancestor closer to 
the common ancestor of both mortierii and ventralis than 
to ventralis itself’. In bill structure it more closely resembled 
G. mortierii than G. ventralis, and was more divergent from 
G. ventralis than G. mortierii in morphology of the fore- 
and hindlimbs (Olson, 1975a). Gallinula disneyi obviously 
could not have been ancestral to either G. ventralis or G. 
hodgenorum, but it is possible that this species was in the 
direct lineage of G. mortierii (see below). 

It is known that G. mortierii is not an insular derivative 
of G. ventralis', these species have been separated for a long 
time (Olson, 1975b). Olson (1975b) considered that G. 
mortierii was not a Tasmanian autochthon, instead probably 
having colonized Tasmania from the mainland during a 
period of low sea level. Tasmania has been alternately 
connected and separated from mainland Australia by 
changes in sea levels during the Tertiary. Although 
oscillations during the Quaternary are well documented, 
there have been other periods in the Tertiary during which 
lower sea levels would have exposed the intervening land. 
Thus the loss of flight in this lineage could have occurred 
in Tasmania before the Quaternary. Both G. disneyi and G. 
mortierii may have evolved in insular Tasmania while that 
island was isolated and then subsequently invaded the 
mainland when the connection to the mainland was re¬ 
established. The fossil record of Tasmania is thus far 
inadequate to determine whether a species of flightless 
Gallinula occurred there in the Tertiary. 

Another possibility is that there was but a single event of 
this kind, which gave rise to G. disneyi. This species, in 
turn, was the direct ancestor of G. mortierii. Retreat and 
restriction of the latter to Tasmania was a Quaternary event. 
This scenario has other possible ramifications. Represent¬ 
atives of many lineages, across a variety of vertebrate 
groups, exhibited marked increases in body size through 
the latter half of the Pliocene, peaking during the 
Pleistocene. These were usually the largest members of their 
respective lineages (megafauna). Subsequently, the 
megafauna elements either died out about 40-50,000 years 
ago or became smaller (dwarfing), continuing as the modern 
representatives. It is possible that the transition of G. disneyi 
to the much larger G. mortierii was part of this phenomenon. 
Any Late Quaternary dwarfing of G. mortierii was minimal, 
however, leaving this species a large-bodied animal. 

Olson (1975b) recognized a chronosubspecies (G. m. 
reperta ) for specimens from Chinchilla, Queensland, on the 
basis of their overall smaller size than modern birds. Baird 
(1984) found considerable overlap in measurements 
between modern and Late Pleistocene specimens of G. 


Boles: Riversleigh flightless gallinule 189 


mortierii, and consequently synonymized the Chinchilla 
material with the living form. A more refined temporal 
division of specimens may yet demonstrate that size 
differences do exist between Pliocene and Late Pleistocene/ 
Holocene specimens. The Chinchilla deposit is now 
considered to be of Early to Middle Pliocene age 
(Woodburne et al., 1985), earlier than previously thought. 
If G. mortierii evolved directly from G. disneyi, then such 
smaller mortierii- type birds would not be unexpected. 

If G. disneyi gave rise to G. mortierii, the rate of such a 
change would have been very marked between the Late 
Miocene and the Pliocene. Gallinula disneyi is known from 
a time span of about 15 million years. The sample size is 
too incomplete to track major morphological changes across 
this period; however, there is no indication of any noticeable 
increase in size. The tarsometatarsus shows no shift towards 
mortierii- like size or robustness between the System A 
White Hunter Site and System C Ringtail Site. Between 
these widely separated times, sufficient anagenetic change 
in this lineage might be expected such that samples would 
exhibit recognisable morphological differences. In G. 
disneyi-mortierii, sudden changes in size might have been 
related to the onset of the aridification of Australia, starting 
in the Late Miocene. A possible argument against a G. 
disneyi-G. mortierii ancestor-descendant relationship is that 
there is no evidence of an ongoing decrease in the coracoid 
and forelimb, as might be expected in a flightless lineage 
of such long duration. 

Other Riversleigh rails 

Two additional specimens of rails have been recovered at 
Riversleigh. One shows differences from the comparable 
elements of G. disneyi and may belong to a different species. 
The other is tentatively referred to this family and represents 
a considerably smaller animal. Both are considered Rallidae 
indeterminate at this time. 

A tibiotarsus (QM L30696) of a rail from Camel Sputum 
Site, from where material referred to G. disneyi has been 
recovered, consists of a left distal fragment retaining much 
of shaft (length as preserved 77.5 mm). It has damage to its 
posterodistal face and most of the condylus medialis. The 
shaft is of comparable in width to that of G. disneyi as 
retained on a proximal fragment (QM L24130). Compared 
with the two distal tibiotarsal fragments of G. disneyi (QM 
L31474, QM L31475) the distal end of this specimen is 
more robust, with a greater width, both actually and 
proportionally relative to the shaft; the condylus lateralis is 
deeper. Morphological differences between this specimen 
and the other tibiotarsi include the position of the condyli; 
rather than being more or less parallel and in line with the 
margins of the shaft, the condyli of the Camel Sputum 
specimen are placed further laterally and medially, 
respectively. This makes the distal end flare outwards from 
the shaft much more. The condylus lateralis is inclined more 
proximomedially-distolaterally. Distal width > 9.3 mm; 
depth of condylus lateralis 8.6 mm. In view of these 
differences in size and morphology, this tibiotarsus is not 
placed with G. disneyi. It may be that these features fall 
within the variation of that species, although it is not evident 
from the other specimens, nor is there an indication of such 
a range of differences in modern species. 


A small, damaged left carpometacarpus (QM L40203), 
still attached to the rocky matrix, from Dunsinane Site 
(System A; Arena, 1997), appears to be that of a rail. 
Because sections of the proximal end are missing, a definite 
identification is precluded. This specimen is substantially 
smaller than the carpometacarpus of G. disneyi, and is 
comparable in size to the living Rallus pectoralis; there is 
no indication that it represents a juvenile It is tentatively 
assigned to the Rallidae. 


Acknowledgments. For access to comparative specimens, I 
thank Richard Schodde and John Wombey, Australian National 
Wildlife Collection, and Alan Tennyson, Museum of New Zealand 
Te Papa Tongarewa. Robert Baird provided valuable discussions 
in the early stages of this study; comments by Storrs Olson and 
Patricia Vickers-Rich considerably improved the content. The 
pictures were taken by the Photographic Department of the 
Australian Museum. The Australian Museum provided a venue 
in which to work. The Riversleigh project has been supported by 
the Australian Research Council, Department of the Environment, 
Sport and Territories, National Estate Programme Grants 
(Queensland), Queensland National Parks and Wildlife Service, 
Australian Geographic Society, Linnean Society of New South 
Wales, ICI, Australian Museum, Queensland Museum (QM), 
University of New South Wales and Pasminco Pty Ltd. 

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Manuscript received 28 March 2003, revised 26 February 2004 and 
accepted 3 March 2004. 

Associate Editor: G.D. Edgecombe. 


© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 191-210. ISSN 0067-1975 


Revision of the Genera 

Sphodropoda, Trachymantis and Zopheromantis 
(Mantodea: Mantidae: Mantinae) 


G.A. Milledge 


Terrestrial Zoology, Australian Museum, 

6 College Street, Sydney NSW 2010, Australia 

grahamm@austmus.gov.au 


Abstract. The genera Sphodropoda Stal, Trachymantis Giglio-Tos and Zopheromantis Tindale are 
revised. Aspects of their biology are discussed and distributions given. One new species of Sphodropoda, 
S. lepida, is described. Sphodropoda moesta Giglio-Tos, S. mjobergi Sjostedt and S. papua are new 
synonyms of S. tristis (Saussure). Sphodropoda dentifrons Stal is transferred to Trachymantis. S. loripes 
Tindale is transferred to Zopheromantis. Z. trimaculata Tindale is a new synonym of Z. loripes. 


Milledge, G.A., 2005. Revision of the genera Sphodropoda, Trachymantis and Zopheromantis (Mantodea: 
Mantidae: Mantinae). Records of the Australian Museum 57(2): 191-210. 


Sphodropoda Stal, Trachymantis Giglio-Tos and Zophero¬ 
mantis Tindale are three closely related mantid genera, 
known only from mainland Australia and, in the case of 
Sphodropoda, Papua New Guinea, the Solomon Islands and 
possibly Fiji. All are shrub and tree dwellers, and most of 
the species inhabit drier regions. 

Stal (1871) erected Sphodropoda as a subgenus of 
Hierodula Burmeister and included a single species, H. 
(Sphodropoda) tristis (Saussure). Westwood (1889) elevated 
Sphodropoda to generic level. Trachymantis was erected 
by Giglio-Tos (1917) for T. obesa Giglio-Tos and 
Zopheromantis by Tindale (1924) for Z. trimaculata Tindale. 
Beier (1935) incorrectly synonymized Zopheromantis with 
Hierodula without giving reasons. Zopheromantis is 
recognized here as a valid genus. 

Materials and methods 

Methods and terminology follow Milledge (1990, 1997). 
The following abbreviations are used for male genitalia: 
apr, apical process of left phallomere; dpr, distal process 
of ventral phallomere; pa, phalloid apophysis. Measure¬ 


ments are in mm and range from the smallest specimen 
examined to the largest, except where an insufficient range 
of material was available, when single measurements are 
given. Abbreviations used for institutions where material is 
held are as follows: AM, Australian Museum, Sydney; 
ANIC, Australian National Insect Collection, Canberra; 
BMNH, The Natural History Museum, London; NHRM, 
Swedish Museum of Natural History, Stockholm; NMV, 
Museum Victoria, Melbourne; SAM, South Australian 
Museum, Adelaide; WAM, Western Australian Museum, 
Perth; ZMHU, Zoologisches Museum der Humbolt- 
Universitat, Berlin. 

Relationships 

These three genera appear to form a closely related group, 
sharing a number of characters including well-developed 
preacetabular spine, facial shield as wide as high, denticules 
between teeth on outer margins of fore femora and bifurcate 
distal process in the male genitalia. Beier (1964) placed 
Sphodropoda and Trachymantis in his Miomantini, a tribe 

www.amonline.net.au/pdf/publications/1442_complete.pdf 


192 Records of the Australian Museum (2005) Vol. 57 


Figs. 1-6. Right fore femur, females, arrows indicate denticles. (1) Iris sp., ventral view; (2) same, ventral view, distal two thirds; (3) 
same retrolateral view, distal two thirds. (4) Sphodropoda quinquedens, ventral view; (5) same, ventral view, distal two thirds; (6) same 
retrolateral view, distal two thirds. Scales = 2 mm 


sharing only the single synapomorphy of denticules between 
the outer spines of the fore femora. He obviously had not 
examined specimens of Zopheromantis as he considered it 
to be a synonym of Hierodula (Mantini), which lacks this 
feature. There is some doubt as to whether the Miomantini 
is monophyletic, as the genera within it display a wide 
variety of forms. The denticules between the outer spines 
in the three genera considered here are really only small 
tubercules, and could be interpreted as a manifestation of the 
relatively tuberculate cuticle they possess. I have examined 
specimens of the European genus Iris Saussure, which Beier 
also included in his Miomantini. The pattern of denticles (Figs. 
1-3) is different from that found in the genera considered here 
(Figs. 4-6). In fact, the denticles are not between the outer 
spines but on the ventral surface of the femur. The original 
authors (Stal, Giglio-Tos and Tindale) considered the genera 


revised here to be related to Hierodula. Although this seems 
more likely than Beier’s grouping, as noted elsewhere 
(Milledge, 1997), further work is required on the mantid 
fauna of other parts of the world before the relationships of 
the Australian fauna can be fully understood. 

Sphodropoda Stal 

Sphodropoda Stal, 1871: 399. Type species Sphodropoda tristis 
Saussure, by original designation. 

Diagnosis. Head about as high as wide, frontal shield 
without dorsal extensions; preacetabular spine strong; inner 
face of first tarsal segment partly or wholly black, ventral 
face of fore femur tuberculate, especially toward outer 
margin; both sexes macropterous (some females slightly 
brachypterous), ventral face of costal area of tegmen at least 


Milledge: revision of three mantid genera 193 


partly red. Can be distinguished from Trachymantis by the 
lack of a tooth on the apex of the frontal shield and from 
Zopheromantis by the hind femur being only about half the 
length of the abdomen. 

Description. Small to moderately large, body robust, male 
macropterous, female macropterous to slightly brachy- 
pterous. Head about as wide as high, not significantly 
compressed anteroposteriorly, apical margin gently arched; 
eyes rounded, not particularly prominent; frontal shield 
slightly wider than high, flat, with distinct ridge laterally 
and apically but not produced into a tooth at apex. 

Pronotum moderately elongate, supracoxal expansion 
slight to distinct, dorsal surface finely to moderately coarsely 
granulate, margins finely denticulate, ventral surface with 
preacetabular spine prominent. Fore coxa as long as or 
longer than metazone of pronotum, often with distinctive 
pattern on inner face and a number of prominent tubercles 
on anterior margin. Fore femur finely tuberculate on ventral 
surface, particularly toward outer margin; with four 
discoidal spines, relative length from longest to shortest 3- 
2-1-4; with four outer and 15 inner spines, claw groove 


situated at about a third the total distance from base. Inner 
face of tarsus largely black. 

Tegmen with costal area opaque, partially red beneath; 
discoidal area entirely opaque to mostly hyaline. Hind wing 
with costal and discoidal area pigmented, anal area largely 
hyaline. Mid and hind legs relatively short, hind femur about 
half the length of abdomen. Abdomen moderately elongate, 
cerci short, slender and cylindrical. Male genitalia with dpr 
bifurcate. 

Notes. All the species in this genus have relatively short 
legs, a feature that appears to be associated with the 
behavioural adaptation of depressing the body against the 
substrate when threatened. This behaviour has been 
observed in three of the species recognized here ( S . tristis, 
S. quinquedens and S. viridis), the behaviour of the other 
(S. lepida) has not been observed. If the substrate is a branch 
or twig this behaviour includes moving to the side of the 
branch facing away from the threat. Similar behaviour 
appears to have evolved independently in other genera of 
Mantidae, e.g., Tarachodula (Preston-Mafham 1990, plate 
33) and Galepsus (Hevers & Liske 1991, plate 21). 


Key to species of Sphodropoda 

1 Internal face of fore coxa with 5-6 transverse white bands 

anteriorly (Fig. 15). S. quinquedens 

-Internal face of fore coxa lacking bands. 2 

2 Anterior margin of fore coxa internally with six to seven whitish 

tubercles (Fig. 12). S. tristis 

—— Anterior margin of fore coxa without such tubercles. 3 

3 Major veins of discoidal area of tegmina uniformly green. S. viridis 

-Major veins of discoidal area of tegmina flushed reddish brown. S. lepida 


Sphodropoda tristis (Saussure) 

Figs. 7-8, 11-13, 17, 42 

Mantis tristis Saussure, 1871: 93. 

Hierodula (Sphodropoda) tristis (Saussure)-Stal, 1871: 399. 
Sphodropoda tristis (Saussure).-Westwood, 1889: 13. 
Sphodropoda moesta Giglio-Tos, 1911: 20; new synonym. 
Sphodropoda mjobergi Sjostedt, 1918: 25; new synonym. 
Sphodropoda papua Beier, 1965: 489-90; new synonym. 

Type material examined. Syntype female of Sphodropoda 
moesta , Cape York, Daemel, cat. no. 3883 (ZMHU). Syntype 
male and syntype female of Sphodropoda mjobergi, 
Kimberley district, N.W. Australia, Mjoberg (NHRM). 

Other specimens examined. Queensland. 1d, Armstrong Creek 
crossing, 13 kmNNW of Guthalungra, 26 Jan 1982, M.S. & B.J. Moulds. 
1 $, Biggenden Bluff, Mt. Walsh National Park, 10 Jan 1984, D. Rugg. 
Id 1 , 1 9, 7 km S of Biggenden, Mt. Walsh National Park, 9-12 Apr 
1971, H. Frauca. 5 9, Bluff Range, near Biggenden, 2-12 May 1971, H. 
Frauca. 2 juv, Bluff Range, 8 km S of Biggenden, 9 Jan 1971, H. Frauca. 
19, 20 km NE of Bundaberg, Apr 1971, H. Frauca. Id, Byfield, 10 
May 1955, Common & Norris. 2d, 19, Carnarvon National Park (Tourist 
Lodge), 25°05'S 148°15’E, 27 & 29 Apr 1979, K.H.L. Key. 1 d, Clermont, 
15 Feb 1975, R.A. Farrow. 3d, 7 km SSW of Clermont, 2 Apr 1977, 
R.C. Lewis. 19,1 juv, near Clohesy River 12 km SW of Kuranda, 7 Leb 
1988, 16°54'S 145°34'E, D.C.L. Rentz. 1 juv, 52 km SE of Cloncurry, 
25 Aug 1960, M.J.D. White. 1 juv, Desailly Creek, 10 km NW of Mt. 


Carbine, 16°30'S 144°55'E, 19-21 May 1981, D.C.F. Rentz. Id, 1 juv, 
8 km W of Dimbulah, 17°09'S 145°02’E, 22 Mar 1988, D.C.F. Rentz. 
1 d, 42 km N of Emerald, 20 Apr 1955, Norris & Common. 1 9, Forty 
Mile Scrub, 55 km SSW of Mt. Garnet, 18°06'S 144°50’E, 8 Dec 1985, 

J. Balderson. 1 d, 3 km N of Greenvale HS, W of Ingham, 5 Apr 1962, 

K. H.L. Key & E.L. Corby. 1 juv, Isla Gorge Lookout, Isla Gorge National 
Park, SSW of Theodore, 25°10'S 150°00'E, 21 Apr 1982, D.C.F. Rentz. 
1 d, 3 km SE of Mary Kathleen, 21 Apr 1962, K.H.L. Key & E.L. Corby. 
Id, Moondoo, 26 Feb 1963, A.L. Dyce & M.D. Murray. 1 juv, 10 km 
SE of Mt. Carbine, 16°37'S 145°12'E, 24 Nov 1981, J. Balderson. 1 juv, 
17 km WSW of Mt. Faulkner, Clermont district, 7 Jan 1965, M.J.D. 
White. Id, Mt. Larcom, 18 Apr 1955, Norris & Common. 2d, Running 
River, 22 km W of Paluma, 11 Feb 1971, J.G. Brooks. Id, near Poona 
Lake, Cooloola National Park, 25°58’S 153°07’E, 4 Apr 1978, D.C.F & 
B.G.F. Rentz. 1 9, 1 km N of Rounded Hill, near Cooktown, 15°17'S 
145°13'E, 5-7 May 1981, D.C.F. Rentz. Id, 23 kmNNE of Taroom, 29 
Mar 1977, R.C. Lewis. 1 d, 9 km WNW of Taroom, 25°34'S 149°45’E, 

1 Feb 1981, D.C.F. Rentz & D.T. Gwynne. 1 juv, 7 km E of Toowoomba, 

2 Jan 1956, M.J.D. White. Id, Townsville, 7 Jan 1968, P. Ferrar. Id, 
Townsville, 31 Dec 1967, P. Ferrar. 19, Watalgan Range, near 
Bundaberg, 3 Jul 1971, H. Frauca. 1 9, Watalgan Range, S of Rosedale, 
Mar 1971, H. Frauca. 1 d, Waverley Creek, Bruce Highway, 10 km S of 
St. Lawrence turnoff, 27 Jan 1988, M.S. & B.J. Moulds (All ANIC). 
19, Augathella, 25°48'S 146°35’E, 18 Jan 1993, G. Milledge. 19, 16 
km ESE of Burke & Wills Junction, 19°17'S 140°29'E, 15 Jan 1993, G. 
Milledge. Id, 2 9 1km NNE of Collins Weir, W of Atherton, 17°15'S 
145°17'E, 10 Feb 1989, G. Milledge. Id, 8 km E of Emuford, 30 Dec 
1989, M.S. & B.J. Moulds. 59, Georgetown, 12-14 Apr 1991, G. 
Milledge. Id, Georgetown, 16 Apr 1991, G. Milledge. Id, 12 kmE of 
Georgetown, 12 Apr 1991, G. Milledge. 19,6 km S of Normanton, 


194 Records of the Australian Museum (2005) Vol. 57 


Figs. 7-8. Sphodropoda tristis. (7) Male, dorsal; (8) female dorsal. Scale = 10 mm. 


17°44'S 141°05'E, 10 Jan 1993, G. Milledge. Id, Oaky CreekRd, 8 km 
W of Cooktown, 15°29'S 145°10'E, 8 Apr 1991, G. Milledge. Id, 
Portland Roads, Iron Range, 29 Oct 1991, J. Hasenpusch. 1 $, 0.5 km S 
of Wills Creek, 9.5 km NE of Normanton, 17°38'S 141°09'E, 11 Jan 
1993, G. Milledge (all NMV). 

New South Wales. 1 juv, 29 km ENE of Coonabarabran, 31°08’S 
149°33E, 25 Nov 1983, D.C.F. Rentz & M.S. Harvey. 19, Dungay Creek, 
13 km W of Kempsey, I.H. Parberry. 1 juv, 5 km SE of Merrygoen, 18 
Jan 1961, M.J. D.White. 1 d, 15 km WNW of Monia Gap, 2 Feb 1964, 
M.J.D. White. 1 juv, 10 km W of Temora, 12 Jan 1955, M.J.D. White 
(all ANIC). 1 9, Clarence River, Mr Wilcox (NMV). 

Northern Territory. 1 9,43 km SE of Adelaide River, 4 Nov 1966, A. & R. 
Mesa. 2d, Barrow Creek Telegraph Station, 20 Mar 1955, K.H.L. Key. 1 
juv, Berrimah, 10 Mar 1972, J.C. Wombey. 19, 38 km ENE of Birrindudu 
HS, 18°15’S 129°45E, 11 Jul 1969, C. Simpson. 1 juv, Blackfellows Station 


to Burnside, 2-3 Apr 1929, T.G. Campbell. 19, Borroloola, McArthur River, 
16 Jun 1929, T.G. Campbell. Id, 22 km WSW of Borroloola, 16°08'S 
136°06E, 16 Apr 1976, Key & Balderson. Id, 36 km SW of Borroloola, 
16°19'S 136°05E,4Nov 1975, M.S. Upton. 1 d, 46 km SSW of Borroloola, 
16°28'S 136°09E, 28 Oct 1975, M.S. Upton. 1 juv, Ca im an Creek, Coburg 
Peninsula, 11°14’S 132°12E, 13 Feb 1977, R.C. Lewis. 19, Caranbirini 
Waterhole, 33 km SW of Borroloola, 16°16’S 136°05E, 21 Apr 1976, Key 
& Balderson. 1 juv, 7 km SW of Coolibah HS, 15°34’S 30°54E, 28 Jun 
1968, M. Mendum. 1 9 , Darwin, 25 Apr 1972, E.C. Abbey. 1 d, 8 kmNNW 
of Elliott, 17°29'S 133°30E, 14 Oct 1972, M.S. Upton. 19, Howard Springs, 
30 Dec 1986, M.S. & B.J. Moulds. Id, Humpty Doo, 29-30 Jan 1939, 
E.B. Boerema. Id, Mataranka, 26 Mar 1955, K.H.L. Key. 19, McArthur 


Milledge: revision of three mantid genera 195 


Figs. 9-10. Sphodropoda quinquedens. (9) Male, dorsal; (10) female dorsal. Scale = 10 mm. 


River HS, 80 km SW of Borroloola, 16°39'S 135°51'E, 13 May 1973, M.S. 
Upton & J.E. Feehan. 1 $, 16 km WSW of McArthur River HS, 16°45’S 
135°44E, 13 May 1975, Balderson & Freeman. Id, 15 kmE of Mt. Cahill, 
12°52’S 132°50E, 7 Mar 1973, K.H.L. Key. 1 6 , October Creek, Carpentaria 
Hwy, 180 km E of Daly Waters, 11 Jan 1986, M.S. & B.J. Moulds. Id, 
Plenty Hwy, 268 km ENE of Alice Springs, 22°47'S 136° 18E, 14 Oct 1978, 
M.S. Upton. Id, 30 km ENE of Soudan HS, 19°55'S 137°15'E, 25 Apr 
1976, Key & Balderson. Id, Standley Chasm, 43 km W of Alice Springs, 9 
Feb 1966, Britton, Upton & Mclnnes. 1 d, Stuart Hwy, 58 km NW of Alice 
Springs, 23°11’S 133°44’E, 28 Oct 1988, D.C.F. Rentz. 1 juv, Tanami 
Borehole, 19°59’S 129°42E, Jul-Sep 1971, J. Hodgson. 2d, Tindal, 14°31'S 
132°22'E, 1-20 Dec 1967, W.J.M. Yestjens. 1 8 , Tindal, 2 Dec 1967, W.J.M. 
Vestjens. 1 d, 13 km W of Top Springs, 24 Oct 1965, Blackith & Mesa. 1 d, 


Uluru Motel, Ayers Rock, 25°21'S 131°03’E, 4 Nov 1980, K.H.L. Key (all 
ANIC). Id, 31 km S of Alice Springs, 27 Sep 1987, G. Milledge. 19, 
Darwin, Jul-Aug 1912, Prof Spencer. 1 9, Uluru Camp Site, Horn Centenary 
Expedition, 25°23’10"S 131°00'46"E, 24 Oct 1994, G. Milledge, 19, 
Watarrka National Park, Mar 1995, G. Milledge (all NMV). 

South Australia. 3d, 8.5 km WSW of Calperum HS, 34°05'S 140°38’E, 
2 Mar 1995, Cardale, Colloff & Pullen (ANIC). 

Victoria. 19, Inglewood. Id, Mallee District, 3 Mar 1914, C. French. 
Id, Mallee District. 19, 22.3 km N of Millewa South Bore, 34°35’S 
141°03’E, 17-21 Feb 1987, G. Milledge. 19, 16.8 km SSW of 
Murrayville, 35°25’S 141°09’E, 23 Feb 1986, G. Milledge. 19, 
Wyperfield National Park, 3 Mar 1964, H.E. Tarr (all NMV). 


196 Records of the Australian Museum (2005) Vol. 57 


Figs. 11-16. Sphodropoda spp. (11) S. tristis, female head, anterior view; (12) same, fore femur, internal view; (13) same, male 
genitalia, dorsal view. (14) S. quinquedens, female head, anterior view; (15) same, fore leg, internal view; (16) same, male genitalia, 


dorsal view. Scales = 2 mm. 

Western Australia. 1$, Balgo Hills, 13 Oct 1985, M. Golding. 3d, 38 
km WNW of Balladonia Motel, 22 Feb 1980, D.C.F. 7 B.G.F. Rentz. 
Id, Brogo Hill, 160 km S of Halls Creek, 10 Sep 1985, M. Golding. 
Id, 42 km ESE of Broome, 16 Apr 1963, L.J. Chinnick. Id, 145 km 
ESE of Broome, 18°55'S 123°27’E, 8 Aug 1976, I.F.B. Common. 19, 
186 km ESE of Broome, 18°53’S 123°43'E, 11 Aug 1976, I.F.B. Common. 


Id, 5 km SSW of Cape Bertholet, 17°17'S 122°10’E, 21 Apr 1977, D.H. 
Colless. Id, 8 km S of Cape Bertholet, 17°19’S 122°10'E, 16 Apr 1977, 
D.H. Colless. 19, Halls Creek, 29 Sep 1953, Brittan. Id, Kimberley 
Research Station, via Wyndham, 15 Aug 1955, E.C.B. Langfield. Id, 
Kimberley Research Station, via Wyndham, 28 Dec 1956, E.C.B. 
Langfield. 19, Kimberley Research Station, 21-22 Apr 1958, L.J. & 


Milledge: revision of three mantid genera 


Fig. 17. Sphodropoda tristis, variation in male genitalia, cf figure 13, arrows indicate pa rotated to lateral view, (a) 
WA—near Wyndham; (b) Qld—near Cooktown; (c) Qld—near Greenvale HS; ( d) WA—near Broome; (e) NT— 
October Ck; (/) Qld—Townsville; (g) WA—near Midstream HS; (h) NT—near Alice Springs; (i) Qld—Byfield; (j) 
WA—near Balladonia; (k) Vic—Mallee district; (/) Qld—near Taroom. 


198 Records of the Australian Museum (2005) Vol. 57 


M.F. Chinnick & J. Walker. 1 S, 1 km NNE of Millstream HS, 21°35'S 
117°04'E, 3 Apr 1971, Upton & Mitchell. 3 c?, 2 km ENE of Millstream 
HS, 21°35'S 117°04'E, 22 & 30 Oct & 4 Nov 1970, M.S. Upton & J.E. 
Feehan. 1 $, Mining Camp, Mitchell Plateau, Kimberley district, 14°49'S 
125°50’E, 9-19 May 1983, D.C.F. Rentz & J. Balderson. 12, 50 km 
SW of Sandfire Flat, Broome-Port Hedland Rd, 29 Oct 1978, M.S. & 
B.J. Moulds. 1 juv, 8 km SW of Walsh Point, Admiralty Gulf, 14°37’S 
125°48’E, 17 May 1983, D.C.F. Rentz & J. Balderson. 1 2, 2 km NNE 
of Wyndham Port P.O., 15°27’S 128°06’E, 16 Apr 1985, K.H.L. Key. 
1 <?, Wyndham, 25 Apr 1930, T.G. Campbell (all ANIC). 1 juv, 217 km SE 
of Broome, 15 Sep 1924, A.S. Cudmore. 1 juv, Forrest River Mission, 8-12 
Sep 1953 (both NMV). 1 c? , Bamboo Creek, 20°56'S 120° 13E, 22 Jan 1974, 
A.M. & M.J. Douglas. 12, Beverley Springs HS, 16°43'S 125°27'E, 19 
May 1979, B.G. Muir, lc?. Cape Range, May 1965, G.W. Kendrick, lc?, 
Derby, 1962, G. Beamish. 1 c? , Greys Camp, Rudall River, near Larrys Creek, 
May 1971, K. Clarke & P. Moore. 1 c? , 13 km ESE of Mooka HS, 24°58’S 
1 14°49’E, 9-11 May 1981, B. Hanich & T.F. Houston, lc?, 51 km SSW of 
Norseman, 8 Mar 1984, T.F. Houston. 12, Salmon Gums, lc?, Walganna 
Rock, Austin Downs, 27°43'S 117°28'E, 27 Aug 1987, R.P. McMillan. 12, 
Winjana Gorge, campsite 13, 12 Apr 1970, Lemley Expedition (all WAM). 

Diagnosis. This species can be distinguished from others 
in the genus by the pattern of tubercles and colouration on 
the inner face of the foreleg (Fig. 12). 

Description. Body (Figs. 7-8) smallish to moderately large. 
Colour brown with mottled tegmina, green with mottled 
tegmina or uniform green. Ridge on frontal shield with blunt 
point at apex. Prothorax with numerous scattered tubercles 
on dorsal surface, particularly in female; lateral margin 
finely denticulate; metazone dark grey ventrally, sometimes 
with orange patch between coxal insertions. Fore leg (Fig. 
12) with inner face of coxa dark grey with 6-7 contrasting 
cream tubercles on anterior margin; inner face of femur 
orange, with 3-4 small black spots at bases of spines; inner 
face of first tarsal segment mostly black, of other tarsal 
segments black apically. Wings of male longer than 
abdomen, as long as or slightly shorter than abdomen in 
female; ventral surface of costal area of tegmen rosy red, at 
least in basal half; discoidal area completely opaque in 
female, only partly so in male, usually with dark patch on 
distal margin of stigma (absent in uniformly green 
specimens and small males of other colour forms); hind 
wing with costal and discoidal area reddish to yellowish, 
anal area slightly smoky, particularly toward apex, in female, 
mostly hyaline in male. Abdomen with median black spot 
on proximal margin of ventral sclerites 2-5. Male genitalia 
(Figs. 13, 17) with relatively short stout apr, pa and dpr 
rather variable, pa compact, shagreened, L to C shaped when 
viewed dorsally, with pointed, laterally directed posterior 
process and sometimes with blunt pointed, dorsally directed 
process; dpr terminated with two short, pointed projections 
of variable form. 

Measurements (mm). Body length, S 38-68, 9 41-72. 
Head width, 6 7-9, $ 8-10. Head depth, S 4-7, 9 6-9. 
Pronotum length, S 9-18, 9 11-19. Pronotum width, S 
3-6, 9 4-8. Fore coxa length, S 6-11, 9 8-13. Fore femur 
length, S 7-13, $ 10-15. Hind femur length, S 7-13, $ 
9-14. Hind tibia length, S 7-13, 9 9-14. Tegmen length, 
6 26-46, 9 26-37. 

Immature stages. First two instars dark coloured and ant¬ 
like in appearance and behaviour, the abdomen being curled 
dorsally when active. Later instars more like adult in 
behaviour and appearance. Ootheca pale grey or cream, and 
squat looking. Chapman & Balderson (1984) recorded a 
female of this species from Brisbane as laying its oothecae 


in the ground. This behaviour may not be obligatory however, 
as I have collected oothecae from the “paper” bark of melaleuca 
trees in the Cairns region. I have also had females in captivity 
which laid oothecae on the roof and sides of the container, 
although oviposition was not observed. 

Distribution and habits. Found through most of mainland 
Australia (Fig. 42), also southeast Papua New Guinea and 
the Solomon Islands (Beier 1965). The locality of the female 
type specimen, which cannot be located (Balderson 1984), 
is recorded as the Fiji Islands (Saussure 1871). This record, 
however, must remain doubtful until further specimens are 
obtained. This species is a shrub and tree dweller, usually 
found on stems or trunks. When disturbed, individuals move 
to the other side of the stem and flatten themselves against 
the substrate. Females are capable of at least short flights. 

Remarks. This widespread species is quite variable in size 
and to some extent in the form of the male genitalia (Fig. 
17). Specimens from the north west of the continent tend to 
be smaller (S. mjobergi form) while those from the northeast 
tend to be larger (typical S. tristis ). However, after examining 
numerous specimens I cannot find any group of characters 
that would separate one population from the rest. 
Morphological characters useful in delineating other species 
within the genera considered here (e.g., pattern on inside 
of foreleg, head shape) show little or no variation. Thus S. 
mjobergi is considered a synonym of S. tristis. I have not 
been able to locate the holotype of S. papua. It is apparently 
not in the New Guinea Dept, of Agriculture & Livestock 
collection, although several paratypes are (F. Dori, pers. 
comm.), nor is it in the Bishop Museum collection (G. 
Nishida, pers. comm.). Beier’s (1965) description and 
photograph suggests this species falls well within the range 
of variation observed for Australian S. tristis and it is 
therefore considered synonymous. I have examined a 
syntype of S. moesta Giglio-Tos and there is nothing to 
indicate his species is different from S. tristis. As suggested 
by Tindale (1923), it is merely the brown colour form of S. 
tristis. The syntype specimen of S. moesta examined was 
listed as the holotype by Balderson (1984). The original 
description by Giglio-Tos, however, clearly lists two 
specimens, the second from “the Island of Australia”. This 
second specimen cannot be located in the ZMHU (M. Ohl, 
pers. comm.). I have also examined two syntypes of S. 
mjobergi and it is obviously the north western form of S. 
tristis. There has been some confusion as to the authorship 
of the name of this species. Although Saussure described 
the species he attributed the name tristis to Brunner. Up 
until recently all subsequent authors except Westwood 
(1889) attributed the name to Saussure. Balderson et al. 
(1998), however, attributed the name to Brunner in Saussure, 
but, as there is no indication that Brunner was responsible 
for the description, Saussure must be considered the author. 

Sphodropoda quinquedens (Macleay) 

Figs. 4-6, 9-10, 14-16, 43 

Mantis quinquedens Macleay, 1826: 454. 

Hierodula quinquedens (Macleay).-Saussure, 1871: 78. 
Sphodropoda quinquedens (Macleay).-Kirby, 1904: 242. 


Milledge: revision of three mantid genera 199 


Material examined. Lectotype female, Australia (ANIC). 

Other specimens examined. Queensland. 1 $, Black Mountain 
NP, 15°39'05"S 145°13'13"E, 16 May 2000, G. Milledge (AM). Id, 
Albatross Hotel, Weipa, 12°38'S 141°52’E, 6 May 1981, K.H.L. Key. 
1$, Annan River, 3 km SW of Black Mt., near Cooktown, 15°41’S 
145°12'E, 26-27 Apr 1981, D.C.F. Rentz. Id, 44 km N of Cairns, 10 
Dec 1982, J.T. Doyen. 1 $, 8 km W of Dimbulah, 17°09’S 145°02’E, 22 
Mar 1988, D.C.F. Rentz. Id, Emu Creek, 27 km SW of Dimbulah, 
17°20'S 144°57’E, 25-26 Nov 1981, J. Balderson. 1$, Iron Range, 12 
Jul 1969, J.C. Le Souef. Id, McKenzie River crossing, 75 km NNE of 
Dingo, 17 Jan 1987, M.S. & B.J. Moulds. 1 juv, Shiptons Flat, near 
Cooktown, 15°47'S 145°14'E, 16-18 May 1981, D.C.F. Rentz. 1 d, Silver 
Plains HS, Cape York Pen., 25 Apr 1968, J.F. Wassell (all ANIC). Id, 
Annan River crossing, 2 km S of Cooktown, 4 Apr 1991, G. Milledge. 
1 $, Brisbane, 12 Oct 1914, J. Frost. 1 d , 1 km NNE of Collins Weir, W 
of Atherton, 17°15'S 145°17'E, 10 Feb 1989, G. Milledge. Id, 8 km E 
of Emuford, 30 Dec 1989, M.S. & B.J. Moulds. 1 d , Jardine River ferry 
crossing, 50 km S of Bamaga, 11°05'S 142°18'E, K. Walker. 1 9, Poison 
Creek Rd, 10 km W of Cooktown, 4 Apr 1991, G. Milledge (all NMV). 

Western Australia. 1 juv, 8 km SW of Walsh Point, Admiralty Gulf, 
14°37'S 125°48'E, 17 May 1983, D.C.F. Rentz & J. Balderson (ANIC). 

Diagnosis. Sphodropoda quinquedens can be separated 
from the other species of this genus by the distinctive pattern 
on the inside face of the fore leg (Fig. 15), the colour 
markings on the ventral surface of the costal area of the 
tegmen and the form of the male genitalia (Fig. 16). 

Description. Body (Figs. 9-10) moderately large. Colour 
brown with mottled tegmina, green with mottled tegmina 
or uniform green. Ridge on frontal shield with blunt point 
at apex. Prothorax with numerous scattered tubercles on 
dorsal surface, particularly in female; lateral margin finely 
denticulate, slightly lamellate in prozone. Fore leg (Fig. 15) 
with inner face of coxa purplish in posterior half, anterior 
half orange brown with 6 contrasting pointed cream 
tubercles on anterior margin which extend basally as ridges 
across surface; inner face of femur without markings; inner 
face of first tarsal segment mostly black, of other tarsal 
segments black apically. Wings of both sexes longer than 
abdomen; ventral surface of costal area of tegmen rosy red 
on inner half, outer half black, whole area crossed with 
slender white bands and spots; discoidal area completely 
opaque in female, only partly so in male; hind wing with 
costal and discoidal area yellowish, anal area slightly smoky, 
particularly toward apex, in female, mostly hyaline in male. 
Male genitalia (Fig. 16) with apr relatively elongate and 
narrow distally; pa compact, shagreened, with anterior 
pointed lateral projection and broad blunt posterior 
projection; dpr with anterior projection much longer than 
posterior one. 

Measurements (mm). Body length, 8 64, $ 70. Pronotum 
length, 8 16, $ 21. Pronotum width, 8 6, $ 9. Fore coxa 
length, 8, 11, 9 15. Fore femur length, 8 13, 9 18. Hind 
femur length, 8 10, 9 14. Hind tibia length, 8 10, 9 16. 
Tegmen length, 8 46, 9 46. 

Immature stages. Early stages and ootheca unknown. 

Distribution and habits. Found in north eastern Queensland 
and one record from north Western Australia (Fig. 43). This 
species displays cryptic behaviour similar to that described 
above for S. tristis. This behaviour has been noted previously 
in S. quinquedens (Heath & Cowgill, 1989). 


Sphodropoda viridis Tindale 
Figs. 18-19, 36-38, 44 
Sphodropoda viridis Tindale, 1923: 446. 

Material examined. Holotype male, Mount Painter, Flinders Range, 
South Australia, H.G. Stokes, I 14061 (SAM). —Queensland. 2d, 27 
km SE of Cunnamulla, 27 Mar 1972, R.C. Fewis (ANIC). Id, Gilbert 
River, 27 Mar 1990, J. Hasenpusch. 2 d , Georgetown, 13 & 14 Apr 1991, 
G. Milledge (all NMV). 1 9 , Cunnamulla, H. Hardcastle (SAM).— New 
South wales. Id, 1 9, Trangie, 21 Apr 1956, F.J. Chinnick (ANIC).— 
Northern Territory. 1 9 , Barrow Creek Tel.Sta., 20 Mar 1955, K.H.F. 
Key. 5d, Clay Pan Well, 38 km NW of Tanami, 12 Apr 1963, F.J. 
Chinnick. 1 9,7 km SW of Coolibah HS, 15°34'S 130°54'E, 6 Jul 1968, 
M. Mendum (all ANIC). 1 d , Tourist Camp site, Ormiston Gorge National 
Park, Horn Centenary Expedition, 23°38'03"S 132°43'23''E, 10 Mar 
1995, P. Swinkels (NMV).—SOUTH AUSTRALIA. 1 d , Mt. Painter, Flinders 
Range, H.G. Stokes (SAM).— Victoria. 1 9 , Mildura Gun Club, 27 Mar 
1990, P. Robertson (NMV). —Western Australia. Id, 17 km NE of 
Cane River HS, 21°56'S 115°39'E, 27 Apr 1971, Key, Upton & Mitchell. 
Id, 3 km NNW of Mandora HS, SSW of Broome, 17 Apr 1963, F.J. 
Chinnick. Id, 1 km N of Midstream HS, 21°35'S 117°04'E, 15 Apr 
1971, Upton & Mitchell. Id, 1 9, 43 km NW of Wittenoom, 22 Apr 
1963, F.J. Chinnick. 19,5 km N of Wittenoom, 21 Apr 1963, F.J. 
Chinnick. 1 9, 48 km ESE of Wittenoom, 25 Apr 1963, F.J. Chinnick 
(all ANIC). Id, Hancock Gorge, 15 km S of Wittenoom, 15 May 1980, 
G. Marney & P. Duncan. 1 9, Minilya. 1 9,2.5 km E of Marandoo Camp, 
22°38'S 118°06’E, 5-19 May 1980, T.F. Houston (all WAM). 

Diagnosis. This species can be distinguished by the uniform 
green colouration, lack of markings on the inner face of the 
fore coxa (Fig. 37) and the form of the male genitalia (Fig. 
38). 

Description. Body (Figs. 18-19) medium to rather large. 
Colour uniform green. Ridge on frontal shield with blunt 
point at apex. Prothorax with a few scattered tubercles on 
dorsal surface, more so in female; lateral margin finely 
denticulate. Fore leg (Fig. 37) with inner face of coxa 
without markings, anterior margin with 7-10 denticles; inner 
face of femur with two small dark spots at base of spines; 
inner face of all tarsal segments black. Wings of male longer 
than abdomen, those of female as long as or longer than 
abdomen; ventral surface of costal area of tegmen reddish; 
discoidal area completely opaque in female, only partly so 
in male; hind wing with costal and discoidal area yellowish, 
anal area mostly hyaline. Abdomen with median black spot 
on proximal margin of ventral sclerites 2-5. Male genitalia 
(Fig. 38) apr of moderate length; pa with smooth, pointed 
elongate anterior projection, shagreened and blunt 
posteriorly; distal projections of dpr short and broad, one 
directed anterolaterally, the other posterodorsally. 

Measurements (mm). Body length, 8 48-64, 9 44-60. 
Pronotum length, 8 12-18, 9 14-19. Pronotum width, 8 
3.5-6, $ 5-6.5. Fore coxa length, 8 9-11, $ 9-12.5. Fore 
femur length, 8 9-12, 9 10-14. Hind femur length, 8 8- 
12, 9 9-12. Hind tibia length, 8 8.5-14, 9 11-14. Tegmen 
length, 8 36-47, 9 30-40. 

Immature stages. Immature stages and ootheca unknown. 

Distribution and habits. Found across the drier parts of 
the continent (Fig. 44). This species displays cryptic 
behaviour similar to that described for S. tristis and S. 
quinquedens. 


200 Records of the Australian Museum (2005) Vol. 57 


Figs. 20-23, 43 

Material examined. Holotype 6 , Uluru campsite, Horn Centenary 
Expedition, 25°23'10''S 131°00'46''E, Northern Territory, 25 Mar 1995, 
G. Milledge (NMV). Paratypes (7 <3 3): Id, Ayers Rock, Northern 
Territory, 16 Feb 1967, M.S. Upton; 6d <3, 56 km Wof Amata, Musgrave 
Ranges, 26°09’S 130°35’E, South Australia, 20-21 Jan 1982, D.C.F. & 
B.G.F. Rentz & R. Honeycutt (all ANIC). 


Description. Male only, female unknown. Medium sized 
species. Body colour partly green and partly purplish brown. 
Head (Fig. 21) purplish brown; narrower than other species 
in the genus, ridge on frontal shield without blunt point at 
apex. Prothorax purplish brown, paler at margins; with a 
few scattered tubercles on dorsal surface; lateral margin 


Milledge: revision of three mantid genera 201 


finely denticulate. Fore legs green; inner face of coxa 
without markings (Fig. 22), anterior margin with 7 denticles; 
inner face of fore femur with three small dark spots at base 
of spines; inner face of all tarsal segments black. Mid and 
hind legs green. Wings about as long as abdomen; dorsal 
and ventral surface of costal area of tegmen purplish, 


Figs. 20-23. Sphodropoda lepida, male. (20) Dorsal view; (21) head, anterior 
view; (22) fore leg, internal view; (23) genitalia, dorsal view. Scales = 2 mm. 


stronger beneath, with narrow whitish marginal band; 
discoidal green along anterior margin, major veins flushed 
with purplish brown giving banded appearance, remainder 
hyaline; hind wing with costal and discoidal area greenish, 
anal hyaline. Abdomen with median black spot on proximal 
margin of ventral sclerites 2-5. Male genitalia (Fig. 23) with 
short, broad apr; pa with smooth, pointed anterior 
projection, shagreened posteriorly with pointed lateral and 
blunt dorsal projections; apr with anterior projection 
moderately elongate, posterior one extremely short. 


202 Records of the Australian Museum (2005) Vol. 57 


Figs. 24-25. Trachymantis obesa. 
(24) Male, dorsal; (25) female, 
dorsal. Scale =10 mm. 


Measurements (mm, paratype male). Body length, S 46. 
Pronotum length, S 12. Pronotum width, S 4. Fore coxa 
length, S 9. Fore femur length, S 11. Hind femur length, 
S 8.5. Hind tibia length, $ 10. Tegmen length, S 32. 

Immature stages. Immature stages and ootheca unknown. 

Distribution and habits. At present, only known from 
central Australia (Fig. 43). Habits unknown but likely to be 
similar to other members of the genus. 


Trachymantis Giglio-Tos 

Trachymantis Giglio-Tos, 1917: 47. Type species Trachymantis 
obesa Giglio-Tos, by monotypy. 

Ngawala Tindale, 1923: 447. Type species Ngawala dentifrons 
(Stal), by original designation. 


Milledge: revision of three mantid genera 203 


Diagnosis. Head about as high as wide or higher than wide, 
frontal shield with pronounced tooth at apex; preacetabular 
spine strong; ventral face of fore femur tuberculate, 
especially toward outer margin; male macropterous, female 
brachypterous, ventral face of costal area of tegmen partially 
dark grey, mid and hind legs normal. Can be distinguished 


ins: 


204 Records of the Australian Museum (2005) Vol. 57 


Figs. 28-33. Trachymantis spp. (28) Trachymantis obesa, male head, anterior 
view; (29) same, fore leg, internal view; (30) same, male genitalia, dorsal 
view. (31) Trachymantis dentifrons, female head, anterior view; (32) same, 
fore leg, internal view; (33) same, male genitalia, dorsal view. Scales = 2 mm. 


Milledge: revision of three mantid genera 


from Sphodropoda and Zopheromantis by the presence of 
the pointed projection at the apex of the frontal shield. 

Description. Body rather large and robust, male macro- 
pterous, female brachypterous. Head about as wide as high 
or higher than wide, not significantly compressed 
anteroposteriorly, apical margin gently arched; eyes 
rounded, not particularly prominent; frontal shield slightly 
to considerably wider than high, flat, with distinct ridge 
laterally and apically produced into a tooth at apex. 
Pronotum moderately elongate, supracoxal expansion 
distinct, dorsal surface finely granulate, mostly in prozone, 


.... .y*.jl 


206 Records of the Australian Museum (2005) Vol. 57 


Figs. 36-41. Sphodropoda viridis and Zopheromantis loripes. (36) Sphodropoda viridis, female head, anterior view; (37) same, fore 
leg, internal view; (38) same, male genitalia, dorsal view; (39) Zopheromantis loripes female head, anterior view; (40) same, fore leg, 
internal view showing variation in coxal pattern; (41) same, male genitalia, dorsal view. Scales = 2 mm. 


margins finely denticulate, ventral surface with preacetab- 
ular spine prominent. Fore coxa as long as metazone of 
pronotum, with or without markings, a number of denticles 
on anterior margin. Fore femur finely tuberculate on ventral 
surface, particularly toward outer margin; with four discoidal 
spines, relative length longest to shortest 3-2-1-4; with four 
outer and 15-18 inner spines, claw groove situated at about 

Key to species 

1 Facial shield much higher than wide. 

—— Facial shield about as high as wide. 


a third the total distance from base. Tegmen with costal 
area opaque, partially grey beneath; discoidal area entirely 
opaque to mostly hyaline. Hind wing with costal and 
discoidal area pigmented, anal area largely hyaline. Mid 
and hind legs normal, hind femur more than half the length 
of abdomen. Abdomen moderately elongate, cerci short, 
slender and cylindrical. Male genitalia with dpr bifurcate. 

of Trachymantis 

. T. obesa 

. T. dentifrons 


Milledge: revision of three mantid genera 207 


Trachymantis obesa Giglio-Tos 

Figs. 24-25, 28-30, 45 

Trachymantis obesa Giglio-Tos, 1917: 47. 

Material examined. Holotype female, Hermannsburg, Central 
Australia, H.J.Hillier, 1911-311 (BMNH).—QUEENSLAND: 2d, 3 km 
W of Bundeena HS, SW of Quilpie, 12 Feb 1972, R.C. Lewis. 1 9, 14 
km S of Quilpie, 14 Nov 1971, R.C. Lewis. 13,5 km N of Eromanga, 3 
Nov 1967, R.C. Lewis (all ANIC).—New South Wales: 2d, 18 km 
WNW of Bourke, 9 Feb 1972, R.C. Lewis (ANIC).—Northern 
Territory: Id, Wigley Waterhole, 8 km N of Alice Springs, 16 Feb 
1966, Britton, Upton & Mclnnes. 19, 22 km NNW of Kulgera HS, 8 
Oct 1955, L.J. Chinnick. 1 juv, 37 km N of Alice Springs, 17 Nov 1966, 
White, Blackith & Blackith (all ANIC). 1 9, Kathleen Creek, Watarrka 
National Park, 24°20'38"S 131°40'54"E, 21 Oct 1994, G. Milledge 
(NMV).—Western Australia: 2d, lOkmNEofMenzies, 14 Jan 1989, 
M.S. & B.J. Moulds. Id, 55 km ESE of Mount Magnet, 18 Jan 1989, 
M.S. & B.J. Moulds. Id, 110 km S of Mount Magnet, 19 Jan 1989, 
M.S. & B.J. Moulds. 2d, 25 km E of Sandstone, 17 Jan 1989, M.S. & 
B.J. Moulds. 

Diagnosis. Can be distinguished from T. dentifrons by the 
facial shield being higher than wide. 

Description. Body (Figs. 24-25) rather large. Colour 
uniform green or bluish green. Head (Fig. 28) higher than 
wide; frontal shield distinctly higher than wide with ridge 
produced into prominent point at apex. Pro thorax with fine 
scattered tubercles on dorsal surface, mostly in prozone, 
more so in female; lateral margin finely denticulate. Fore 
leg (Fig. 29) with inner face of coxa bluish, apricot on distal 
quarter, anterior margin with five black spots and 11-12 
small denticles; inner face of fore femur apricot in lower 
half with large black patch proximally and three narrow 
black bands distally; inner face of fore tibia orange. Wings 
of male longer than abdomen, those of female slightly 
shorter than abdomen; ventral surface of costal area of 
tegmen dark bluish grey with narrow apricot marginal band; 
discoidal area completely opaque in female, only partly so 
in male; hind wing with costal and discoidal area yellowish 
green, anal area mostly hyaline. Male genitalia (Fig. 30) 
with very short blunt apr; pa with single, elongate hook¬ 
like projection; distal projections of dpr elongate, narrow 
and of similar length. 

Measurements (mm). Body length, 8 54-70, 9 12-15. 
Pronotum length, <3 17-20, 9 24-26. Pronotum width, 8 
4.5-6, 9 8. Fore coxa length, 8 14-17, 2 16-18. Fore 
femur length, 8 12-14, 9 21-22. Hind femur length, 8 
11-15, $ 18. Hind tibia length, 8 12-16, $ 20. Tegmen 
length, 8 34-46, 9 34-37. 

Immature stages. Later instar nymphs similar in appearance 
to adults, early instars and ootheca unknown. 

Distribution and habits. Recorded (Fig. 45) from the drier 
parts of Queensland, New South Wales, Northern Territory 
and Western Australia. The single specimen I have collected 
was found in a large Acacia shrub. Whilst in captivity it did 
not display cryptic behaviour as found in Sphodropoda. 

Trachymantis dentifrons (Stal, 1877) n.comb. 

Figs. 26-27, 31-33, 46 

Hierodula ( Sphodropoda ) dentifrons Stal, 1877: 56. 
Sphodropoda dentifrons (Stal), Kirby, 1904: 242. 

Ngawala dentifrons (Stal), Tindale, 1923: 447. 


Material examined. Two syntype females, Australia, Boucard 
(NHRM). 

Other material. New South Wales: 13, 3 km NW of Bramah HS, 
NW of Balranald, 34°24'S 143°14’E, 24 Oct 1983, D.C.F. Rentz & M.S. 
Harvey. 13, 26 km S of Pooncarie, 21 Jan 1980, M.J.D. White (both 
ANIC).—Northern Territory: 13, 17 km SSE of Mt. Harris, 25 Aug 
1955, L.J. Chinnick (ANIC). 23, Uluru Camp Site, Horn Centenary 
Expedition, 25°23T0"S 131 o 00’46"E, 24 & 25 Oct 1994, G. Milledge. 
13, Uluru Mallee site, Horn Centenary Expedition, 25°24’07"S 
130°59'19"E, 26 Mar 1995, G. Milledge. 13, Kathleen Creek, Mallee 
site, Horn Centenary Expedition, Watarrka NP, 24°20'54"S 131°40’36"E, 
24 Mar 1995, P. Lilywhite (all NMV).—SOUTH AUSTRALIA: 2 juv, Lowan 
Stn., 7 km S of Sherlock, 4 Mar 1953 & 20 Dec 1954, L.J. Chinnick. 2 
juv, Lowan Stn., 7 km S of Sherlock, 9 Mar 1953. 13, 56 km W of 
Amata, Musgrave Ranges, 26°09’S 130°35’E, 20-21 Jan 1982, D.C.F. 
Rentz & R. Honeycutt. 23, 1 $, 20 km NE of Kimba, 33°07’S 136°38’E, 
5 Oct 1982, D.C.F. Rentz. 13, Brachina Gorge, Flinders Ranges, 9 Nov 
1987, J.C. Car dale. 2 juv, 13 km WNW of Minnipa, 22 Oct 1968, J. 
Balderson & M.S. Upton. 1 juv, 5 km SSE of Minnipa, 30 Oct 1969, 
Key & Upton. 13, 18 km SSW of Pinnaroo, 35°24’S 140°48'E, 25 Oct 
1983, D.C.F. Rentz & M.S. Harvey. 13, 1.5 km ESE of Ooldea, 3 Oct 
1968, Key, Upton & Balderson (all ANIC). 12, Overland Railway, 40 
km W of Kychering Soak, Dec 1908, Mr Chandler (NMV). 13, Darke’s 
Peak, R.G. Walsh (SAM).—Victoria: 1 2, Big Desert, P. Robertson. 
12, Big Desert, 35°46’S 140°58’E, 15 Apr 1987, P. Johnson. 1 2, N of 
Chinamans Well, Big Desert, 2 Mar 1982, A.J. Coventry. 1 juv, 
Grampians, Nov 1885. 1 2, 5.4 km E of Hattah, 34°47'S 142°20’E, 24 
Jan 1986, G. Milledge. 13, 15 km SSW of Hattah, 34°54'S 142°15’E, 

24 Jan 1986, G. Milledge. 1 2, 19.2 km SW of Hattah, 34°54’S 142°09’E, 
21 Jan 1987, G. Milledge. 1 2,20.8 km SE of Hattah, 34°52'S 142°28'E, 

25 Jan 1987, G. Milledge. 13, Inglewood. 1 m, 0.5 km W of Lake Hattah, 
28 Jan 1986, G. Milledge. 1 2, Lake Hattah, 18 Oct 1985, G. Milledge. 

12, Lake Hattah, 8 Apr 1982, G. Milledge. 13, Mallee District, 3 Mar 
1914, C. French. 12, 12.8 km N of Millewa South Bore, 34°40’S 
141°04’E, 27 Mar 1986, G. Milledge. 1 2, 3.1 km N of Millewa South 
Bore, 34°45'S 141°04E, 14 Nov 1985, G. Milledge. 1 2, 0.6 km N of 
Millewa South Bore, 34°46'S 141°04’E, 21 Feb 1986, G. Milledge. 13, 
7 km SSW of Morkalla, 34°26’S 141°08'E, 15 Nov 1985, G. Milledge. 
12,6 km SW of Murray Valley Hwy & Annuello Rd junction, 34°50’S 
142°35’E, 10 Feb 1986, G. Milledge. 23, 16.8 km SSW of Murrayville, 
35°25’S 141°10’E, 18 Feb 1987, G. Milledge. 22, 16.8 km SSW of 
Murray ville, 35°25’S 141°10’E, 23 Feb 1987, G. Milledge. 1 2, 16.8 km 
SSW of Murrayville, 35°25'S 141°10’E, 19 Feb 1986, G. Milledge. 12, 
7 km SW of Nowingi, 34°39'S 142° 16E, 16 Oct 1985, G. Milledge. 
12,1 juv, Ouyen, 26 Jun 1912, W.A. Hall. 13, Ouyen, 26 Apr 1911, 
S.S. Scaree. 13, Round Swamp, Big Desert, 35°42’S 141°43E, 28 Mar 
1985, G. Milledge. 13,6.9 km N of Round Swamp, Big Desert, 35°41'S 
141°43’E, 28 Mar 1983, G. Milledge. 3 2, The Springs, Murrayville 
Track, Big Desert, 28 Mar 1985, G. Milledge. 2 2, Waithe Fauna Reserve, 
19 & 22 May 1986, G. Milledge. 1 2,4 km N of Wedderburn, 36°23’S 
143°37’E, 18 Nov 1989, G. Milledge. 13, Woomelang, 4 Dec 1902, Mr 
Hill (all NMV).—Western Australia: 1 2, Nedlands, Jan 1941, K.R. 
Norris. 13, Eyre Tower Microwave Station, 22 km ESE of Cocklebiddy, 
32°08'S 126°17’E, 7 Oct 1982, D.C.F. Rentz. 1 juv, 23 km ESE of 
Cocklebiddy, 32°08’S 125°18’E, 25 Oct 1977, M.S. Upton & J.E. Feehan. 

13, Cocklebiddy Repeater Stn., 36 km SW of Caiguna, 3 Nov 1984, 
D.C.F. Rentz. 23, Lake Douglas, 12 km SW of Kalgoorlie, 13 Jan 1989, 
M.S. & B.J. Moulds. 13, Kununurra, 16-19 Sep 1979, L.P Kelsey. 22, 
13 km SSE of Dongara, 28°21'S 115°00E, 12 Sep 1981, D.C.F. Rentz. 
12, Cape Naturaliste Lighthouse, 33°32’S 115°01'E, 15 Feb 1980, D.C.F. 
Rentz. 1 2, 19 km S of Menangina HS, 85 km E of Menzies, 24 Feb 

1983, M.J. Whitten. 1 2,27 km S of Balladonia Motel, 32°35'S 123°36'E, 
13 Feb 1978, D.C.F. Rentz & M.J.D. White. 1 2,31 km S of Balladonia 
Motel, 9 Feb 1980, D.C.F & B.G.F. Rentz. 13, 36 km WNW of 
Balladonia Hotel, 5 Mar 1982, M.J.D. White. 13 60 km SSW of 
Norseman, 32°38'S 121°29’E, 17 Nov 1969, Key & Upton. 12, 
Cockleshell Gully, 19 km NNE of Jurien, 30°09'S 115°07'E, 25 Oct 

1984, D.C.F. Rentz. 12, Crawley, 15 Mar 1934, K.R. Norris. 23, 33 
km ENE of Perth, 30 Jan 1967, M.S. Upton. 1 juv, 37 km SW of Mt. 
Ragged, 33°40’S 123°11'E, 2 Nov 1977, M.S. Upton & J.E. Feehan. 
23, 5 km SW of Mt. Ragged, 12 Nov 1969, Key & Upton. 13, 20 km 
SSE of Yanchep, 9 Sep 1981, D.C.F. Rentz. 1 2, 15 km WSW of Fraser 
Range HS, 32°04'S 122°39’E, 6 Sep 1981, D.C.F. Rentz. 1 2,27 km W 
of Eneabba, 29°49'S 114°59'E, 9 Sep 1981, D.C.F. Rentz. 13, Fitzgerald 
National Park, 12 km NE of Hopetoun, 33°51'S 120°09'E, 10 Feb 1980, 
D.C.F. & B.G.F. Rentz (all ANIC). 13, Junga Dam, 23 km ESE of 


208 Records of the Australian Museum (2005) Vol. 57 


Kalbarri, 6 Jan 1969, Kalbarri Survey. 1 3 , Miling, Jan 1976, T. Verbakel. 
lc?, 7.5 km E of Yuinmery HS, 28°34'S 119°01'E, 11-19 Feb 1980, T.F. 
Houston eta/., 1 juv, Roebourne, 1922. 1 juv, Cunderdin, Jul-Aug 1913. 
1 juv, Fitzgerald River Reserve, Jul 1970, A. Baynes. 1 juv, 13.6 km 
SSW of Mt Jackson, 30°59'S 119°07E, 5-11 Sep 1979, T.F. Houston et 
al., 1 juv, Corrigin. 1 juv, 1.6 km W of Mundaring Weir, 6 Mar 1963, 
J.Dell. 1 juv, 30 km N of Bullfinch, 30°59’S 119°07'E, 7 Sep 1979, T.F. 
Houston et al., 1 juv, Kanowna, 30°36'S 121°36'E, 16 Oct 1968, R.R 
McMillan. 1 juv, Carollgouda Well, Kalbarri area, 30 Nov 1968, Hale 
School (all WAM). 

Diagnosis. Can be distinguished from T. obesa by the facial 
shield being approximately as high as wide. 

Description. Body (Figs. 26-27) rather large. Colour 
uniform green or bluish green. Head (Fig. 31) and frontal 
shield approximately as high as wide with ridge of frontal 
shield produced into prominent point at apex. Prothorax 
with fine scattered tubercles on dorsal surface, more so in 
female; lateral margin finely denticulate. Fore leg (Fig. 32) 
with inner face of coxa without markings, anterior margin 
with 9-11 small denticles; inner face of fore femur with 
large dark patch proximal to claw groove (not apparent in 
some specimens) and three small dark spots distal of claw 
groove; femora of mid and hind legs often orange red in 
proximal half. Wings of male longer than abdomen, those 
of female shorter than abdomen; ventral surface of costal 
area of tegmen dark bluish grey in anterior half; discoidal 
area completely opaque in female, only partly so in male; 
hind wing with costal and discoidal area yellowish green, 
anal area hyaline. Abdomen of female with posterior margin 
of fourth and particularly fifth stemite curved ventrally. Male 
genitalia (Fig. 33) with moderately elongate apr, narrowed 
distally; pa with anterior hook-like projection and posterior 
knob-like projection; distal projections of dpr of moderate 
length, anterior one longer than posterior one. 

Measurements (mm). Body length, S 53-75, ? 59-75. 
Pronotum length, S 19-20, ? 21-26. Pronotum width, S 

5.5- 6, 9 7-9. Fore coxa length, S 12-15, $ 15-19. Fore 
femur length, S 13-16, 9 18-22. Hind femur length, S 

14.5- 15, 9 17-22. Hind tibia length, S 15-16, 9 17-23. 
Tegmen length, S 42-54, 9 28-38. 

Immature stages. First two instars dark coloured and ant¬ 
like in behaviour and appearance. Ootheca squat, cream to 
sandy orange in colour. Appears to be mostly laid on the 
ground, usually attached to a piece of timber. The peculiar 
form of abdominal sternites 4-5 in the female may play a 
role in ootheca deposition. 

Distribution and habits. Found across the arid regions of 
southern and central Australia (Fig. 46). Appears to be most 
commonly associated with mallee vegetation where the blue 
green body colour blends well with the leaves of certain 
mallee eucalypts. I have observed numerous individuals of 
this species, in nature and captivity, and never seen the 
cryptic behaviour displayed by members of the genus 
Sphodropoda. Adult females are commonly caught in pitfall 
traps, probably in the process of seeking oviposition sites. This 
would also suggest they are not capable of effective flight. 

Remarks. The original description appears to be based on 
one specimen although this is not specified. One of the two 
female syntypes is labelled type, the other cotype. These 
are printed labels and there is no indication that they were 
placed there by the author. Therefore, I have decided not to 


designate a lectotype. A syntype specimen is illustrated by 
Sjostedt (1930, plate 8, fig. 1) 

Zopheromantis 

Zopheromantis Tindale, 1924: 550. Type species Zopheromantis 

trimaculata Tindale, by monotypy. 

Diagnosis. Head approximately as high as wide, frontal 
shield without pronounced tooth at apex; preacetabular spine 
strong; ventral face of fore femur tuberculate, especially 
toward outer margin; male macropterous, female brachy- 
pterous, ventral face of costal area of tegmen red, mid and 
hind legs normal. Can be distinguished from Trachymantis 
by the absence of the apical tooth on the frontal shield and 
from Sphodropoda by the length of the mid and hind legs, 
the femur of the hind leg being distinctly longer than half 
the length of the abdomen. 

Description. Body small to medium in size, male 
macropterous, female brachypterous. Head about as wide 
as high, not significantly compressed anteroposteriorly, 
apical margin gently arched; eyes rounded, not particularly 
prominent; frontal shield slightly wider than high, flat, with 
distinct ridge laterally and apically, forming blunt point at 
apex. Pronotum moderately elongate, rather slender, 
supracoxal expansion distinct, dorsally sparsely granulate 
in prozone, metazone almost smooth, margins finely 
denticulate in female largely entire in male, ventral surface 
with preacetabular spine prominent. Fore coxa as long as 
metazone of pronotum, with or without markings, a number 
denticles on anterior margin. Fore femur finely tuberculate 
on ventral surface, particularly toward outer margin; with 
four discoidal spines, relative length longest to shortest 
3214; with four outer and 16-17 inner spines, claw groove 
situated at about a third the total distance from base. Tegmen 
with costal area opaque, red beneath; discoidal area entirely 
opaque to mostly hyaline. Hind wing with costal and 
discoidal area pigmented, anal area largely hyaline. Mid 
and hind legs normal, hind femur more than half the length 
of abdomen. Abdomen moderately elongate, cerci short, 
slender and cylindrical. Male genitalia with dpr bifurcate. 

Zopheromantis loripes (Tindale, 1923) n.comb. 

Figs. 34-35, 39-41, 47 

Sphodropoda loripes Tindale, 1923: 446. 

Zopheromantis trimaculata Tindale, 1924: 551, new synonym. 
Hierodula trimaculata Beier, 1935: 82. 

Material examined. Fectotype male (here designated) of 
Sphodropoda loripes, Cunnamulla, Queensland, H. Hardcastle, 1-14071 
(SAM). Holotype female of Zopheromantis trimaculata, Teetulpa, South 
Australia, Jan. 1904, G. Farrand, 1-14590 (SAM).— Queensland: 13, 
Coopers Creek, 10.7 km NE of Windorah, 25°22'18"S 142°44'4rE, 4 
Apr 1991, G.Y. Maynard & G. Davis. 1 S, 33 km W of Quilpie, 16 Mar 

1964, F.J. Chinnick. 13, 18 km NE of Thargomindah, 15 Mar 1964, 
F.J. Chinnick. 13, 23 km SW of Toompine, 10 Apr 1970, R.C. Fewis 
(all ANIC). 19, Augathella, 25°48’S 146°25F, 18 Jan 1993, G. Milledge 
(NMV).— New SOUTH WALES: 33,18kmWNWofBourke,9Feb 1972, 
R.C. Fewis. 5 9,6 km W of Cobar, 23 Mar 1972, White & Jaworska. 
1 9,7 km W of Cobar, 20 Feb 1963, F.J. Chinnick. 13,2 juv, 7 km W of 
Cobar, 1 Feb 1964, M.J.D. White. 1 9, 15 km SW of Enngonia, 14 Jan 

1965, M.J.D. White. 13, 37 km SE of Wanaaring, 10 Feb 1972, R.C. 
Fewis. 13,1 km W of Warri gate, 11 Feb 1972, R.C. Fewis. 1 juv, 17 
km SE ofWilcannia, 31 Jan 1964, M.J.D. White (all ANIC).— NORTHERN 
Territory: 23, 37 km S of Alice Springs, 15 Feb 1966, Britton, Upton 
& Mclnnes. 13, 103 km NNW of Alice Springs, 12 Feb 1966, Britton, 


Milledge: revision of three mantid genera 209 


Figs. 42-47. Distributions of Australian mantids: (42) Sphodropoda tristis; (43) • S. quinquedens, ▲ S. lepida; 
(44) S. viridis; (45) Trachymantis obesa\ (46) T. dentifrons; (47) Zopheromantis loripes. 


Upton & Mclnnes. Id, Barrow Creek Tel.Sta., 20 Mar 1955, K.H.L. 
Key. Id, 3 km S of Barrow Creek, 13 Feb 1966, Britton, Upton & 
Mclnnes. 1$, 25 km NW of Mount Wedge HS, 10 Apr 1963, L.J. 
Chinnick. 1 d, 5 km W of Mulga Park HS, 24 Mar 1963, L.J. Chinnick. 
Id, 26 km WSW of Mulga Park HS, 26°00'S 131°25'E, 18 Jan 1982, 
D.C.F., B.G.F. Rentz. 1 d, 25 km NE of Narwietooma HS, 9 Apr 1963, 
L.J. Chinnick & R. Honeycutt. Id, Taylors Creek, 47 km N of Barrow 


Creek township, 22 Jan 1984, M.S. & B.J. Moulds. Id, Wauchope, 21 
Mar 1955, K.H.L. Key (all ANIC). 2d, Kathleen Creek, Watarrka 
National Park, Horn Centenary Expedition, 24°20'38"S 131°40’54"E, 
14 Mar 1995, G. Milledge (both NMV).— South Australia: 1 9,7 km 
W of Iron Knob, 28 Mar 1964, M.J.D. White. Id, Marla Bore, 50 km W 
of Welbourne Hill, 23 Mar 1963, L.J. Chinnick (both ANIC).— Western 
Australia: 2d, 23 km WSW of Barradale, 22°56'S 114°45'E, 30 Mar 


210 Records of the Australian Museum (2005) Vol. 57 


1971, Upton & Mitchell. 1$, Canegrass, 8 Feb 1981, M.J.D. White. 
1 $, 5 km NE of Kanowana ruins, near Broad Arrow, 18Feb 1981,M.J.D. 
White. Id, 5 km NE of Kanowana ruins, near Broad Arrow, 24 Feb 
1981, M.J.D. White. 19, Malcolm, 21 Feb 1979, M.J.D. White. 19,1 
km S of Malcolm, 28°47'S 121°31F, 19 Feb 1978, D.C.F.Rentz& M.J.D. 
White. 1 d, 6 km SE of Mt. Boyce, near Yerilla HS, SE of Eeonora, 15 
Feb 1983, M.J.D. White. 2 juv, Mt. Eeonora, near Eeonora, 13 Feb 1981, 
M.J.D. White. 19, Mt. Leonora, near Leonora, 10 Feb 1983, M.J.D. 
White. 1 9, 3 km NNE of Mt. Ross, NW of Leonora, 28°41’S 121°03'E, 
18 Feb 1978, D.C.F. Rentz & M.J.D. White. 1 d, New Yamarna HS, S of 
Minnie Creek, near Virginia Range, 31 Jan 1967, M.J.D. White. 1 9,43 
km NW of Wittenoom, 22 Apr 1963, L.J. Chinnick (all ANIC). 19, 
Middalya, 4 Feb 1902, T. Watt (NMV). 2d, Kathleen Valley, 1963, T. 
Moriarty. 1 d, Lake Violet. 1 d, Laverton. Id, 2.5 km N of Mt. Linden, 
29°19'S 122°25'E, 17-23 Mar 1979, T.F. Houston. Id, 19, Nolba, 
28°22'S 114°52'E. Id, Warburton Ranges, 25 Mar 1963, H. De Graff. 
Id, 7.5 km E of Yuinmery HS, 28°34'S 119°01'E, 11-19 Feb 1980, T.F. 
Houston. 1 d, 9 km SE of Yuinmery HS, 28°34'S 119°01'E, 25 Mar 1979, 
T.F. Houston (all WAM). 

Description. Body (Figs. 34-35) size varies from small to 
medium. Body colour uniform green or bluish green. Head 
(Fig. 39) approximately as high as wide; frontal shield 
slightly wider than high, more so in male. Fore leg (Fig. 
40) with inner face of coxa without markings or with one 
to three dark spots, anterior margin with 7-8 small tubercles; 
inner face of femur without markings; inner face of tarsal 
segments dark at distal end. Wings of male longer than 
abdomen, those of female slightly shorter than abdomen; 
ventral surface of costal area of tegmen red with narrow 
pale band on anterior margin; discoidal area completely 
opaque in female, only partly so in male; hind wing with 
costal and discoidal area yellowish green, anal area hyaline. 
Male genitalia (Fig. 41) with short, stout apr, pa with single, 
blunt pointed anterolateral projection, dorsal edge slightly 
folded; distal projections of dpr moderately elongate and 
narrowed. 

Measurements (mm). Body length, 3 37-50, 9 39-58. 
Head width, 3 6-7, 9 7-8. Head depth, 3 3.5-5, 9 4.5- 
6.5. Pronotum length, 3 11-13, 9 15-22. Pronotum width, 
3 3-4.5, 9 4-6. Fore coxa length, 3 7-9, 9 11-14. Fore 
femur length, 3 8.5-12, 9 12-16. Hind femur length, 3 
8.5-12, 9 12-18. Hind tibia length, 3 9.5-12, 9 13-19. 
Tegmen length, 3 27-35, 9 19-29. 

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

Distribution and habits. Found in the arid regions of the 
continent (Fig. 47). The few specimens of this species that 
I have collected have been found in Acacia shrubs. None 
displayed the cryptic behaviour associated with Sphodro- 
poda. 

Remarks. The female syntype of Sphodropoda loripes 
described by Tindale is, in fact, a female of Sphodropoda 
viridis. Therefore I have designated the male syntype as 
the lectotype. 


Acknowledgments. I would like to thank M. Moulds (AM), D. 
Rentz (ANIC), J. Marshall (BMNH), K. Johanson (NHRM), K. 
Walker (NMV), J. Forrest (SAM), T. Houston (WAM) and M. 
Ohl (ZMHU) for the loan of specimens. S. Lindsay (AM) prepared 
the S.E.M. images. 


References 

Balderson, J., 1984. Catalogue of Australian Mantodea. CSIRO 
Technical Paper no. 23. 

Balderson, J., D.C.F. Rentz & A.M.E. Roach, 1998. Mantodea. 
In Zoological Catalogue of Australia, Archaeognatha, 
Zygentoma, Blattodea, Isoptera, Mantodea, Dermaptera, 
Phasmatodea, Embioptera, Zoraptera, ed. W.W.K. Houston 
& A. Wells, pp. 251-277, vol. 23. Melbourne: CSIRO 
Publishing, Australia. 

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

Beier, M., 1964. Blattopteroidea: Mantodea. Bronns Klassen und 
Ordungen des Tierreichs (5) (3) 6: 849-970. Leipzig: 
Akademische Verlagsgesellschaft Geest & Portig K.-G. 

Beier, M., 1965. Die Mantodeen Neu-Guineas. Pacific Insects 7 
(3): 473-502. 

Chapman, R.C., & J. Balderson, 1984. Ovipositional behaviour 
of Sphodropoda tristis Saussure (Mantodea: Mantidae). 
Australian Entomological Magazine 11(1): 5-7. 

Giglio-Tos, E., 1911. Mantidi Esotici. II. Le Polispilote. Societa 
entomologica Italiana, Bollettino 42: 3-38. 

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

Heath, G.L., & G. Cowgill, 1989. Rearing and Studying the 
Praying Mantids. Leaflet no. 36, Amateur Entomologists’ 
Society, London. 

Hevers, J., & H. Liske, 1991. Lauernde Gefahr. Das Leben der 
Gottesanbeterinnen. Braunschweig: Staatliches Naturhistor- 
isches Museum. 

Kirby, W.F., 1904. A Synonymic Catalogue of Orthoptera. Volume 
1. London: British Museum (Natural History). 

Macleay, W.S., 1826. In Narrative of a Survey of Inter-tropical 
and Western Coasts of Australia Performed between the Years 
1818 and 1822 by Captain Phillip P. King, vol. 2, by PP King. 
London: John Murray. 

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

Milledge, G., 1997. Revision of the tribe Archimantini (Mantodea: 
Mantidae: Mantinae). Memoirs of the Museum of Victoria 
56(1): 1-63. 

Preston-Mafham, K., 1990. Grasshoppers and Mantids of the 
World. London: Blanford. 

Saussure, H., 1871. Melanges Orthopterologiques. Fascicule 3. 
Memoires de la Societe de Physique et d’Histoire naturelle de 
Geneve 21: 1-214. 

Sjostedt, Y., 1918. Results of Dr E. Mjoberg’s Swedish scientific 
expeditions to Australia 1910-1913. 17. Mantidae and 
Phasmidae. Arkiv fur Zoologi 11(19): 1-60. 

Sjostedt, Y., 1930. Orthopterentypen im Naturhistorischen 
Reichsmuseum zu Stoc kh olm. I. Mantidae. Arkiv fur Zoologi 
21A(32): 1-43. 

Stal, C., 1871. Orthoptera quaedam africana. Ofversigt af Konglia 
Vetenskaps-Akademiens forhandlingar 3: 375-401. 

Stal, C., 1877. Systema Mantodeorum. Bihang till Konglia 
Svenska Vetenskaps-Akademiens Handlingar 4(10): 1-91. 

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

Tindale, N., 1924. Review of Australian Mantidae. Part 2. Records 
of the South Australian Museum 2: 547-552. 

Westwood, J.O., 1889. Revisio Insectorum Familie Mantidarum, 
Speciebus Novis aut Minus Cognitis Descriptis et Delineatis. 
London: Gurney and Jackson. 

Manuscript received 19 August 2003, revised 4 March 2004 and accepted 

7 April 2004. 

Associate Editor: M.S. Moulds. 


© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 211-220. ISSN 0067-1975 


A New Middle Devonian Arthrodire (Placoderm Fish) 
from the Broken River Area, Queensland 


Gavin C. Young 


Department of Earth and Marine Sciences, 
Australian National University, Canberra ACT 0200, Australia 

gy oung @ ems. anu .edu.au 


Abstract. Associated bones of the trunk-armour belonging to a large brachythoracid arthrodire are 
described as Confractamnis johnjelli n.gen. and n.sp. The specimen comes from strata of probable 
Eifelian age in the Broken River Group of Queensland. It shows a range of derived brachythoracid 
features including reduction of dermal ornament, and strong development of the dermal neck joint 
articulation. The posterior dorsolateral and posterior lateral plates of the trunk armour were high and 
narrow, and the anterior lateral plate had a distinctive narrow bilobed dorsal angle as indicated by the 
shape of its overlap area on the anterior dorsolateral plate. A provisional reconstruction suggests that the 
bones came from a large fish over 2 m in length. A cross section of the trunk armour is compared with 
the large arthrodire Taemasosteus from the Early Devonian of Burrinjuck, New South Wales. A possible 
close relative of this new taxon has been illustrated from the Early Devonian of Morocco, supporting 
other indications of resemblance in placoderm fish assemblages between these regions of east and north 
Gondwana during the Early-Middle Devonian. 


Young, Gavin C., 2005. A new Middle Devonian arthrodire (placoderm fish) from the Broken River area, 
Queensland. Records of the Australian Museum 57(2): 211-220. 


Over 20 years ago Professor J.S. Jell (University of 
Queensland) made a small collection of limestone samples 
apparently containing bones from several localities in the 
Devonian outcrops of the Broken River area of Queensland 
(Fig. 1). They were sent to Professor K.S.W. Campbell 
(Australian National University), who passed on remains 
of placoderms (Devonian armoured fishes) to the author 
for preparation and study. 

Acetic acid preparation, which completely removes 
bones (calcium phosphate) from limestone (calcium 
carbonate), revealed a small collection of six placoderm 
specimens from the ten original samples. Two samples 
belonged to one specimen (ANU VI028, described below). 
Three other samples contained only vertebrate microfossils 
(small scales and plates), which have been well documented 
from many horizons in the Broken River Devonian sequence 
by other authors (e.g., De Pomeroy, 1995, 1996; Turner et 


al., 2000). The six placoderm specimens belong to two 
orders, the Antiarchi and the Arthrodira, which were the 
most successful of some seven orders within the class 
Placodermi (Denison, 1978; Young, 1986; Goujet& Young, 
1995). The two antiarch specimens from Broken River were 
the best preserved, with articulated armours and some small 
bones of the jaws, cheek, and sclerotic ring surrounding 
the eye, not previously well documented for the group. They 
were described as two new genera (Nawagiaspis and 
Wurungulepis ) by Young (1990). 

The other four specimens can be assigned to the Arthrodira, 
by far the most diverse placoderm order, with its major 
subgroup, the Brachythoraci, comprising nearly 60% of about 
170 genera within the Arthrodira (Carr, 1995). These specimens 
are much more fragmentary, probably because they belonged 
to considerably larger fishes, all representatives of the 


www.amonline.net.au/pdf/publications/1443_complete.pdf 


212 Records of the Australian Museum (2005) Vol. 57 


Fig. 1. (A) Location of the Broken River area in Queensland, Australia. ( B ) Geological map of the collecting area 
(modified from Turner et al., 2000: fig. 2), showing the locality (Grid Reference 640 460) for the specimens 
described in this paper. 


brachythoracid arthrodires. In marine environments of the Late 
Devonian the brachythoracids included probably the largest 
predators of their time (Young, 2003c). 

The major radiation of brachythoracid subgroups had 
apparently already occurred by the Middle Devonian, and 
the diverse faunas that occupied shallow marine environ¬ 
ments of eastern Australia during the Early-Middle Devonian 
have revealed primitive representatives that are important in 
resolving the origins and interrelationships of major 
brachythoracid subgroups (e.g., Lelievre, 1995; Mark-Kurik 
& Young, 2003; Young et al., 2001; Young, 2004a). 

Two other brachythoracid specimens from the Broken River 
Devonian sequence have recently been described: a new species 
of the genus Atlantidosteus Lelievre, 1984a, a large 
brachythoracid first recorded from the Early Devonian of 
Morocco (Young, 2003a), and a new genus and species 
Doseyosteus talenti Young, 2004b. The two specimens 
documented in this paper are the last to be described from the 
original J.S. Jell collection. An isolated suborbital plate of 
another arthrodire was illustrated by Turner et al. (2000: fig. 
8.7), and an isolated toothplate was ascribed to IPtyctodus sp. 
(placoderm order Ptyctodontida) by Turner & Cook (1997). 
Other vertebrate macro-remains documented from the Broken 
River Devonian sequence include a lungfish skull (S. Turner, 
pers. comm.) and jaw remains of an onychodontid (Turner et 
al., 2000: fig. 6). Apart from these osteichthyan (bony fish) 
remains, placoderm bones, mainly from brachythoracid 
arthrodires, seem to be most common amongst unprepared 
vertebrate samples from Broken River (S. Turner, pers. comm.). 

Locality and age of the samples 

The stratigraphic occurrence of various placoderm remains 
in the Broken River sequence was documented by Young 
(1993, 1996), De Pomeroy (1995, 1996), and Turner et al. 
(2000). Dating the sedimentary sequence is based mainly 


on conodonts (Mawson & Talent, 1989; Sloan etal., 1995). 
Locality and age information for all previously described 
placoderm taxa from the original J.S. Jell collection was 
reviewed by Young (2004b), to clarify conflicting 
information previously published. 

The specimens described below were sent to Canberra 
as three samples, two with the field number 58/Ll (ANU 
V1028), and one labelled 58/L2 (ANU V1031). According 
to information provided by J.S. Jell (letter of 17 April, 1980), 
they both came from University of Queensland locality 
L4399, north bank of the Broken River, Grid Reference 640 
460 on the Burges 1:100 000 sheet (see Lig. 1). He suggested 
a Middle Devonian (?Eifelian) age for this locality. On the 
map of the area published by Sloan et al. (1995: fig. 2) the 
locality lies within the outcrop referred to as “undiffer¬ 
entiated Broken River Group”. 

The antiarch Wurungulepis denisoni Young, 1990 came 
from the same locality (erroneously given as L4339 by 
Young, 1990: 45). On this evidence, a “Wurungulepis - 
Atlantidosteus fauna”, of assumed Eifelian age, was listed 
in the macrovertebrate zonation of Young (1993, 1996). 
However De Pomeroy (1995: 480) assigned Wurungulepis an 
older age—the late Emsian serotinus Conodont Zone—citing 
a personal communication of J.A. Talent. This information 
was repeated by Turner et al. (2000: 498). 

No conodont data were obtained during acid preparation 
of the specimens described below, so their precise position 
relative to the standard conodont zonation is uncertain. 
Young (2004b) concluded that the original age assessment 
of Eifelian for the antiarch Wurungulepis, from the same 
locality, was most consistent with current knowledge of the 
stratigraphic distributions of the Antiarchi, a large and 
diverse group comprising some 45 named genera and 154 
species. Various advanced features of the brachythoracid 
arthrodire described below are consistent with this 
interpretation. 


Young: A new Middle Devonian Arthrodire 213 


Abbreviations 

Repositories for material are indicated by specimen prefixes 
as follows: ANU V, Department of Earth and Marine 
Sciences, Australian National University, Canberra; MCD, 
Moroccan specimens in Museum national d’Histoire 
naturelle, Paris. 

Standard abbreviations for placoderm dermal bones are 
used in the text and figures, with bone overlap areas 
designated as “contact face” ( cf) on the inner surface, or 
“overlap area” ( oa ) on the outer surface. These and other 
anatomical abbreviations used in the figures are as follows: 
ADL, anterior dorsolateral plate; AL, anterior lateral plate; 
cd, glenoid condyle of dermal neck-joint; cf.ADL, area 
overlapping ADL plate; cf.PDL, area overlapping PDL plate; 
cf.PL, area overlapping PL plate; dla, dorsolateral axis 
separating dorsal and lateral walls of the trunk armour; 
fo.PL, fossa on PDL plate receiving dorsal corner of PL 
plate; gr.AL, anterior groove for AL plate; IL, interolateral 
plate; laf articular fossa for dermal neck-joint; lie, main 
lateral line sensory canal; MD, median dorsal plate; oa.ADL, 
area overlapped by ADL plate; oa.AL, area overlapped by 
AL plate; oa.MD, area overlapped by MD plate; oa.PDL, 
area inserting into fossa of PDL plate; o.PDL, area of PDL 
plate overlapped by ADL; o.PL, area of PL plate overlapped 
by ADL plate; pap, para-articular process; PDL, posterior 
dorsolateral plate; PL, posterior lateral plate; pr.sg, 
subglenoid process; ri.cd, ridge crossing inner surface of 
articular condyle; ri.p, posterior ridge; ri.vt, vertical ridge; 
SP, spinal plate; th, thickening; vg, vascular grooves; vs.cd, 
ventral surface of articular condyle. 

Systematic palaeontology 

Class Placodermi McCoy, 1848 
Order Arthrodira Woodward, 1891 
Suborder Brachythoraci Gross, 1932 

Confractamnis johnjelli n.gen. and n.sp. 

1992 eubrachythoracid nov-Young et al., 1993: 247 (pars). 

Name. Lrom the Latin confractus (broken) and amnis 
(stream or river), with reference to the Broken River. The 
species name recognizes the collector, Prof. John Jell, 
University of Queensland, who has conducted research in 
the Devonian of the Broken River area over many decades. 

Diagnosis. A large brachythoracid attaining a length of at 
least 2 m; trunk armour high, with anterior lateral, posterior 
dorsolateral, and posterior lateral plates all dorsoventrally 
elongated. Median dorsal plate enclosing a midline angle 
of about 150° between left and right laminae. Anterior 
dorsolateral plate crossed by single sensory canal groove 
close and subparallel to the lateral margin of the median 
dorsal plate. Dorsal corner of anterior lateral plate narrow, 
rounded, and bilobed, with 25-30° angle between main 
margins. Posterior dorsolateral plate extensively overlapped 
by anterior dorsolateral plate; posterior lateral plate with 
high and narrow exposed part, and elongate anterior overlap 
for the anterior dorsolateral plate. External surface of dermal 
bones smooth, or with very fine, closely spaced tubercul- 
ation of low relief. 


Remarks. Because the skull roof is unknown in this new 
taxon, only trunk armour characters are available for 
assessing its affinities. It is clearly not a coccosteid, in which 
the ADL plate is crossed by an additional ventral sensory 
canal groove. The absence of the ventral sensory groove is 
one of four characters by which Carr (1991: 382) 
characterized the most derived pachyosteomorph subgroup 
within the Brachythoraci (comprising the Dinichthyidae and 
Aspinothoracidi). Within the former family, Eastmanosteus 
differs from the new taxon in possessing tubercular 
ornament, and both Eastmanosteus and Dunkleosteus have 
an extensive exposed part of the ADL plate above the 
sensory groove. Levisosteus Otto, 1999 is a poorly known 
brachythoracid showing possible affinity with Dunkleosteus, 
and indicating that ornament reduction may have occurred 
by the Eifelian, but its trunk armour is completely unknown, 
so no other comparisons are possible with the specimens 
described here. The aspinothoracids are characterized by 
various skull features in addition to the absence of a spinal 
plate in the trunk armour (Carr, 1991). There is no 
information on whether this small bone was present in 
Confractamnis n.gen., but it is likely that the spinal was 
lost independently in several groups, as suggested by 
Denison (1984). One group in which this had occurred by 
the Middle Devonian is the Heterostiidae. Heterostius 
resembles Confractamnis n.gen. in large size, reduction of 
ornament, and position of the lateral line groove near the 
dorsal exposed edge of the ADL plate. Heterostius, however, 
differs in extreme trunk armour reduction, with the AL plate 
fused to the ADL plate, whereas the AL was clearly a 
separate bone in Confractamnis n.gen. The Emsian forms 
Tityosteus, Taemasosteus and Antineosteus also resemble 
Confractamnis n.gen. in the dorsal position of the sensory 
groove on the ADL plate, but differ in the dorsal 
configuration of the AL plate (as indicated by the shape of 
its overlap on the ADL), in the shape of the PDL and PL 
plates, in the much flatter MD plate ( Tityosteus and 
Antineosteus ), and in the transversely elongate articular condyle 
attached along its length to the ADL plate ( Antineosteus ; a 
character defining the Homostiidae). Close affinity with 
Atlantidosteus (known from two species, one in Morocco 
and one in the Broken River sequence) can be excluded on 
the assumption that this taxon was a homostiid, with a 
dorsoventrally compressed trunk armour. One isolated ADL 
plate from the Emsian of Morocco resembles Confractamnis 
n.gen. in its pointed articular condyle, the narrow rounded 
dorsal angle of the overlap area for the AL plate, and the 
position of the sensory groove close and subparallel to the 
lateral margin of the MD. This specimen may represent a 
closely related taxon, but it differs from Confractamnis 
n.gen. in its smaller size, coarse tubercular ornament, more 
acute angle between the dorsal lamina of the ADL plate 
and the long (transverse) axis of the articular condyle, and 
the lack of a bilobed dorsal corner on the AL plate. 

Material. ANU V1028 (holotype), a large left ADL plate 
with part of the MD plate attached, associated with left PDL 
and PL plates (all incomplete); ANU V1031, a very 
incomplete MD plate. 

Locality. University of Queensland locality L4399, north 
bank of the Broken River, Grid Reference 640 460 on the 
Burges 1:100 000 sheet (see Lig. 1); field numbers 58/Ll 
(V1028) and 58/L2 (V1031). 


214 


Records of the Australian Museum (2005) Vol. 57 


Fig. 2. Confractamnis johnjelli n.gen. and n.sp. ANU V1028 (holotype). Bones of the trunk armour in external view (all incomplete). 
(A) Left ADL plate, with part of the MD plate and a fragment of the AL plate attached; ( B ) left PDL plate, with part of the PL plate 
attached; (C) left PL plate. All specimens acid-prepared, and whitened with ammonium chloride for photography. 


Horizon and age. According to information provided by 
Prof. J.A. Talent to A. Basden (pers. comm., 28 August 
1995), the holotype of Wurungulepis, from the same locality 
as the material described here, came from strata that were 
pre-Dosey Limestone in the sequence, and equivalent to 
the Bracteata Formation and Lomandra Limestone. Outcrop 
in this vicinity is referred to as “undifferentiated Broken 
River Group” on the most recent published geological map 
(Sloan et al., 1995: fig. 2). Prof. J.S. Jell (letter of 17 April, 
1980) suggested a Middle Devonian (?Eifelian) age for this 
locality, and the sequence is shown spanning the Emsian- 
Eifelian boundary by Sloan et al. (1995: fig. 3). 

Description. The holotype (ANU V1028) includes remains 
of four bones from the trunk armour of a very large 
brachythoracid. The left anterior dorsolateral (ADL) plate 
is particularly massive (Figs. 2A, 3), with the bone some 
35 mm thick at the base of the articular condyle (cd) for the 
dermal neck-joint. The ventral part of the ADL is missing, 


and the posterior margin is broken and incomplete, but the 
fragmented posterior parts reduce in thickness to a little 
more than 1 mm, with part of the actual margin preserved. 
Together with the evidence of adjacent bones it is possible 
to determine the general shape of the ADL plate, except for 
the extent of the ventral margin. 

The external surface of the ADL (Fig. 2A) shows two 
deeply incised overlap areas for the median dorsal (MD) 
and anterior lateral (AL) plates (oa.MD, oa.AL). A broken 
fragment of the left anterolateral corner of the MD plate is 
still attached to the specimen, but slightly displaced. Its 
thickness (13 mm), so close to the lateral margin, gives some 
indication of the large size of this bone (cf. ANU V1031 
below). The rest of this normally massive bone was either 
weathered away, or perhaps was contained in another 
uncollected limestone block. The external surface of the 
MD fragment appears smooth, as does the exposed surface 
of the ADL. On close examination, some parts show a faint 
tuberculation of closely spaced fine tubercles (about 15 per 


Young: A new Middle Devonian Arthrodire 215 


ri.p 


A 

cf.PDL 


pr.sg 


cf.PL 


th pr.sg 


Fig. 3. Confractamnis johnjelli n.gen. and n.sp. 
ANU V1028 (holotype). Incomplete left ADL 
plate of the trunk armour in internal (A) and 
anterior ( B ) views. Specimen whitened with 
ammonium chloride for photography. 


20 mm 


cm near the sensory groove, decreasing in size to 25 per cm 
on the most ventral preserved surface). 

The groove for the main lateral line sensory canal (11c) 
crosses the ADL plate from the region of the articular 
condyle, as in all arthrodires with a developed dermal neck- 
joint. It runs, however, close and subparallel to the edge of 
the overlap for the MD, which is a point of difference to 
many other brachythoracids. The overlap area for the AL 
plate is set in about 10 mm along a deep anterior groove, 
which braced the AL against the anterior edge of the ADL 
plate (gr.AL, Fig. 2A). The overlap shows that the AL had a 
narrow rounded bilobed dorsal corner, which forms a much 
more acute angle (about 25-30°), and is quite different in 
shape to the triangular overlap of Eastmanosteus or 
Dunkleosteus (dorsal angle 45-50°). Near the ventral 
preserved edge the overlap area expands as a posterior 
embayment, in which a fragment of the overlying AL plate 
is preserved (AL, Fig. 2A). At its margin, this embayment 
slopes gradually to the external surface, in contrast to the 
thick and undercut margins of the deeply incised main part 
of the overlap area (oa.AL). These show that immediately 
dorsal to the embayment the posterior edge of the AL was 
enclosed by the ADL to a depth of about 6 mm. The undercut 
surface is exposed by the broken margin of the overlap area, 
to reveal a strong ridge (ri.p, Fig. 2A). 


ri.cd 


MD 


The articular condyle for the dermal neck-joint in 
Confractamnis n.gen. is a very strongly developed 
projection from the anterior margin of the ADL (cd, Figs. 
2, 3). In mesial view, the condyle shows a triangular cross- 
section. The ventral side of the triangle, with typical 
“siebknochen” texture of spongy bone (invested with 
articular cartilage in life), is partly visible in internal view 
(vs.cd, Fig. 3A). The posterior side of the triangle forms 
the smooth inner surface, which is slightly convex about a 
low ridge crossing the condyle from near the mesial 
termination to the thickened part of the ADL (ri.cd). An 
anterior view of the ADL plate (Fig. 3B) shows the strong 
dorsal support for the condyle (the “condylus ridge” of 
Heintz, 1934: 73), and the mesial termination of the condyle 
as a rounded point, with its anterodorsal surface (anterior 
side of triangle) being flat to slightly concave, and also with 
“siebknochen” texture (sieve-like bone). The actual articular 
surface is higher laterally than mesially (see Fig. 5C), a 
condition also noted in both Dunkleosteus and Homostius 
by Heintz (1934: 73). Lelievre (1995) used this as a character 
to group some primitive brachythoracids from Morocco. 
Beneath the condyle is a prominent subglenoid process 


216 Records of the Australian Museum (2005) Vol. 57 


(pr.sg), which would have articulated against the para¬ 
articular process of the skull roof. 

The inner surface of the ADL plate shows extensive 
posterior contact faces for the PDL and PL plates (cf.PDL, 
cf.PL, Fig. 3A). The distinct anterior margin of the very 
extensive PDL contact face is marked by vascular grooves 
(vg) with radiating orientation from the ossification centre 
of the bone (situated anteriorly in the region of the articular 
condyle attachment). Ventrally this margin becomes the 
anterior margin for the PL contact face. A fainter ridge near 
the posterior border of the ADL plate (ri.p) represents the 
anterior margin for the more dorsal part of the PL contact 
face. The anterior half of the inner ADL surface is convex, 
where the bone is massively thickened (th) to form a broad 
dorsoventral ridge, decreasing ventrally where the internal 
convexity lies beneath the deeply incised overlap for the 
AL plate on the external surface. The posterior part of the 
inner surface in ANU V1028 is flat to concave, and inflected 
dorsoventrally at an angle of about 140° about the 
dorsolateral ridge, at the level of the sensory groove on the 
external surface. The PDL plate has a similar inflection 
(“dla” arrow, Fig. 2B). 

Associated left PDL and PL plates of Confractamnis 
n.gen. are also partly preserved in the holotype. They can 
be placed against the ADL plate to co nfir m their life position. 
The PDL (Fig. 2B) is missing much of its dorsal part, which 
was overlapped by the MD plate. Only a narrow posterior 
strip of the external bone surface is preserved, and most of 
the bone comprises an internal lamina overlapped 
extensively by both ADL and PL plates. A thin vertical ridge 
(ri.vt), which was entirely internal, separates the two 
overlaps (oa.ADL, oa.PL). The edges of the bone are 
extremely thin, with many fine fractures which collapsed 
during acid preparation, and could be only partly 
reconstructed from fragments. The margins of the bone, 
however, are indicated by the extent of the contact face 
inside the ADL plate (cf.PDL, Fig. 3A), which shows that 
the degree of overlap was more extensive than in many other 
brachythoracids. The external surface of the PDL plate may 
have expanded dorsally (PDL, Fig. 4A), and presumably 
was crossed by a posterior continuation of the lateral line 
groove (11c). Its distinctive shape can be inferred from the 
marked dorsoventral elongation of the internal ridge (ri.vt, 
Fig. 2B), and the extensive overlap surfaces (o.PDL, Fig. 4A). 
The narrow dorsal comer of the PL plate is still attached to the 
PDL (PL, Fig. 2B). There is no sign of the internal fossa which 
received this comer in Dunkleosteus (fo.PL, Fig. 6B). 

The inner surface of the PDL plate is smooth and 
concave, inflected about the dorsolateral angle at a level 
that corresponds to that on the ADL (dla, Fig. 2B). The 
bone margins show inner vascular grooves like those on 
the inside of the ADL (vg, Fig. 3A), which may indicate 
increased blood supply adjacent to the bone margins during 
periods of growth. The PDL fits inside the ADL such that 
the vertical ridge (ri.vt, Fig. 2B) abuts against a slight ridge 
inside the posterior margin of the ADL plate (ri.p, Fig. 3 A). 

The incomplete PL plate of Confractamnis n.gen. (Fig. 
2C) is a remarkably high and narrow splint of bone, 
comprising an anterior overlap area, and high and narrow 
posterior exposed part. The posterior margin of the preserved 
part is partly broken, but the form of the inner surface shows 
that not much is missing. The PL plate slots into the grooved 
overlap behind the vertical ridge on the PDL (where part of 


/ 


Fig. 4. (A) Confractamnis johnjelli n.gen. and n.sp. Reconstmction 
of left lateral side of the trunk armour, based on ANU VI02 8 
(holotype). ( B-D ) Isolated left ADL plate from the Early Devonian 
of Morocco, in external ( B ), anterior (C), and internal (D) views. 
Specimen MCD 62, figured by Lelievre (1984b: pi. 6F-H). 

the PL is still attached; oa.PL, Fig. 2B). With both bones in 
place inside the ADL, the convex internal surface of the PL 
forms a thickened posterior border to the articulated armour. 
Its overlap area is sandwiched between the PDL and ADL 
plates, with its anterior edge against the posterior ridge 
dorsally (ri.p, Fig. 3A), and ventrally extending past the 
PDL to fit into a more deeply incised contact face (cf.PL). 
This arrangement is summarized in Fig. 4A. The ventral 
part of the PL plate is unknown. 

One small fragment of the AL plate is also preserved 
attached to the external surface of the ADL in ANU V1028. 
Otherwise this bone is unknown, except for the distinctive 
shape of its dorsal part (AL, Fig. 4A), as indicated by the 
overlap area on the ADL. 

The only other arthrodire specimen from locality UQL 
4399 also belonged to a brachythoracid, and is provisionally 
referred to Confractamnis n.gen. ANU V1031 is an 
incomplete MD plate that lacks most diagnostic characters. 
However the external surface is largely smooth, with similar 
texture to the small portion of MD attached to the ADL 
plate in ANU V1028, which is consistent with it belonging 
to a smaller individual of the same taxon. The more complete 


Young: A new Middle Devonian Arthrodire 217 


Fig. 5. Brachythoracid trunk armour restorations. (A, B) 
Taemasosteus (Early Devonian, Burrinjuck, NSW). (A) Posterior 
view of skull (from White, 1978: fig. 79); ( B ) anterior view of 
trunk armour, restored from individual bones, using Harrytoombs¬ 
ia as a model, as illustrated by Miles & Dennis (1979: fig. 9). (C) 
Confractamnis johnjelli n.gen. and n.sp.; trunk armour restoration, 
anterior view, based on Taemasosteus and Harrytoombsia. 

left lamina is gently curved, 95 mm from the midline to the 
preserved lateral edge, and 50 mm across between broken 
anterior and posterior margins. As all margins are 
incomplete, and no contact faces for the ADL and PDL 
plates can be discerned on the internal surface, there is 
insufficient information for a reasonable estimate of total 
length and overall shape. The bone may have been slightly 
broader than long, consistent with the shape of the MD 
overlap area on the ADL plate of the holotype (MD, Fig. 
4A). This suggests a somewhat angular shape, and a straight 
to gently curved lateral margin. 

The MD plate of ANU VI031 evidently came from an 
individual only about half the size of the holotype, but the 
bone reaches about 10 mm thickness close to the midline. 
The incomplete carinal process is 70 mm long and 50 mm 
deep as preserved. Its anteroventral edge thins to about 1 mm, 
and probably not much is missing. The posteroventral edge is 
expanded to a thickness of about 8 mm, but is incomplete in 
lacking the knob-like or grooved termination that is normally 
developed on the carinal process. The angle between the 
carinal process and the left lamina is about 75°. 


Reconstruction 

Arriving at a reliable three-dimensional reconstruction of the 
trunk armour of a brachythoracid from isolated bones is 
difficult. It took over 120 years from the first attempts (Miller, 
1841) for a reliable reconstruction of the armour of Coccosteus 
(Miles & Westoll, 1968: fig. 44), this reconstmction depending 
on earlier attempts for other forms (e.g., Dunkleosteus Heintz, 
1932: fig. 68; Homostius Heintz, 1934: fig. 45). 

With the discovery of exceptional three-dimensional 
preservation in the acid-prepared arthrodires from Gogo, 
Western Australia, it was possible to check the reliability of 
such restorations with actual specimens. Trial and error in 
the Natural History Museum, London, showed that if bones 
were glued together with a tight fit on the clearly defined 
overlap areas, the left and right sides did not join up—only 
by leaving a small space around the edges of bone overlaps 
could a symmetric reconstruction be achieved (Dr R.S. 
Miles, pers. comm.). The first illustration of an actual 
reconstructed specimen from the uniquely preserved Gogo 
fauna was the holotype of Harrytoombsia by Miles & 
Dennis (1979: fig. 9). 

The ADL plate on its own is one of the most informative 
in attempting a reconstruction from isolated bones, because 
the axis of articulation on the condyle must have been 
horizontal for the dermal neck-joint to function in the living 
animal. This provides some constraint on the cross-sectional 
shape of the trunk armour, a point first exploited by Heintz 
(1934: fig. 47) in comparing Dunkleosteus with the 
dorsoventrally compressed armour of Homostius. There is, 
however, always a degree of uncertainty regarding the 
precise orientation of the axis of articulation, because the 
condyle in life was invested with articular cartilage, so its 
true shape is not preserved. Thus, it is possible that the ADL 
may have been slightly more steeply inclined (implying a 
higher trunk armour) than depicted in the reconstruction (Fig. 
5C), although the preserved angles of the MD plate in ANU 
VI031 are generally consistent with this reconstruction. 

Using the trunk armour in the holotype of Harrytoombsia 
as a model (Miles & Dennis, 1979: fig. 9), the armour of 
Confractamnis n.gen. may have been some 37 cm in height, 
and over 30 cm across. As there is no control of the width 
across the articular condyles of the ADL, the armour could 
have been considerably larger by comparison with 
Harrytoombsia (which has proportionately smaller 
condyles), but it is unlikely to have been any smaller. Such 
a fish would have been at least 206 cm long, judging by 
proportions in Coccosteus cuspidatus, where whole animals 
including the tail are preserved, and total length of the fish 
is some 5.5 times trunk armour height (see Miles & Westoll, 
1968: figs. 44, 48). 

Discussion 

The trunk armour bones of Confractamnis n.gen. just 
described reveal many characters by which they can be 
distinguished from other arthrodires. On the ADL plate, the 
groove for the main lateral line sensory canal (lie, Fig. 4A) 
crosses the bone close and subparallel to the edge of the 
overlap for the MD, which is a point of difference to many 
other brachythoracids. In Coccosteus and Harrytoombsia 
(Miles & Westoll, 1968: fig. 30; Miles & Dennis,' 1979: fig. 
4) there are two branches, the main one passing postero- 
ventrally across the ADL plate. In Dunkleosteus and 


218 Records of the Australian Museum (2005) Vol. 57 


Fig. 6. Left trunk armour bones of the Late Devonian brachythoracid Dunkleosteus, arranged to show overlap relations (not to scale). 
(. A,D ) ADL plate in lateral and internal views (reversed images from Heintz, 1932: figs. 46-n>47); ( B ) PDL plate, and (C) PL plate, 
both in external view (from Heintz, 1932: figs. 49, 54). 


Eastmanosteus (Heintz, 1932: fig. 46; Dennis-Bryan, 1987: 
fig. 22A), there is some distance between the groove and 
the MD overlap, with similar exposed areas of the ADL 
plate both above and below the groove (Fig. 6A). In 
Holonema, the arrangement is different again, with the 
straight sensory groove crossing the ADL close to and 
subparallel with the ventral overlap for the AL plate (Miles, 
1971: figs. 62, 63). 

The forms most similar to Confractamnis n.gen. in the 
position of the sensory groove are Taemasosteus and 
Tityosteus from the Early Devonian (White, 1978: fig. 102; 
Gross, 1960: fig. IB), in which it passes straight back from 
the level of the condyle, running subparallel to the edge of 
the MD overlap. However the ADL plate in both of these 
taxa has a triangular ventral overlap area for the AL plate, 
of similar shape to that of Dunkleosteus (Lig. 6A). The large 
Middle Devonian brachythoracid Heterostius also has a 
dorsally placed sensory groove on the ADL (Denison, 1978: 
fig. 48B), but in this case the AL plate is reduced to a massive 
splint of bone fused to the ADL, a unique specialization of 
this family. The prominently projecting articular condyle 
in ANU V1028 shows that Confractamnis n.gen. could not 
have had a dorsoventrally compressed armour, thus 
excluding a close relationship with Atlantidosteus, another 
large arthrodire documented from both Morocco and the 
Broken River Sequence, and referred to the Homostiidae 
(Lelievre, 1984a; Young, 2003a). 

Two examples of the ADL plate from the late Emsian of 
Morocco resemble Confractamnis n.gen. in the unusual 
shape of the AL overlap area. In the holotype of 
Antineosteus, the ADL has a sensory groove that is more 
ventrally placed with respect to the MD overlap (Lelievre, 
1984b: fig. 16A). The articular condyle in this form is 
developed completely differently, with an elongate 
attachment to the front margin of the bone indicating a 
depressed body form typical of the family Homostiidae. 
Another left ADL (MCD 62), from the same locality (near 
Akka), was figured without further comment as “Homostiid 
sp.” by Lelievre (1984b, pi. 6L-H). It is closely similar to 
ANU V1028 in the highly angular overlap for the AL, and 


the orientation of the sensory groove, which has a straight 
course, running close to and subparallel with the ventral 
edge of the overlap for the MD plate (Lig. 4B). This taxon 
is clearly not conspecific with Confractamnis n.gen., since 
the specimen has much coarser tuberculate ornament, even 
though it is only about half the size of the ADL plate in the 
holotype. It also probably differs in having the exposed 
ornamented part extending ventrally past the edge of the 
AL overlap (but the corresponding margin is broken in ANU 
V1028). The articular condyle in the Moroccan specimen 
(cd, Lig. 4B-D) has a similar pointed mesial end in anterior 
view to Confractamnis n.gen., but is more elongate. 

The thickened dorsal attachment of the condyle to the main 
body of the ADL plate in Confractamnis is quite different to 
that of Taemasosteus (Lig. 5), and a range of other 
brachythoracids where this has been illustrated (e.g., Heintz, 
1932: fig. 68, 1934: fig. 45; Lelievre etai, 1981: figs. 7, 15). 
The thickenings on the inner surface of the ADL in 
Confractamnis (Lig. 3A) are similarly developed to a much 
smaller ADL plate belonging to the homostiid Cavanosteus 
from Burrinjuck (Young, 2004c: fig. 6), but it is not clear if 
this is a general or primitive brachythoracid feature. 

The PDL plate of Confractamnis n.gen., although 
incompletely preserved, also demonstrates a distinctive 
morphology compared to other brachythoracids. Its margins, 
as indicated by the extent of the contact face inside the ADL, 
show that the degree of overlap was much more extensive 
than in forms like Taemasosteus, Antineosteus or Heintz- 
ichthys (White, 1978: fig. 105; Lelievre, 1984b: fig. 16B; 
Carr, 1991: fig. 6B). Heintzichthys, Coccosteus and 
Eastmanosteus have a typical brachythoracid PDL plate of 
approximately equilateral triangular shape, with three 
overlap surfaces, essentially as in Dunkleosteus (Lig. 6B). 
An internal fossa for the dorsal corner of the PL plate (fo.PL) 
was interpreted by Carr (1991:383) as one of five characters 
uniting Eastmanosteus and Dunkleosteus in the family 
Dinichthyidae, but this is absent in all Emsian-Eifelian 
brachythoracid taxa known so far. 

Only the dorsal part of the PL plate is preserved in 
Confractamnis n.gen., but again it indicates a distinctive 


Young: A new Middle Devonian Arthrodire 219 


shape. In most other brachythoracids this bone has more 
equilateral proportions, but with similar overlap relations 
to surrounding bones. In Heintzichthys for example (Carr, 
1991: fig. 10), the PL plate overlaps the PDL, and the ADL 
plate overlaps its anterodorsal part, which is sandwiched 
between the ADL on the outside, and the overlap surface of 
the PDL on the inside, essentially as in Dunkleosteus (Fig. 
5C). Brachythoracids typically show a similar connection 
dorsally with the PDL plate, which carries a narrow dorsal 
notch to receive the dorsal angle of the PL, in Eastmanosteus 
and Dunkleosteus developed as a fossa (fo.PL, Fig. 6B). In 
Antineosteus the PL is only known from a small narrow 
dorsal part in the type specimen (H. Lelievre, pers. comm.), 
and from its contact face on the PDL plate, which shows a 
narrow dorsal notch. It is clear that the overlapped area of 
ANU V1028 was much more extensive, indicating that the PL 
plate of Confractamnis n.gen. was much higher and narrower 
than in Antineosteus. In Taemasosteus, as restored by White 
(1978: figs. 103-105), the PL plate overlaps the PDL, but sits 
entirely behind, and is not overlapped by, the ADL plate. It 
therefore has only one ventral overlap area, for the AL plate. 

In some derived brachythoracids the PL plate may be 
much reduced, or completely lost. This was assumed to be 
the case for Homostius by Heintz (1934), until it was 
recorded (with a SP plate, also assumed to be missing in 
this taxon) from a new Estonian locality (Karski). The PL 
of Homostius, as described by Mark-Kurik (1993), 
resembles that of Confractamnis n.gen. in its high and 
narrow shape, but differs in the fact that the external part 
expands rather than narrows dorsally. It also carries a sensory 
groove, even though the normal condition in brachy¬ 
thoracids is for the sensory groove to cross the ADL and 
PDL plates. A shift in the position of the sensory canal may 
relate to the broad, dorsoventrally compressed body form 
of Homostius (Heintz, 1934: fig. 49). 

The MD plate in Confractamnis n.gen. is interpreted, 
mainly from its overlap on the ADL plate of the holotype, 
to have been slightly broader than long, somewhat angular 
in shape, and with a straight to gently curved lateral margin. 
Several other brachythoracids have similar MD plates, 
including Homostius and Antineosteus (Heintz, 1934; 
Lelievre, 1984b). In Dunkleosteus, Taemasosteus and 
Tityosteus the MD plate has a more elongate and rounded 
lateral profile (Heintz, 1932: fig. 44; White, 1978: fig. 94; 
Otto, 1992: fig. 6a). The 75° angle between the carinal 
process and the left lamina is less arched than in 
Taemasosteus (about 50°; White, 1978: fig. 95), but not as 
flat as in Homostius or Tityosteus (e.g., Heintz, 1934; Gross, 
1960). Krasnov & Mark-Kurik (1982) identified an isolated 
MD plate from the Emsian of the Minusinsk area of Russia 
as a new species Tityosteus orientalis. This was also less 
arched than in Confractamnis n.gen., with an angle of about 
60° between the carinal process and one lamina of the MD. 

The attempted reconstruction of the trunk armour of 
Confractamnis n.gen. (Fig. 5C) can be compared with a 
similar reconstruction of the Emsian form Taemasosteus 
from Burrinjuck, where additional information from the 
skull roof (White, 1978: fig. 79) provides a constraint on 
the width between the articular condyles (Fig. 5A-B). Both 
these taxa may have had bodies with an overall fusiform 
shape, in contrast to the dorsoventrally compressed shape 
of Homostius and related taxa. However it is possible that 
relative height of the neck-joint articulation varied, and was 


carried higher on the armour in Coccosteus or Harry- 
toombsia, which may be the reason that the main sensory 
groove across the external surface of the ADL plate has a 
characteristic downward course in these taxa, presumably 
to align with the middle part of the flank as it ran back onto 
the tail of the fish (cf. Miles, 1971: fig. 108). 

Conclusions and summary 

The new taxon Confractamnis n.gen., from assumed Eifelian 
strata in the Broken River sequence of Queensland, shows 
closest affinity amongst known forms to some arthrodire 
material from the late Emsian of Morocco. This suggests 
the same distribution pattern as that indicated for the 
homostiid arthrodire Atlantidosteus, represented by two 
species with a disjunct distribution in the Devonian of 
Morocco and Queensland (Young, 2003a). Faunal 
connections, and continuity of shallow tropical to 
subtropical marine environments along the eastern and 
northern margins of the Gondwana supercontinent, are 
indicated by this pattern. On most current reconstructions 
the northern Gondwana margin had a palaeolatitude between 
0-30°S (e.g., Young, 2003b: fig. 2). The only other likely 
connection (constrained by palaeolatitude) would be across 
the proto-Pacific Ocean, assuming this large ocean existed 
in the early-middle Palaeozoic (cf. Nur & Ben-Avraham, 
1981). An alternative proposal, based on the evidence of 
Devonian fish distributions, is that palaeogeographic change 
during the middle Palaeozoic involved increasing proximity 
between Gondwana and Laurussia (Euramerica), with first 
shallow marine, and then non-marine fish dispersal between 
the two continental blocks during the Middle and Late 
Devonian (Young et al., 2000a,b; Young & Moody, 2002; 
Young, 2003b). 


Acknowledgments. Professors J.S. Jell (Univ. Qld) and K.S.W. 
Campbell (ANU) are thanked for providing the specimen for study. 
Mr R.W. Brown (Geoscience Australia) assisted in acid 
preparation, and Mr A. Haupt did some German translations. 
Professor J.A. Talent and Dr A. Basden (Macquarie University) 
advised and discussed at length the provenance and age of Broken 
River fish material. Comparison with European and Moroccan 
arthrodire material was facilitated by a visiting professorship at 
the Museum national d’Histoire naturelle, Paris, in 1999. Professor 
D. Goujet is thanked for arranging this, and for the provision of 
facilities, and together with Dr H. Lelievre and Dr P. Janvier 
discussed at length placoderm morphology and relationships. Dr 
Lelievre arranged for arthrodire casts to be sent to Canberra for 
comparative study. B. Harrold is thanked for providing essential 
computer support at ANU, and V. Elder is thanked for curation 
and data management of the fossil fish collection. Dr E. Mark- 
Kurik and Dr R. Carr discussed arthrodire phylogeny, and Dr 
Carr arranged for a visit to Cleveland, Ohio, for study of large 
arthrodire material. Financial support was provided in Canberra 
by ANU Faculties Research Fund Grants F01083 and F02059, 
and overseas by the Alexander von Humboldt Foundation, for a 
Humboldt Award in Berlin (2000-2001), and assistance with travel 
to Flagstaff and Cleveland, USA (2000). I thank Prof. H.-P Schultze 
for provision of facilities in the Museum fur Naturkunde, Berlin. 
Prof. P. De Deckker is thanked for provision of facilities in the Earth 
and Marine Sciences Dept., ANU. Drs Herve Lelievre and Bob Carr 
are thanked for helpful reviews of the manuscript. This research was 
a contribution to IGCP Projects 328, 406, 410, and 491. 


220 Records of the Australian Museum (2005) Vol. 57 


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Zoological Journal of the Linnean Society 103: 349-390. 

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Manuscript received 9 June 2003, revised 23 January 2004 and accepted 
30 January 2004. 

Associate Editor: G.D. Edgecombe. 


© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 221-236. ISSN 0067-1975 


Amphipods of the Genera 
Ceradocus, Dulichiella, Melita and Nuuanu 
(Crustacea: Melitidae) 
from Mauritius, Indian Ocean 


Chandani Appadoo 1 and Alan A. Myers 2 * 


1 Department of Biological Sciences, 

Faculty of Science, University of Mauritius, Reduit, Mauritius 

chandani @ uom. ac. mu 

2 Department of Zoology, Ecology and Plant Science, 

National University of Ireland Cork, Lee Maltings, Prospect Row, Cork, Ireland 

alanmyers @ crustacea.net 


Abstract. Taxonomic descriptions and figures are provided for five new species of Melitidae ( Ceradocus 
greeni n.sp., Dulichiella cuvettensis n.sp., Melita corticis n.sp. Melita setimera n.sp. and Nuuanu rectimana 
n.sp.) from collections made in the shallow coastal waters of Mauritius. 


Appadoo, Chandani, & Alan A. Myers, 2005. Amphipods of the genera Ceradocus, Dulichiella, Melita and 
Nuuanu (Crustacea: Melitidae) from Mauritius, Indian Ocean. Records of the Australian Museum 57(2): 221-236. 


In the current study a new species of Ceradocus, C. greeni 
n.sp. is described, bringing the number of species of the 
genus known from Mauritius to three. The two other species, 
C. hawaiensis J.L Barnard (1955) and C. mahafalensis var. 
incisa Ledoyer (1978), were reported by Ledoyer (1978). 
A new species of Dulichiella, D. cuvettensis n.sp. is 
recognized, previously wrongly ascribed to D. appendic- 
ulata (Say, 1818) by Ledoyer (1978) as well as by Appadoo 
& Steele (1998) and Nuuanu rectimana n.sp. is described 
bringing the number of species of this genus known from 
the island to two, the other being Nuuanu amikai J.L. 
Barnard, reported by Ledoyer (1978). Only one species of 
the genus Melita was previously recorded from Mauritius, 
Melita zeylanica (Appadoo & Steele, 1998), here attributed 
to a new species, Melita corticis n.sp. A second new species 
Melita setimera n.sp. is now known. 

* author for correspondence 


Material and methods 

Amphipods were collected from algae, seagrass and coral 
rubble from 24 sites around the island of Mauritius (19°59- 
20°32'S 57°18'-57 0 47'E, Indian Ocean) and from lie 
D’Ambre (20 o 01'-20 o 02.2’S 57°4r-57°42.2’E), a small 
island on the northeast coast within the lagoon from 
February 1998 to February 2000. The sites were visited at 
low tide and samples were collected from the intertidal and 
shallow subtidal zones. Algae and rubble were collected by 
scraping them off their substrates using a small hand trowel. 
Amphipods were extracted using the formalin-wash method 
as formalin is an irritant that causes the animals to release 
hold of the substrates (Barnard, 1976). 

Some of the substrates were also collected by snorkelling 
and diving from depths not exceeding 2 to 3 m. The 

www.amonline.net.au/pdf/publications/1444_complete.pdf 


222 Records of the Australian Museum (2005) Vol. 57 


substrates were then transferred to a plastic bag as soon as 
they were scraped off and amphipods were extracted using 
the above mentioned method once on shore. 

Prior to dissection the body length of amphipods was 
recorded by holding it straight and measuring the distance 
along the dorsal side of the body from the base of the first 
antennae to the base of the telson. A stereomicroscope with 
a micrometer scaled eyepiece was used to take the 
measurement. Drawings were made using a Nikon 
compound microscope equipped drawing tube attachment. 
Type material is deposited in the Australian Museum (AM). 
All other material is kept in the first author’s collection. 
The terminology for cuticular extensions and setae follows 
that of Watling (1989). Geo-spatial coordinates were 
read from a map of scale 1:25 000. 

Abbreviations used in figures. A, Antenna (1-2); 

C, Coxa; D, Dactylus (3-7); Ep, Epimeron; G, 
Gnathopod (1-2); L, Lower lip; Md, Mandible; 

Mx, Maxilla (1-2); Mxp, Maxilliped; P, 

Pereopod (3-7); PI, Pleonite (1-3); p, palp; T, 

Telson; U, Uropods (1-3); It, left; r, right. 


Fig. 1. Ceradocus greeni n.sp., male, 6.2 mm, 
female, 4.9 mm, AM P60866, Flic-en-Flac. 
Scales: a = 1 mm (whole animal), b = 0.2 mm 
(Ul, U2 and U3), c = 0.2 mm (female G1 and 
G2), d = 0.1 mm (T), e = 0.05 mm (outer margin 
of U2). 


Taxonomic section 

Ceradocus (Denticeradocus) greeni n.sp. 

Figs. 1-3 

Ceradocus sp. 1 Appadoo & Steele 1998: 639. 

Type material. Holotype S, 5.2 mm, AM P60865, at depths of 
0.5-2 m living on coral rubble and Pocockiella variegata, Flic-en-Flac 
(20°16.5'S 57°21.7'E), Mauritius, C. Appadoo, 9 November 1998. 
Paratypes: 2d d, 22 9, AM P60866, same data as holotype; 1 9 from 


I \ 

l \ 


A ^ \ 


Appadoo & Myers: Melitidae and gammarellidae from Mauritius 223 


coral rubble and Pocockiella variegata, Flic-en-Flac, 3 March 1998; 1 
juv. from coral rubble and Pocockiella variegata, Flic-en-Flac, 9 
November 1998; Id, 42 2, 2 juv. from coral rubble, Padina sp. and 
Pocockiella variegata, Flic-en-Flac, 5 April 1999; 7 d d , 5 9 9 from coral 
rubble, Padina and Pocockiella variegata, Flic-en-Flac, 10 December 
1999; 3dd, 32 9, 1 juv. from coral rubble, Padina sp., Pocockiella 
variegata and Turbinaria ornata, Flic-en-Flac, 27 January 2000. 

Description. Male length, 6.2 mm. Head with subocular 
notch; eyes round with discrete ommatidia. Antenna 1 


peduncle article 1 with stout robust setae on posterior 
margin; article 2 longer than 1; article 3, 0.3x article 1; 
accessory flagellum 5-articulate; primary flagellum 14- 
articulate. Antenna 2 peduncle 3x as long as flagellum; gland 
cone of article 2 extending to 0.7x the length of article 3; 
article 4 slightly longer than article 5; flagellum 11- 
articulate. Mandible palp 3-articulate, article 1 with medial 
cusp, article 2 longest and 2.5x article 1, with long setae on 


224 Records of the Australian Museum (2005) Vol. 57 


lateral margins and short setae on medial face, article 3 
shortest and 0.6x the length of article 1. Maxilla 1 inner 
plate triangular with plumose setae; outer plate with terminal 
serrated robust setae; palp 2-articulate with double-row of 
terminal setae. Maxilla 2 outer plate with distal setae, inner 
plate with distal setae, inner marginal setae and an oblique 
row of setae. Lower lip with rounded mandibular lobes. 
Maxilliped palp 4-articulate, article 2 longest and 2.2x 
article 1. Gnathopod 1 coxa 1.3x as long as broad, 
anterodistal margin produced into a lobe, posterodistal 
margin notched; basis slender, slightly less than three times 
as long as broad, anterior margin with short robust setae, 
posterior margin with long setae; carpus with groups of 
medial setae, anterior margin sparsely setiferous, posterior 
margin densely setose; propodus subequal in length to 
carpus with a few medial patches of setae, palmar margin 
oblique with short setae; dactylus with short setae on inner 
margin. Gnathopod 2 asymmetrical. Small gnathopod 2 (left 
or right) coxa subrectangular, slightly longer than broad, 
distal margin with a notch and a few setae; basis 2.5x as 
long as broad, posterior margin with strong patches of setae; 


merus anteroventral and posterodistal corners sharply 
produced; propodus 2x as long as broad, 1.3x carpus, palm 
oblique with short robust setae and a few groups of long 
setae; dactylus slender, inner margin with short setae. Large 
gnathopod 2 (left or right) coxa subrectangular, ventral 
margin notched and with 2 setae; basis 2.5x as long as broad; 
merus posterodistal corner sharply produced; carpus 
triangular, slightly more than 1.5x as broad as long; 
propodus robust, slightly over 1.5-1.75x as long as broad, 
palm oblique, distal margin with a process close to base of 
dactylus with stout robust setae followed by an excavation 
and a triangular process with fine setae and robust setae at 
the posterodistal margin; dactylus robust and fitting into 
the triangular process of propodus. Pereopod 3 coxa 
subrectangular, almost as broad as long, ventral margin with 
1 long seta and a few short-setae at the anteroventral corner; 
basis slender with short robust setae on anterior margin and 
groups of long setae on posterior margin; propodus subequal 
to carpus; dactylus with a distinct unguis and anterior margin 
with one plumose seta and posterior margin with 3 setae. 
Pereopod 4 coxa posteriorly excavate, 0.7x as long as broad 


Appadoo & Myers: Melitidae and gammarellidae from Mauritius 225 


with one long seta and a few short setae on the anterodistal 
corner, other features similar to pereopod 3. Pereopod 5 
slender, coxa bilobed; basis 1.8x times as long as broad, 
anterodistal and posterodistal margins sharply produced, 
anterior margin with robust setae, posterior margin weakly 
serrated and with small setae; propodus subequal to carpus; 
dactylus slender. Pereopod 6 slender; basis similar to that 
of pereopod 5, but posterior margin more deeply serrated; 
propodus subequal to carpus, anterior margin with strong 
patches of setae; dactylus slender and similar to that of 
pereopod 5. Pereopod 7 similar to pereopod 6, but basis is 
distally less produced and narrower than basis of pereopod 
6; propodus slightly longer than carpus. Pleonites 1-3 
strongly toothed. Epimeron 2 with one tooth on posterodistal 
margin and a few irregular teeth on distal margin. Epimeron 
3 with 2 teeth on posterodistal margin and 5 teeth on distal 
margin. Urosomite 1 and 2 each with 7 dorsal teeth. Uropod 
1 peduncle, 1.3x outer ramus with a stout robust seta on 
medial outer margin; outer ramus slightly shorter than inner 
ramus. Uropod 2 peduncle 0.7x inner ramus; inner and outer 
rami subequal, armed with robust setae, margins of rami 
with very short fine robust setae. Uropod 3 peduncle inn er 
margin with numerous robust setae; rami spatulate, equal 
in length to each other, twice as long as peduncle; outer 
ramus outer margin with long robust setae; inner ramus outer 
margin with numerous robust setae; rami with stout terminal 
robust setae. Telson deeply cleft; telsonic lobes well 
separated, with a pair of plumose setae on outer margin, 
notched at apex, with outer tooth produced and inner tooth 
vestigial, apices with one long and a few short setae. 

Female: length, 4.9 mm. Gnathopod 1 coxa 1.2x as long 
as broad, anterodistal margin produced, posterodistal margin 
with a notch; basis slender, posterior margin with long setae, 
anterior margin with short robust setae; merus produced at 
posterodistal corner; carpus subequal to propodus with 
groups of medial setae, ventral margin setose; propodus 2.3x 
as long as broad, anterior margin with 5 groups of setae, 
palm oblique with fine setae and short robust setae and stout 
robust setae on medial face. Gnathopod 2 coxa 1.2x as long 
as broad, distal margin with a notch and a few setae; basis 
3.lx as long as broad, anterior margin with robust setae, 
posterior margin with groups of long setae; carpus 0.75x 
length of propodus; propodus 2.2x as long as broad, palm 
with short robust setae and patches of fine setae, with stout 
robust setae on inner medial face. 

Habitat. In the subtidal at depths of 0.5 to 2 m, occurring 
mostly on coral rubble and the associated brown alga, 
Pocockiella variegata. 

Remarks. Ceradocus greeni n.sp. is assigned to the 
subgenus Denticeradocus because pleonites 1-3 are 
multidentate dorsally. This species is distinguished from 
Ceradocus hawaiensis J.L. Barnard (1955) recorded from 
Mauritius by Ledoyer (1978), by having the larger male 
gnathopod 2 with an oblique palm lacking many tooth-like 
processes. The species differs from Ceradocus mahafalensis 
Ledoyer (1978) var. incisa, reported from Mauritius, which 
also has an oblique palm in the larger male gnathopod 2, by 
the broadly sinuous palmar border, with a distal process 
with robust setae as opposed to a palmar margin with a deep 
medial incision. Urosomites 1 and 2 each have 7 teeth in 
Ceradocus greeni n.sp. instead of 5 and 4 respectively in 
Ceradocus mahafalensis var. incisa and C. mahafalensis 


from Madagascar (Ledoyer, 1979). 

Ceradocus greeni shares with C. spiniferus Ledoyer 
(1973), C. tattersalli Ledoyer (1982) and C. serratus (Bate, 
1862), the multidentate pleonites and oblique palm in the 
large male gnathopod 2. However, the shape of the larger 
gnathopod 2 propodus palm separates it from these three 
species. Ceradocus serratus lacks the smooth excavation 
and the triangular process, C. spiniferus has a convex palmar 
margin and a small U-shaped excavation and C. tattersalli 
lacks the triangular process and has a palm with numerous 
robust setae. 

Two other species of Ceradocus with multidentate 
pleonites 1-3 and oblique palm in the male gnathopod 2 
are Ceradocus (Denticeradocus) oxydus Berents (1983) and 
Ceradocus (Denticeradocus) yandala Berents (1983). The 
shape of the large male gnathopod 2 is the distinguishing 
feature. Ceradocus oxydus lacks an excavation in the palmar 
margin which is convex with numerous robust setae. 
Ceradocus greeni differs from C. yandala by having a gentle 
excavation on the male gnathopod 2 without any mid-palmar 
sinus, C. yandala has a quadrate mid-palmar sinus. 

Type locality. Flic-en-Flac, Mauritius. 

Distribution. Mauritius. 

Etymology. This species is named after Prof. John Green 
of Memorial University of Newfoundland for his help in 
the field to one of authors (CA) during an initial study on 
amphipods from Mauritius in 1995. 

Dulichiella cuvettensis n.sp. 

Fig. 4 

Melitaappendiculata- Ledoyer, 1978:282; Appadoo & Steele, 1998: 

639. (Not Gammarus appendiculatus Say, 1818: 377-379). 

Type material. HOLOTYPE 3, 3.3 mm, AM P67233, from Sargassum 
sp. at depth less than 1 m, La Cuvette (20°00'S 57°34.2'E), Mauritius, 
C. Appadoo, 12 October 1999. Paratypes: 13, 1$, from Sargassum 
sp., La Cuvette (20°00'S 57°34.2'E), 14 May 1998. 1 2 from Turbinaria 
sp., Bain Boeuf (19°59'S 57°36'E), 15 May 1998; 13, 39 2 from 
Acanthophora spicifera, Anse la Raie (19°59.5'S 57°37.5'E), 15 May 
1998; 53 3, 3 2 9 and 4 juv. from Sargassum binderi, Bain Boeuf, 16 
June 1998; 13,22 2 from Sargassum sp. and Padina sp., lie D’Ambre 
(20°02’S 57°40'E), 12 November 1998; 13 from Sargassum sp. and 
Ulva reticulata. La Cuvette, 5 May 1999; 13 from Padina sp. and 
Halimeda sp., Grand Baie (20°0.5’S 57°34’E), 5 May 1999; 13, 12, 
from mixture of Padina sp., Pocockiella variegata and Sargassum sp., 
Bain Boeuf; 23 3 and 12, AM P67234, from Sargassum sp. and 
Pocockiella variegata, Bain Boeuf, 12 October 1999. 

Description. Male length, 4 mm. Head without subocular 
notch; eyes round with well-developed ommatidia. Antenna 
1 poorly setiferous, peduncle article 1 with 3 stout robust 
setae on ventral margin; article 2 longest, 1.5x article 1; 
article 3, 0.3x the length of article 1; accessory flagellum 
4-articulate, primary flagellum 35-articulate. Antenna 2 
weakly setiferous, peduncular article 4 subequal to 5, 
flagellum 14-articulate. Mandible palp slender, article 1 with 
a small tooth; article 3 slightly longer than article 2. Maxilla 
1 palp, article 1 with long setae on distal margin; inner plate 
with 2 apical plumose setae. Gnathopod 1 coxa 1.6x as long 
as broad, posterodistal margin with a notch; basis slender, 
4x as long as broad; propodus slightly 0.7x length of carpus, 
palmar margin with short and long setae; dactylus normal. 
Gnathopods 2 dissimilar (left and right). Larger gnathopod 


226 Records of the Australian Museum (2005) Vol. 57 


Fig. 4. Dulichiella cuvettensis n.sp., male, 4 mm, female, 4.8 mm, La Cuvette. Scales: a = 0.2 mm (Ep and r male 
G2), b = 0.1 mm (male G1 and It male G2), c - 0.2 mm (female G1 and G2), d = 0.\ mm (T), e = 0.05 mm (Mdp). 


2 basis without setae, and 2.3x as long as broad; carpus 
reduced, 3.3x as long as broad; propodus robust, distally 
expanded, palmar margin transverse, with 3 well-developed 
medial protuberances; dactylus broad throughout its length. 
Small gnathopod 2, coxa slightly less than 2x as long as 
broad, distal margin with short setae; basis slender, 3.5x as 


long as broad, with setae on anterior and posterior margins; 
merus posterodistal margin acute; propodus slightly shorter 
than carpus, subrectangular, palm oblique, dactylus fitting 
palm. Pereopod 3 coxa subrectangular with a small notch 
on posterodistal margin; basis anterior margin with stout 
short setae; propodus 1.5x length of carpus; dactylus with 


Appadoo & Myers: Melitidae and gammarellidae from Mauritius 227 


bifid tip. Pereopod 4 similar to pereopod 3, but coxa without 
posterodistal notch and weakly excavate posteriorly. 
Pereopod 5 basis subrectangular, slightly more than 2x as 
long as broad, anterior and posterior margins parallel, 
anterior margin with numerous robust setae, posterior 
margin weakly serrated with short setae; dactylus bifid. 
Pereopod 6 similar to pereopod 5 but basis more slender; 
merus and propodus with strong groups of setae on anterior 
and posterior margins. Pereopod 7 basis anterior margin 
straight, posterior margin slightly convex; other features 
similar to pereopod 6. Pleonites 1 and 2 with 7 teeth, median 
tooth and the last tooth on either side shorter than others. 
Pleonite 3 with 7 teeth, median tooth small, other teeth on 
either side of this median tooth are successively longer than 
one another. The concavities of the teeth of pleonites have 
small setae. Urosomite 1 produced into acute teeth on dorsal 
surface. Urosomite 2 with a small robust seta and small 
tooth on dorsal surface. Urosomite 3 with a small dorsal 
tooth. Epimeron 1 with one robust seta on distal margin, 
posterodistal margin rounded. Epimeron 2 posterodistal 
margin slightly produced, posterior margin smooth, distal 
margin bears 3 robust setae. Epimeron 3 posterodistal 
margin produced into an acute tooth, posterior margin with 
a very small tooth, distal margin with three robust setae. 
Uropod 1 slender, with robust setae, rami subequal to each 
other and slightly longer than peduncle; inner margin of 
inner ramus with very fine short setae. Uropod 2 outer ramus 
slightly shorter than inner ramus; peduncle 0.75x inner 
ramus; inner margin of inner ramus similar to that of uropod 

l. Uropod 3 outer ramus 1.8x peduncle, 2-articulate, article 
1 truncate, article 2 pointed; inner ramus vestigial consisting 
of a small oval lobe with one robust seta. Telson cleft to 
about three-quarter its length, telsonic lobes produced at 
apex. Telson with three groups of robust setae, located 
subapically and medially on inner and outer margins. 

Female: length, 4.8 mm (mature, with eggs). Gnathopod 
1 coxa 0.75x as long as broad, posterodistal margin with 
notch and setae; basis with setae on anterior and posterior 
margins; merus with a triangular process at anterodistal 
margin; propodus palm oblique, palm with long setae. 
Gnathopod 2 coxa subrectangular about as long as broad; 
basis anterior and posterior margins setose; ischium anterior 
margin sinuous; propodus slightly longer than carpus, palm 
oblique with setae on margins. 

Remarks. Dulichiella cuvettensis n.sp. differs from D. 
appendiculata (Say, 1818) in having epimeron 1 with a 
smoothly rounded posteroventral margin (rather than with 
a small acute spine) and the propodus disto-lateral margin 
with three (as apposed to two) subacute teeth. Dulichiella 
cuvettensis n.sp. is most similar to D. australis (Haswell, 
1879) but differs from that species in the strongly setose 
uropod 3 outer ramus as well as in the rounded postero¬ 
ventral corner of epimeron 1. 

Habitat. This species was collected in depths of less than 1 

m. It occurs mostly on brown algae especially Sargassum 
sp. and was collected from sites on the north coast of the 
island. 

Type locality. La Cuvette, Mauritius. 

Distribution. Mauritius. 

Etymology. Named after the type locality. 


Melita corticis n.sp. 

Figs. 5-6 

Melita zeylanica Appadoo & Steele, 1998: 639. 

Type material. Holotype 3,4.3 mm, AM P60867,0-1 m depth, living 
on a mixture of Ulva lactuca and Ulva reticulata, le Bouchon (20°28'S 
57°40.5'E), C. Appadoo, 27 October 1998. Paratypes: 13, 3$ 9, AM 
P60868, same data as holotype; 23 3,169 9, and lOjuv. from Ulva lactuca 
and Ulva reticulata , Le Bouchon, 16 May 1998; 23 3, 29 9,3 juv. from 
Ulva lactuca and Ulva reticulata, Le Bouchon, 27 October 1998. 

Description. Male length, 6.2 mm. Head with subocular 
notch, eyes round, a ring of clear ommatidia surrounding a 
dark central core. Antenna 1 weakly setiferous, article 2, 
1.3x article 1, article 3, 0.5x article 1; accessory flagellum 
3-articulate; primary flagellum 16-articulate (possibly 
regenerating in this specimen), flagellum can be 27- 
articulate (observed from additional material). Antenna 2 
weakly setiferous, peduncular article 5 subequal to 4, 
flagellum 8-articulate. Mandible palp article 3 slightly 
longer than 2, article 1, 0.3x article 3; article 2 with two 
groups of setae on posterior margin, article 3 with a few 
lateral and terminal setae. Maxilla 1 inner plate with 8 
plumose apical setae; Lower lip with rounded mandibular 
lobes. Gnathopod 1 coxa 1.4x as long as broad with short 
setae on ventral margin; basis 3x as long as broad with a 
strong patch of setae on anterodistal margin; carpus 1.5x 
length of propodus; propodus with transverse palm and 
forming a hood above dactylus; dactylus with medial 
protrusion on posterior margin. Gnathopod 2 coxa 
subrectangular 1.5x as long as broad, with setae on distal 
margin; basis 2.9x as long as broad, with a few groups of 
long setae on anterior margin; merus slightly produced 
ventrodistally; carpus 1.2x as broad as long; propodus 
subrectangular, 1.6x as long as broad, palmar margin weakly 
convex, palm rounded, with short stout setae and slender 
setae; dactylus broad throughout its length and slightly 
tapered at tip and closing across inner face of propodus. 
Pereopod 3 coxa subrectangular, 1.6x as long as broad, with 
very short setae on ventral margin; propodus and carpus 
subequal; dactylus with distal unguis. Pereopod 4 coxa 
deeply excavate posteriorly; other features similar to 
pereopod 3. Pereopod 5 coxa about 1.2x as long as broad, 
anterior margin with robust setae, posterior margin weakly 
serrated with short setae; dactylus short and robust with 
terminal unguis. Pereopod 6 coxa lobular; basis subovate, 
1.4x as long as broad, anterior margin with stout setae, 
posterior margin serrated with stout setae; propodus 2x 
length of carpus; other features similar to pereopod 5. 
Pereopod 7 basis, 1.3x as long as broad, anterior margin 
with numerous robust setae, posterior margin more convex 
and weakly serrated; other features similar to pereopod 6. 
Epimera 2 and 3 posterior margin weakly serrated, distal 
margins with a few stout setae. Urosomite 1 smooth. 
Urosomite 2 with two stout robust setae on each side. 
Uropods 1-2, rami subequal to each other and shorter than 
peduncle. Uropod 3 inner ramus rudimentary, with one 
robust seta; outer ramus 1-articulate, spatulate, 2.5x the 
length of peduncle, with robust setae and slender setae. 
Telson cleft to base, lobes with pointed apex; each lobe with 
two robust setae on distal inner margins and one on the 
outer margin; 1 or 2 robust setae present about half-way 
along inner margin. 


Records of the Australian Museum (2005) Vol. 57 


Fig. 5. Melita corticis n.sp., male, 6.2 mm, AM P60868, Le Bouchon. Scales: a = 0.4 mm (Hd, Ep, male G2, male 
C6), b = 0.2 mm (male Gl, enlargement of male G2), c = 0.05 mm (enlargement of male Gl), d = 0.1 mm (Mdp). 


Appadoo & Myers: Melitidae and gammarellidae from Mauritius 


229 


Fig. 6. Melita corticis n.sp., male, 6.2 mm, female, 3.3 mm, AM P60868, Le Bouchon. Scales: a = 0.2 mm (P3 and 
P5-P7), b = 0.2 mm (female G2 and female C6), c - 0.05 mm (female G1 and T), d = 0.2 mm (U3). 


230 Records of the Australian Museum (2005) Vol. 57 


Table 1. The character states in the Melita zeylanica group of species. 


Antenna 2 

Gnathopod 1 

Gnathopod 2 

U1 peduncle 

U3 inner 

Urosomite 2 


(male) dactylus 

; carpus 

basofacial 

ramus terminal 

dorsal 


proximal process 

robust seta 

robust setae 

robust setae 

M. zeylanica Stebbing, 1904 

weakly setose 

not swollen 

compressed 

absent 

1 

4-6 

M. zeylanica kauerti Barnard, 1972 

weakly setose 

swollen 

not compressed 

present 

3-4 

6 

M. setiflagella Yamato, 1988 

densely setose 

swollen 

not compressed 

present 

3 

6 

M. corticis n.sp. 

weakly setose 

swollen 

not compressed 

present 

1 

4 


Female: length, 3.3 mm (mature with eggs). Gnathopod 
1 coxa subrectangular, 1.6x as long as broad; basis about 
three times as long as broad; carpus 1.3x length of propodus; 
propodus palm transverse, palmar margin with short setae; 
dactylus large at base and tapered at tip. Gnathopod 2, coxa 
2x as long as broad; propodus subrectangular, 1.3x length 
of carpus with stout setae and slender setae on palm. 
Pereopod 6, coxa with a large hook-like anterior lobe. 

Remarks. The present material falls into the group of Melita 
that lacks a second article on the outer ramus of uropod 3 
(see Ledoyer, 1982: 568). It resembles M. pahuwai Barnard 
(1970) from Hawaii in having only one robust seta instead 
of 3 in the inner face of male gnathopod 1 propodus; in 
having the robust setae on the palmar margin of the female 
gnathopod 2 shorter than the inner facial robust setae rather 
than vice-versa and in lacking robust setae on the proximal 
outer margins of the telson. 

It appears to be particularly closely related to Melita 
zeylanica Stebbing, 1904, M. zeylanica kauerti J.L. Barnard 
1972 and M. setiflagella Yamato 1988. 

It can be distinguished from M. setiflagella Yamato 
(1988) by the well-developed circular eyes (smaller, slightly 
reniform eyes in M. setiflagella ) by antenna 2 peduncular 
articles 5 and 4 being subequal (peduncular article 5 shorter 
than 4 in M. setiflagella) by antenna 2 being poorly 
setiferous and 8-articulate (densely setiferous and 15- 
articulate in M. setiflagella ), and by female coxa 6 being 
without scale-like denticles. 

It differs from Melita zeylanica Stebbing (1904) in the 
presence of an anterodistal bulge near the base of the 
dactylus in the male gnathopod, in the non-compressed 
carpus of the gnathopod 2 in females, and in the presence 
of a robust basofacial seta on the peduncle of uropod 1, the 
latter feature, however, may have been overlooked by 
Stebbing (1904). Unlike Melita zeylanica kauerti Barnard 
(1972: 235, fig. 139-140) it has 2 (rather than 3) dorsolateral 
robust setae on urosomite 2 and one (rather than four) robust 
seta in the apex of the inner ramus of uropod 3. Barnard 
(1972) notes that Sri Lankan material of Melita zeylanica 
has one robust seta on uropod 3 inner ramus as in present 
material. 

These four species form a group of related forms. The 
current material compares most closely with Melita zeylanica 
kauerti but that species is closer to M. setiflagella than it is to 
the present material. Also Melita zeylanica kauerti differs 
more from Melita zeylanica than it does from the present 
material. This material is considered to represent a new 
species that can be distinguished from its close congeners 
by the combination of characters shown in Table 1. 

Habitat. Known only from Le Bouchon at depths of less 


than 1 m. The site is characterized by low salinity, green- 
algal growth and some estuarine conditions, which is in 
agreement with the general occurrence of the genus in 
brackish waters (Bousfield, 1973). 

Type locality. Le Bouchon, Mauritius. 

Distribution. Mauritius. 

Etymology. From the Latin cortex meaning a cork, in 
reference to the name of the type locality. 

Melita setimera n.sp. 

Figs. 7-8 

Type material. Holotype 4, 4.1 mm, AM P60869, 0-1 m depth, 
living on a mixture of Centroceras clavulatum, Hypnea sp., Gracilaria 
corticata, Enteromorpha flexuosa and Sargassum densifolium, Tamarin 
(20°19.5'S 57°22'E), Mauritius, C. Appadoo, 11 October 1999. 
Paratypes: 243,49$, AM P60870, same data as holotype; 1 9 from 
Acanthophora spicifera, Souillac (20°31'S 57°30.7'E), 10 November 
1998; 14,19 from Padina sp. and Halimeda sp., Grand Baie (20°0.5'S 
57°34'E), 5 May 1999; 244, 19, 1 juv. from mixture of Padina sp., 
Enteromorpha flexuosa, Hypnea sp., Amphiroa sp. and Caulerpa 
sertulariodes, Tamarin, 18 June 1999; 14, 3 9 9, 3 juv. from Amphiroa 
flagellisima and Padina, Ulva lactuca and Enteromorpha sp. and ash- 
coloured sand, Tamarin, 2 August 1999; 944, 129 9,9 juv. from mixture 
of Centroceras clavulatum, Hypnea sp., Gracilaria corticata, Enteromorpha 
flexuosa and Sargassum densifolium, Tamarin, 11 October 1999. 

Description. Male length, 4.3 mm. Head lacking subocular 
notch; eyes subround. Antenna 1 peduncle article 1 with 
stout robust setae on ventral margin; article 2, 1.2x article 
1; article 3 slightly less than half length of article 1; 
accessory flagellum 2-articulate, primary flagellum 20- 
articulate. Antenna 2 article 5 subequal to 4, flagellum 9- 
articulate. Mandible palp article 3 slightly longer than article 
2, article 1, 0.5x the length of article 3. Maxilla 1 inner 
plate with 6 apical plumose setae. Lower lip with rounded 
mandibular lobes. Gnathopod 1 coxa subrectangular, 1.6x 
as long as broad, distal margin with very short setae; basis 
slender, 2.9x as long as broad, with very dense patches of 
setae on anterior margin; propodus 0.6x length of carpus, 
with a hood over the dactylus; dactylus broad at base, with 
a small medial expansion and tapering tip. Gnathopod 2 
coxa subrectangular, 1.4x as long as broad; basis slightly 
expanded about 2.2x as long as broad, with dense long setae 
on anterior margin and a few patches of setae on posterior 
margin; carpus cup-shaped, 1.5x as broad as long; propodus 
1.3x times as long as broad, palmar margin slightly oblique, 
palmar border broadly sinuous; dactylus slender and 0.6x 
the length of propodus, dactylus closing across the medial 
face of propodus. Pereopod 3 coxa subrectangular, 1.5x as 
long as broad with short setae on ventral margin; basis 
slender, anterior margin concave, 3.5x as long as broad; 


Appadoo & Myers: Melitidae and gammarellidae from Mauritius 231 


Fig. 7. Melita setimera n.sp., male, 4.3 mm, female, 3.8 
mm, AM P60870, Tamarin. Scales: a = 0.4 mm (Hd and 
Ep), b = 0.2 mm (male Gl, G2 and female G1 and G2), 
c = 0.05 mm (enlargement of male Gl). 


propodus subequal to carpus; dactylus with terminal unguis 
and one seta on anterior margin. Pereopod 4 coxa excavate on 
posterior margin; other features as pereopod 3. Pereopod 5 
basis, 1.5x as long as broad, anterior margin with stout robust 
setae, posterior margin weakly serrated and with short setae; 
dactylus with 1 seta on anterior margin. Pereopod 6 basis 1.4x 
as long as broad, otherwise like that of pereopod 5; merus and 


carpus with dense patches of long setae on anterior margins 
and short robust setae on posterior margin. Pereopod 7 similar 
to pereopod 6 except dense patches of setae on merus and 
carpus on the posterior margins and short robust setae on 
anterior margins. Epimeron 1 with small posterodistal tooth. 
Epimeron 2 weakly toothed at posterodistal margin. Epimeron 
3 weakly toothed on posterodistal margin. Urosomite 1 with 


232 Records of the Australian Museum (2005) Vol. 57 


Fig. 8. Melita setimera n.sp., male, 4.3 mm, female, 3.8 mm, AM P60870, Tamarin. Scales: a = 0.2 mm (P3, C4, 
Ul,2, and female C6), b = 0.2 mm (P5 to P7), c = 0.05 mm (U3, T). 


acute dorsal tooth, urosomite 2 with one robust seta on mid¬ 
dorsal surface. Uropod 1 peduncle with basofacial robust seta, 
rami slender, 0.9x peduncle. Uropod 2 rami subequal to 
peduncle. Uropod 3 inner ramus rudimentary, with one or two 
terminal robust setae; outer ramus 1-articulate, 2x length of 
peduncle, spatulate, and with short robust setae and long 


fine setae. Telson apices pointed, each lobe with two robust 
setae on outer margin and two on medial hump. 

Female: length, 3.8 mm (mature, with eggs). Gnathopod 
1 coxa 2x as long as broad; basis slender, with patches of 
setae on proximal and distal anterior margins and on medial 
posterior margin; propodus palm transverse; dactylus 


Appadoo & Myers: Melitidae and gammarellidae from Mauritius 233 


Fig. 9. Nuuanu rectimana n.sp., male, 5.2 mm, female, 4.2 mm, AM P60906, Albion. Scales: a = 1 mm (whole animal), 
b = 0.2 mm (female G1 and male G1 and G2), c - 0.2 mm (male G2) and d - 0.2 mm (enlargement of male G2). 


normal. Gnathopod 2 coxa 2.lx as long as broad; basis 
slender, 3x as long as broad; propodus slightly longer than 
carpus, palm oblique, with long setae on palmar margin 
and anterior margin. Pereopod 6, coxa bilobed without any 
finger-like protuberance. Female pereopods 6 and 7 without 
dense setae on merus and carpus. 


Remarks. This species most closely resembles Melita 
simplex Myers (1985) from Fiji, in having an acute dorsal 
tooth on urosomite 1 and a non sexually-dimorphic female 
coxa 6. Melita setimera male gnathopod 1, however, has a 
densely setose distal margin on the basis, the propodus is 
shorter than the carpus and it has a lobe above the dactylus. 


234 Records of the Australian Museum (2005) Vol. 57 


Fig. 10. Nuuanu rectimana n.sp., male, 5.2 mm, AM P60906, Albion. Scales: a = 0.4 mm (P3, C4, P5 to P7, A1 to 
A2), b = 0.2 mm (U1 to U3), c = 0.1 mm (Mxp), d = 0.1 mm (Md, Mxl, D3, D5), e = 0.05 mm (T). 


The male gnathopod 2 propodus of M. setimera has parallel 
margins and the palmar border is broadly sinuous whereas 
the ventral margin of male gnathopod 2 of M. simplex is 
evenly convex. Pereopods 6 and 7 merus and carpus are 
densely setose in M. setimera, but this character state is 


unknown in Melita simplex. The telson apices of M. simplex 
have a terminal robust seta that is lacking in M. setimera. 

Melita setimera can be distinguished from all other 
species of Melita from Mauritius, by the presence of the 
dorsal tooth on urosomite 1 on both males and females. 


Appadoo & Myers: Melitidae and gammarellidae from Mauritius 235 


Males are easily distinguished by the broadly sinuous palmar 
margin of gnathopod 2 and the presence of dense long setae 
on merus of pereopods 6 and 7. 

Habitat. Melita setimera was collected only at Tamarin at 
depths of less than 1 m. The site has some freshwater 
influence due to a river flowing in the vicinity. 

Type locality. Tamarin, Mauritius. 

Distribution. Mauritius. 

Etymology. From the Latin saeta = bristle coupled with 
merus referring to the strongly setose merus of male 
pereopods 6 and 7. 

Nuuanu rectimana n.sp. 

Figs. 9-10 

Nuuanu sp. 1 (Appadoo & Steele, 1998). 

Type material. Holotype 6, 4 mm, AM P60905, 0.5-1 m depth, 
living on mixture of Enteromorpha flexuosa, Laurencia papillosa, 
Halodule uninervis, Poste La Fayette (20°08.2'S 57°44.5'E), Mauritius, 
C. Appadoo, 7 February 2000. Paratypes: 1 d, 39 9, AM P60906, 0.5 
m, living among Gracilaria salicornia, Souillac (20°31'S 57°30.7'E), 
Mauritius, 25 March 1999; Id, 1 juv. from green filamentous algae and 
Halodule uninervis Albion (20° 13'S 57°23.7’E), 12 May 1998; Id from 
Sargassum sp., Bain Boeuf (19°59'S 57°36'E), 15 May 1998; 1 d from 
Sargassum sp., Bain Boeuf, 16 June 1998; 1 juv. from Jania sp. and 
Valonia sp., Bain Boeuf, 28 July 1998; 1 juv. from Pocockiella variegata 
and Valonia sp., Balaclava (20°03.7'S 57°30.7'E), 10 September 1998; 
1 juv. from Gracilaria salicornia, Souillac, 25 March 1999; 2dd, 
119 9,5 juv. from Laurencia papillosa and Cladophora sp., Albion, 20 
April 1999; 1 juv. from Sargassum sp., Amphiroa sp., Pocockiella 
variegata and Cymodocea sp., Bain Boeuf, 16 June 1999; Id, 2 juv. 
from Sargassum sp., La Cuvette, 12 October 1999; 19 from coral rubble, 
Padina sp. and Pocockiella variegata, Flic-en-Flac, 10 December 1999; 
19 from mixture of Padina sp., Turbinaria sp., Sargassum sp., 
Pocockiella variegata, Bain Boeuf, 24 January 2000; 1 juv. from mixture 
of Enteromorpha flexuosa, Laurencia papillosa and Halodule uninervis, 
Poste La Fayette, 7 February 2000. 

Description. Male length, 4.0 mm. Head with lateral 
cephalic lobe notched; ommatidia of eyes sparse. Antenna 
1 poorly setiferous, peduncle article 2 slightly longer than 
article 1; article 3, 0.5x article 1; accessory flagellum 4- 
articulate, primary flagellum 30-articulate. Antenna 2 poorly 
setiferous, article 5, 0.8x article 4, flagellum 20-articulate. 
Mandible palp slender, subfalcate, article 2 longest; article 
3,0.8x article 2 and 2. lx article 1; article 3 with short setae 
on medial margin and three long terminal setae. Maxilla 1 
palp 2-articulate, article 2 with stout blunt robust setae at 
tip; inner plate with a small protuberance at apex and with 
long marginal setae. Maxilla 2 inner plate with oblique setal 
row. Gnathopod 1 coxa 1.5x as long as broad, posterodistal 
margin with a notch and distal margin with very short setae; 
basis slender, 3x as long as broad, posterior margin with a 
strong patch of setae; carpus slender, subrectangular, 3.5x 
as long as broad; propodus 0.6x length of carpus; palmar 
margin oblique with few setae; dactylus short, fitting palm. 
Gnathopod 2 coxa subrectangular 1.6x as long as broad, 
posterodistal margin with notch and distal margin with very 
few short setae; basis 2.5x as long as broad; carpus over 
three and half times as broad as long; propodus subrect¬ 
angular, posterior margin straight, defined by a small hump 
at the proximal end and well-developed blunt projections 
embedded at the distal end, densely setose, palm obsolete, 
dactylus 0.6x length of propodus, robust and broad 


throughout its length. Pereopod 3 coxa subrectangular, 2.2x 
as long as broad, posterodistal margin with notch; dactylus 
with two long stout setae and one small slender seta close 
to apical unguis. Pereopod 4 coxa 1.7x as long as broad, 
posterior margin excavate; other features similar to pereopod 
3. Pereopod 5 basis 1.3x as long as broad, anterior margin 
convex and with stout robust setae, posterior margin 
serrated, convex proximally and concave distally; propodus 
subequal to carpus; dactylus with 2 long setae and one 
slender seta at unguis. Pereopod 6 basis about 1.5x as long 
as broad, similar to that of pereopod 5 except posterior 
margin is more castelloserrate. Pereopod 7 basis broadly 
expanded, about as long as broad, anterior margin with 
robust setae, posterior margin strongly convex, castellate 
with short setae; propodus slightly longer than carpus. 
Pleonites 1 and 2 with well-developed dorsal tooth. Epimera 
1 to 3 subrectangular, distal margins smooth. Uropod 1 rami 
subequal to each other and 0.8x the length of peduncle. 
Uropod 2 rami subequal to each other, and slightly longer 
than peduncle, with stout robust setae. Uropod 3 peduncle 
1.5x as long as broad; outer ramus 2-articulate, article 1 
truncate with a stout robust setae on mid-lateral margin and 
two stout distal robust seta; article 2 produced with a stout 
robust seta; inner ramus short and sub-falcate; inner margins 
of both rami with very fine short setae. Telson cleft to 80% 
its length; telson lobes with broadly rounded apex, each 
with one plumose seta and two short slender setae. 

Female: length, 4.2 mm (mature, oostegites with setae). 
Gnathopod 1 coxa 1.7x as long as broad, posterodistal margin 
with a notch and short setae; basis slender, 3.5x as long as 
broad; anterior margin with patches of setae, posterior margin 
with one strong patch of setae; propodus 0.7x length of carpus; 
palmar margin oblique, with short setae; dactylus stout, fitting 
palm. Gnathopod 2 coxa 2. lx as long as broad, posterodistal 
margin with notch and short setae; basis slender and 3.6x as 
long as broad; propodus 1.2x length of carpus; propodus 
slightly less 3.5x as long as broad, palmar margin oblique, 
defined by a stout robust seta; dactylus stout, fitting palm. 

Remarks. The genus Nuuanu, established by Barnard 
(1970), belongs to the “ Gammarella ” group, recently revised 
by Lowry & Watson (2002). Males of Nuuanu rectimana 
n.sp. can easily be distinguished from Nuuanu amikai 
Barnard (1970), recorded by Ledoyer (1978) from 
Mauritius, by the presence of a well-developed flat-topped 
processes on the propodus of gnathopod 2. Other differences 
include the shape, setation and spination of the telson. 
Nuuanu rectimana n.sp. telson is symmetrical with two 
robust setae and one plumose seta on each lobe whereas in 
Nuuanu amikai Barnard (1970: 167, fig. 105) the telson is 
asymmetrical, with one robust seta and two plumose setae 
on one lobe and one plumose seta on the other. In addition 
in Nuuanu rectimana n.sp. the inner lobe of uropod 3 is 
slender, subfalcate and about half the left of the outer lobe 
whereas in N. amikai (Barnard 1970: 168, fig. 106) the inner 
lobe is triangular and less than half the length of the outer 
ramus. Nuuanu rectimana n.sp. differs from Nuuanu 
numbadi Barnard (1974: 39, fig. 27) from Australia by 
having more distally tapered telson lobes with one robust 
and two plumose setae as compared to more distally broad 
telson lobes with two robust and one plumose seta. Another 
difference is that epimera 2 and 3 are less acute in N. 
rectimana n.sp. than in N. numbadi. 


236 Records of the Australian Museum (2005) Vol. 57 


Habitat. Nuuanu rectimana was collected in depths of less 
than 1 m, from seagrass (Halodule uninervis or Cymodocea 
sp.) mixed with other algae and coral rubble at Albion, Poste 
La Fayette and Bain Boeuf. 

Type locality. Poste La Fayette, Mauritius. 

Distribution. Mauritius. 

Etymology. The species is named from the Latin rectus 
meaning straight and manus meaning hand, referring to the 
straight posterior margin of the propodus of the male 
gnathopod 2. 


Acknowledgments. We are grateful to the University of 
Mauritius and the Tertiary Education Commission for their support 
in carrying out the current study. Thanks also due to University 
of Mauritius (Higher Technical Education Plan) for fully 
sponsoring visits of one of us (C.A.) to University College Cork, 
Ireland. We are also deeply indebted to Prof. I. Fagoonee for his 
support in carrying out this study. We thank Prof. J. Davenport 
and the staff of the Department of Zoology at University College 
Cork, for their hospitality and support. The authors are also 
grateful to Dr J.K. Lowry of the Australian Museum for allowing 
us access to unpublished manuscripts and for critical appraisal of 
the manuscript. 


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tropics: a review. Micronesica 12(1): 169-176. 


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Grand recif de Tulear (Madagascar) (Crustacea, Amphipoda). 
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Ledoyer, M., 1982. Crustaces Amphipodes Gammaridiens famille 
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Madagascar 59(1): 1-598. 

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group, with a new species from the Andaman Sea (Crustacea, 
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Special Publication 23: 197-212. 

Myers, A.A., 1985. Shallow-water, coral reef and Mangrove 
Amphipoda (Gammaridea) of Fiji. Records of the Australian 
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Manuscript received 26 January 2001, revised 24 September 2003 and 

accepted 14 November 2003. 

Associate Editor: G.D.F. Wilson. 


© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 237-302. ISSN 0067-1975 


New and Little-known Melitid Amphipods 
from Australian Waters 
(Crustacea: Amphipoda: Melitidae) 


J.K. Lowry* and R.T. Springthorpe 


Australian Museum, 6 College Street, Sydney NSW 2010, Australia 
jimlowry@crustacea.net • rogers@austmus.gov.au 


Abstract. During a recent review of Australian melitid amphipods (Lowry, Berents & Springthorpe, 
2001), a number of problems and undescribed species were revealed. These problems and others 
uncovered during the course of this study are addressed here. The Melitidae, as currently conceived, is 
considered to contain at least five natural groups: the Ceradocopsis group; the Ceradocus group; the 
Eriopisa group; the Nuuana group and the Melita group. In the Ceradocus group Austromaera n.gen. is 
established for Maera mastersii Haswell, 1879a, that is redescribed based on syntype material and new 
collections from Port Jackson. Ceradocus circe n.sp. is described based on specimens from Tasmania, 
originally reported by Chilton (1921b) as C. rubromaculatus (Stimpson, 1856). Maera boecki Haswell, 
1879 and Maera hamigera Haswell, 1879 ( sensu stricto ), both originally described from Port Jackson, 
are redescribed and these species as well as Maera octodens Sivaprakasam, 1968 are transferred to the 
genus Linguimaera Pirlot, 1934, recently re-established by Krapp-Schickel (2003). Linguimaera 
schickelae n.sp. is described from the Sydney area. Maera grijfini Berents, 1983, is tentatively placed in 
the genus Maeropsis Chevreux, 1919. Mallacoota subcarinata is redescribed based on syntypes and 
new collections and four new Australian species (M. chandaniae n.sp., M. euroka n.sp., M. kameruka 
n.sp. and M. malua n.sp.) are described, two of which have previously been mis-identified as M. 
subcarinata. Mallacoota nananui Myers, 1985, is reported from Australia for the first time. Miramaera 
thetis n.gen., n.sp. is established for the specimens mis-identified by Stebbing, 1910a as Maera inaequipes. 
We describe a new species of Parelasmopus, P. sowpigensis n.sp., from Port Jackson, New South Wales. 
Quadrivisio sarina n.sp. is described from near McKay, Queensland, the first record of Quadrivisio 
Stebbing, 1907 in Australian waters. In the Eriopisa group Victoriopisa australiensis (Chilton, 1923) is 
redescribed and illustrated. A second species, Victoriopisa marina n.sp., is described from estuarine and 
marine habitats in New South Wales and Victoria. In the Melita group Dulichiella australis (Haswell, 
1879a) is redescribed based on syntype material and D. pacifica n.sp. is described based on collections 
from the Great Barrier Reef, the southwestern Pacific Ocean and the South China Sea. Melita ophiocola 
n.sp. is described from Port Jackson, living in association with the brittlestar Ophionereis schayeri. 


Lowry, J.K., & R.T. Springthorpe, 2005. New and little-known melitid amphipods from Australian waters 
(Crustacea: Amphipoda: Melitidae). Records of the Australian Museum 57(2): 237-302. 


* author for correspondence 


www.amonline.net.au/pdf/publications/1463_complete.pdf 


238 Records of the Australian Museum (2005) Vol. 57 


Melitid amphipods are a diverse and abundant part of the 
Australian amphipod fauna. Including this work and the 
recent work of Krapp-Schickel (2003), there are 86 known 
Australian marine and freshwater species (Table 1). The 
majority of the marine component comes from the 
southeastern and southwestern parts of the country. Many 
species are yet to be discovered from tropical areas. Nearly 
75% of the Australian melitid fauna has been described since 
the early 1970s and many of the earlier species were 
redescribed during this time. Lowry et al. (2001) reviewed 
the marine component of this fauna and provided 
redescriptions, pictures and an interactive key to all marine 
species at http://www.crutacea.net. The new species from 
that review are described here and some of the more 
problematical species are redescribed. 


The Haswell species 

The first descriptor of the Australian amphipod fauna, W. A. 
Haswell, described 10 species of melitids (Table 1), mainly 
from Port Jackson (Haswell, 1879a,b). Based on the original 
descriptions, these amphipods have never been easy to 
identify, and the status of his types, which could be used 
for redescriptions, has always been a problem (Springthorpe 
& Lowry, 1994). Sheard (1936) and J.L. Barnard (1972a) 
redescribed Ceradocus ramsayi (Haswell, 1879a) and 
Maera mastersii (Haswell, 1879a), but the material Sheard 
(1936) considered to be M. mastersii has recently been 
described as Linguimaera tias Krapp-Schickel, 2003 and 
the material of J.L. Barnard (1972a) has been described as 
Linguimaera leo Krapp-Schickel, 2003. J.L. Barnard 
(1972a) redescribed Mallacoota diemenensis (Haswell, 


Table 1. Australian Melitidae, * freshwater species. 


Ceradocopsis group 

Ceradocopsis hamondi Moore, 1988 

Ceradocus group 

Austromaera mastersii (Haswell, 1879a) 
Ceradocus circe n.sp. 

Ceradocus dooliba J.L. Barnard, 1972a 
Ceradocus hawaiensis J.L. Barnard, 1955 
Ceradocus oxyodus Berents, 1983 
Ceradocus ramsayi (Haswell, 1879a) 

Ceradocus rubromaculatus (Stimpson, 1856) 
Ceradocus sellickensis Sheard, 1939 
Ceradocus serratus (Bate, 1862) 

Ceradocus woorree Berents, 1983 
Ceradocus yandala Berents, 1983 
Elasmopus alalo Myers, 1986 
Elasmopus bollonsi Chilton, 1915 
Elasmopus crenulatus Berents, 1983 
Elasmopus hooheno J.L. Barnard, 1970 
Elasmopus menurte J.L. Barnard, 1974 
Elasmopus pocillimanus (Bate, 1862) 
Elasmopus spinicarpus Berents, 1983 
Elasmopus warra Kelaher & Lowry, 2002 
Elasmopus yunde J.L. Barnard, 1974 
Hoho carteta (J.L. Barnard, 1972a) 

Hoho hirtipalma Lowry & Fenwick, 1983 
Hoho marilla (J.L. Barnard, 1972a) 
Linguimaera boecki (Haswell, 1879b) 
Linguimaera bogombogo Krapp-Schickel, 2003 
Linguimaera caesaris Krapp-Schickel, 2003 
Linguimaera garitima Krapp-Schickel, 2003 
Linguimaera hamigera (Haswell, 1879b) 
Linguimaera kellissa Krapp-Schickel, 2003 
Linguimaera leo Krapp-Schickel, 2003 
Linguimaera octodens (Sivaprakasam, 1968) 
Linguimaera schickelae n.sp. 

Linguimaera thomsoni (Miers, 1884) 
Linguimaera tias Krapp-Schickel, 2003 
Maeracoota sp. Krapp-Schickel & Ruffo, 2001 
Maeropsis griffini (Berents, 1983) 

Mallacoota balara Berents, 1983 
Mallacoota chandaniae n.sp. 

Mallacoota diemenensis (Haswell, 1879a) 
Mallacoota euroka n.sp. 

Mallacoota kameruka n.sp. 

Mallacoota malua n.sp. 

Mallacoota nananui Myers, 1985 


Mallacoota subcarinata (Haswell, 1879b) 
Miramaera thetis n.sp 
Parapherusa crassipes (Haswell, 1879b) 
Parelasmopus echo J.L. Barnard, 1972a 
Parelasmopus sowpigensis n.sp. 

Parelasmopus suensis (Haswell, 1879b) 
Parelasmopus ya J.L. Barnard, 1972a 
Premaera thetis n.sp. 

Quadrimaera quadrimana (Dana, 1852) 
Quadrimaera reishi (J.L. Barnard, 1979) 
Quadrimaera serrata (Schellenberg, 1938) 
Quadrimaera viridis (Haswell, 1879b) 

Quadrivisio sarina n.sp. 

Eriopisa group 

*Nedsia chevronia Bradbury, 2002 
*Nedsia douglasi Barnard & Williams, 1995 
*Nedsiafragilis Bradbury & Williams, 1996 
*Nedsia halletti Bradbury, 2002 
*Nedsia humphreysi Bradbury & Williams, 1996 
*Nedsia hurlberti Bradbury & Williams, 1996 
*Nedsia macrosculptilis Bradbury & Williams, 1996 
*Nedsia sculptilis Bradbury & Williams, 1996 
*Nedsia stefania Bradbury, 2002 
*Nedsia straskraba Bradbury & Williams, 1996 
*Nedsia urifimbriata Bradbury & Williams, 1996 
*Norcapensis mandibulis Bradbury & Williams, 1997 
*Nurina poulteri Bradbury & Eberhard, 2000 
Victoriopisa australiensis (Chilton, 1923) 
Victoriopisa marina n.sp. 

Nuuana group 

Gammarella berringar (J.L. Barnard, 1974) 

Nuuanu merringannee (J.L. Barnard, 1974) 

Nuuanu mokari J.L. Barnard, 1974 
Nuuanu numbadi J.L. Barnard, 1974 

Melita group 

*Brachina invasa Barnard & Williams, 1995 
Dulichiella australis (Haswell, 1879a) 

Dulichiella pacifica n.sp. 

Melita jestiva (Chilton, 1884) 

Melita kauerti J.L. Barnard, 1972a 
Melita matilda J.L. Barnard, 1972a 
Melita myersi Karaman, 1987 
Melita oba J.L. Barnard, 1972a 
Melita ophiocola n.sp. 

*Melita plumulosa Zeidler, 1989 


Lowry & Springthorpe: Australian melitid amphipods 239 


1879a) and Quadrimaera viridis (Haswell, 1879b) and 
Berents (1983) redescribed Parelasmopus suensis (Haswell, 
1879b). Parapherusa crassipes (Haswell, 1879b) is 
considered to be a well known, distinctive species. In this 
paper we redescribe Austro maera mastersii (Haswell, 1879a), 
Dulichiella australis (Haswell, 1879a), Linguimaera boecki 
(Haswell, 1879b), Linguimaera hamigera (Haswell, 1879b) 
md Mallacoota subcarinata (Haswell, 1879b). 

The Dulichiella complex 

Once LeCroy (2000) redescribed Dulichiella appendiculata 
(Say, 1818) from near the type locality (LeCroy, pers. 
comm.), it was clear that material from Australia did not 
belong to that species. Ledoyer (1986) gave an excellent 
summary of the problems associated with available names 
for species of Dulichiella. Based on this information we 
were able to locate and borrow material of D. appendiculata 
(Say, 1818), St Catherines Island, Georgia, USA, D.fresnelii 
(Audouin, 1826), Great Bitter Lake, Suez Canal and D. 
anisochir (Kroyer, 1845) Rio de Janeiro, Brazil. Dana’s 
types are lost and his species (Dulichiella validus (Dana, 
1852), Singapore; Dulichiella setipes (Dana, 1852), Rio de 
Janeiro; Dulichiella pilosus (Dana, 1852), Rio de Janeiro) 
are only identifiable at generic level. Comparison of our 
material with these species showed that D. australis is a valid 
species and that an undescribed species (D. pacifica n.sp.) 
occurred in northeastern Australia, the southwest Pacific and 
the South China Sea. A separate paper (Lowry & Springthorpe, 
in prep.) revises Dulichiella on a world-wide basis. 

The Maera complex 

Krapp-Schickel & Ruffo (2000) recently established the 
genus Quadrimaera and transferred four of the nine 
Australian species, previously considered to be Maera , to 
this genus (Table 1). Krapp-Schickel (2003) recently re¬ 
established the genus Linguimaera Pirlot, 1936 and 
described seven new Australian species in the genus. But 
five remaining Australian species are still unassigned. 

In the original description of Maera mastersii Haswell, 
1879a, the illustrations of uropod 3 and the telson differ 
significantly from those presented by Sheard (1936) and 
J.L. Barnard (1972a). We examined syntype material and 
new material, both from Port Jackson, that agree with the 
description and illustrations of Haswell (1879a). Based on 
this material we redescribe the species and place it in the 
new genus Austromaera. Within Australia, the material 
considered by Sheard (1936) as Maera mastersii has been 
described as Linguimaera tias Krapp-Schickel, 2003 and that 
of J.L. Barnard (1972a) has been described as Linguimaera 
leo Krapp-Schickel, 2003. Krapp-Schickel (2003) has also re¬ 
established the Torres Strait species, Maera thomsoni (Miers, 
1884) (synonymized withM mastersiiby Haswell, 1885), and 
transferred it to Linguimaera. 

Two main characters define Linguimaera Pirlot, 1936 
(Krapp-Schickel, 2003). The first is the second gnathopods 
of the male that are always asymmetrical, so that one is 
similar to the female second gnathopod and the other is 
enlarged and morphologically dissimilar, as is typical of 
mate-guarding amphipods. The second character is a serrate 
posterior margin on epimeron 3 (weakly serrate in M. 
hamigera). Among the Australian species in our study, 
Maera boecki Haswell, 1879, M. hamigera Haswell, 1879 


and M. octodens Sivaprakasam, 1968, all have these 
characteristics. They are here transferred to the genus 
Linguimaera. In addition, a new species, Linguimaera 
schickelae n.sp., is described from the Sydney area. 

The original description of Maera boecki Haswell, 1879b, 
was inadequate and the type material is apparently lost 
(Springthorpe & Lowry, 1994). As a result the species has 
been unidentifiable. Della Valle (1893) referred to it as 
?Maera boeckii. Stebbing (1899) transferred it to Elasmopus 
in his world monograph (Stebbing, 1906). K.H. Barnard 
(1916) appears to have erroneously reported E. boecki from 
South Africa. Since Sheard (1937) placed it in his catalogue 
of Australian Gammaridea, there have been no further 
records or new material. While studying material for the 
Australian Amphipod Project, we discovered material from 
Port Jackson that we are referring to this species. The species 
has all the characteristics of a Linguimaera , except for an 
emarginate telson, which appears to be independently derived. 

The name Maera hamigera Haswell, 1879b (type locality 
Port Jackson), has not been used for an Australian species 
since Stebbing (1910a), but it has been used for species 
living in the Red Sea (Walker, 1909; Lyons & Myers, 1993), 
southern Africa (K.H. Barnard, 1916), Micronesia (J.L. 
Barnard, 1965), the Mediterranean Sea (Karaman & Ruffo, 
1971), Madagascar (Ledoyer, 1982) and Western Samoa 
(Myers, 1997). A microscope slide from the syntype series 
shows the unusual gnathopod 2 of this species. Using this 
evidence we discovered many specimens of this species in 
collections from Twofold Bay on the south coast of New 
South Wales. We redescribe the species here, based on the 
syntypes and the newly discovered material. These results 
indicate that L. hamigera is currently confined to 
southeastern Australia. Material from other areas that also 
appears to be in the genus Linguimaera, and has been 
attributed to this species, needs to be re-examined. Krapp- 
Schickel (2003) recently renamed material, identified as M. 
hamigera, from the Mediterranean, the Red Sea, Madagas¬ 
car and Western Samoa as L. caesaris, but left the material 
from southern Africa and Micronesia unresolved. 

Stebbing (1910a) identified material from the Thetis 
Expedition as Maera inaequipes (Costa, 1851). Even as he 
did, he stated that “the specimens ... do not justify the 
specific name”. We establish the new genus and species 
Miramaera thetis based on material in the Australian 
Museum collections and transfer the New Zealand species, 
Maera tepuni J.L. Barnard, 1972b, to Miramaera. 

Based on the key in Krapp-Schickel (2000) and 
unpublished phylogenetic analyses (JKL) we tentatively 
place Maera grijfini Berents, 1983, in the genus Maeropsis 
Chevreux, 1919. Krapp-Schickel & Ruffo (2001) deduced 
that Maera tenella of Tattersall, 1922 (Wooded Island, 
Abrolhos Islands) is actually a species of Maeracoota 
Myers, 1997. Unfortunately material of this species is not 
available for study. 

The Mallacoota Complex 

Mallacoota J.L. Barnard, 1972, was established for six 
species: M. carteta J.L. Barnard, 1972, M. diemenensis 
(Haswell, 1879), M. insignis (Chevreux, 1901), M. marilla 
J.L. Barnard, 1972, M. odontoplax (Pirlot, 1936) and M. 
subcarinata (Haswell, 1879). At the same time Barnard 
demonstrated several “phenotypes” for M. subcarinata and 
M. carteta. Lowry & Lenwick (1983) subsequently removed 


240 Records of the Australian Museum (2005) Vol. 57 


M. marilla and M. carteta to a new genus, Hoho, and 
described a third species, H. hirtipalma. 

Since then Ortiz has described M. carausui Ortiz, 1976, 
Ledoyer has described M. subinsignis Ledoyer, 1979, M. 
latidactylus Ledoyer, 1982 and M. schellenbergi Ledoyer, 
1984 and transferred (Ledoyer, 1982) Elasmopus latibrach¬ 
ium Walker, 1905 to Mallacoota. Berents (1983) has 
described M. balara and Myers (1985) has described M. 
nananui. Myers (1985) summarized the status of Mallacoota 
and concluded that an in-depth study using a wide range of 
material was needed. Appadoo et al. (2002) began this 
process by redescribing M. insignis, M. latibrachium and 
M. schellenbergi and describing the new species M. caerulea 
from Mauritius. They also excluded M. subcarinata of 
Ledoyer (1978) from that area. 

The original description of Mallacoota subcarinata 
(Haswell, 1879b) was apparently based on a series of small 
adult specimens from Port Jackson. These syntypes are 
lodged in the Australian Museum, but are in poor condition. 
All have the unusual gnathopod 2 palmar shape indicated 
by Haswell’s (1879b) illustrations. Unfortunately this is not 
one of the species with which the name has been 
subsequently associated. To add to this confusion J.L. 
Barnard (1972) described three “phenotypes” in his 
redescription of M. subcarinata. In this paper we re-establish 
the original species concept of M. subcarinata by 
redescribing and illustrating the types and new material from 
near the type locality. We describe new Australian species 
(M. chandaniae n.sp., M. euroka n.sp., M. kameruka n.sp. 
and M. malua n.sp.) some of which have in the past been 
mis-identified as M. subcarinata and comment on records 
of M. subcarinata from geographic areas outside of 
Australia. We also report M. nananui Myers, 1985 from 
Australian waters for the first time. 

Currently Mallacoota contains 16 species (Table 2) half of 
which occur in Australia. The genus is essentially Indo Pacific 
with one species, M. carausui, in the tropical western Atlantic. 

Parelasmopus, Quadrivisio and Victoriopisa 

J. L. Barnard (1972a) redefined Parelasmopus and 
established Ifalukia for a species that did not quite fit the 
Parelasmopus mould. In the same paper he described two 
Australian species and synonymized P. suensis (Haswell, 
1879b) with P. setiger Chevreux, 1901. Strangely he 
continued to use the later name, P. setiger. J.L. Barnard 
(1974) reassessed his position and considered P. suensis 
and P. setiger to be separate species. Finally, Berents (1983) 
redescribed P. suensis from a lectotype male. In this paper 
we describe the fourth Australian species and the first species 
from the southeastern part of the country. This species, 
Parelasmopus sowpigensis, also does not quite fit the 
Parelasmopus or Ifalukia moulds, but it is placed in a slightly 
expanded concept of the genus Parelasmopus. 

Quadrivisio currently contains five species: Q. aviceps 

K. H. Barnard, 1940; Q. bengalensis Stebbing, 1907; Q. 
bousfieldi Karaman & Barnard, 1979; Q. lobata Asari, 1983; 
and Q. lutzi (Shoemaker, 1933) none of which are known 
from Australian waters. We describe a new species, Q. 
sarina, from the Queensland coast. 

Stock & Platvoet (1981) revised the genus Victoriopisa and 
described a new species from Mauritania in the eastern North 
Atlantic. Including the new species described here, Victoriopisa 
now contains seven species: V. atlantica Stock & Platvoet, 1981; 


Table 2. Species of Mallacoota. 


M. balara Berents, 1983 

M. caerulea Appadoo, Myers & Fagoonee, 2002 
M. carausui Ortiz, 1976 
M. chandaniae n.sp. 

M. diemenensis (Haswell, 1879) 

M. euroka n.sp. 

M. insignis (Chevreux, 1901) 

M. kameruka n.sp. 

M. latibrachium (Walker, 1905) 

M. latidactylus Ledoyer, 1982 
M. malua n.sp. 

M. nananui Myers, 1985 
M. odontoplax (Pirlot, 1936) 

M. schellenbergi Ledoyer, 1984 
M. subcarinata (Haswell, 1879b) 

M. subinsignis Ledoyer, 1979 


V. australiensis (Chilton, 1923); V. chilkensis (Chilton, 1921a); 
V. epistomata (Griffiths, 1974a); V. marina n.sp.; V. papiae 
Asari, 1983 and V. ryukyuensis Morino, 1991. In a phylogenetic 
analysis of the Eriopisa group van der Ham & Vonk (2003) 
affirmed the monophyly of Victoriopisa. 

The original description and illustrations of Victoriopisa 
australiensis (Chilton, 1923) were inadequate and the type 
was thought to be lost (Springthorpe & Lowry, 1994). When 
Karaman & Barnard (1979) established the genus 
Victoriopisa they included V. australiensis, but did not 
redescribe it. This is a very distinctive species that lives in a 
restricted habitat. For these reasons we redescribe and 
illustrate this species, based on material from Boambee 
Creek, Sawtell, New South Wales, near the type locality, 
Trial Bay. After our illustration of the Boambee Creek 
specimens were completed the type was located, too late to 
be incorporated into this study. A second species 
(Victoriopisa marina n.sp.) is described, from estuarine and 
marine habitats in New South Wales and Victoria. 

Methods 

The taxonomic descriptions presented in this paper were 
generated from a DELTA (Dallwitz et al., 1993; Dallwitz et 
al., 1998) database of Australian melitid species. Unless 
indicated otherwise, the following attributes are implicit 
throughout the descriptions, except where the characters 
concerned are inapplicable. 

Head. Eyes present; one pair. Antenna 1 peduncular article 
2 not geniculate. Mandible palp present, well developed. 

Pereon. Gnathopod 1 not sexually dimorphic; carpus about 
2x as long as broad; carpus without anterodistal swelling; 
propodus without anterodistal projection, posterodistal margin 
not swollen. Gnathopod 2 left and right gnathopods subequal 
in size; propodus without strong concentration of setae, 
distolateral margin without spines; dactylus closing along palm, 
reaching end of palm, inner margin smooth. Pereopod 5 
dactylus unguis anterior margin without accessory spines; 
carpus and propodus with few (or none) long, slender setae 
along anterior margin. Pereopod 6 coxa anterior lobe ventral 
margin not produced ventrally; carpus and propodus with few 
(or none) long, slender setae along anterior margin; propodus 
not expanded posterodistally; dactylus unguis anterior margin 
without accessory spines. Pereopod 7 basis with posterior 


Lowry & Springthorpe: Australian melitid amphipods 241 


margin smooth or minutely castelloserrate; mems posterodistal 
margin narrowly rounded or subquadrate; propodus not 
expanded posterodistally; dactylus unguis anterior margin 
without accessory spines. 

Pleon. Pleonite 1 without dorsal serrations, without 
dorsodistal spines, not dorsally bicarinate. Pleonite 2 
without dorsal serrations, without dorsodistal spines, not 
dorsally bicarinate. Pleonite 3 without dorsal serrations, 
without dorsodistal spines, not dorsally bicarinate. Epimeron 
1 anteroventral corner without curved spine. Epimeron 3 
posteroventral margin smooth. Urosomite 1 without dorsal 
carina, without a small dorsal hump, without dorsal 
serrations, without spines or gape, not dorsally bicarinate, 
without posterodorsal spine. Urosomite 2 posterior margin 
smooth, without dorsolateral robust setae. Urosomite 3 
without dorsal robust setae. Uropod 1 peduncle with 
basofacial robust seta; without distoventral spur. Telson cleft, 
without robust setae on inner margins. 

A separate generic level DELTA database to species in the 
Maera complex was used to generate generic diagnosis and to 
generate nexus files. A preliminary phylogenetic analysis was 
generated using PAUP 4.0 win 10 in order to analyse 
relationships among genera and species in this complex. 

Material used in this study is lodged in the Australian 
Museum, Sydney (AM). The following abbreviations are 
used on the plates: A, antenna; C, coxa; E , epimeron; G, 
gnathopod; H, head; MD, mandible; MP, maxilliped; MX, 
maxilla; p, palp; P, pereopod; PLN, pleonite; T, telson; U, 
uropod; UR, urosomite; L, left; R, right. 

Taxonomy 

Melitidae Bousfield, 1973 

Lowry & Watson (2002) reviewed the discussion of informal 
groups within the Melitidae. They found a “ Maera- 
Elasmopus group” and a “ Melita-Eriopisa ” group 
recognized by Bousfield (1977) and a “ Nuuanu group” 
recognized by McKinney & Barnard (1977). Barnard & 
Barnard (1983) recognized a Ceradocus group (. Maera- 
Elasmopus group of Bousfield, 1977), a Ceradocopsis 
group, an Eriopisa group, a Nuuana group ( Nuuanu group of 
McKinney & Barnard, 1977), a Melita group ( Melita-Eriopisa 
group of Bousfield, 1977) and a Parapherusa group. 

The Melitidae as conceived by Bousfield (1973) is not 
well defined and we can find no synapomorphy to define 
the whole group. In fact there appears to be very few 
widespread synapomorphies within the group. The 
extremely unequal rami of uropod 3 (the inner ramus is 
scale-like and the outer ramus is at least 3x longer than 
wide) appears to be a synapomorphy that defines a Melita 
and an Eriopisa group. The Eriopisa group has an extremely 
well-developed second article on the outer ramus of uropod 
3, another strong synapomorphy that separates this group 
from the Melita group. A third synapomorphy (two groups 
of small setae guarded by spines on the dorsum of urosomite 
2) occurs throughout the Melita group and the Nuuana 
group, but not in the Eriopisa group. 

The Ceradocopsis, Ceradocus and Parapherusa groups 
all appear to be paraphyletic assemblages at best, with no 
defining synapomorphies. In this paper, for practical 
reasons, we recognize a Ceradocopsis group, a Ceradocus 
group (including Parapherusa ), an Eriopisa group, a 
Nuuana group and a Melita group. 


Ceradocus group 
Austromaera n.gen. 

Type species. Maera mastersii Haswell, 1879a. 

Diagnosis. Head with anteroventral slit; eye ovate. Antenna 
1 accessory flagellum short, significantly less than half 
length of primary flagellum. Mandible palp article 1 not 
produced distally; article 3 short, tapering distally; article 2 
longer than article 3. Maxilla 1 inner plate with mainly apical 
setae. Gnathopod 1 coxa anteroventral corner produced, 
acute. Gnathopod 2 significantly enlarged in male and 
female; left and right gnathopods symmetrical in male (right 
slightly bigger than left); palm acute in male and female, 
male both propodi with well defined corner (greater than 
90°), female both propodi with well defined corner (greater 
than 90°); dactylus with 1 or 2 setae on anterior margin. 
Pereopods 5-7 dactyli simple. Epimeron 2 posteroventral 
corner with 1 small spine. Epimeron 3 posterior margin 
smooth. Uropod 3 rami about 2x peduncle, distally 
subacute, without apical robust setae; outer ramus 1- 
articulate. Telson deeply cleft, lobes apically subacute, with 
robust setae on inner margins, without robust setae on outer 
margins, without apical robust setae. 

Etymology. A combination of the Latin word auster, 
meaning southern, with the Latin stem Maera. 

Species composition .Austromaera mastersii (Haswell, 1879a). 

Remarks. Austromaera is part of the large Maera group, in 
which gnathopod 2 is symmetrical and significantly 
enlarged in males and females, the palms are acute with 
well defined corners and the rami of uropod 3 are distally 
acute or subacute. The main defining characters of the genus 
are: the weakly produced first article of the mandibular palp 
and the distally acute margins of the rami of the third 
uropods. Austromaera is the only member of the Maera 
complex with such strongly setose inner margins on the 
telson. It is excluded from Maera and Miramaera because 
of its ovate eyes, its short accessory flagellum and its weakly 
produced first mandibular palp article. It is excluded from 
Maera and Lupimaera because of its sparsely setose 
gnathopod 2 dactylus. Austromaera occurs in the Maera 
group, but appears to be most similar to the Indian Ocean 
species of Zygomaera that apparently have symmetrical 
second gnathopods. The main difference between these taxa 
is the telson that is emarginate in the Indian Ocean species. 
Currently Austromaera is confined to Australian waters. 

Distribution. Australia. 

Austromaera mastersii (Haswell, 1879a) 

Ligs. 1-4 

Megamaera mastersii Haswell, 1879a: 265, pi. 11, fig. 1.-Haswell, 
1882: 258.-Haswell, 1885: 105. 

Maera mastersii- Stebbing, 1899: 426.-Stebbing, 1906: 439.- 
Stebbing, 1910a: 642.-?Chilton, 1921b: 72 (southern 
Australia). 

Not Moeramastersii- Chilton, 1911: 564 (KermadecIslands) (=L. 
tias Krapp-Schickel, 2003, according to Krapp-Schickel, 
2003). 

Not Maera mastersi- Thomson, 1882: 235 (in part, part = M. 
quadrimana ) (New Zealand).-Sheard, 1936: 177, fig. 3 (South 
Australia) (=L. tias Krapp-Schickel, 2003, according to Krapp- 
Schickel, 2003).-Sheard, 1937: 24 (South Australia) (=L. tias 
Krapp-Schickel, 2003, according to Krapp-Schickel, 2003).- 


242 Records of the Australian Museum (2005) Vol. 57 


Fig. 1 . Austromaera mastersii (Haswell, 1879a), syntype, 9, AM P3487. Port Jackson, New South Wales, Australia. 
Scales for MDp and MX1IP represent 0.1 mm, remainder represent 0.2 mm. 


Sivaprakasam, 1969: 36, fig. 1 a-g [=L. mannarensis 
(Sivaprakasam, 1970), according to Krapp-Schickel, 2003].- 
J.L. Barnard, 1972a: 226, fig. 132 (southern Australia) (=L. 
leo Krapp-Schickel, 2003, according to Krapp-Schickel, 2003).- 
J.L. Barnard, 1972b: 109, figs 55-56 (New Zealand) (=L tias 
Krapp-Schickel, 2003, according to Krapp-Schickel, 2003).- 
Ledoyer, 1979: 77, fig. 44 [=Zygomaera pseudemarginata 
(Ledoyer, 1982) (Madagascar)].-Lowry & Fenwick, 1983: 236 
(New Zealand subantarctic) (=?L. tias Krapp-Schickel, 2003, 


according to Krapp-Schickel, 2003). 

Not Maera mastersii- Chevreux, 1908: 481 (French Polynesia).- 
Stebbing, 1910b: 457 (South Africa).-Chilton, 1916: 367 (New 
Zealand)-Chilton, 1925:317 (Chatham Islands, NewZealand).- 
Hale, 1929: 215, fig. 213 (figure = L. thomsoni of Miers, 1884) 
(South Australia) (=L. tias Krapp-Schickel, 2003, according to 
Krapp-Schickel, 2003).-Hurley, 1954: 603 (New Zealand) (=L. 
tias Krapp-Schickel, 2003, according to Krapp-Schickel, 2003).- 
Griffiths, 1974b: 291 (South Africa) .-J.L. Barnard, 1962:99 (key). 


Lowry & Springthorpe: Australian melitid amphipods 243 


Type material. Syntype, female, ovigerous, AM P3487, 
Port Jackson, New South Wales, Australia, [approx. 
33°50.9'S 151°16.2'E], [from AM Old Collection]. 

Additional material examined. New South Wales: male, 11.8 mm, AM 
P60389; female, 12.5 mm, AM P27286, Bottle and Glass Rocks, Port 
Jackson, 33°50.9'S 151°16.2'E, in and among dense tubes on sediment 
covered rocks, G.D. Fenwick, 29 Aug 1977. Id, AM P60390; 5 
specimens, AM P60391, off Wy-ar-gine Point, Port Jackson, [approx. 
33°49'S 151°15.rE], sand and shell, dredge, Malacological Society, 8 
May 1971. 1 specimen, AM P60392, northeast of Marys Rock, Cook 
Island, 28°11.42'S 153°34.79'E, orange bryozoan, 19 m, R.T. 
Springthorpe, 8 June 1993, stn NSW-816. 

Type locality. Port Jackson, New South Wales, Australia, 
[approx. 33°50.9'S 151°16.2'E]. 

Description. Based on syntype female, AM P3487, male, 
AM P60389 and female AM P27286. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral slit, anteroventral corner subquadrate, with 
acute/subacute spine. Antenna 1 longer than antenna 2; 
peduncular article 1 subequal in length to article 2, with 1 
distal robust seta on posterior margin; flagellum with 34 
articles; accessory flagellum with 6 articles. Antenna 2 
peduncular article 2 cone gland reaching at least to end of 
peduncular article 3; article 4 longer or subequal to article 
5; flagellum with 20 articles Mandible palp article 3 
rectolinear, setose along straight medial margin, longer than 
article 1; article 2 longer than article 3; article 1 not 
produced, shorter than article 2, about twice as long as broad. 
Maxilla 1 inner plate with about 3 setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
acute, posteroventral corner notch absent; merus without 
posterodistal spine; palm extremely acute, slightly convex, 


without posterodistal comer, defined by posterodistal robust 
setae. Gnathopod 2 not sexually dimorphic; subchelate; 
coxa postero ventral corner notch absent; merus with sharp 
postero ventral spine; carpus compressed; propodus without 
medial depression, palm acute, straight, sculptured, with 
sparse robust setae, defined by posterodistal robust setae, 
defined by postero ventral corner; apically acute/subacute. 
Pereopod 5 basis posterior margin straight, posteroventral 
corner broadly rounded. Pereopod 6-7 basis posterior 
margin convex, posteroventral corner broadly rounded. 

Pleon. Epimeron 1 posteroventral corner broadly rounded, 
or with small acute spine. Epimera 1-2 posteroventral 
margin without spines above posteroventral corner. 
Epimeron 3 posterior margin smooth, posteroventral corner 
with small acute spine. Uropod 3 inner ramus subequal in 
length to outer ramus; outer ramus longer (1.2 to 2x length) 
than peduncle, 1-articulate. Telson with robust setae on inner 
margins (1 or 2 per lobe), without apical robust setae, apical 
conical extension absent. 

Habitat. Marine; littoral; in and among dense tubes on 
sediment covered rocks, bryozoans, sand and shell, 7 to 19 
m depth. 

Remarks. Although there are many records of A. mastersii 
in the literature from inside and outside Australia, only 
Haswell (1879a: 265, pi. 11, fig. 1), Sheard(1936: 177, fig. 
3), J.L. Barnard (1972a: 226, fig. 132), J.L. Barnard, 1972b: 
109, figs 55-56 and Ledoyer (1979: 77, fig. 44) illustrated 
their material. Neither Sheard (1936), nor J.L. Barnard 
(1972a,b) showed the distinctive third uropods and telson 
illustrated by Haswell (1879a). Their misidentified species 
have recently been studied by Krapp-Schickel (2003). The 


244 Records of the Australian Museum (2005) Vol. 57 


Fig. 3. Austromaera mastersii (Haswell, 1879a), male, 11.8 mm, AM P60389. Bottle and Glass Rocks, Port Jackson, 
New South Wales, Australia. Scale for MX IIP represents 0.1 mm, remainder represent 0.2 mm. 


material of Ledoyer (1979) was later shown to be a different 
species, now known as Zygomaera pseudemarginata 
(Ledoyer, 1982). Based on currently available information, 
A. mastersii appears to be confined to southeastern and 
possibly southern Australia. Other records attributed to this 
species (at least since Haswell [1885] synonymized L. 
thomsoni with A. mastersii and certainly since the 
illustrations of Sheard [1936]) represent Linguimaera leo 
and L. tias of Krapp-Schickel (2003) plus several unknown 
species. 

Distribution. New South Wales : Bottle and Glass Rocks 
and Wy-ar-gine Point, Port Jackson; Marys Rock, Cook 
Island (all AM). 

Australian geographic areas. Southeastern Australia. 


Ceradocus Costa, 1853 
Ceradocus circe n.sp. 

Figs. 5-7 

Ceradocus rubromaculatu s.-Chilton, 1921b: 71, fig. 9a-c. 

Type material. Holotype 8 “a”, 24.3 mm, AM P60564; 1 paratype, 
$ “a”, 18.6 mm, AM P60565; 1 paratype, 8 “b”,22.9mm,AMP5915; 
8 Paratypes, AM E6543, 16 kms north of Circular Head, Tasmania, 
Australia, [approx. 40°46'S 145°18'E], FIS Endeavour , 1909-1914. 1 
paratype, female, AM E6542, Tasmanian Coast, Australia, FIS 
Endeavour , 1909-1914 

Type locality. 16 kms north of Circular Head, Tasmania, 
Australia, [approx. 40°46'S 145°18'E]. 

Description. Based on holotype 6 “a”, AM P60564, paratype 
$ “a”, AM P60565, and paratype 6 “b”, AM P5915. 


Lowry & Springthorpe: Australian melitid amphipods 245 


Fig. 4. Austromaera mastersii (Haswell, 1879a), male, 11.8 mm, AM P60389. Bottle and Glass Rocks, Port Jackson, 
New South Wales, Australia. Scales represent 0.2 mm. 


Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner with acute/ 
subacute spine. Antenna 1 longer than antenna 2; peduncular 
article 1 subequal in length to article 2, with 4 or more robust 


setae along posterior margin; flagellum with at least 30 
articles; accessory flagellum with 11 articles. Antenna 2 
peduncular article 2 cone gland not reaching to end of 
peduncular article 3; article 4 subequal to article 5; flagellum 


246 Records of the Australian Museum (2005) Vol. 57 


with about 20 articles. Mandible palp article 3 cone-like, 
setose on distomedial margin, subequal to article 1; article 
2 longer than article 3; article 1 produced distally, shorter 
than article 2, about as long as broad. Maxilla 1 inner plate 
setose along entire inner margin. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
acute, posteroventral corner notch absent; merus with sharp 
posterodistal spine; propodus palm acute, straight, defined 
by posterodistal corner, defined by posterodistal robust 
setae. Gnathopod 2 sexually dimorphic; subchelate; coxa 
posteroventral corner notch absent; merus with sharp 
posteroventral spine; carpus compressed; propodus without 
medial depression, without strong concentration of setae, 
palm acute, straight, sculptured, with group of anterodistal 
robust setae, without posterodistal robust setae, defined by 
posteroventral spine; apically acute/subacute. Pereopod 5 
basis posterior margin straight, posteroventral comer narrowly 
rounded or subquadrate. Pereopod 6 coxa anterior lobe ventral 
margin slightly produced, rounded; basis posterior margin 
straight, posteroventral comer with acute or subacute process. 
Pereopod 7 basis posterior margin straight, posteroventral 
comer with acute or subacute process. 

Pleon. Pleonites 1-3 with dorsal serrations. Epimeron 1 
posteroventral corner with small acute spine. Epimera 1-2 
posteroventral margin with 3 large or small spines above 
spine defining posteroventral corner, or without spines 
above posteroventral corner. Epimeron 3 posterior margin 
serrate, posteroventral corner with strongly produced acute 
spine, posteroventral margin serrate. Urosomite 1 with 
dorsal serrations, with spines at midline, no 
conspicuous medial gape. Urosomite 2 
posterior margin serrate. Uropod 3 inner 
ramus subequal in length to outer 
ramus; outer ramus much longer 
(more than 2x length) than 
peduncle; 1-articulate. Telson 
each lobe with 3 or more 
apical/subapical robust 


setae, apical conical extension reaching at least halfway 
along longest seta. 

Female (sexually dimorphic characters). Gnathopod 2 
convex, smooth, without robust setae, defined by 
posterodistal robust setae, defined by posterodistal spine. 

Habitat. Marine; continental shelf. 

Etymology. Named for the wooden schooner Circe , which 
went ashore in 1892, between Entrance Island and the North 
Spit, while attempting to pass through Hells Gates and enter 
Macquarie Harbour, Tasmania. 

Remarks. Among Australian species of Ceradocus, C. circe 
occurs in the group with sparse serrations along the posterior 
margins of epimera 1 and 2. Within this group C. circe 
appears to be the only species with the rami of uropod 3 
more than twice as long as the peduncle (not known for C. 
yandala Berents, 1983 from tropical northeastern Australia). 

Ceradocus orchestiipes A. Costa, 1853 from the 
Mediterranean Sea, has sparse serrations along the posterior 
margins of epimera 1 and 2, long rami on uropod 3 and the 
second gnathopods in the male and the female are very 
similar to those of C. circe. But C. orchestiipes does not 
have dorsal serrations on urosomites 1-3, nor does it have 
a strongly serrate posterodistal corner on epimeron 3. 

Distribution. Tasmania : north of Circular Head; eastern 
slope of Bass Strait (Chilton, 1921b). 

Australian geographic areas. Southeastern Australia. 


Fig. 5. Ceradocus circe n.sp., paratype, male “b”, 22.9 mm, AM 
P5915, 16 kms north of Circular Head, Tasmania, Australia. 


Lowry & Springthorpe: Australian melitid amphipods 247 


Fig. 6. Ceradocus circe n.sp., holotype, male “a”, 24.3 mm, AM P60564, 16 kms north of Circular Head, Tasmania, 
Australia. Scales for Al, A2, Ul, U3, T represent 0.5 mm, remainder represent 0.2 mm. 


Linguimaera Pirlot 

Linguimaera Pirlot, 1936: 309.-Krapp-Schickel, 2003: 258. 

Type species. Linguimaera pirloti Krapp-Schickel, 2003, 
replacement name for misidentified type species. 

Diagnosis. Head with anteroventral slit; eye ovate to 
reniform. Antenna 1 accessory flagellum short, significantly 
less than half length of primary flagellum. Mandible palp 
article 1 not produced distally; article 2 longer than article 
3; article 3 long, rectolinear. Maxilla 1 inner plate with 
mainly apical setae. Gnathopod 1 coxa anteroventral corner 
produced, acute or subacute. Gnathopod 2 significantly 
enlarged in male, not in female; left and right gnathopods 
asymmetrical in male; palm acute in male and female, male 
one propodus with well defined corner, the other propodus 
with poorly defined or no corner, female with poorly defined 
corner or no corner in either propodus; dactylus with 1 or 2 


setae on anterior margin. Pereopods 5-7 dactyli simple. 
Epimeron 3 posterior margin serrate. Uropod 3 rami about 
2x peduncle, distally truncated, apical robust setae short; 
outer ramus 1-articulate. Telson emarginate, without robust 
setae on outer margins, with apical robust setae, apical robust 
setae long. 

Species composition. Linguimaera boecki (Haswell, 
1879b); L. bogombogo Krapp-Schickel, 2003; L. caesaris 
Krapp-Schickel, 2003; L. eugeniae Schellenberg, 1931; L. 
garitima Krapp-Schickel, 2003; L. hamigera (Haswell, 
1879b); L. kellissa Krapp-Schickel, 2003; L. leo Krapp- 
Schickel, 2003; L. mannarensis Sivaprakasam, 1970; L. 
octodens (Sivaprakasam, 1968); L. pirloti Krapp-Schickel, 
2003; L. schickelae n.sp.; L. thomsoni (Miers, 1884); L. tias 
Krapp-Schickel, 2003. 

Distribution. Indo-Pacific and Mediterranean. 


248 Records of the Australian Museum (2005) Vol. 57 


Linguimaera boecki (Haswell) 

Figs. 8-10 

Megamoera boeckii Haswell, 1879b: 336, pi. 21, fig. 6. 

IMaera boeckii.-Della. Valle, 1893: 732. 

Elasmopusboeckii- Stebbing, 1899: 426-Stebbing, 1906: 445.- 
Stebbing, 1910a: 643. 

Maera boecki- Barnard & Barnard, 1983: 623. 

Not Elasmopus boeckii- K.H. Barnard, 1916: 199: pi. 27, figs 
13-14. 

Type material. Neotype: male, 7.5 mm, AM P60619, off Sow and Pigs 
Reef, Port Jackson, New South Wales, 33°50.3'S 151°16.2’E, shelley sand, 
5 m, benthic grab, J.K. Lowry & A.R. Jones, 30 Sep. 1976, stn NSW-184. 


Type locality. Sow and Pigs Reef, Port Jackson, New South 
Wales (33°50.3’S 151°16.2'E), shelley sand, 5 m depth. 

Description. Based on neotype male, AM P60619 and 
female, AM P60947. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner subquadrate. 
Antenna 1 longer than antenna 2; peduncular article 1 
subequal in length to article 2, with 2 robust setae along 
posterior margin; flagellum with 20 articles; accessory 
flagellum with 4 articles. Antenna 2 peduncular article 2 
cone gland not reaching to end of peduncular article 3; 
article 4 longer than article 5; flagellum with 10 articles. 


Lowry & Springthorpe: Australian melitid amphipods 249 


Mandible palp article 3 rectolinear, setose on distomedial 
margin, longer than article 1; article 2 longer than article 3; 
article 1 not produced, shorter than article 2, about twice as 
long as broad. Maxilla 1 inner plate with 3 setae mainly 
terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
acute, posteroventral corner notch absent; merus without 
posterodistal spine; propodus palm acute, straight, without 
posterodistal corner, defined by posterodistal robust setae. 
Gnathopod 2 sexually dimorphic; left and right gnathopods 
unequal in size, subchelate; coxa posteroventral corner notch 
absent; (larger) merus with sharp posteroventral spine; 
carpus compressed; propodus palm angle acute, straight, 
defined by posterodistal spine, with robust setae; dactylus 
apically blunt; (smaller) merus with sharp posteroventral 
spine; carpus long; propodus palm straight, without 
posteroventral spine. Pereopod 5 basis posterior margin 
straight, posteroventral corner narrowly rounded or 
subquadrate. Pereopod 6 coxa anterior lobe ventral margin 
slightly produced, rounded; basis posterior margin straight, 
posteroventral corner narrowly rounded or subquadrate. 
Pereopod 7 basis posterior margin straight, with posterior 
margin smooth or minutely castelloserrate, posteroventral 
corner narrowly rounded or subquadrate. 

Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin with 1 to 2 large 
or small spines above spine defining posteroventral corner. 
Epimeron 3 posterior margin minutely serrate, postero¬ 
ventral corner with small acute spine. Urosomites 1-3 
dorsally smooth. Uropod 3 inner ramus subequal in length 
to outer ramus; outer ramus longer (1.2 to 2x length) than 
peduncle, 1-articulate. Telson deeply emarginate, each lobe 
with 1 apical/subapical robust seta, apical conical extension 
reaching scarcely one third along longest seta. 

Female (sexually dimorphic characters). Gnathopod 2 
carpus long; propodus without medial depression, setose, 
palm acute, straight or slightly concave, smooth, lined with 


robust setae, defined by posterodistal robust setae, defined 
by posteroventral corner; dactylus apically acute/subacute. 

Habitat. Marine; littoral, shelley sand, 5 m depth. 

Remarks. Linguimaera boecki appears to be most similar 
to L. hamigera. They differ from other members of the genus 
in having rather short rami on uropod 3 and L. boecki is the 
only species of Linguimaera with an emarginate telson. 

Distribution. New South Wales : Port Jackson (Haswell, 
1879b; AM). 

Australian geographic areas. Southeastern Australia. 

Linguimaera hamigera (Haswell) 

Figs. 11-13 

Moera hamigera Haswell, 1879b: 333, pi. 21, fig. 1. 

Maera hamigera-Stebbing, 1888: 1790-Della Valle, 1893:723- 
Stebbing 1906: 437.-Barnard & Barnard, 1983: 623. 

Not Maera hamigera- Walker 1909: 335, pi. 43, fig. 5, pi. 3- 
Stebbing 1910a: 600.-K.H. Barnard, 1916: 196, pi. 27, figs 
11-12.-Chilton, 1921b: 73.-J.L. Barnard, 1965: 507, fig. 16 
(in part).-Karaman & Ruffo, 1971: 152, figs 21-23.-Ledoyer, 
1982: 523, figs 196-197.-Karaman, 1982: 312, fig. 211.- 
Lyons & Myers, 1993: 587, fig. lO.-Myers, 1997: 109. 

Type material. Syntype, 6, AM P3477, Port Jackson, New South 
Wales, Australia, [approx. 33°51'S 151°16'E], [AM Old Collection]. 

Type locality. Port Jackson, New South Wales, Australia, 
(33°51’S 151°16’E). 

Additional material examined. New South Wales: 1 specimen, 
AM P57328, 100 m north west of Split Solitary Island, 30°14.0'S 
153°10.8'E, sponge, 15-17 m, R.T. Springthorpe, 7 March 1992, stn 
NSW-683. 1 specimen, AM P57329, 50 m west of Split Solitary Island, 
30°14.0’S 153°10.8’E, rocks with brown & red algae (coralline & 
Halimeda sp.), 15-17 m, P. Hutchings & C. Rose, 7 March 1992, stn 
NSW-692. 1 specimen, AM P56712, Coffs Harbour Jetty, Coffs Harbour, 
30°18.4'S 153°08.5'E, arborescent sponge on jetty pilings, 7 m, S.J. 
Keable, 9 March 1992, stn NSW-735. 1 specimen, AM P25467, 5.5-6.5 


250 Records of the Australian Museum (2005) Vol. 57 


km off Wattamolla, 34°10'S 151°11'E, mud, 99-108 m, E.R. Waite on 
HMCS Thetis, 22 March 1898, stn 57. Id, AM P60608, Munganno 
Point, Twofold Bay, 37°06.2'S 149°55.7'E, subtidal rock platform, wharf 
pile, S J. Keable & E.A. Bamber, 12 December 1984, stn M2,3. 1 $, AM 
P60609, same locality. Tasmania: 1 specimen, E6546, eastern slope of 
Bass Strait, [approx. 39°00'S 148°40'E], FIS Endeavour , 1909-1914. 1 
specimen, AM P25468, same locality. 

Description. Based on syntype, <3, AM P3477, 3, AM 
P60608, and female, AM P60609. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner subquadrate. 
Antenna 1 longer than antenna 2; peduncular article 1 
slightly longer than or subequal in length to article 2, with 
1 robust seta on posterior distal margin; flagellum with 26 
articles; accessory flagellum with 4 articles. Antenna 2 
peduncular article 2 cone gland not reaching to end of 
peduncular article 3; article 4 slightly longer than article 5; 
flagellum with 9 articles. Mandible palp article 3 rectolinear, 
setose along straight medial margin, longer than article 1; 
article 2 longer than article 3; article 1 not produced, shorter 
than article 2, about as long as broad. Maxilla 1 inner plate 
with setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
acute, posteroventral corner notch absent; merus without 
posterodistal spine; propodus palm acute, slightly convex, 


defined by posterodistal corner, defined by posterodistal 
robust setae. Gnathopod 2 sexually dimorphic; subchelate; 
left and right gnathopods unequal in size; coxa postero- 
ventral corner notch absent; (larger) merus with subquadrate 
postero ventral corner; carpus compressed; propodus palm 
angle nearly transverse, concave, defined by posterodistal 
spine, with robust setae; dactylus apically blunt; (smaller) 
merus with sharp posteroventral spine; carpus short, or long; 
propodus palm straight, without posteroventral spine. Pereopod 
5 basis posterior margin slightly convex, posteroventral comer 
broadly rounded. Pereopod 6 coxa anterior lobe ventral margin 
slightly produced, rounded, or not produced ventrally; basis 
posterior margin straight, posteroventral corner broadly 
rounded. Pereopod 7 basis posterior margin slightly convex, 
with posterior margin smooth or minutely castelloserrate, 
posteroventral comer broadly rounded. 

Pleon. Epimeron 1 posteroventral corner broadly rounded. 
Epimera 1-2 posteroventral margin without spines above 
posteroventral corner. Epimeron 3 posterior margin smooth 
or minutely serrate, posteroventral corner with strongly 
produced acute spine. Uropod 3 inner ramus subequal in 
length to outer ramus; outer ramus longer (1.2 to 2x length) 
than peduncle, 1-articulate. Telson each lobe with 3 or more 
apical/subapical robust setae, apical conical extension 
reaching scarcely one third along longest seta. 


Lowry & Springthorpe: Australian melitid amphipods 251 


Female (sexually dimorphic characters). Gnathopod 2 robust setae, defined by posteroventral corner; dactylus 
merus with sharp posteroventral spine; carpus short, or long; apically acute/subacute. 

propodus without medial depression, palm acute, straight, Habitat. Marine; littoral; subtidal rock platforms, 0 to 7 m 
sculptured, with sparse robust setae, defined by posterodistal depth. 


252 Records of the Australian Museum (2005) Vol. 57 


Remarks. Since Haswell (1879b) described Linguimaera 
hamigera (as Maera hamigera ) from Port Jackson, only 
Stebbing (1910a) has reported it from Australian waters— 
several specimens of about 5 mm length from off 
Wattamolla, which he unfortunately did not illustrate. 
However, it has been reported by Walker (1909) from the 
Red Sea, K.H. Barnard (1916) from Southern Africa, J.L. 
Barnard (1965) from Micronesia, Karaman & Ruffo (1971) 
from the Mediterranean Sea, Myers (1997) from Western 
Samoa and Ledoyer (1982) from Madagascar. Ledoyer 
(1982) completely illustrated and described his material. 
Based on this species concept he suggested that the Maera 
sp. A of J.L. Barnard, 1970, is also L. hamigera , extending 
its distribution to Hawaii and that L. mannarensis 
(Sivaprakasam, 1970) is a synonym of L. hamigera , thus 
extending its distribution into India, a proposition not 
accepted by Krapp-Schickel (2003). 

All reports subsequent to Stebbing (1910a) must be 
considered as erroneous. As currently known, A. hamigera 
is confined to southeastern Australia. Illustrated species in 
the literature (Walker, 1909; K.H. Barnard, 1916; J.L. 
Barnard, 1965; Karaman & Ruffo, 1971; Ledoyer, 1982) 
refer to one or more unnamed species. For instance Karaman 
& Ruffo (1971) illustrated a cleft telson without setae on 
the inner margins, whereas Ledoyer’s (1982) specimens 
have well-developed robust setae along the inner margins. 

Linguimaera hamigera is most similar to L. boecki, but 
easily distinguished from that species by its cleft telson. 

Distribution. New South Wales : Coffs Harbour (AM); Port 


Jackson (Haswell, 1879b); Munganno Point, Twofold Bay 
(AM). Tasmania : eastern slope of Bass Strait. 

Australian geographic areas. Southeastern Australia. 

Linguimaera schickelae n.sp. 

Figs. 14-16 

Type material. Holotype, 6 , 15.4 mm, AM P60620, Little 
Bay, New South Wales, Australia, [approx. 33°59'S 
151°15'E], under stones between tide marks, G.P. Whitley, 
18 April 1924. 2 paratypes, AM P60618; 1 paratype $, 
14.1 mm, AM P60948; Clovelly Pool, Clovelly, New South 
Wales, Australia, [approx. 33°55'S 151°16'E], from under 
stones, 7 m, PC. Terrill, 12 June 1979. 

Type locality. Little Bay, New South Wales, Australia, 
[approx. 33°59'S 151°15’E]. 

Additional material examined. New South Wales: 1 
specimen, AM P62902, northern cove of Boondelbah Island, 
Port Stephens, 32°42.28'S 152°13.47'E, airlift under small 
boulders, 19.6 m, R.T. Springthorpe, P.B. Berents & A. 
Murray, 28 May 1998, stn NSW-1401. 

Description. Based on holotype male, AM P60620 and 
paratype female, AM P60948. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner subquadrate, 
with acute/subacute spine. Antenna 1 longer than antenna 
2; peduncular article 1 subequal in length to article 2 or 
shorter than article 2, with 1 robust seta on posterior 
margin (distal); flagellum with about 26 articles; 
accessory flagellum with 7 articles. Antenna 2 
peduncular article 2 cone gland not reaching 
to end of peduncular article 3; article 4 
longer than article 5; flagellum with 
about 14 articles. Mandible palp 
article 3 rectolinear, setose 


Fig. 11. Linguimaera hamigera (Haswell, 
1879b), male, AM P60608, Munganno Point, 
Twofold Bay, New South Wales, Australia. 


Lowry & Springthorpe: Australian melitid amphipods 253 


Fig. 12. Linguimaera hamigera (Haswell, 1879b), male, AM P60608, Munganno Point, Twofold Bay, New South 
Wales, Australia. Scales for MDp, MX IIP represent 0.1 mm, remainder represent 0.2 mm. 


along straight medial margin, longer than article 1; article 
2 longer than article 3; article 1 not produced, shorter than 
article 2, about as long as broad. Maxilla 1 inner plate with 
setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
acute, posteroventral comer notch present; merus with sharp 
posterodistal spine; carpus about 3x as long as broad; palm 
acute, straight, defined by posterodistal corner, defined by 
posterodistal robust setae. Gnathopod 2 sexually dimorphic; 
left and right gnathopods unequal in size; coxa postero¬ 
ventral corner notch absent; (larger) subchelate; mems with 
sharp posteroventral spine; carpus short; palm angle acute, 
concave, defined by posterodistal spine, with robust setae; 
dactylus apically acute; (smaller) subchelate; mems with 
sharp posteroventral spine; carpus short; palm slightly 
concave, defined by posteroventral spine. Pereopod 5 basis 
posterior margin slightly concave or straight, posteroventral 
corner with acute or subacute process. Pereopod 6 coxa 
anterior lobe ventral margin slightly produced, rounded; 


basis posterior margin slightly concave, basis posteroventral 
corner narrowly rounded or subquadrate. Pereopod 7 basis 
posterior margin straight, posteroventral corner narrowly 
rounded or subquadrate. 

Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin with 1 to 2 large 
or small spines above spine defining posteroventral corner. 
Epimeron 3 posterior margin serrate, posteroventral corner 
with small acute spine. Uropod 3 inner ramus subequal in 
length to outer ramus; outer ramus longer (1.2 to 2x length) 
than peduncle; 1-articulate. Telson with robust setae on inner 
margins (about 4 per lobe), each lobe with 2 apical/subapical 
robust setae, apical conical extension reaching scarcely one 
third along longest seta. 

Female (sexually dimorphic characters). Gnathopod 2 
propodus palm straight, sculptured, lined with robust setae. 

Etymology. Named for Traudl Krapp-Schickel, who has 
contributed greatly to the revision of the Maera complex. 


254 Records of the Australian Museum (2005) Vol. 57 


Fig. 13. Linguimaera hamigera (Haswell, 1879b), male, AM P60608, female, AM P60609 Munganno Point, 
Twofold Bay, New South Wales, Australia. Scales represent 0.2 mm. 


Fig. 14. Linguimaera schickelae n.sp., holotype, 
male, 15.4 mm, AM P60620, Little Bay, New South 
Wales, Australia. 


Lowry & Springthorpe: Australian melitid amphipods 255 


Fig. 15. Linguimaera schickelae n.sp., holotype, male, 15.4 mm, AM P60620, Little Bay, New South Wales, 
Australia; scales for MDp, MX IIP, MX2 represent 0.2 mm, remainder represent 0.5 mm. 


256 Records of the Australian Museum (2005) Vol. 57 


Fig. 16. Linguimaera schickelae n.sp., holotype, male, 15.4 mm, AM P60620, paratype, female, AM P60948, 
Little Bay, New South Wales, Australia. Scales represent 0.5 mm. 


Habitat. Marine; littoral; under stones between tide marks 
and from 7 m depth. 

Remarks. Linguimaera schickelae appears to be most 
similar to L. mannarensis (Sivaprakasam, 1970) and L. tias 
Krapp-Schickel, 2003. They all have the short tapering third 
article of the mandibular palp and the well defined corner 


of gnathopod 2 palm that is greater than 90°. Linguimaera 
schickelae and L. mannarensis are the only species in which 
the posterodistal comer of epimeron 2 has three small spines. 

Distribution. New South Wales: Boondelbah Island, Port 
Stephens; Little Bay; Clovelly (all AM). 

Australian geographic areas. Southeastern Australia. 


Lowry & Springthorpe: Australian melitid amphipods 257 


Linguimaera thomsoni (Miers, 1884) 

Megamaera thomsoni Miers, 1884: 318, pi. 34, fig. B. 
Maera mastersii Haswell, 1885: 105 (in part). 

Type material. Apparently lost. 

Type locality. Torres Strait. 

Description. Based on Miers, 1884: 318, pi. 34, fig. B. 


Remarks. Linguimaera thomsoni is a poorly described 
species. Krapp-Schickel (2003) was able to separate L. 
thomsoni from L. pirloti by the first coxa that is 
anteroventrally rounded (acute in L. pirloti ) and the palm 
of gnathopod 2 that has a large excavation in L. thomsoni 
(two smaller excavations in L. pirloti ). 

Distribution. Queensland : Albany Island, Prince of Wales 
Channel, Thursday Island, Torres Strait (Miers, 1884). 


Head. Lateral cephalic lobes broad, rounded. Antenna 1 
longer than antenna 2; peduncular article 1 shorter than 
article 2, with 1 robust seta on posterior margin; flagellum 
with 30+ articles. Article 4 subequal to article 5; antenna 2 
flagellum with 10+ articles. 


Australian geographic areas. Northeastern Australia. 

Mallacoota J.L. Barnard, 1972 
Mallacoota chandaniae n.sp. 


Pereon. Gnathopod 1 coxa anteroventral corner produced, 
rounded; merus with sharp posterodistal spine; carpus about 
3x as long as broad; palm extremely acute, concave, without 
posterodistal corner. Gnathopod 2 subchelate; merus with 
sharp posteroventral spine; carpus long; propodus without 
medial depression, setose, palm acute, convex, sculptured, 
with sparse robust setae, defined by posteroventral spine; 
apically acute/subacute. Pereopod 5-7 basis posterior 
margin concave or straight, posteroventral corner narrowly 
rounded or subquadrate. 


Figs. 17-19 

Type material. Holotype, 3 “a”, 11 mm, AM P59021; 1 paratype, 
3 “b”, 9.1 mm, AM P62996, at end of sugar loading jetty 5 km long, 
Lucinda, Queensland, Australia, [approx. 18°31'S 146°19'E], pylon 
scrapings, 7 m, Frank Hoedt, CRIMP survey, August 1999, stn A138. 5 
paratypes, 3 “c”, 9.5 mm, 9 “a” 10.8 mm, 3 9$, AM P59020, Evans 
Landing, Weipa, Queensland, Australia, [approx. 12°35'S 141°36'E], 
pylon scrapings, 0.5 m, Frank Hoedt, CRIMP survey, October 1999, stn 
A220. 

Type locality. At end of sugar loading jetty 5 km long, 
Lucinda, Queensland, Australia, [approx. 18°31'S 
146°19'E], pylon scrapings. 


Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin with 1 to 2 large 
or small spines above spine defining posteroventral corner. 
Epimeron 3 posterior margin serrate, posteroventral corner 
with small acute spine. Uropod 3 rami distally acute/ 
subacute; inner ramus subequal in length to outer ramus; 
outer ramus longer (1.2 to 2x length) than peduncle, 1- 
articulate. Telson with robust setae on inner margins, each 
lobe with 1 apical/subapical robust seta, apical 
conical extension absent. 

Habitat. Marine; 7 to 16 m depth. 


Description. Based on holotype male, AM P59021, paratype 
male, AM P62996 and paratype female, AM P59020. 

Head. Lateral cephalic lobes broad, truncated, with 
anteroventral notch or slit, anteroventral corner rounded. 
Antenna 1 longer than antenna 2; peduncular article 1 
subequal in length to article 2, with 4 
or more robust setae along post¬ 
erior margin; flagellum 


Fig. 17. Mallacoota chandaniae n.sp., holo¬ 
type, male “a”, 11 mm, AM P59021, Lucinda, 
Queensland, Australia. 


258 Records of the Australian Museum (2005) Vol. 57 


with about 26 articles; accessory flagellum with 4 articles. 
Antenna 2 peduncular article 2 cone gland not reaching to 
end of peduncular article 3; article 4 longer than article 5; 
flagellum with about 11 articles. Mandible palp article 3 
rectolinear, with setae mostly terminal, longer than article 
1; article 2 subequal to article 3; article 1 not produced, 
shorter than article 2, about twice as long as broad. Maxilla 
1 inner plate with 2 terminal setae. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
slightly rounded, posteroventral corner notch absent; merus 
without posterodistal spine; palm acute, convex, without 
posterodistal corner, defined by posterodistal robust setae. 


Gnathopod 2 sexually dimorphic; subchelate; coxa 
posteroventral corner notch absent; merus with subquadrate 
posteroventral comer; carpus compressed; propodus without 
medial depression, with strong setal bunch, palm slightly 
acute, sinusoidal, sculptured, with group of anterodistal 
robust setae, without posterodistal robust setae, defined by 
posteroventral spine; dactylus apically blunt. Pereopod 5 
basis posterior margin straight, posteroventral corner 
broadly rounded. Pereopod 6 coxa anterior lobe ventral 
margin slightly produced, rounded; basis posterior margin 
straight, posteroventral corner broadly rounded. Pereopod 
7 basis posterior margin straight, posteroventral corner 
narrowly rounded or subquadrate. 


Lowry & Springthorpe: Australian melitid amphipods 259 


Fig. 19. Mallacoota chandaniae n.sp., holotype, male “a”, 11 mm, AM P59021, * paratype, male “b”, 9.1 mm, AM 
P62996, Lucinda, Queensland, Australia; paratype, female, “a” 10.8 mm, AM P59020, Weipa, Queensland, Australia. 


Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner subquadrate. Urosomite 1 
dorsally bicarinate. Uropod 3 rami distally truncated; inner 
ramus subequal in length to outer ramus; outer ramus longer 
(1.2 to 2x length) than peduncle, 1-articulate. Telson each lobe 


with 3 or more apical/subapical robust setae, apical conical 
extension reaching at least halfway along longest seta. 
Female (sexually dimorphic characters). Gnathopod 2 
merus with sharp posteroventral spine; carpus short; setose, 
convex, smooth, with sparse robust setae, defined by 
posterodistal robust setae, without posteroventral corner; 
dactylus apically acute/subacute. 


260 Records of the Australian Museum (2005) Vol. 57 


Habitat. Marine; littoral; living on encrusted wharf pilings. 

Etymology. Named for Chandani Appadoo, in recognition 
of her work on the melitid amphipods of the Indian Ocean. 

Remarks. Mallacoota chandaniae is a very distinctive 
species. For instance no Australian species has a strictly 
subquadrate epimeron 3 whereas seven species outside of 
Australia have this characteristic. Only two species within 
this group, M. schellenbergi Ledoyer, 1984 and the M. 
subcarinata of Myers, 1985 have a deeply cleft telson 
similar to M. chandaniae, but neither of these has the almost 
transverse palm of the male gnathopod 2. 

Distribution. Queensland : Weipa; Lucinda (both AM). 

Australian geographic areas. Northeastern Australia. 

Mallacoota euroka n.sp. 

Figs. 20-22 

Mallacoota subcarinata-J .L. Barnard, 1972a: 247, fig. 144.- 
Barnard & Barnard, 1983: 632 (in part). 

Type material. Holotype, A, 10.1 mm, AM P60561; 1 
paratype, $, AM P60562; 8 paratypes, AM P27034, 
between Troubridge Light and Cape Jervis, South Australia, 
[approx. 35°20'S 137°40'E], sponges on mud bottom, 20 
m, D. Blake & H. Larsen, 14 March 1978. 


Type locality. Between Troubridge Light and Cape Jervis, 
South Australia, [approx. 35°20'S 137°40'E], sponge on mud 
bottom, 20 m. 

Description. Based on holotype male, AM P60561 and 
paratype female, AM P60562. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner rounded. 
Antenna 1 longer than antenna 2; peduncular article 1 
subequal in length to article 2, with 4 or more robust setae 
along posterior margin; flagellum with about 29 articles; 
accessory flagellum with 3-4 articles. Antenna 2 peduncular 
article 2 cone gland reaching at least to end of peduncular 
article 3; article 4 longer than article 5, or article 4 subequal 
to article 5; flagellum with about 10 articles. Mandible palp 
article 3 rectolinear, with setae mostly terminal, longer than 
article 1; article 2 shorter than article 3; article 1 not 
produced, subequal to article 2, about twice as long as broad. 
Maxilla 1 inner plate with setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
rounded, posteroventral comer notch present; merus without 
posterodistal spine; propodus palm acute, convex, defined 
by posterodistal corner, without posterodistal robust setae. 
Gnathopod 2 sexually dimorphic; subchelate; coxa 
posteroventral corner notch present; merus with sharp 
posteroventral spine; carpus compressed; propodus without 
medial depression, with strong setal bunch, palm acute, 


Lowry & Springthorpe: Australian melitid amphipods 261 


Fig. 21. Mallacoota euroka n.sp., holotype, male, 10.1 mm, AM P60561, between Troubridge Light and Cape 
Jervis, South Australia. Scales for MDp, MX1IP represent 0.1 mm, scales for Al, A2 represent 0.5 mm, remainder 
represent 0.2 mm. 


straight, sculptured, with sparse robust setae and with group 
of anterodistal robust setae, without posterodistal robust 
setae, defined by posteroventral spine; dactylus apically 
blunt. Pereopod 5 basis posterior margin concave, postero¬ 
ventral corner narrowly rounded or subquadrate. Pereopod 
6 coxa anterior lobe ventral margin slightly produced, 
rounded; basis posterior margin concave, posteroventral 
corner narrowly rounded or subquadrate. Pereopod 7 basis 
posterior margin convex, with posterior margin smooth or 
minutely castelloserrate, posteroventral corner broadly 
rounded. 

Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner with strongly produced acute 
spine. Urosomite 1 dorsally bicarinate. Urosomite 2 posterior 
margin smooth. Uropod 3 inner ramus subequal in length 
to outer ramus; outer ramus longer (1.2 to 2x length) than 
peduncle, 1-articulate. Telson each lobe with 3 or more 
apical/subapical robust setae, apical conical extension 
reaching scarcely one third along longest seta. 

Female (sexually dimorphic characters). Gnathopod 2 
carpus short; setose, smooth, without robust setae, defined 


by posterodistal robust setae, defined by posteroventral 
corner; dactylus apically acute/subacute. 

Habitat. Marine; littoral; sponges on mud bottom, 20 m 
depth. 

Etymology. Named for the schooner Euroka, built in 
Brisbane Waters and sunk off the Sandon River mouth, south 
of Clarence Head, New South Wales, in 1875. 

Remarks. This species was originally described as 
Mallacoota subcarinata phenotype A by J.L. Barnard 
(1972a). It has a large posteroventral spine on epimeron 3, 
like M. subcarinata and M. diemenensis. Mallacoota euroka 
differs from M. diemenensis in having no dorsal spines on 
pleonites 1 to 3. It differs from M. subcarinata in the number 
of robust setae on the first peduncular article of antenna 1 
(five in M. euroka and one to three in M. subcarinata ), in having 
posteroventral notches on coxae 1 and 2 and in having a convex 
posterior margin on the basis of pereopod 7. 

Distribution. Victoria: Port Phillip (J.L. Barnard, 1972a). 
South Australia: Cape Jervis (AM). 

Australian geographic areas. Southern Australia. 


262 Records of the Australian Museum (2005) Vol. 57 


Fig. 22. Mallacoota euroka n.sp., holotype, male, 10.1 mm, AM P60561, paratype female, AM P60562, between 
Troubridge Light and Cape Jervis, South Australia. Scale represents 0.5 mm. 


Mallacoota kameruka n.sp. 

Figs. 23-25 

Elasmopus subcarinatus- Stebbing, 1888: 1019, pi. 98 (plates 
labelled as E. persetosus). 

Mallacoota subcarinata- J.L. Barnard, 1972a: 247, fig. 145-Bamard 
& Barnard, 1983: 632 (in part)-Hutchings et al., 1989: 362. 

Type material. Holotype, 8 “a”, 10.0 mm, AM P60491; 1 
paratype, $, 10.0 mm, AM P60492; 10 paratypes, 5 8 8, 5 $ $, AM 
P60493; 1 paratype, 8 “b”, AM P60494, northeast of Marys Rock, 
Cook Island, New South Wales, Australia, 28°11.42'S 153°34.79'E, 
orange bryozoan, 19 m, R.T. Springthorpe, 8 June 1993, stn NSW-816. 

Additional material examined. New South Wales: 41 8 8 
(including juveniles), 80$ 9 (incl. juveniles), AM P54972, type locality. 


13 specimens, AM P57672, 100 m north west of Julian Rocks, Byron 
Bay, 28°36.8'S 153°37.8'E, red alga eDeliseapulchra, 16 m, S.J. Keable, 
4 March 1992, stn NSW-648. 1 specimen, AM P56677, hand collected 
at low tide northern shore under Fred Hansen Bridge, Boambee Creek, 
Sawtell, 30°20.4'S 153°05.5'E, exposed mud flat, Australian Museum 
party, 8 March 1992, stn NSW-717. 4 specimens, AM P5724, Port 
Stephens, [approx. 32°42'S 152°06'E], dredged, A. Musgrave, 30 August 
1920. 1 specimen, AM P47047, west side of Box Head, Broken Bay, 
33°33'S 151 °21'E, coralline algae in low intertidal zone exposed to the 
south, A. Murray, R.T. Springthorpe & H.E. Stoddart, 11 April 1981, 
stn NSW-2. 28 8, 29 9, AM P60495, Port Jackson, [approx. 33°51'S 
151°16'E], 15 specimens, G926, Jervis Bay, [approx. 35°03'S 150°44'E], 
T. Whitelegge. Id, AM P52785, Moe’s Rock, South of Jervis Bay, 
35°09'S 150°45'E, foliose bryozoan, 18 m, R.T. Springthorpe & J.K. 
Lowry, 29 June 1981, stn NSW-55. 3 9 9, AM P52784, southern end of 
Lighthouse Reef, Ulladulla, New South Wales, 35°22.14'S 150°29.31'E, 
bryozoan ?Orthoscuticella sp., 16 m, PB. Berents, K.B. Attwood, 30 


Lowry & Springthorpe: Australian melitid amphipods 263 


264 Records of the Australian Museum (2005) Vol. 57 


Fig. 25. Mallacoota kameruka n.sp., holotype, male “a”, AM P60491, paratype female, 10.0 mm, AM P60492, 
Marys Rock, Cook Island, New South Wales, Australia. Scales represent 0.5 mm. 


April 1997, stn NSW-1267. Id, AM P47054, Merimbula Wharf, 
Merimbula, 36°53.92'S 149°55.64'E, mixed red and brown algae, 8 m, 
K.B. Attwood, 18 May 1995, NSW-1103. 7 specimens, AM P63381, 
Murrumbulga Point, Twofold Bay, New South Wales, 37°04.7'S 
149°53.1'E, subtidal rock platform, S.J Keable, A. Paul, L. Walker, 29 
March 1985, stn Q8/9. Queensland: 29 9, AM P3493, Port Denison, 


[approx. 20°03'S 148°15'E], [AM Old Collection], 1$, P 47055, Boat 
Rock, North Stradbroke Island, 27°25.1'S 153°33.28'E, bryozoans, 
hydrozoans & brown algae, 28 m, R.T. Springthorpe, 3 June 1993, stn 
QLD-853. Victoria: Id, AM P3494, Griffiths Point, [approx. 38°32'S 
145°22'E], [AM Old Collection], Western Australia: 2 specimens, AM 
P41234, 300 m southeast of Penguin Island, Warnbro Sound, 32°18.5'S 


Lowry & Springthorpe: Australian melitid amphipods 265 


115°41.6'E, seagrass: Amphibolis griffithii, 3.5 m, P. Hutchings et al., 
7-9 November 1990, stn A. 

Type locality. Marys Rock, Cook Island, New South Wales, 
Australia, 28°11.42'S 153°34.79'E, on abryozoan, 19 m. 

Description. Based on holotype male, AM P60491 and 
paratype female, AM P60492. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner rounded. 
Antenna 1 longer than antenna 2; peduncular article 1 
subequal in length to article 2, with 3 robust setae along 
posterior margin; flagellum with at least 24 articles; 
accessory flagellum with 3-4 articles. Antenna 2 peduncular 
article 2 cone gland reaching at least to end of peduncular 
article 3; article 4 longer than article 5; flagellum with about 
11 articles. Mandible palp article 3 rectolinear, setose along 
straight medial margin, longer than article 1; article 2 shorter 
than article 3; article 1 not produced, shorter than article 2, 
about twice as long as broad. Maxilla 1 inner plate with 
about 3 setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
rounded, posteroventral corner notch absent; merus without 
posterodistal spine; propodus palm acute, convex, without 
posterodistal corner, defined by posterodistal robust setae. 
Gnathopod 2 sexually dimorphic; subchelate; coxa 
posteroventral corner notch absent; merus with sharp 
posteroventral spine; carpus compressed; propodus without 
medial depression, with strong setal bunch, palm acute, 
straight, sculptured, with group of anterodistal robust setae, 
without posterodistal robust setae, defined by posteroventral 
spine; dactylus apically falcate. Pereopod 5 basis posterior 
margin convex, posteroventral corner broadly rounded; 
carpus and propodus with many long, slender setae along 
anterior margin. Pereopod 6 coxa anterior lobe ventral 
margin slightly produced, rounded; basis posterior margin 
convex, posteroventral corner broadly rounded; carpus and 
propodus with many long, slender setae along anterior 
margin. Pereopod 7 basis posterior margin convex, with 
posterior margin smooth or minutely castelloserrate, 
posteroventral corner broadly rounded. 

Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner subquadrate or with small 
acute spine. Urosomite 1 dorsally bicarinate. Uropod3 inner 
ramus subequal in length to outer ramus; outer ramus longer 
(1.2 to 2x length) than peduncle, 1-articulate. Telson each 
lobe with 3 or more apical/subapical robust setae, apical 
conical extension absent. 

Female (sexually dimorphic characters). Gnathopod 2 
carpus short; propodus setose, smooth, with sparse robust 
setae, defined by posterodistal robust setae, defined by 
posteroventral corner; dactylus apically acute/subacute. 

Habitat. Marine; littoral; living among bryozoans and red 
algae, 16 to 19 m depth. 

Etymology. Named for the coastal steamer Kameruka, 
wrecked on Pedro Reef, off Moruya, New South Wales, in 
1897. 


Remarks. This species was originally described as 
Mallacoota subcarinata phenotype B by J.L. Barnard 
(1972a). It appears to be the same as the Challenger 
specimens Stebbing (1888) described from off Melbourne. 
Mallacoota kameruka is similar to M. subcarinata and M. 
malua in the absence of posteroventral notches on coxae 1 
and 2. It differs from all Australian species in having convex 
posterior margins of pereopods 5-7. 

Distribution. Queensland: Port Denison; North Stradbroke 
Island (both AM). New South Wales: Marys Rock, Cook 
Island; Julian Rocks, Byron Bay; Boambee Creek, Sawtell; 
Port Stephens; Broken Bay; Port Jackson; Jervis Bay; 
Ulladulla; Merimbula (all AM); Munganno Point, Twofold 
Bay (Hutchings et al ., 1989). Victoria : Off Melbourne 
(Stebbing, 1888); Griffiths Point (AM); Port Phillip (J.L. 
Barnard, 1972a). Western Australia: Point Peron; Rottnest 
Island; Cottesloe Beach (all J.L. Barnard, 1972a); Warnbro 
Sound (AM). 

Australian geographic areas. Northeastern, southeastern, 
southern and southwestern Australia. 

Mallacoota malua n.sp. 

Ligs. 26-28 

Type material. Holotype, 3 “a”, 10 mm, AM P56679; 10 paratypes, 
33 3,7$ 2, AM P60485; Coffs Harbour Jetty, Coffs Harbour, New South 
Wales, 30°18.4'S 153°08.5'E, arborescent sponge on jetty pilings, 7 m, 
S.J. Keable, 9 March 1992, stn NSW-735; 22 paratypes, 83 3, 14$ $, 
AM P56678, type locality, Pyura praeputialis on jetty pilings, 8 m, PB. 
Berents & S.J. Keable, 9 March 1992, stn NSW 733; 1 paratype, $, 8.5 
mm, AM P60486; 1 PARATYPE, 3 “c”, 7.1 mm, AM P60490, type locality, 
coral scrapings on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, 
stn NSW 726. 

Additional material examined. New South Wales: 13, 2$ $, AM 
P56674, 50 m west of Split Solitary Island, 30°14.0'S 153°10.8'E, 
Herdmania momus, rocks, sponges & ascidians, 15-17 m, PA. Hutchings 
& C.L. Rose, 7 March 1992, stn NSW-677. 1 $, AM P56676, Boambee 
Creek, Sawtell, 30°20.8'S 153°05.6E, silty sand submerged at low tide, 
0.3 m, E. Albertson & S. Keable, 8 March 1992, stn NSW-714. 433, 
12$ $, AM P57220, Coffs Harbour Jetty, Coffs Harbour, 30°18.4'S 
153°08.5'E, worm tubes encrusted with sponge on jetty pilings, 6 m, 
R.T. Springthorpe, 9 March 1992, stn NSW-725. 1 specimen, AM 
P57221, same locality, coral scrapings on jetty pilings, 6 m, R.T. 
Springthorpe, 9 March 1992, stn NSW-738. 5 3 3, 6$ $, AM P57301, 
same locality, finger sponge on jetty pilings, 4 m, R.T. Springthorpe, 9 
March 1992, stn NSW-734. 13, 4$ $, AM P57302, same locality, coral 
scrapings on jetty pilings, 6 m, R.T. Springthorpe, 9 March 1992, NSW 
726. 7 $ $, AM P58230, outer end of Kurnell Pier, Botany Bay, 34°00.2'S 
151°12.5'E, pylon scrapings, 7 m, NSW Fisheries/CRIMP Survey, 21 
October 1998, stn BB KP1 Pl-7. 13 (“b”), AM P60487, same locality. 
23 3, 1 $, AM P63071, same locality, pylon scrapings, 3 m, NSW 
Fisheries/CRIMP Survey, 21 October 1998, BB KP1 P2-3. 13, AM 
P58231, Kurnell Pier (near shoreline). Botany Bay, 34°00.5'S 151°12.7'E, 
pylon scrapings, 3 m, NSW Fisheries/CRIMP Survey, 21 October 1998, 
stn BB KP2 P3-3. 233, 1 $, AM P60488, north east corner of Clark 
Island, Port Jackson, 33°51.85'S 151°14.47'E, red alga, 2 m, I. Takeuchi 
& D. Bray, 17 April 1996, stn NSW-1250. 633, 5$ $, AM P60623, 
Port Jackson, 33°51'S 151°16’E, [AM Old Collection], 

Type locality. Coffs Harbour Jetty, New South Wales, 
Australia, 30°18.4'S 153°08.5'E, sponges, ascidians on 
pilings, 7 m. 

Description. Based on holotype male “a”, AM P56679, 
paratype female, AM P60486, paratype male “c”, AM 
P60490 and male “b”, AM P60487. 


266 Records of the Australian Museum (2005) Vol. 57 


Head. Lateral cephalic lobes broad, truncated, with 
anteroventral notch or slit, anteroventral corner rounded. 
Antenna 1 longer than antenna 2; peduncular article 1 
subequal in length to article 2, with 3 robust setae along 
posterior margin; flagellum with at least 23 articles; 
accessory flagellum with 3-4 articles. Antenna 2 peduncular 
article 2 cone gland reaching at least to end of peduncular 
article 3; article 4 longer than article 5, or article 4 subequal 
to article 5; flagellum with about 12 articles. Mandible palp 
article 3 rectolinear, with setae mostly terminal, longer than 
article 1; article 2 subequal to article 3; article 1 not 
produced, shorter than article 2, about twice as long as broad. 
Maxilla 1 inner plate with about 4 setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
rounded, posteroventral corner notch absent; merus without 
posterodistal spine; propodus palm acute, convex or straight, 
defined by posterodistal corner, defined by posterodistal 
robust setae. Gnathopod 2 sexually dimorphic; subchelate; 
coxa posteroventral corner notch absent; merus with sharp 
posteroventral spine; carpus compressed; propodus without 
medial depression, with strong setal bunch, palm acute, 
straight, sculptured, with sparse robust setae and with group 
of anterodistal robust setae, without posterodistal robust 
setae, defined by posteroventral spine; dactylus apically 
falcate. Pereopod 5 basis posterior margin slightly concave, 
posteroventral corner narrowly rounded or subquadrate. 
Pereopod 6 coxa anterior lobe ventral margin slightly 
produced, rounded; basis posterior margin slightly concave 
or straight, posteroventral corner narrowly rounded or 
subquadrate; propodus expanded posterodistally to from a 
hood-like projection. Pereopod 7 basis posterior margin 


straight, with posterior margin smooth or minutely 
castelloserrate, posteroventral corner narrowly rounded or 
subquadrate; propodus expanded posterodistally to from a 
hood-like projection. 

Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner subquadrate or with small 
acute spine. Urosomite 1 dorsally bicarinate. Uropod3 inner 
ramus subequal in length to outer ramus; outer ramus longer 
(1.2 to 2x length) than peduncle, 1-articulate. Telson each 
lobe with 3 or more apical/subapical robust setae, apical 
conical extension absent. 

Female (sexually dimorphic characters). Gnathopod 2 
carpus short; propodus setose, slightly convex, smooth, with 
sparse robust setae, defined by posterodistal robust setae, 
without posteroventral corner; dactylus apically acute/ 
subacute. 

Habitat. Marine; littoral; among ascidians, sponges and red 
algae on wharf pilings, 0 to 17 m depth. 

Etymology. Named for the wooden steamer Malua, built 
in Botany Bay and sunk at the mouth of Port Hacking, New 
South Wales, in 1886. 

Remarks. Mallacoota malua is most closely related to 
Mallacoota kameruka. They differ in the shape of the bases 
of pereopods 5-7, and in the sculpturing on the palm of 
gnathopod 2. Mallacoota malua is the only species with 
posterodistal hood-like projections on pereopods 6 and 7. 


Lowry & Springthorpe: Australian melitid amphipods 267 


Fig. 27. Mallacoota malua n.sp., holotype, male “a”, 10 mm, AM P56679, Coffs Harbour Jetty, New South Wales, 
Australia. Scales for MDp, MX1IP, MX2 represent 0.1 mm, remainder represent 0.2 mm. 


Distribution. New South Wales : Split Solitary Island; Coffs 
Harbour; Boambee Creek, Sawtell; Clark Island, Port 
Jackson; Kurnell, Botany Bay (all AM). 

Australian geographic areas. Southeastern Australia. 

Mallacoota nananui Myers 

lElasmopus subcarinatus Chilton, 1915 (in part), 325, fig. 5. 
Mallacoota subcarinata.-LL. Barnard, 1972b: 114, figs 59-60. 
Mallacoota nananui Myers, 1985: 121, fig. 95.-Myers, 1986: 
1389, fig. 8. 

Type material. Holotype: 8, 5.7 mm, AM P35209; paratypes, 
18 8, 15 9 9, AM P35210; lagoon, Nananui Ra, Viti Levu, Fiji [17°15'S 
178°12'E], on Halimeda sp., A.A. Myers, 7 October, 1979, stn 53. 

Material examined. New South Wales: about 95 specimens, AM 
P64661, between Comet Hole and the reef, Lord Howe Island lagoon, 


[approx. 31°30.5'S 159°03'E], associated with red and brown algae, 2- 
3 m, J.K. Lowry & G.D. Fenwick, 10 May 1977, stn LHA-11.11 
specimens, AM P64662, reef front west of Signal Point, Lord Howe 
Island Lagoon, [approx. 31°30.5'S 159°03'E], Caulerpa, 1.5-2.0m, G.D. 
Fenwick, 11 May 1977, stn LHA-15. 

Type locality. Lagoon, Nananui Ra, Viti Levu, Fiji [approx. 
17°15’S 178°12’E]. 

Habitat. Marine; littoral; among red and brown algae in 
lagoons, 0 to 3 m depth. 

Remarks. This is the first record of M. nananui from 
Australian waters. 

Distribution. New South Wales: Lord Howe Island (AM). 
Extrinsic distribution. Fiji; Nuie; New Zealand. 
Australian geographic areas. Southeastern Australia. 


268 Records of the Australian Museum (2005) Vol. 57 


Fig. 28. Mallacoota malua n.sp., holotype, male “a”, 10 mm, AM P56679, paratype, female, 8.5 mm, AM P60486, 
*paratype, male “c”, 7.1 mm, AM P60490, Coffs Harbour Jetty, New South Wales, Australia; ** male “b”, AM 
P60487, Botany Bay, New South Wales Australia. Scales represent 0.5 mm. 


Lowry & Springthorpe: Australian melitid amphipods 269 


Mallacoota subcarinata (Haswell) 

Figs. 29-33 

Megamoera sub-carinata Haswell, 1879b: 335, pi. 21, fig. 4. 
Elasmopus subcarinatus Stebbing, 1906: 441 -Stebbing, 1910a: 
602.-Chilton, 1921b: 76. 

Not Megamoera sub-carinata.-Chilton, 1885: 1039. 

Not Moera petriei Thomson, 1882: 236, pi. 18, fig. 3.-Chilton, 
1883: 82, pi. 2, fig. 4.-Chilton, 1885: 1039. 

Not Moera sub-carinata- Chilton, 1884: 230.-Thomson & 
Chilton, 1886: 146. 

Not Moera subcarinata.-Chilton, 1885:1039.-Thomson, 1889:261. 
Not Elasmopus subcarinatus-Stebbing, 1888: 1019, pi. 98 (plate 
labelled as E. persetosus ). Chilton, 1892: 261.-Walker, 1904: 
275, pi. 5, fig. 34.-Stebbing, 1906: 441.-Walker, 1909: 335.- 
Stebbing, 1910a: 602.-Stebbing, 1910b: 457.-Thomson, 1913: 
243-Chilton, 1915: 321, figs l-6.-Stephensen, 1931: 11.- 
K.H. Barnard, 1935: 286.-Pirlot, 1936: 317, figs 136-145.- 
K.H. Barnard, 1937: 160. 

Not Maerasubcarinata.-K.H. Barnard, 1940: 460-Nayar, 1966: 149. 


Not Mallacoota subcarinata.-].L. Barnard, 1972a: 247, figs 144- 
145 .-J.L. Barnard, 1972b: 114, figs 59-60.-Lowry, 1974: 112, 
125 figs 9b,d (key).-Ledoyer, 1978: 281, fig. 32.-Barnard & 
Barnard, 1983: 632.-Ledoyer, 1984:71,72 figs 34-35.-Myers, 
1985: 121, fig. 96.-Myers, 1986: 1390, fig. 9.-Hutchings et 
al., 1989: 362.-Myers, 1995: 38. 

Type material. 1 syntype, S , 9.6 mm, AM G5390; 1 syntype, $, 10.4 
mm, AM P63972; 2 SYNTYPES, AM P63973, Port Jackson, New South Wales, 
Australia, [approx. 33°51'S 151°16'E, [AM Old Collection]; 3 SYNTYPES, 
AM P3492, Port Stephens, 32°42'S 152°06’E, [AM Old Collection], 
[specimens not located, March 1993 (Springthorpe & Lowry, 1994)]. 

Additional material examined. New South Wales: 2 specimens, 
AM P22484, east of Mona Vale, 33°41'S 151°19’E, 16 m, Australian 
Museum Shelf Benthic Survey, 11 May 1972. 2 specimens, AM P22479, 
east of Long Reef, 33°44'S 151°22'E, 38 m, Australian Museum Shelf 
Benthic Survey, 24 Aug 1972. 5 specimens, AM P22480, same locality, 
36 m, Australian Museum Shelf Benthic Survey, 11 May 1972. 2 
specimens, AM P22481, same locality, 32 m, Australian Museum Shelf 
Benthic Survey, 28 May 1972. 1 specimen, AM P22482, same locality, 
15 m, Australian Museum Shelf Benthic Survey, 28 April 1972. 2 


Fig. 29. Mallacoota subcarinata (Haswell, 1879b), syntype male, 9.6 mm, syntype, female, 10.4 mm, AM G5390, 
Port Jackson, New South Wales, Australia. Scales represent 0.2 mm. 


270 Records of the Australian Museum (2005) Vol. 57 


Fig. 30. Mallacoota subcarinata (Haswell, 1879b), syntype, male, 9.6 mm, AM G5390, syntype, female, 10.4 
mm, AM P63972, Port Jackson, New South Wales, Australia. Scales represent 0.5 mm. 


specimens, AM P22483, same locality, Australian Museum Shelf Benthic 
Survey, 1972. 1 8 “a” 10.4 mm, AM P22487, east of Long Reef, 33°44'S 
151°22’E, 40 m, Australian Museum Shelf Benthic Survey, 29 June 1972. 
1 8 “b” 5.7 mm, AM P60563, same locality. 9 specimens, AM P22476, 
east of North Head, Port Jackson, 33°49'S 151°18'E, host sponge: Halme 
gigantea, 25 m, Australian Museum Shelf Benthic Survey, 26 February 
1974, transect 07. 1 specimen, AM P22477, east of North Head, Port 
Jackson, 33°49.5'S 151°18'E, 32 m, Australian Museum Shelf Benthic 
Survey, 23 May 1972. 5 specimens, AM P5860, Balmoral, Port Jackson, 
[approx. 33°49.7'S 151°15.1’E], T. Whitelegge, [AM Old Collection], 1 
specimen, AM P22485, east of South Head, Port Jackson, 33°50°S 
151 0 18'E, host sponge: Halme gigantea , 21m, Australian Museum Shelf 
Benthic Survey, February 1972, transect 11.5 specimens, AM P63072, 
outer end of Kurnell Pier, Botany Bay, 34°00.2'S 151°12.5'E, pylon 


scrapings, 7 m, NSW Fisheries/CRIMP Survey, 21 October 1998, stn 
BB KP1 P2-7. 3 specimens, AM P63073, Kurnell Pier (near shoreline), 
Botany Bay, 34°00.5'S 151°12.7'E, pylon scrapings, 3 m, NSW Fisheries/ 
CRIMP Survey, 21 October 1998, stn BB KP2 P2-3. 6 specimens, AM 
P2497, 3-4 km off Botany Bay, [approx. 34°05’S 151°15'E], mud, 91- 
95 m, E.R. Waite on HMCS Thetis, 11 March 1898, stn. 37. 1 specimen, 
AM P2498, 4.5-5 km off Jibbon Point, [approx. 34°07.5'S 151°12’E], 
sand, mud, 84-101 m, E.R. Waite on HMCS Thetis , 12 March 1898, stn. 
38. 8 specimens, AM P2494, 5.5-6.5 km off Wattamolla, [approx. 
34°10’S 151°11'E], mud, 99-108 m, E.R. Waite on HMCS Thetis, 22 
March 1898, stn 57. 1 specimen, AM P2499,11-12.5 km off Wollongong, 
[approx. 34°27'S 151°04'E], sand, mud, rock, 102 m, E.R. Waite on 
HMCS Thetis, 18 March 1898, stn. 48. 18, AM P63117, Jervis Bay, 
[approx. 35°03'S 150°44'E], T. Whitelegge. 2 specimens, AM P62905, 


Lowry & Springthorpe: Australian melitid amphipods 271 


Fig. 31. Mallacoota subcarinata 
(Haswell, 1879b), male “a”, 10.4 mm, 
AM P22487, east of Long Reef, New 
South Wales, Australia. 


Burrill Rock, south of Warden Head, 35°23.39'S 150°28.24'E, 
gorgonacean, 24 m, R.T. Springthorpe, 7 May 1997, stn NSW-1349. 9 
specimens, AM P35986, Munganno Point, Twofold Bay, 37°06.2'S 
149°55.7'E, subtidal rock platform, 0-7 m, P. Hutchings, 10 October 
1984, stn M3. Tasmania: 1 $ , E6549, Tasmanian Coast, FIS Endeavour, 
1909-1914. 2 specimens, E6550, eastern slope of Bass Strait, [approx. 
39°00'S 148°40'E], FIS Endeavour, 1909-1914. 3 specimens, AM P5936, 
same locality. 

Type locality. Port Jackson, New South Wales (33°51'S 
151°16'E) and Port Stephens, New South Wales, Australia, 
(32°42'S 152°06'E). 

Description. Based on male syntype, AM G5390, female 
syntype, AM P63972, male “a”, AM P22487 and male “b”, 
AM P60563. 

Head. Lateral cephalic lobes broad, truncated, with 
anteroventral notch or slit, anteroventral corner rounded. 
Antenna 1 longer than antenna 2; peduncular article 1 
subequal in length to article 2, with 1-3 robust setae on 
posterior; flagellum with about 31 articles; accessory 
flagellum with 4-5 articles. Antenna 2 peduncular article 2 
cone gland reaching at least to end of peduncular article 3; 
article 4 subequal to article 5; flagellum with about 12 
articles. Mandible palp article 3 rectolinear, with setae 
mostly terminal, longer than article 1; article 2 subequal to 
article 3; article 1 not produced, shorter than article 2, about 
twice as long as broad. Maxilla 1 inner plate with 4-5 setae 
mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
rounded, posteroventral corner notch absent; merus without 
posterodistal spine; propodus palm acute, convex, without 
posterodistal corner, defined by posterodistal robust setae. 
Gnathopod 2 sexually dimorphic; subchelate; coxa 


posteroventral corner notch absent; merus with sharp 
posteroventral spine; carpus compressed; propodus without 
medial depression, with strong setal bunch, palm extremely 
acute, convex or sinusoidal, sculptured, with group of 
anterodistal robust setae, without posterodistal robust setae, 
with or without posteroventral corner; dactylus apically 
blunt or falcate. Pereopod 5 basis posterior margin straight 
or slightly concave, posteroventral comer narrowly rounded 
or subquadrate. Pereopod 6 coxa anterior lobe ventral 
margin slightly produced, rounded; basis posterior margin 
concave, posteroventral corner serrate. Pereopod 7 basis 
posterior margin slightly concave or straight, with posterior 
margin smooth or minutely castelloserrate, posteroventral 
corner narrowly rounded or subquadrate. 

Pleon. Epimeron 1 posteroventral corner narrowly rounded 
or subquadrate. Epimera 1-2 posteroventral margin without 
spines above posteroventral corner. Epimeron 3 posterior 
margin smooth, posteroventral corner with strongly 
produced acute spine. Urosomite 1 dorsally bicarinate. 
Uropod 3 inner ramus subequal in length to outer ramus; 
outer ramus longer (1.2 to 2x length) than peduncle, 1- 
articulate. Telson each lobe with 3 or more apical/subapical 
robust setae, apical conical extension reaching scarcely one 
third along longest seta or absent. 

Female (sexually dimorphic characters). Gnathopod 2 
carpus short; propodus setose, convex, smooth, with sparse 
robust setae, defined by posterodistal robust setae; dactylus 
apically acute/subacute. 


272 Records of the Australian Museum (2005) Vol. 57 


Fig. 32. Mallacoota subcarinata (Haswell, 1879b), male “a”, 10.4 mm, AM P22487, east of Long Reef, New 
South Wales, Australia. Scales for MDp, MX IIP, MX2, represent 0.1 mm, remainder represent 0.2 mm. 


Habitat. Marine; littoral and continental shelf; subtidal rock 
platforms, sand, mud, sponges and from jetty pilings, 3.5 
to 108 m depth. 

Remarks. Mallacoota subcarinata Haswell, 1879b appears 
to be confined to Australian waters. All extrinsic records 
need to be carefully studied. The species appears to be most 
similar to those species with a strong posteroventral corner 
spine on epimeron 3. In Australian waters this includes M. 
diemenensis and Mallacoota euroka. Mallacoota diemen- 
ensis differs from all other Australian species in having 
dorsal carinae on pleonites 1 to 3. Mallacoota subcarinata 
differs from Mallacoota euroka in having only one to three 
robust setae along the posterior margin of antennal 
peduncular article 1 and in the posterior margin of the basis 
of pereopod 7 that is straight in M. subcarinata and convex 
in M. euroka. 


Distribution. New South Wales : east of Long Reef; east of 
Port Jackson (both AM); Port Jackson (Haswell, 1879b); off 
Botany Bay; off Jibbon; off Wattamolla; off Wollongong (all 
Stebbing, 1910a); Jervis Bay; Ulladulla; Munganno Point, 
Twofold Bay (all AM). Tasmania : Bass Strait (Chilton, 1921b). 

Australian geographic areas. Southeastern Australia. 

Miramaera n.gen. 

Type species. Miramaera thetis n.sp. 

Diagnosis. Head without anteroventral notch; eye ovate to 
reniform. Antenna 1 accessory flagellum long, nearly half 
to more than half length of primary flagellum. Mandible 
palp article 1 strongly produced distally; article 2 longer 
than article 3; article 3 long, rectolinear. Maxilla 1 inner 
plate with mainly apical setae. Gnathopod 1 coxa 
anteroventral corner produced, acute or subacute. 


Lowry & Springthorpe: Australian melitid amphipods 273 


Fig. 33. Mallacoota subcarinata (Haswell, 1879b), male “a”, 10.4 mm, AM P22487, *male “b”, 5.7 mm, AM 
P60563, east of Long Reef, New South Wales, Australia. Scales represent 0.2 mm. 


Gnathopod 2 significantly enlarged in male and female; 
left and right gnathopods symmetrical in male; palm acute 
in male and female, both male propodi with well defined 
corner (greater than 90°), both female propodi with well 
defined corner (greater than 90°); dactylus with 1 or 2 setae 
on anterior margin. Pereopods 5-7 dactyli simple. Epimeron 
3 posterior margin smooth. Urosomite 1 dorsal and posterior 
margins smooth. Uropod 3 rami distally truncated, about 
1.5x to 3 or more t im es peduncle, apical robust setae long 
or short; outer ramus 1-articulate. Telson deeply cleft, lobes 
truncated with apical cusps, with or without robust setae on 
outer margins, with short apical robust setae. 


Species composition. Miramaera tepuni (J.L. Barnard, 
1972b); Miramaera thetis n.sp. 

Etymology. A combination of the Latin word mirus, 
meaning wonderful, with the Latin stem Maera. 

Remarks. Miramaera is excluded from the Linguimaera 
group because of its symmetrical second gnathopods. It is 
excluded from Quadrimaera because of the strong apical 
notch on the anteroventral margin of the head, the acute 
palms on the propodi of male gnathopod 2, the simple 
dactyli on pereopods 5-7 and the telsonic lobes that are 
truncated with apical cusps. It may be most similar to genera 


274 Records of the Australian Museum (2005) Vol. 57 


in the Maera group ( sensu stricto ), but it differs from that 
group in the setation of the dactyli of the second gnathopods. 
The significant differences between Miramaera and 
Lupimaera are that in Miramaera the posterior margin of 
epimeron 3 is smooth, the rami of uropod 3 are longer than 
the peduncle and the telsonic lobes have apical cusps and 
short robust setae. Miramaera differs from Maeropsis in 
having acute palms and a well defined corner of more than 
90° on both male and female propodi of gnathopod 2. 

Distribution. Australia; New Zealand. 

Miramaera thetis n.sp. 


Head. Lateral cephalic lobes broad, rounded, without notch 
or slit, anteroventral corner with acute/subacute spine. 
Antenna 1 longer than antenna 2; peduncular article 1 
slightly shorter than article 2, with 4 or more robust setae 
along posterior margin; flagellum with 16 articles; accessory 
flagellum with 9 articles. Antenna 2 peduncular article 2 
cone gland not reaching to end of peduncular article 3; 
article 4 longer than article 5; flagellum with 8 or 9 articles. 
Mandible palp article 3 rectolinear, setose on distomedial 
margin, longer than article 1; article 2 longer than article 3; 
article 1 produced distally, shorter than article 2, about twice 
as long as broad. Maxilla 1 inner plate with 3 setae mainly 
terminal. 


Figs. 34-36 

Maera inaequipes.-Stebbing, 1910a: 599.-Sheard, 1937: 24. 

Type material. Holotype, S “a”, 7.9 mm, AM P62798; many 
PARATYPES, AM P27035; 1 paratype, 9, 6.7 mm, AM P62799; 1 
PARATYPE, S “b”, 7.8 mm, AM P62800; 10 paratypes, AM P62801, 
between Troubridge Light and Cape Jervis, South Australia, 35°20'S 
137°40'E, sponges on mud bottom, 20 m, D. Blake & H. Larsen, 14 
March 1978. 

Additional material examined. New South Wales: 1 specimen, 
AM P2492, 8-9.5 km off Coogee, 33°57’S 151°21.5'E, fine sand, 91 m, 
E.R. Waite on HMCS Thetis, 15 March 1898, stn 44. South Australia: 1 
ovigerous female, E6544, 24 km north west of Cape Jervis, [approx. 
35°26'S 137°55’E], 31 m, FIS Endeavour, 1909-1914. 

Type locality. Between Troubridge Light and Cape Jervis, 
South Australia, [approx. 35°20'S 137°40'E], sponges on 
mud bottom, 20 m. 


Pereon. Gnathopod 1 coxa anteroventral corner produced, 
acute, posteroventral corner notch absent; merus without 
posterodistal spine; carpus with anterodistal swelling; palm 
acute, convex, without posterodistal corner, defined by 
posterodistal robust setae. Gnathopod 2 sexually dimorphic; 
subchelate; coxa posteroventral corner notch absent; merus 
with sharp posteroventral spine; carpus compressed; 
propodus without medial depression, palm acute, convex, 
sculptured, lined with robust setae, defined by posterodistal 
robust setae, defined by posteroventral spine; apically acute/ 
subacute. Pereopod 5-6 basis posterior margin slightly 
concave, posteroventral corner broadly rounded. Pereopod 
7 basis posterior margin slightly convex, with posterior 
margin castelloserrate, posteroventral corner broadly 
rounded. 


Description. Based on holotype male, AM P62798 and 
paratype female, AM P62799. 


Pleon. Epimeron 1 posteroventral corner broadly rounded. 
Epimera 1-2 posteroventral margin without spines above 
posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral comer subquadrate. Uropod 
3 rami distally tmncated; inner ramus subequal 
in length to outer ramus; outer ramus 
longer (1.2 to 2x length) than peduncle, 
1-articulate. Telson each lobe with 2 
apical/subapical robust setae, apical 
conical extension reaching scarcely 
one third along longest seta. 

Female (sexually dimorphic 
characters). Gnathopod 2 
palm smooth. 


Fig. 34. Miramaera thetis n.sp., paratype, male “b”, 7.8 mm, AM P62800, 
between Troubridge Light and Cape Jervis, South Australia. 


Lowry & Springthorpe: Australian melitid amphipods 275 


Fig. 35. Miramaera thetis n.sp., holotype, male, 7.9 mm, AM P62798, between Troubridge Light and Cape Jervis, South Australia. 


Habitat. Marine; sponges on mud bottom, sand; littoral, 
continental shelf; 20 to 100 m depth. 

Etymology. Named for the Thetis Expedition, which 
collected the marine fauna off the coast of Sydney during 
1898. 

Remarks. Miramaera thetis differs significantly from 
Miramaera tepuni (J.L. Barnard, 1972b), the only other 


species in the genus, in the rami of uropod 3 that are only 
half the length of those of P. tepuni. 

Distribution. New South Wales : off Coogee and off 
Wollongong (both Stebbing, 1910a). South Australia: Cape 
Jervis (AM) 

Australian geographic areas. Southeastern and southern 
Australia. 


276 Records of the Australian Museum (2005) Vol. 57 


Parelasmopus Stebbing, 1888 
Parelasmopus sowpigensis n.sp. 

Figs. 37-40 

Type material. Holotype, S, 8.9 mm, AM P60496; 1 paratype, 9, 
7.1 mm, AM P60497; 16 paratypes, 3c? <J, 139 9, AM P60498; 21 
paratypes, 13 c? <?, 8 9 9, AM P60499, off Sow and Pigs reef, Port Jackson, 
New South Wales, Australia, 33°50.3'S 151°16.2’E, shelley sand, 5 m, benthic 
grab, J.K. Lowry & A.R. Jones, 30 September 1976, stn NSW-184. 


Type locality. Sow and Pigs Reef, Port Jackson, New South 
Wales, 33°50.3’S 151°16.2’E, shelley sand, 5 m. 

Description. Based on holotype male, AM P60496 and 
paratype female, AM P60497. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner with acute/ 
subacute spine. Antenna 1 longer than antenna 2; peduncular 
article 1 subequal in length to article 2, with 3 robust setae 


Lowry & Springthorpe: Australian melitid amphipods 277 


along posterior margin; flagellum with 22 articles; accessory 
flagellum with 4 articles. Antenna 2 peduncular article 2 
cone gland reaching at least to end of peduncular article 3; 
article 4 longer than or subequal to article 5; flagellum with 
6 articles. Mandible palp article 3 rectolinear, setose on 
distomedial margin, subequal to or shorter than article 1; 
article 2 shorter than article 3; article 1 curved, swollen 
distally, longer than article 2, at least 3x as long as broad. 
Maxilla 1 inner plate with 2 setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner not 
produced, posteroventral comer notch absent; mems without 
posterodistal spine; propodus palm acute, convex, without 
posterodistal corner, defined by posterodistal robust setae. 
Gnathopod 2 sexually dimorphic; subchelate; coxa 
posteroventral corner notch absent; merus with sharp 
posteroventral spine; carpus short; propodus without medial 
depression, without strong concentration of setae, palm 
nearly transverse, straight, smooth, with group of 
anterodistal robust setae, defined by posterodistal robust 
setae, without posteroventral comer; dactylus apically blunt. 
Pereopod 5 basis posterior margin straight, posteroventral 
corner narrowly rounded or subquadrate. Pereopod 6 coxa 
anterior lobe ventral margin slightly produced, rounded; 
basis posterior margin slightly concave, posteroventral 
corner broadly rounded, narrowly rounded or subquadrate. 
Pereopod 7 basis posterior margin straight, with posterior 
margin smooth or minutely castelloserrate, posteroventral 
corner narrowly rounded or subquadrate. 

Pleon. Epimeron 1 posteroventral comer narrowly rounded or 
subquadrate. Epimera 1-2 posteroventral margin without 
spines above posteroventral corner. Epimeron 3 posterior 
margin smooth, posteroventral comer with small acute spine. 
Urosomite 1 dorsally bicarinate. Uropod 3 inner ramus 
subequal in length to outer ramus; outer ramus longer (1.2 to 
2x length) than peduncle, 1-articulate. Telson each lobe with 3 


or more apical/subapical robust setae, apical conical extension 
reaching scarcely one third along longest seta. 

Female (sexually dimorphic characters). Gnathopod 2 
carpus long; propodus palm acute, convex, lined with robust 
setae, defined by posteroventral corner; dactylus apically 
acute/subacute. 

Habitat. Marine; littoral; shelley sand, 5 m depth. 

Etymology. Named for the Sow and Pigs Reef, the type 
locality. 

Remarks. This species fits Parelasmopus because of the 
highly distinctive mandibular palp with its very long first 
article and very short second article, but it differs from the 
generic definition in not having a serrate posteroventral 
margin on epimeron 3, nor does it have dorsally bicarinate 
first and second pleonites. J.L. Barnard (1972a) established 
Ifalukia for one species of Parelasmopus that lacked dorsal 
carinae on the pleonites and urosomites, and lacked 
posteroventral serrations on epimeron 3. Parelasmopus sow- 
pigensis is most similar to Ifalukia , but differs in having a 
dorsally bicarinate first urosomite. It therefore strictly fits 
neither genus. If the mandibular palp is the main synapo- 
morphy defining Parelasmopus, then it appears that some 
species in the complex have either lost or never had dorsal 
carinae or serrate ventral margins. Until the phylogenetic 
implications of these questions can be analysed it is best to 
maintain a broad concept of the genus. 

Parelasmopus sowpigensis is therefore a distinctive 
species differing from other Australian species in having 
dorsally smooth first and second pleonites and smooth 
ventral margins on epimeron 3. 

Distribution. New South Wales: Sow and Pigs Reef, Port 
Jackson (AM). 

Australian geographic areas. Southeastern Australia. 


278 Records of the Australian Museum (2005) Vol. 57 


Fig. 38. Parelasmopus sowpigensis n.sp., holotype, male, 8.9 mm, AM P60496, Sow and Pigs, Port Jackson, New 
South Wales, Australia. Scales for mouthparts represent 0.1 mm, remainder represent 0.2 mm. 


Quadrivisio Stebbing, 1907 
Quadrivisio sarina n.sp. 

Figs. 41-43 

Type material. Holotype, $, 7.4 mm, AM P60482; 1 paratype, 
6", 8.1 mm, AM P60484; 1 PARATYPE, $, AM P60483, Armstrong Beach, 
Sarina, Queensland, Australia, [approx. 21°46'S 149°29'E], sand beach, 
mid-tide level, N. Hacking, 18 December 1994. 

Type locality. Armstrong Beach, Sarina, Queensland, 


Australia, [approx. 21°46'S 149°29'E], sand beach, mid¬ 
tide level. 

Description. Based on holotype female, AM P60482 and 
paratype male, AM P60484. 

Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner rounded. 
Antenna 1 shorter than antenna 2; peduncular article 1 longer 
than article 2, with 2 robust setae along posterior margin; 


Lowry & Springthorpe: Australian melitid amphipods 279 


Fig. 40. Parelasmopus sowpigensis 
n.sp., holotype, male, 8.9 mm, AM 
P60496, Sow and Pigs, Port Jackson, 
New South Wales, Australia. Scales 
represent 0.2 mm. 


280 Records of the Australian Museum (2005) Vol. 57 


flagellum with at least 14 articles; accessory flagellum with 
6 articles. Antenna 2 peduncular article 2 cone gland reaching 
at least to end of peduncular article 3; article 4 subequal to 
article 5; flagellum with 15 articles. Mandible palp article 3 
rectolinear, with setae mostly terminal, longer than article 1; 
article 2 subequal to or shorter than article 3; article 1 
produced distally, shorter than article 2, about as long as 
broad. Maxilla 1 inner plate setose along entire inner margin. 

Pereon. Gnathopod 1 coxa anteroventral corner not 
produced, posteroventral comer notch absent; mems without 
posterodistal spine; propodus palm nearly transverse, 
convex, without posterodistal corner, defined by postero¬ 
distal robust setae. Gnathopod 2 sexually dimorphic; 
subchelate; coxa posteroventral corner notch absent; mems 
with rounded posteroventral corner, or with subquadrate 
posteroventral corner; carpus compressed; propodus with 
medial depression, without strong concentration of setae, 
palm extremely acute, convex, smooth, lined with robust 
setae, defined by posterodistal robust setae, without 
posteroventral corner; dactylus apically acute/subacute. 
Pereopod 5-6 basis posterior margin convex, posteroventral 
corner narrowly rounded or subquadrate. Pereopod 7 basis 
posterior margin convex, with posterior margin smooth or 
minutely castelloserrate, posteroventral corner narrowly 
rounded or subquadrate. 


Etymology. Named for the Queensland town of Sarina, near 
the type locality of the species. 

Remarks. This is the first record of Quadrivisio in 
Australian waters. It is most similar to Q. bengalensis 
Stebbing, 1907, in having robust setae on the inner margins 
of the telson. Quadrivisio sarina differs from that species 
in the shape of the basis of pereopod 7 and in the shape of 
the palm of gnathopod 2. The antennae and rami of uropod 
3 are generally less setose in Quadrivisio sarina. 

Distribution. Queensland: Armstrong Beach, near Sarina (AM). 
Australian geographic areas. Northeastern Australia. 

Eriopisa group 

Victoriopisa Karaman & Barnard, 1979 
Victoriopisa australiensis (Chilton) 

Figs. 44-46 

Niphargus australiensis Chilton, 1923: 80, fig. l.-Sheard, 1937: 24. 
Victoriopisa australiensis Stock, 1980: 383-Stock & Platvoet, 
1981: 30 (key).-Barnard & Barnard, 1983: 670.-Karaman, 
1984: 58.-Jones, et al., 1986: 541.-Jones, 1987: 623.- 
Hutchings et al., 1989: 362. 

Type material. Holotype, AM P5852, in South West Creek, 400 m 
from the sea, Macleay River, South West Rocks, Trial Bay, New South 
Wales, [approx. 30°53'S 153°03’E], tidal lagoon, J.R. Kinghorn, 1920. 


Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner with small acute spine. 
Urosomite 1 with posterodorsal spine. Urosomite 2 with 
two groups of 1-3 small dorsolateral robust setae. Uropod 
3 inner ramus subequal in length to outer ramus; outer ramus 
longer (1.2 to 2x length) than peduncle, 1-articulate. Telson 
with 1 robust seta per lobe on inner margins, each lobe with 
3 or more apical/subapical robust setae, apical conical 
extension absent. 

Female (sexually dimorphic characters). Gnathopod 2 
merus with sharp posteroventral spine or with subquadrate 
posteroventral corner; propodus without medial depression, 
defined by posteroventral corner or without posteroventral 
corner. 


Additional material examined. New South Wales: 4 specimens, 
AM P63955, near boat ramp, Hickey Island, Clarence River, New South 
Wales, Australia, 29°26.039'S 153°21.552°E (GPS), sand silt, van Veen 
grab, 3 m, P. Hutchings et al., 25 February 2003, stn NSW-2120. 1 
specimen, AM P56683, near Fred Hansen Bridge Boambee Creek, 
Saw tell, 30°20.8’S 153°05.6'E, seagrass ( Zostera ) and mud, 0.2 m, 
Australian Museum party, 8 March 1992, stn NSW-718. 13, 11.1 mm, 
AM P56684, 50 m upstream from Fred Hansen Bridge, Boambee Creek, 
Sawtell, New South Wales, Australia, 30°20.8'S 153°05.6’E, seagrass 
(Zostera), 0.5 m, sweep net at low tide, Australian Museum party, 8 
March 1992, stn NSW-719. 1 specimen, AM P60599, 50 m upstream 
from Fred Hansen Bridge, Boambee Creek, Sawtell, 30°20.8'S 
153°05.6'E, seagrass (Zostera), 0.5 m, sweep net at low tide, Australian 
Museum party, 8 March 1992, stn NSW-719. 2 specimens, AM P54276, 
Wallis Fake, [approx. 32°17'S 152°30'E], M. Fincoln-Smith, stn Zostera 
Bay 5. Many specimens, AM P30831-P30971, near mangroves, north 
west side Fullerton Cove, Hunter River, [approx. 32°50'S 151°46.5F], 
soft mudflat, 0-3.5 m, Australian Fittoral Society, NSW Division, 
between 8 June 1975 and 12 November 1977, 
transects Al-4, Bl-3, C6-7. 1 specimen, 


Lowry & Springthorpe: Australian melitid amphipods 281 


Fig. 42. Quadrivisio sarina n.sp., holotype, female, 7.4 mm, AM P60482, Armstrong Beach, Sarina, Queensland, 
Australia. Scales for mouthparts represent 0.1 mm, remainder represent 0.2 mm. 


Survey, 8 October 1985, stn 6, 6. 5 specimens, AM P55282, 1 km 
upstream of Thackeray Street footbridge, Parramatta River, New South 
Wales, 33°49.24'S 151°02.17'E, muddy sand, 3.3 m, P. Berents & party, 
7 December 1994, Site 2, Rep 1, Upper Parramatta River Dredging 
Survey, 1992-1994. 4 specimens, AM P54282, Brays Bay, Parramatta 
River, 33°50.0'S 151°05.5'E, Coast & Wetlands Society Survey, 8 October 
1985, stn 1, 8. 2 specimens, AM P54284, Homebush Bay, Parramatta 
River, 33°50'S 151°05'E, Coast & Wetlands Society Survey, 8 October 
1985, stn 5, 6. 50 specimens, AM P41818, south end of Homebush Bay, 
New South Wales, 33°50.03’S 151°04.73’E, 2.5 m, Van Veen grab, P. 


Berents & party, 26 November 1992, site 25. 2 specimens, AM P54277, 
Cabarita, Parramatta River, 33°51’S 151°07’E, L. Garcia, 8 September 
1988, stn D7.rl. 1 specimen, AM P55249, Back Creek mangrove, Tuross 
Lake, 36°03.67’S 150°06.43'E, mud, Australian Museum Eurobodalla 
Shire Estuary Survey, 16 September 1974. 2 specimens, AM P36699, 
Shadrachs Creek, Twofold Bay, 37°04.8'S 149°52.5’E, Zostera, infauna, 
S. Keable & A. Reid, 26 June 1985, stn L2. 

Type locality. South West Rocks, Trial Bay, New South 
Wales, [approx. 30°53’S 153°03'E], tidal lagoon. 


282 Records of the Australian Museum (2005) Vol. 57 


Fig. 43. Quadrivisio sarina n.sp., holotype, female, 7.4 mm, AM P60482, paratype male, 8.1 mm, AM P60484, 
Armstrong Beach, Sarina, Queensland, Australia. Scales represent 0.5 mm. 


Description. Based on male, AM P56684. 

Head. Eyes absent; lateral cephalic lobes absent, lacking 
notch or slit, anteroventral corner rounded. Antenna 1 longer 
than antenna 2; peduncular article 1 subequal in length to 


article 2, without robust setae along posterior margin; 
peduncular article 2 geniculate with article 3; flagellum with 
33 articles; accessory flagellum with 2 articles. Antenna 2 
peduncular article 2 cone gland not reaching to end of 
peduncular article 3; article 4 shorter than article 5; 


Lowry & Springthorpe: Australian melitid amphipods 283 


than article 2, about as long as broad. Maxilla 1 inner plate 
setose along entire inner margin. 

Pereon. Gnathopod 1 coxa anteroventral comer produced, 
rounded, posteroventral comer notch absent; mems without 
posterodistal spine; propodus palm nearly transverse, convex, 
defined by posterodistal comer, defined by posterodistal robust 
setae. Gnathopod 2 not sexually dimorphic, subchelate; coxa 
posteroventral comer notch absent; mems with rounded or 


subquadrate posteroventral corner; carpus short; propodus 
without medial depression, palm acute, concave, smooth, with 
sparse robust setae, defined by posterodistal robust setae, 
defined by posteroventral corner; dactylus apically acute/ 
subacute. Pereopods 5-6 basis posterior margin straight, 
posteroventral comer narrowly rounded or subquadrate; carpus 
and propodus with many long, slender setae along anterior 
margin. Pereopod 7 basis posterior margin convex, with 


Fig. 45. Victoriopisa australiensis (Chilton, 1923), male, 11.1 mm, AM P56684, Boambee Creek, Sawtell, 
New South Wales, Australia. Scales for MD, MX1IP represent 0.2 mm, remainder represent 0.5 mm. 


284 Records of the Australian Museum (2005) Vol. 57 


posterior margin smooth or minutely castelloserrate, 
posteroventral comer broadly rounded; mems posterodistal 
margin broadly rounded. 

Pleon. Epimeron 1 posteroventral comer narrowly rounded or 
subquadrate. Epimera 1-2 posteroventral margin without 
spines above posteroventral corner. Epimeron 3 posterior 
margin smooth, posteroventral comer with small acute spine 


or with strongly produced acute spine. Uropod 1 peduncle with 
2 basofacial robust setae. Uropod 3 inner ramus scale-like, 
much shorter than outer ramus; outer ramus much longer (more 
than 2x length) than peduncle, 2-articulate. Telson without 
apical robust setae, apical conical extension absent. 

Habitat. Estuarine; littoral; mangrove mud flats and 
seagrasses; 0 to 3 m depth. 


Lowry & Springthorpe: Australian melitid amphipods 285 


Remarks. The holotype (AM P5852) was originally 
deposited in Australian Museum, but assumed to be lost 
(Springthorpe & Lowry, 1994). Consequently another 
specimen from near the type locality (AM P56684) was 
illustrated and used for the description. Since then the type 
specimen has been located. 

Victoriopisa australiensis and V. marina differ most 
obviously from each other as follows. In V. australiensis 
the male gnathopod 2 has a short carpus and the palm of 
the propodus is broadly excavate and not bordered by robust 
setae, whereas in V. marina the carpus is compressed and 
the palm is narrowly excavate and bordered by robust setae. 
Victoriopisa australiensis has a large, broadly expanded 
basis and merus on pereopod 7 with many slender setae 
distally. In V. marina the basis and merus are narrower and 
the distal part of the pereopod is not nearly as setose. 

Distribution. New South Wales : Southwest Creek, Trial Bay 
(Chilton, 1923); Boambee Creek, Sawtell; Wallis Lake; 
Fullerton Cove, Hunter River; Hawkesbury River (Jones et 
al., 1986; Jones, 1987); Pittwater; Parramatta River; Botany 
Bay; Tuross; Twofold Bay (all AM). 

Australian geographic areas. Southeastern Australia. 

Victoriopisa marina n.sp. 

Figs. 47-49 

Victoriopisa sp. 2 Jones et al., 1986: 541 
Victoriopisa sp. Jones, 1987: 623. 

Type material, holotype, 6, 8.1 mm, AM P61227, Cobblers (Bate 
Bay), New South Wales, Australia, [approx. 34°07'S 151°10'E], 65-70 
m, grab, Ecology Lab, October 1990, stn T3-135. 1 paratype, 2, 7.4 
mm, AM P41961, 800 m southwest of airport runway, Botany Bay, New 
South Wales, Australia, 33°58.33'S 151°10.22'E, 7 m, Australian Museum 
party, 7 April 1992, stn NSW-771. 

Additional material examined. New South Wales: l S , l $, am 

P53976, Royal Motor Yacht Club, Pittwater, 33°39.2’S 151°18.0E, fine 
mud, 12 m, C. Rose, December 1992, stn RMYC A5. 1 S, AM P61360, 
between Juno Head and Hungry Beach, Hawkesbury River, 33°34’S 
151°16E, muddy sand, 10 m, Smith-Mclntyre grab, A.R. Jones & C. 
Watson-Russell, 7 August 1979, stn HES 1-3. 

Type locality. Cobblers (Bate Bay), New South Wales, 
Australia [approx. 34°07’S 151°10’E], 65-70 m. 

Description. Based on holotype male, AM P61227 and 
paratype female, AM P41961. 


Head. Eyes absent; lateral cephalic lobes absent, 
lacking notch or slit, anteroventral corner rounded. 
Antenna 1 longer than antenna 2; peduncular 
article 1 shorter than article 2, without robust 


setae along posterior margin; peduncular article 2 geniculate 
with article 3; flagellum with 30 articles; accessory 
flagellum with 2 articles. Antenna 2 peduncular article 2 
cone gland not reaching to end of peduncular article 3; 
article 4 shorter than article 5; flagellum with 2-3 articles. 
Mandible palp article 3 rectolinear, with setae mostly 
terminal, longer than article 1; article 2 subequal to or longer 
than article 3; article 1 produced distally, shorter than article 
2, about twice as long as broad. Maxilla 1 inner plate setose 
along entire inner margin. 

Pereon. Gnathopod 1 coxa anteroventral corner produced, 
rounded, posteroventral corner notch absent; merus without 
posterodistal spine; propodus palm acute, convex, defined 
by posterodistal corner, defined by posterodistal robust 
setae. Gnathopod 2 not sexually dimorphic; subchelate; 
coxa posteroventral corner notch absent; merus with 
subquadrate posteroventral corner; carpus compressed; 
propodus without medial depression, without strong 
concentration of setae, palm acute, concave or sinusoidal, 
sculptured, with sparse robust setae and with group of 
anterodistal robust setae, defined by posterodistal robust 
setae, defined by posteroventral corner; dactylus apically 
acute/subacute. Pereopod 5 basis posterior margin straight, 
posteroventral corner narrowly rounded or subquadrate. 
Pereopod 6 coxa anterior lobe ventral margin slightly 
produced, rounded; basis posterior margin straight, 
posteroventral corner narrowly rounded or subquadrate. 
Pereopod 7 basis posterior margin convex, with posterior 
margin smooth or minutely castelloserrate, posteroventral 
corner broadly rounded. 


Fig. 47. Victoriopisa marina n.sp., holotype, 
male, 8.1 mm, AM P61227, Cobblers, Bate Bay, 
New South Wales, Australia. 


286 Records of the Australian Museum (2005) Vol. 57 


Fig. 48. Victoriopisa marina n.sp., holotype, male, 8.1 mm, AM P61227, Cobblers, Bate Bay, New South Wales, 
Australia. Scales for Al, A2, U3 represent 0.5 mm, remainder represent 0.2 mm 


Pleon. Epimeron 1 posteroventral corner broadly rounded. 
Epimera 1-2 posteroventral margin without spines above 
posteroventral corner. Epimeron 3 posterior margin smooth, 
posteroventral comer with small acute spine or with strongly 
produced acute spine. Urosomite 1-3 dorsally smooth. 
Uropod 3 inner ramus scale-like, much shorter than outer 
ramus; outer ramus much longer (more than 2x length) than 
peduncle, 2-articulate. Telson without apical robust setae, 
apical conical extension absent. 

Habitat. Marine and estuarine; littoral and continental shelf; 
fine to sandy mud, 7 to 70 m depth. 

Remarks. Victoriopisa marina is most similar to species of 
Victoriopisa with slender seventh pereopods. It differs from 
all of these species in the extremely broadened palm of 
gnathopod 1 and the shape of the palm of gnathopod 2. 

Distribution. New South Wales : Hawkesbury River (Jones 
et al., 1986; Jones, 1987); Pittwater; Botany Bay; Cobblers, 
Bate Bay (all AM). 

Australian geographic areas. Southeastern Australia. 


Melita group 
Dulichiella Stout, 1912 

Dulichiella australis (Haswell) 

Figs. 50-53 

Melita australis Haswell, 1879a: 264, pi. 9, figs 6-7-Haswell, 
1882: 252. 

Melitafresnelii- Stebbing, 1906:423 (inpart).-Stebbing, 1910a: 

596-597, 642.-Chilton, 1921b: 70.-Hale, 1927: 314. 

Melita appendiculatus- Stebbing, 1906: 428 (in part). 
Dulichiella australis.-Karamm & Barnard, 1979: 152.-Barnard 
& Barnard, 1983: 668.-Hutchings et al., 1989: 362. 

Type material. 5 SYNTYPES, AM G5393; 1 Syntype, AM P3495, 
Port Jackson, New South Wales, Australia, [approx. 33°51'S 151°16'E], 
[AM Old Collection] 

Additional material examined. Queensland: many specimens, 
AM P61134, Lizard Island, Queensland, 14°40'S 145°28'E, October 
1978. New South Wales: 3 specimens, AM P55002, northeast of Marys 
Rock, Cook Island, 28°11.42'S 153°34.79'E, tan sponge with large osculi, 
18 m, G.D.F. Wilson, 8 June 1993, stn NSW-810. Several specimens, 
AMP56651; 1 specimen, AM P57284, 100 m north west of Julian Rocks, 


Lowry & Springthorpe: Australian melitid amphipods 287 


Byron Bay, 28°36.8'S 153°37.8'E, rock with finger sponge, 15 m, E.L. 
Albertson, R.T. Springthorpe & G.D.F. Wilson, 3 Marchl992, stn NSW- 
635. 2 specimens, AM P57684, same locality, erect plate sponge, 15 m, 
E.L. Albertson, R.T. Springthorpe & G.D.F. Wilson, 3 Marchl992, stn 
NSW-639. 5 specimens, AM P57285, same locality, mixed sponges, 17 
m, stn NSW-642. 7 specimens, AM P56650, 100 m north west of Split 
Solitary Island, 30°14.0’S 153°10.8'E, plate coral, solitary ascidian, 
gorgonian & sponges, 15-17 m, G.D.F. Wilson, 7 March 1992, stn NSW- 
687. 4 specimens, AM P57105, same locality, under rock ledges, 15-17 
m, R. Gentle (URG), 7 March 1992, stn NSW-697. 4 specimens, AM 
P57286, same locality, lace bryozoan, 17 m, R.T. Springthorpe & S.J. 
Keable, 7 March 1992, stn NSW-681. 5 specimens, AM P57287, same 
locality, sponge, 15-17 m, R.T. Springthorpe, 7 March 1992, stn NSW- 
683. 1 specimen, AM P57288, same locality, mixed red algae, 17 m, 
S.J. Keable, 7 March 1992, stn NSW-693. 3 specimens, AM P27266, 
Split Solitary Island, [approx. 30°15'S 153°12'E], 10 m, J. Marshall, 23 
March 1978. 5 specimens, AM P56653, Coffs Harbour Jetty, Coffs 
Harbour, 30°18.4'S 153°08.5'E, orange sponge on jetty pilings, 6 m, 
R.T. Springthorpe, 9 March 1992, stn NSW-729. Several specimens, 
AM P56654, same locality, Pyura praeputialis on jetty pilings, 8 m, 
P.B. Berents & S.J. Keable, 9 March 1992, stn NSW-733. Many 
specimens, AM P56655, same locality, arborescent sponge on jetty 


pilings, 7 m, S.J. Keable, 9 March 1992, stn NSW-735. 24 d d, 269 9, 
AM P57103, same locality, worm tubes encrusted with sponge on jetty 
pilings, 6 m, R.T. Springthorpe, 9 March 1992, stn NSW-725. 1d, AM 
P57104, same locality, coral scrapings on jetty pilings, 6 m, R.T. 
Springthorpe, 9 March 1992, stn NSW-738. 11 9 s, 5 d d, AM P57289, 
same locality, coral scrapings on jetty pilings, 6 m, R.T. Springthorpe, 9 
March 1992, stn NSW-726. 2 specimens, AM P57290, same locality, 
Diopatra tubes at base of jetty pilings, 6.5 m, S.J. Keable & R.T. 
Springthorpe, 9 March 1992, stnNSW-730. 289 2,16d d, AMP57291, 
same locality, finger sponge on jetty pilings, 4 m, R.T. Springthorpe, 9 
March 1992, stn NSW-734. Many specimens, AM P61141, same locality, 
Diopatra tubes at base of jetty pilings, 8.5 m, S.J. Keable, 9 March 
1992, stn NSW-728. 1 specimen, AM P56652, 150 m downstream of 
Fred Hansen Bridge, Boambee Creek, Sawtell, 30°20.4'S 153°05.5'E, 
exposed flat of sandy mud, 1.2 m, van Veen grab, Australian Museum 
party, 8 March 1992, stn NSW-724. 25 specimens, AM P3496, Port 
Stephens, [approx. 32°42'S 152°06'E], [AM Old Collection], 1 specimen, 
AM P23158, 500 m east of Burwood Beach, 32°57.52'S 151 0 44.72’E, 
coarse sand bottom, 14 m, Shipek collection, Australian Museum Hunter 
District Water Board Survey, 18 December 1975, stn HDWBS06030101, 
transect 3. 1 d, AM P53923, west side of Box Head, Broken Bay, 33°33'S 
151°2TE, bryozoan on rocky substrate with small crinoid Antedon 


288 Records of the Australian Museum (2005) Vol. 57 


incomoda, 15 m, J.K. Lowry & R.T. Springthorpe, 22 November 1982, 
stn NSW-158. Many specimens, AM P53924, same locality, bryozoan, 
stn NSW-159. Many specimens, AM P53925, same locality, sponges, 
stnNSW-165.4 specimens, AM P53926, same locality, sponge, stnNSW- 
166. Several specimens, AM P53927, same locality, stn NSW-167. Many 
specimens, AM P53928; Id, AM P60600; 19, AM P60601, same 
locality, sponge Echinoclathria sp., stn NSW-173. 4 specimens, AM 
P22493, east of Long Reef, [approx. 33°44'S 151°22’E], 40 m, Australian 
Museum Shelf Benthic Survey, 29 June 1972. 2 specimens, AM P22494, 
same locality, 36 m, 11 May 1972. 5 specimens, AM P22495, same 
locality, 38 m, 24 August 1972. Many specimens P22488, east of North 
Head, Port Jackson, [approx. 33°49’S 151°20’E], host sponge: cf 
Teichonella labrinthica, 21 m Australian Museum Shelf Benthic Survey, 
20 February 1973, transect 9. 1 specimen, AM P22489, same locality, 
host sponge: Polymastea craticia. 1 specimen, AM P22490, same locality, 
host sponge: Halme gigantea, 25 m, 26 February 1974, transect 07. 3 
specimens, AM P22497, same locality, host sponge: Polymastea craticia, 

19 m, 19 February 1973, transect 10. 3 specimens, AM P22498, same 
locality, host sponge: Halme gigantea, 19 m, 19 February 1973 transect 
10. 1 specimen, AM P22499, same locality, sponge 19 m, 19 February 
1973 transect 10. Many specimens, AM P22501, same locality, host 
sponge: cf Teichonella labrinthica, 19 m, 19 February 1973 transect 10. 

20 specimens, AM P22491, east of North Head, Port Jackson, [approx. 
33°49.5’S 151°18F], 28 m, Australian Museum Shelf Benthic Survey, 
13 December 1971. 1 specimen, AM P22492, same locality, among 
sewage outfall, 12 May 1972. Several specimens, AM PI 8198, off Sydney 
Heads, [approx. 33°50'S 151°28’E], 119 m, [AM Old collection]. Many 
specimens, AM P22496, east of South Head, Port Jackson, [approx. 
33°50'S 151°18'E], host sponge: Halme gigantea, 21 m, Australian 
Museum Shelf Benthic Survey, February 1974, transect 11.3 specimens, 
AM P5334, Port Jackson, [approx. 33°51’S 151°16'E], W.A. Haswell, 
1918. 8 specimens, AM P61140, north east corner of Clark Island, Port 
Jackson, 33°51.85'S 151°14.47F, red algae, 2 m, I. Takeuchi & D. Bray, 
17 April 1996, stn NSW-1250. 2dd, AM P63067, Brotherson Dock 
Berth 2, Botany Bay, 33°58.2'S 151°12.6'E, pylon scrapings, 7 m, NSW 
Fisheries/CRIMP Survey, 22 October 1998, stn BB BD2 P2-7. 1 
specimen, AM P63068, Bulk Liquids Berth, Botany Bay, 33°58.5'S 
151°12.6'E, pylon scraping, 7 m, NSW Fisheries/CRIMP Survey, 19 
October 1998, stn BB BLB P3-7. 2 9 9, AM P63069, Channel Marker 
4, Botany Bay, 33°59.3’S 151°12.6’E, pylon scraping, 7 m, NSW 
Fisheries/CRIMP Survey, 21 October 1998, stn BB CH4 Pl-7. 2 
specimens, AM P63083, same locality, pylon scraping, 3 m, NSW 
Fisheries/CRIMP Survey, 21 October 1998, stn BB CH4 Pl-3. 3 
specimens, AM P58236, outer end of Kurnell Pier, Botany Bay, 34°00.2’S 


151°12.5'E, pylon scrapings, 7 m, NSW Fisheries/CRIMP Survey, 21 
October 1998, stn BB KP1 Pl-7. 1 9, AM P63070, Kurnell Pier (near 
shoreline), Botany Bay, 34°00.5'S 151°12.7'E, pylon scrapings, 3 m, 
NSW Fisheries/CRIMP Survey, 21 October 1998, stn BB KP2 P2-3. 1 
specimen, AM P2486, 3-4 km off Botany Bay, [approx. 34°05’S 
151°15'E], mud, 91-95 m, E.R. Waite on HMCS Thetis, 11 March 1898, 
stn 37. 1 specimen, AM P2487, 5.5-6.5 km off Wattamolla, [approx. 
34°10'S 151°U'E], mud, 99-108 m, E.R. Waite on HMCS Thetis, 22 
March 1898, stn 57. 2 specimens, AM P63548, Boat Basin, Wollongong 
Harbour, 34°25.35'S 150°54.4’E, pylon scrapings, 3 m, NSW Fisheries/ 
CRIMP Survey, 17 May 2000, stn PK WHB Pl-3. 1 specimens, AM 
P63549, same locality, pylon scrapings, 0.5 m, NSW Fisheries/CRIMP 
Survey, 17 May 2000, stn PK WHB P2-0. Many specimens, AM P63550 
same locality, pylon scrapings, 3 m, NSW Fisheries/CRIMP Survey, 17 
May 2000, stn PK WHH. 1 specimens, AM P44306, off Wollongong, 
34°26.54’S 150°57.98F, Globigerina ooze, 50 m, baited trap, J.K. Lowry 

6 K. Dempsey on MV Robin E, 27-28 March, 1994, SEAS project, stn 
NSW-939. Many specimens, AM P63543, Inflammable Liquids Berth, 
Port Kembla Outer Harbour, 34°27.95’S 150°54.25'E, pylon scrapings, 

7 m, NSW Fisheries/CRIMP Survey, 13 May 2000, stn PK ILB P2-7. 4 
specimens, AM P63546, east end No.6 Jetty, Port Kembla Outer Harbour, 
34°28.25'S 150°54.1'E, pylon scrapings, 3 m, NSW Fisheries/CRIMP 
Survey, 14 May 2000, stn PK J60 Pl-3. 2 specimens, AM P63547, 
same locality, pylon scrapings, 7 m, NSW Fisheries/CRIMP Survey, 14 
May 2000, stn PK J60 Pl-7. Many specimens, AM P63544, south end 
No.3 Jetty, Port Kembla Outer Harbour, 34°28.6’S 150°54.5'E, pylon 
scrapings, 3 m, NSW Fisheries/CRIMP Survey, 16 May 2000, PK J30 
P2-3. 2 specimens, AM P63545, same locality, pylon scrapings, 7 m, 
NSW Fisheries/CRIMP Survey, 16 May 2000, stn PK J30 P2-7. Id, 
19, AM P53870, off Moona Moona Creek, Jervis Bay, 35°02.9'S 
150°41.0'E, from surface of ascidian Herdmania momus, in Ecklonia 
bed, 4.5 m, PB. Berents, 13 November 1981, stn NSW-249. 2 specimens, 
AM P53871, same locality, Ecklonia holdfasts, 5 m, P. Berents, 15 August 
1981, stn NSW-87. 1 specimen, AM P54274, Jervis Bay, 35°03'S 
150°44'E, sponge, scallop beds, 17 m, P. Berents, 13 August 1981. Many 
specimens, AM P30799; many specimens, AM P30800, south east of 
Tathra Head, 36°45-48’S 150°02-03'E, 64 m, trawl, FRV Kapala, 10 
June 1980, stn K80-07-02. 2dd, AM P36221, Murrumbulga Point, 
Twofold Bay, 37°04.7'S 149°53.1'E, subtidal rock platform, 2-9 m, S. 
Keable & E. Bamber, 11 December 1984, stn Ql. 1 specimen, AM 
P18318, Twofold Bay, 37°05'S 149°55F, W.A. Haswell. 108 9 9,92dd, 
AM P35980, Munganno Point, Twofold Bay, 37°06.2’S 149°55.7’E, 
subtidal wharf pile, 6 m, S. Keable, 10 October 1984, stn M5. Tasmania: 
1 d, AM P60602; many specimens, AM P61142, north side of Esperance 


Lowry & Springthorpe: Australian melitid amphipods 289 


represent 0.5 mm, remainder 
represent 0.2 mm. 


Point, D’Entrecasteaux Channel, Tasmania, 43°19.5'S 147°05.5'E, lace 
bryozoan, 13 m, S.J. Keable, J.K. Lowry & R.T. Springthorpe, 18 April 
1991, stn TAS-186. 3 specimens, AM P61143, same locality, 13 m, S.J. 
Keable, J.K. Lowry & R.T. Springthorpe, 18 April 1991. 3 9 9, AM 
P60603, same locality, sponges, lace bryozoan, red algae and IVittaticella 
sp., 13 m, S.J. Keable, J.K. Lowry & R.T. Springthorpe, 18 April 1991, 
stn TAS-187. South Australia: E6541, 3 specimens, Sanders Bank, 
Kangaroo Island, [approx. 35°50’S 137°15'E], 51 m, LIS Endeavour , 
1909-1914. 3 specimens, AM P5923, same locality. 1 specimen, E4849, 
Spencer’s Gulf, [approx. 34°00’S 137°00’E],LIS Endeavour , 1909-1914. 


Type locality. Port Jackson, New South Wales, Australia, 
[approx. 33°51’S 151°16’E]. 

Description. Based on a male, 10 mm, AM P60600 and a 
female, AM P60601. 

Head. Lateral cephalic lobes broad, truncated, lacking notch 
or slit, anteroventral corner subquadrate. Antenna 1 longer 
than antenna 2; peduncular article 1 shorter than article 2, 
with 4 or more robust setae along posterior margin; 


290 Records of the Australian Museum (2005) Vol. 57 


flagellum with at least 40 articles; accessory flagellum with 
6 articles. Antenna 2 peduncular article 2 cone gland 
reaching at least to end of peduncular article 3; article 4 
subequal to article 5; flagellum with about 12 articles. Mandible 
palp article 3 rectolinear, setae along both margins and terminal, 
longer than article 1; article 2 shorter than article 3; article 1 
produced distally, shorter than article 2, about as long as broad. 
Maxilla 1 inner plate with setae mainly terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner not 
produced, posteroventral corner notch present; merus 
without posterodistal spine; propodus palm nearly 
transverse, slightly convex or straight, defined by 
posterodistal corner, without posterodistal robust setae. 
Gnathopod 2 sexually dimorphic; left and right gnathopods 
unequal in size; coxa posteroventral corner notch present; 
(larger) chelate; merus with rounded posteroventral corner; 
carpus compressed; propodus, distolateral margin with 3 
rounded indistinct spines; palm angle obtuse, straight, 
posterodistal spine absent, without robust setae; dactylus 
apically blunt; (smaller) subchelate; merus with sharp 


posteroventral spine; carpus long; propodus palm straight, 
without posteroventral spine. Pereopod 5 basis posterior 
margin straight, posteroventral corner narrowly rounded or 
subquadrate; carpus and propodus with many long, slender 
setae along anterior margin; dactylus unguis anterior margin 
with accessory spine. Pereopod 6 coxa anterior lobe ventral 
margin slightly produced, rounded; basis posterior margin 
straight, posteroventral corner narrowly rounded or 
subquadrate; carpus and propodus with many long, slender 
setae along anterior margin; dactylus unguis anterior margin 
with accessory spine. Pereopod 7 basis posterior margin 
straight or slightly subsigmoidal, with posterior margin 
smooth or minutely castelloserrate, posteroventral corner 
narrowly rounded or subquadrate; dactylus unguis anterior 
margin with accessory spine. 

Pleon. Pleonites 1-3 with dorsal serrations, with dorsodistal 
spine. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner with strongly produced acute 


Lowry & Springthorpe: Australian melitid amphipods 291 


Fig. 53. Dulichiella australis (Haswell, 1879a), male, 10.0 mm, AM P60600, female, AM P60601, west side of 
Box Head, Broken Bay, New South Wales, Australia. Scales represent 0.5 mm. 


spine. Urosomite 1 with broad dorsal midline gape, rarely 
bearing midline spines, dorsally bicarinate. Urosomite 2 
posterior margin serrate, with two groups of 1-3 small 
dorsolateral robust setae. Uropod 3 inner ramus scale-like, 
much shorter than outer ramus; outer ramus much longer 
(more than 2x length) than peduncle, 2-articulate. Telson 
without apical robust setae, apical conical extension absent. 

Female (sexually dimorphic characters). Gnathopod 2 
subchelate; merus with sharp posteroventral spine; carpus 
short; propodus without medial depression, palm slightly 
acute, straight or slightly concave, smooth, without robust 
setae, defined by posterodistal robust setae, defined by 
posteroventral corner; dactylus apically acute/subacute. 
Pereopod 7 basis posterior margin straight. 

Habitat. Marine; littoral and continental shelf; living among 
bryozoans, sponges, ascidians and algae, 4.5 to 120 m depth. 

Remarks. There are at least two distinctive species of 
Dulichiella along the east coast of Australia. Their ranges 
overlap along the Great Barrier Reef. Dulichiella australis 


and D. pacifica are easily distinguished by the spines on 
the distolateral margin of male gnathopod 2. In D. australis 
there are 3 rounded, rather indistinct spines and in D. 
pacifica there are 4 acute distinct spines. In addition 
pereopods 6 and 7 are more setose in D. australis. 

Distribution. Queensland: Lizard Island (AM). New South 
Wales: Julian Rocks, Byron Bay; Marys Rock, Cook Island; 
Split Solitary Island; Coffs Harbour; Boambee Creek, 
Sawtell (all AM); Tacking Point (Haswell, 1879a); Manning 
River (Stebbing, 1910a); Port Stephens; off Burwood Beach; 
Box Head, Broken Bay; off Long Reef (all AM); Port 
Jackson (Haswell, 1879a); Clark Island, Port Jackson; 
Botany Bay; off Wattamolla; off Wollongong; Jervis Bay 
(all AM); Twofold Bay (Hutchings et al., 1989). Victoria: 
Western Port (NMV). Tasmania : Esperance Point, 
D’Entrecasteaux Channel (AM). South Australia: Sanders 
Bank, Kangaroo Island (Chilton, 1921b); Spencers Gulf 
(AM). 

Australian geographic areas. Northeastern Australia, 
southeastern Australia, southern Australia. 


292 Records of the Australian Museum (2005) Vol. 57 


Dulichiella pacifica n.sp. 

Figs. 54-57 

Dulichiella appendiculata- Berents, 1983: 111, fig. 9.-Ledoyer, 
1986: 187, fig. 9S. 

Type material. Holotype, 8 “a”, 4.4 mm, AM P61112; 1 paratype, 
9 ,5.9 mm, AM P61113; 9 paratypes, 1 8 , 5 ovigerous females, 3 non- 
ovigerous females, AM P61117, between Tandai Point and Koilo Point, 
Guadalcanal, Solomon Islands, 9°22.5'S 159°52.2'E, coral rubble, 10 
m, R.T. Springthorpe, 24 September 1991, SI-3; 1 paratype, 8 “b”, 4.1 
mm, AM P61114; 4 PARATYPES, males, AM P61115, Tandai Point, 
Guadalcanal, Solomon Islands, 9°23'S 159°52.5'E, sand with low algal 
turf, 20 m, hand dredge, R.T. Springthorpe, 11 October 2001, SI-36; 1 
PARATYPE, 8, AM P61116, same locality, black finger sponges from 
rubble bottom, 16 m, R.T. Springthorpe, 11 October 2001, SI-38. 

Additional material examined. Queensland: 1 specimen, AM 
P30123, outer slope, Yonge Reef, 14°36’S 145°38'E, reef rock with 
Halimeda and Lithothamnion, 36 m, PB. Berents & P.A. Hutchings, 9 
January 1975, stn 75 LIZ D-l. 6 specimens, AM P30124, Watsons Bay, 
Lizard Island, 14°40'S 145°28E, Halophila, Caulerpa, Udotea & drift 
algae, 7 m, J.K. Lowry & P.A. Hutchings, 29 September 1978, stn LI-2. 
5 specimens, AM P30125, fringing reef between Bird Islet and South 
Island, Lizard Island, 14°40’S 145°28’E, Halophila , mixed algae, 
sediment from grass beds off reef, 24 m, J.K. Lowry, 9 October 1978, 
stn LI-27.2 specimens, AM P30126, same locality, PC. Terrill, 9 October 
1978, stn LI-28. New Caledonia: 1 specimen, AM P47550, off IlotMaitre, 
New Caledonia, 22°19.35’S 166°25.85’E, dead branching coral, 21 m, 
ORSTOM divers, 10 November 1995, stn NCL-72. 3 specimens, AM 
P47556; 7 specimens, AM P47563, same locality, large sea fan covered 
in sponges & other epiflora & epifauna, air lift, 21 m, J.K. Lowry, 10 
November 1995, stn NCL-65. 1 specimen, AM P47522, Hot Maitre, New 
Caledonia, 22°19.61’S 166°24.07’E, Padina -like alga, 10.5 m, G. 
Bargibant, ORSTOM, 14 November 1995, stn NCL-98. Several 
specimens, AM P47528, same locality, coralline algal “reef’, 10.5 m, 
G. Bargibant, ORSTOM, 14 November 1995, stnNCL-99. 5 specimens, 
AM P48322, Hot Maitre, New Caledonia, 22°20.57'S 166°25.43’E, red 
alga, 20 m, I. Takeuchi, 7 November 1995, stn NCL-38. 3 specimens, 
AM P48332, same locality, dead coral, 20 m, ORSTOM divers, 7 
Novemberl995, NCL-40. 10 specimens, AM P47334; 2 specimens, AM 
P47347, between Ilot Maitre and Croissant reef, New Caledonia, 
22°19.7’S 166°23.3’E, Sargassum sp., 10 m, I. Takeuchi, 6 November 
1995, stn NCL-28, 30. 1 specimen, AM P47506, between Ilot des 
Goelands and Grand Recif Abore, New Caledonia, 22°24.10’S 
166°20.90’E, Halimeda sp. fine red alga, 10 m, J.K. Lowry, 16 November 
1995, stn NCL-111. 1 specimen, AM P47590, between He Nge & Seche 
Croissant, New Caledonia, 22°19.41'S 166°20.89E, purple bryozoan, 
Iodyctium buchneri, 20 m, ORSTOM divers, 9 November 1995, stn NCL- 


57. 1 specimen, AM P47967, same locality, sediment sample (sand), 20 m, 
J.K. Lowry, 9 November 1995, stn NCL-56. 1 specimen, AM P48354, 200 
m off Poe Plage, New Caledonia, 21°36.41'S 165°22.73'E, Sargassum, 1-2 
m, J.K. Lowry, 19 November 1995, stn NCL-212.4 specimens, AM P48379, 
fringing reef Thio, New Caledonia, dead coral, 1 m, A.A. Myers, 20 
November 1995, stn NCL-217. Several specimens, AM P48460, same 
locality, “felt like” alga, 1 m, A.A. Myers, 20 Nov 1995, NCL-218. 
Singapore: \8, AM P61135, Pulau Sakia, Singapore, [approx. 01°16'N 
103°42'E], Smith-Mclntyre grab, C.S.C. Lee, University of Singapore, 23 
May 1991, stn E, ZRC-1991-16108. Many specimens, AM P61136, Changi 
Floating Fish Farm, Singapore, [approx. 01°24’N 103°58'E], 1 m, J.B. 
Sigurdsson, 7 May 1997, associated with biofouling. 

Type locality. Between Tandai Point & Koilo Point, 
Guadalcanal, Solomon Islands, (9°22.5’S 159°52.2'E), coral 
rubble. 

Description. Based on holotype male “a”, AM P61112, 
paratype female, AM P61113, and paratype male “b”, AM 
P61114. 

Head. Lateral cephalic lobes broad, truncated, lacking notch 
or slit, anteroventral corner subquadrate. Antenna 1 slightly 
longer than antenna 2; peduncular article 1 shorter than 
article 2, with 3 robust setae along posterior margin; 
flagellum with about 28 articles; accessory flagellum with 
4 articles. Antenna 2 peduncular article 2 cone gland 
reaching at least to end of peduncular article 3; article 4 
subequal to article 5; flagellum with 12 articles. Mandible 
palp article 3 rectolinear, setae along both margins and 
terminal, longer than article 1; article 2 subequal to article 
3; article 1 produced distally, shorter than article 2, about 
as long as broad. Maxilla 1 inner plate with setae mainly 
terminal. 

Pereon. Gnathopod 1 coxa anteroventral corner not 
produced, posteroventral corner notch present; merus 
without posterodistal spine; propodus palm acute, convex, 
without posterodistal corner, defined by posterodistal robust 
setae. Gnathopod 2 sexually dimorphic; left and right 
gnathopods unequal in size; coxa posteroventral corner 
notch present; (larger) chelate; merus with rounded 
posteroventral corner; carpus compressed; propodus, 
distolateral margin with 4 acute distinct spines; palm angle 


Fig. 54. Dulichiella pacifica n.sp., paratype, male “b”, 4.1 mm, AM P61114, Tandai Point, Guadalcanal, Solomon Islands. 


Lowry & Springthorpe: Australian melitid amphipods 293 


Fig. 55. Dulichiellapacifica n.sp., holotype, male “a”, 4.4 mm, AM P61112, Tandai Point, Guadalcanal, Solomon 
Islands. Scales for MDp, MX1IP represent 0.1 mm, remainder represent 0.2 mm. 


obtuse, straight, posterodistal spine absent, without robust 
setae; dactylus apically blunt; (smaller) subchelate; merus 
with sharp posteroventral spine; carpus long; propodus palm 
straight, without posteroventral spine. Pereopod 5 basis 
posterior margin straight, posteroventral corner narrowly 
rounded or subquadrate; dactylus unguis anterior margin 
with accessory spine. Pereopod 6 coxa anterior lobe ventral 
margin slightly produced, rounded; basis posterior margin 
slightly concave, posteroventral corner narrowly rounded 
or subquadrate; dactylus unguis anterior margin with 
accessory spine. Pereopod 7 basis posterior margin straight, 
with posterior margin smooth or minutely castelloserrate, 
posteroventral corner narrowly rounded or subquadrate; 
dactylus unguis anterior margin with accessory spine. 

Pleon. Pleonites 1-3 with dorsal serrations, with dorsodistal 
spine. Epimeron 1 posteroventral corner with small acute 


spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner with strongly produced acute 
spine. Urosomite 1 with broad dorsal midline gape, rarely 
bearing midline spines, dorsally bicarinate. Urosomite 2 
posterior margin serrate. Uropod 3 inner ramus scale-like, 
much shorter than outer ramus; outer ramus much longer 
(more than 2x length) than peduncle, 2-articulate. Telson 
without apical robust setae, apical conical extension absent. 

Female (sexually dimorphic characters). Gnathopod 2 
subchelate; merus with sharp posteroventral spine; carpus 
long; propodus without medial depression, palm acute, 
concave, smooth, with sparse robust setae, defined by 
posterodistal robust setae, defined by posteroventral corner; 
dactylus apically acute/subacute. Pereopod 7 basis posterior 
margin sub sigmoidal. 


294 Records of the Australian Museum (2005) Vol. 57 


Fig. 56. Dulichiella pacifica n.sp., holotype, male “a”, 4.4 mm, AM P61112, paratype, female, 5.9 mm, AM 
P61113, Tandai Point, Guadalcanal, Solomon Islands. Scales represent 0.2 mm. 


Habitat. Marine; littoral; coral rubble, sponges, sand with 
low algal turf, 10 to 20 m depth. 

Etymology. Named for its widespread distribution in the 
Pacific Ocean. 

Remarks. This distinctive species was illustrated from 
Senegal by Ledoyer (1986). It is common in Singapore and 
occurs along the archipelago that runs from New Guinea to 
Fiji. A closely related, undescribed species has been reported 
from Florida and Bermuda by LeCroy (2000). See remarks 
under D. australis for differences between these species. 

Distribution. Queensland : Yonge Reef, Great Barrier Reef; 
Lizard Island (Berents, 1983; AM). 

Extrinsic distribution. Senegal, Singapore, New Guinea, 
Solomon Islands, New Caledonia. 

Australian geographic areas. Northeastern Australia. 


Melita Leach, 1814 
Melita ophiocola n.sp. 

Figs. 58-61 

Type material. Holotype, 9, 9.6 mm, AM P55192, west side. Bottle 
and Glass Rocks, Port Jackson, 33°50.9'S 151°16.15'E, 4 m, J.K. Lowry 
& R.T. Springthorpe, 23 March 1982, stn NSW-100. Paratypes (114): 
1 paratype, 6, AM P46263; 1 paratype, 9, AM P46264, type locality, 
10 m, J.K. Lowry & R.T. Springthorpe, 21 January 1982, stnNSW-97; 8 
paratypes, AM P25456, type locality, 9 m, G.D. Lenwick, 29 August 
1977; 1 paratype, AM P55193; 1 paratype, AM P55194; 4 paratypes, 
AM P55199; 9 paratypes, AM P55200, type locality; 27 paratypes, AM 
P55195; 5 paratypes, AM P55196; 4 paratypes, AM P55197; 1 paratype, 
AM P55198, type locality, 4 m, 23 April 1982, stn NSW-101; 2 paratypes, 
AM P55236, type locality, 26 Leb 1982, stn NSW-99; 18 paratypes, AM 
P55241; 18, AM paratypes, AM P55242, type locality, 4 m, 26 November 
1982, stn NSW-177; 11 paratypes, AM P55245, type locality, 4 m, R.T. 
Springthorpe & P.N. Weber, 24 December 1982, stn NSW-267; 3 
paratypes, AM P55240, south of Vaucluse Point, Port Jackson, New South 


Lowry & Springthorpe: Australian melitid amphipods 295 


Wales, Australia, 33°52'S 151°17'E, under rocks, shelley substrate, in 
association with ophiurid Ophionereis schayeri, 2 m, J.K. Lowry & R.T. 
Springthorpe, 22 June 1982, stn NSW-107. 

Additional material examined. New South Wales: 1 2, AM P62903, 
northern cove of Boondelbah Island, Port Stephens, 32°42.28'S 
152°13.47'E, under small boulders, 19.6 m, 


AM P31161 (4); AM P31162 (3); AM P31163 (1); AM P31164 (1); AM 
P31165 (2); AM P31166 (15); AM P31167 (14); AM P31168 (2); AM 
P31169 (1); AM P31170 (1); AM P31171 (4); AM P31172 (2); AM 
P31173 (11); AM P31174 (7); AM P31175 (8); AM P31176 (12); AM 
P31177 (2); AM P31178 (2), Little Box Head, Broken Bay, [approx. 
33°33'S 151°16E], sandy-muddy, very fine silt, 13 m, S. Arnam & R. 
Springthorpe, 18 November 1980. 68 specimens, AM P31142 (7); 


296 Records of the Australian Museum (2005) Vol. 57 


Fig. 59. Melita ophiocola n.sp., holotype, female, 9.6 mm, AM P55192, Bottle and Glass Rocks, Port Jackson, 
New South Wales, Australia. Scales for MD, MX1, MX2, MP represent 0.1 mm, remainder represent 0.2 mm. 


AM P31159 (4); AM P31160 (6), 90 m south of Fairlight Pool, Port 
Jackson, 33°48.1'S 151°16.3'E, under stones, muddy sand, shell, in 
association with ophiuroid Ophionereis schayeri, 10 m, S. Arnam & R. 
Springthorpe, 13 October 1980.29 $, AM P45267, 50 m north of Cannae 
Point, Port Jackson, 33°49'S 151°, on brittle star: Ophionereis schayeri, 
5 m, PB. Berents, P. Castro, G. Towner, 21 June 1995, stn NSW-1138. 
67 specimens, AM P31134 (8); AM P31135 (2); AM P31136 (47); AM 
P31137 (1); AM P31138 (1); AM P31139 (5); AM P31140 (1); AM 
P31141 (2), type locality, 0.5 m, S. Arnam, J.K. Lowry & R.T. 
Springthorpe, 29 September 1980. 19 specimens, AM P55243 (17); AM 
P55244 (2), south of Vaucluse Point, Port Jackson, 33°50.9'S 


151°16.15’E, rocky with sediment patches, in association with ophiuroid 
Ophionereis schayeri, 2 m, J.K. Lowry & R.T. Springthorpe, 27 May 
1982, stn NSW-105. 7 specimens, AM P55234 (4); AM P55235 (3), 
South of Vaucluse Point, Port Jackson, 33°52’S 151°17'E, under rocks 
on soft substrate, in association with ophiuroid Ophionereis schayeri, 4 
m, R.T. Springthorpe & A. Murray, 30 September 1982, stn NSW-156. 
Id, AM P55233, type locality. 2 specimens, AM P55237, type locality, 
26 February 1982, stn NSW-99. 4 specimens, AM P55238 (2); AM 
P55239 (2), type locality, 10 m, 21 January 1982, stn NSW-97. 6 
specimens, AM P55246, type locality, 4 m, R.T. Springthorpe & P.N. 
Weber, 24 December 1982, stn NSW-267. 6 specimens, AM P36648, 


Lowry & Springthorpe: Australian melitid amphipods 297 


Murrumbulga Point, Twofold Bay, 37°04.7'S 149°53.1'E, subtidal rock 
platform, 3 m, S. Keable, 9 October 1984, stn Q2. Id, AM P55117, 
Munganno Point, Twofold Bay, 37°06’S 149°56’E, subtidal rock platform, 
6 m, S.J. Keable, 10 October 1984, stn M6. 

Type locality. Bottle and Glass Rocks, Port Jackson, New 
South Wales, Australia, 33°50.9’S 151°16.15'E, living in 


association with the ophiuroid Ophionereis schayeri under 
rocks, 4 m. 

Description. Based on holotype female, AM P55192 and 
paratype male, AM P55193. 


298 Records of the Australian Museum (2005) Vol. 57 


Fig. 61. Melita ophiocola n.sp., paratype male and female, AM P55240, in association with ophiuroid Ophionereis 
schayeri, south of Vaucluse Point, Port Jackson, New South Wales, Australia. Illustration by Sharne Weidland. 


Head. Lateral cephalic lobes broad, rounded, with 
anteroventral notch or slit, anteroventral corner rounded. 
Antenna 1 longer than antenna 2; peduncular article 1 longer 
than article 2, with 4 or more robust setae along posterior 
margin; flagellum with about 30 articles; accessory 
flagellum with about 5 articles. Antenna 2 peduncular article 
2 cone gland not reaching to end of peduncular article 3; 
article 4 subequal to article 5; flagellum with about 13 
articles. Mandible palp article 3 rectolinear, setose along 
straight medial margin, longer than article 1; article 2 
subequal to article 3; article 1 not produced, shorter than 
article 2. Maxilla 1 inner plate with setae mainly terminal. 


Pereon. Gnathopod 1 coxa anteroventral corner produced, 
rounded, posteroventral corner notch absent; merus without 
posterodistal spine; carpus about 3x as long as broad; 
propodus palm acute, slightly convex, defined by 
posterodistal corner, defined by posterodistal robust setae. 
Gnathopod 2 not sexually dimorphic; subchelate; coxa 
posteroventral corner notch absent; merus with sharp 
posteroventral spine; carpus compressed; propodus without 
medial depression, without strong concentration of setae, 
palm broadly rounded, convex, smooth, with sparse small 
robust setae, defined by posterodistal robust setae, without 
posteroventral corner; dactylus apically acute/subacute. 


Lowry & Springthorpe: Australian melitid amphipods 299 


Pereopod 5-6 basis posterior margin straight, posteroventral 
corner broadly rounded; dactylus unguis anterior margin 
with 2 or more accessory spines. Pereopod 7 basis posterior 
margin slighlty convex, with posterior margin smooth or 
minutely castelloserrate, postero ventral corner broadly 
rounded; dactylus unguis anterior margin with 2 or more 
accessory spines. 

Pleon. Epimeron 1 posteroventral corner with small acute 
spine. Epimera 1-2 posteroventral margin without spines 
above posteroventral corner. Epimeron 3 posterior margin 
smooth, posteroventral corner with strongly produced acute 
spine, posteroventral margin smooth or minutely serrate. 
Urosomite 1 with spines at midline, no conspicuous medial 
gape, with posterodorsal spine. Uropod 3 inner ramus scale¬ 
like, much shorter than outer ramus; outer ramus much 
longer (more than 2x length) than peduncle, 2-articulate. 
Telson each lobe with 3 apical/subapical robust setae, apical 
conical extension reaching at least halfway along longest 
seta. 

Female (sexually dimorphic characters). Pereopod 6 coxa 
anterior lobe ventral margin slightly produced, rounded. 

Habitat. Marine; living in association with the ophiuroid 
Ophionerieis schayeri under rocks. 

Depth zone. Littoral (1 to 13 m). 

Remarks. Melita ophiocola is an unusual and distinctive 
species of the genus. It is an obligate commensal of the 
brittle star Ophionereis schayeri (Muller & Troschel). 
Preliminary results of experimental data indicate that the 
largest adult amphipods tend to occur in heterosexual pairs 
on the host and occasionally more than one pair may occur. 
Over 80% of individuals sampled were juveniles and there 
can be as many as 50 young per host. Juveniles occur 
throughout the year. 

Mate selection experiments involving live animals 
showed that a significant number of large adults initially 
released on an overcrowded host move to another host and 
form heterosexual pairs (Peter Castro, pers. comm.). This 
apparent mobility may explain the high infection rates of 
over 90% observed in the field. 

In a highly unusual situation for species of Melita, there 
is virtually no sexual dimorphism between males and 
females. We assume that this is because the female and male 
are always present together on the host and so there is no 
need for precopulatory amplexus and its associated 
morphological modifications. 

Ophionereis schayeri is found in shallow water beneath 
large rocks and in crevices. It is more active at night when 
it may forage in the open. It moves by sinuous flexing of its 
arms and feeds on detritus that is transported to the mouth 
by the tube feet. It is assumed that M. ophiocola feeds on 
the detritus collected by its host. Melita ophiocola has a 
banded colour pattern of purple and cream dorsally with 
purple gnathopods and pereopods tending to cream distally. 
This is similar to the colour pattern of the aboral surface of its 
host. Melita ophiocola is able to move freely over the body 
and arms of the brittle star but lives mostly on the oral surface 
of the arms between the two rows of tube feet. 

Distribution. New South Wales: Boondelbah Island, Port 
Stephens; Broken Bay; Port Jackson; Twofold Bay (all AM). 


Acknowledgments. We thank Chandani Appadoo (University 
of Mauritius) for help with the illustrations of Mallacoota 
chandaniae. We thank Sarah LeCroy (Gulf Coast Research 
Laboratory, Ocean Springs, USA), Jens Oleson (Zoological 
Museum, Copenhagen) and Peter Dworschak (Natural History 
Museum, Vienna) for the loan of valuable material of Dulichiella. 
We thank Alan Myers (National University of Ireland) and John 
Bradbury (University of Adelaide) for their constructive comments 
on our manuscript. 


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Manuscript received 29 April 2003, revised 25 November 2003 and 

accepted 26 November 2003. 

Associate Editor: S.T. Ahyong. 


© Copyright Australian Museum, 2005 

Records of the Australian Museum (2005) Vol. 57: 303-320. ISSN 0067-1975 


Revision of Two Prioniodontid Species (Conodonta) 
from the Early Ordovician Honghuayuan Formation of 

Guizhou, South China 


Yong Yi Zhen 1 *, Jianbo Liu 2 , and Ian G. Percival 3 


1 Australian Museum, 6 College Street, Sydney NSW 2010, Australia 
yongyi@austmus.gov.au 

2 Department of Geology, 

Peking University, Beijing 100871, P.R. China 

jbliu@pku.edu.cn 

3 Geological Survey of New South Wales, Department of Primary Industries, 

State Geoscience Centre, 947-953 Londonderry Road, Londonderry NSW 2753, Australia 

ian. percival @ minerals. ns w. gov. au 


Abstract. The septimembrate conodonts Acodus triangularis (Ding in Wang, 1993) and Prioniodus 
honghuayuanensis n.sp., are described and illustrated from Guizhou, South China where both species 
are widely distributed in Lower Ordovician strata. The adenticulate A. triangularis, which ranges through 
the middle and upper parts of the Honghuayuan Formation, is morphologically more primitive than the 
denticulate P. honghuayuanensis, which is present from the uppermost Honghuayuan Formation into 
the lower part of the succeeding Dawan Formation. Prioniodus honghuayuanensis, elements of which 
were previously ascribed to Oepikodus communis (Ethington & Clark, 1964), appears to be closely 
related to a species of Prioniodus from the basal Whiterockian of Utah, North America. 


Zhen, Yong Yi, Jianbo Liu & Ian G. Percival, 2005. Revision of two prioniodontid species (Conodonta) from 
the Early Ordovician Honghuayuan Formation of Guizhou, South China. Records of the Australian Museum 57(2): 
303-320. 


Conodonts from the Honghuayuan Formation of Guizhou 
Province, South China were first studied by An (1987) in 
the Honghuayuan and Ganxi sections (Fig. 1). In the 
Honghuayuan section, An (1987) recognized three conodont 
assemblages with the lowermost consisting only of coniform 
species. The middle assemblage is characterized by the 
occurrence of Serratognathus diversus An, 1981. The upper 
assemblage is much higher in diversity, with the appearance 
of a number of pectiniform species of Prioniodus, 
Bergstroemognathus and Rhipidognathus. Similar faunas 
were also recorded from the Honghuayuan Formation of 

* author for correspondence 


Yanhe, northern Guizhou (An, 1987; X.Y. Chen et al., 1995), 
and are widely distributed in the Honghuayuan Formation 
and age equivalent units in South China (An et al., 1985; 
An, 1987; Wang, 1993). 

More recently, conodont samples were collected from 
Lower Ordovician sections in Guizhou and other parts of 
South China with the aim of revising the faunas to provide 
support for a more precise biostratigraphic correlation and 
age alignment, both regionally and internationally. This 
revision will assist our understanding of the origin, radiation 
and phylogeny of the prioniodontid and related clades 

www.amonline.net.au/pdf/publications/1448_complete.pdf 


304 Records of the Australian Museum (2005) Vol. 57 


Fig. 1. Locality maps of the study areas. (A) China showing the location of Guizhou Province. ( B ) northeast Guizhou showing the 
distribution of the Ordovician rocks (after Gao, 1976) and the locations of two sampled sections for this study: 1) Honghuayuan 
Section of Tongzi; 2) Ganxi Section ofYanhe. (C) simplified geological map of the area in the vicinity of Honghuayuan village, 
showing the location of the Honghuayuan Section (Geology information modified from 1:200 000 Geology Map of Tongzi by 
Guizhou Regional Geological Survey, 1978). 


through the late Early Ordovician. The present contribution 
is the first of several publications arising from this ongoing 
research project. 

Regional geological setting and lithostratigraphy 

The Honghuayuan Formation was initially named the 
Honghuayuan Limestone by Chang & Sheng (1958) based 
on 40 m of grey, thick-bedded limestone at Honghuayuan, 
7 km south of Tongzi County town, Guizhou (Fig. 1B,C). 
Zhang et al. (1964) renamed the unit the Honghuayuan 
Formation, which has been widely used to describe upper 
Lower Ordovician strata in South China (Wang et al., 1996). 
The formation conformably overlies the Tongzi Formation. 
The contact with the overlying Meitan Formation is 
conformable and gradational (Fig. 1C). 

The Honghuayuan Formation in the type section is a 
medium- to thick-bedded skeletal grainstone and packstone 
with thin-bedded to nodular skeletal wackestone (Fig. 2). 
Lime mudstone and shales are rare. Skeletal debris is mostly 
composed of cephalopods, brachiopods, trilobites, 
Archaeoscyphia, and bryozoans. The high biotic diversity 
suggests deposition in a well-circulated neritic environment. 


Abundant skeletal grainstones and cross bedding indicate 
that the formation was mostly deposited in a shoreward 
environment in the high-energy, shallow sub tidal zone (Fig. 
2), although lower-energy, deep subtidal sediments, e.g., 
nodular argillaceous lime mudstone, are also present. 
Deposition of carbonates of the Honghuayuan Formation 
ceased due to sea level rise, to be replaced by an increasing 
supply of terrigenous muds, and then greyish green mudstone 
(the Meitan Formation) was deposited in Guizhou (Fig. 1C). 

The Honghuayuan Formation is widely distributed in 
Guizhou (Fig. IB) as well as on the Yangtze Platform with 
similar lithofacies in the Honghuayuan section (Wang et 
al., 1996). Near shore facies of the formation deposited 
adjacent to eroded land (or island) areas are more dolomitic 
than the type section. Some patch reefs, which are mainly 
formed by abundant Calathium and less common Archaeo¬ 
scyphia (Zhu et al., 1995) were reported from the 
Huanghuachang section in Yichang, Hubei. The thickness 
of the Honghuayuan Formation on the Yangtze Platform is 
commonly 20 to 50 m, varying from a few metres to more 
than one hundred metres. In Guizhou, the thickness increases 
to more than 130 m northeastwards from the “Qianzhong 
Oldland” (an erosional land area at the time) (Fig. IB). 


Zhen et al Early Ordovician prioniodontid species 305 


AFI999& 
,THH1 
JAFIS97 
^AI=1995 
=^AFI993 
' THH2 
^AFI99Z & 
THH3 


- AFI9S6 


- TUNS 


3 

£ 

I 

=6; 
V) 1 

3, 

S' 

fQ I 
C 

S' 


Q. 

(A 

(/> 

C 

QJ 

e 

<TJ 

£ 

ftJ 

3 

■g 

O) 

c 

o 

-c 

■§ 

o 

c 

o 

q: 


r 5m 


L 0 m 


Shale/mud slone 
Argillaceous lime mudstone 
Nodular argillaceous 
skeletal wackestone 
Skeletal wackestone 
intraclastic packstone 
Skeletal packstone 
Skeletal gra instone 
Intraclaslic grainstone 


Fig. 2. Stratigraphic section through the Honghuayuan Formation 
at Honghuayuan, Tongzi, Guizhou Province showing the sampling 
positions and the stratigraphic ranges of Serratognathus diversus 
and the two species studied. 


Age and correlation of the Honghuayuan faunas 

The most characteristic feature of the conodont faunas from 
the Honghuayuan Formation is the abundance of Serrato¬ 
gnathus diversus in the middle part of the formation (Fig. 
2). This distinctive conodont species is apparently restricted 
in distribution to South China, the Tarim Basin (Zhao et al., 
2000), and probably Australia (R.S. Nicoll, pers. comm.) 
and other biogeographically related tectonic units (Zhen et 
al. in Webby et al., 2000; Zhen & Percival, 2003). An (1981, 
1987) proposed the S. diversus Zone (divided into lower 
and upper subzones) based on the wide distribution of this 
species in South China, and correlated the lower subzone 
to the P. proteus Zone and the upper subzone to the P. 
elegans Zone of Baltoscandian successions. The S. diversus 
Zone was also approximately correlated with the T. 
approximatus graptolite Zone, evidenced by the occurrence 
of the T. approximatus fauna in deeper water clastic 
graptolitic facies laterally equivalent to the carbonate facies 
of the Honghuayuan Formation (X. Chen et al., 1995). 

An (1981, 1987) also recognized the communis Zone 
ranging from the top of the Honghuayuan Formation to the 
basal part of the overlying Dawan Formation in South China, 


and correlated it with the communis Zone of North American 
Mid-continent successions. Although the age determination 
for the Honghuayuan Formation established by An (1981, 
1987) is still current, the present study has demonstrated 
that the so-called “ Baltoniodus communis ” from South 
China of previous authors represents a new species of 
Prioniodus, P. honghuayuanensis named herein. It is neither 
conspecific nor congeneric with Oepikodus communis 
(Ethington & Clark, 1964), an index zonal species of North 
American Mid-continent successions. Prioniodus honghua¬ 
yuanensis in fact exhibits close relationships with a younger 
unnamed species of Prioniodus from the basal Whiterockian 
succession of the Ibex area, Utah. The morphological 
changes (documented herein) from adenticulate Acodus 
species to denticulate Prioniodus indicate the likely origin 
of Prioniodontid clade. Prioniodus honghuayuanensis may 
have evolved from an adenticulate species revised herein 
as Acodus triangularis (Ding in Wang, 1993), which first 
occurs in the middle of the Honghuayuan Formation 
(THH10, Fig. 2) and is also associated with P. honghua¬ 
yuanensis at the top of the Formation. Morphologically 
Acodus triangularis resembles A. longibasis McTavish, 
1973 from the Emanuel Formation of Western Australia, 
which also represents the primitive adenticulate prionio¬ 
dontid species, but is probably of slightly younger age. 

Material and methods 

Twenty conodont samples (with prefix THH) were collected 
in 2000 from the Honghuayuan Formation in a 32 m thick 
limestone succession exposed on the hill slope to the 
southeast of Honghuayuan village (Fig. 1), about 10 km 
south of the Tongzi County. Acodus triangularis occurs in 
four samples (out of fifteen which yielded conodonts), 
whereas Prioniodus honghuayuanensis n.sp. was recovered 
from two samples at the top of the Formation (Table 1). All 
samples were completely dissolved in 10% acetic acid, with 
residues separated and concentrated using sodium 
poly tungstate. Subsequently, 38 large samples (with prefix 
AFI) were recollected from the Honghuayuan Formation 
on this section (location coordinates: 28°04.27'N 106° 
50.9l'E) to the southeast of the village. Both species were 
recovered from five samples at the top of the Honghuayuan 
Formation (Table 1). 

In the Ganxi Section, measured along the road from 
Ganxi village to Shichangao, southwest of Yanhe County 
Town (Fig. 1, location coordinates: 28°22.72'N 108° 
25.70'E), the Honghuayuan Formation consists of 71.8 m 
of thick-bedded limestones. Out of 27 productive samples 
collected from this section (with prefix YTH), only one from 
the top of the Honghuayuan Formation yielded Acodus 
triangularis and Prioniodus honghuayuanensis (Table 1). 

All photographic illustrations shown in Figs 3-9 are SEM 
photomicrographs captured digitally (numbers with the 
prefix IY are the file names of the digital images). Figured 
specimens bear the prefix AMF and are deposited in the 
collections of the Palaeontology Section at the Australian 
Museum in Sydney. Authorship of the new species is 
attributable solely to Zhen. Conodont terminology and 
notation employed in this contribution are conventional as 
defined in the Treatise Part W (Clark et al., 1981), except 
for the M elements (makellate), whose orientation, 
morphology and terminology was introduced by Nicoll 
(1990, 1992). 


306 Records of the Australian Museum (2005) Vol. 57 


Table 1 . Distribution of Acodus triangularis and Prioniodus honghuayuanensis n.sp., in ten samples of the 
Honghuayuan Formation at the Honghuayuan Section of Tongzi, and in one sample from the top of the Honghuayuan 
Formation at the Ganxi Section of Yanhe, Guizhou Province, South China. 


Acodus triangularis Prioniodus honghuayuanensis 


Samples 

Pa 

Pb 

M 

Sa 

Sb 

Sc 

Sd 

Total 

Pa 

Pb 

?Pc 

M 

Sa 

Sb 

Sc 

Sd 

Total 

AFI999 

2 

_ 

1 

_ 

_ 

_ 

_ 

3 

2 

1 

1 

2 

2 

4 

3 

4 

19 

AFI997 

13 

5 

6 

5 

4 

2 

7 

42 

9 

4 

— 

7 

13 

12 

11 

16 

72 

AFI995 

6 

— 

— 

2 

1 

1 

3 

13 

1 

1 

— 

1 

2 

2 

6 

9 

22 

AFI993 

26 

26 

18 

7 

2 

3 

11 

93 

34 

11 

— 

38 

26 

53 

27 

103 

292 

AFI992 

1 

1 

2 

6 

7 

9 

2 

28 

3 

— 

— 

2 

— 

4 

— 

2 

11 

THH1 

THH2 

THH3 


A 

1 

3 


C 


1 

— 

1 

5 

4 

4 

2 

2 

2 

4 

22 

_ 

_ 

_ 

1 

1 

_ 

_ 

1 

3 

THH8 

1 

1 

— 

— 

— 

— 

1 

3 


THH10 

4 

— 

1? 

2 

1 

2 

1 

11 


YTH1 

1 

— 

— 

— 

— 

1? 

— 

2 

3 

1 

— 

— 

1 

5 

2 

3 

15 

Total 

59 

37 

34 

26 

17 

20 

29 

222 

52 

18 

1 

51 

45 

80 

50 

138 

435 


Systematic palaeontology 
Class Conodonta Pander, 1856 
Acodus Pander, 1856 

Type species. Acodus erectus Pander, 1856. 

Remarks. The generic definition of Acodus and other 
related genera like Tripodus Bradshaw, 1969 and Tropodus 
Kennedy, 1980 has been a subject of disagreement among 
conodont workers (e.g., Lindstrom in Ziegler, 1977; Sweet, 
1988, Kennedy, 1980; Albanesi in Albanesi et al., 1998; 
Johnston & Barnes, 2000; X.H. Chen et al., 2003). It seems 
likely that there are more than one genus involved in these 
species embraced under a rather broad definition of Acodus 
given by Zhen et al. (2004). However, a serious revision of 
this species group, which will certainly need a monographic 
treatment, is far beyond the scope of the present project, 
and the level of our current understanding of these species 
does not yet support such a major revision. Acodus, defined 
as a multielement genus by Lindstrom (in Ziegler, 1977), is 
deeply entrenched in the Ordovician conodont literature. It 
was widely distributed in the Early and Middle Ordovician, 
and played a crucial part in the origin and early evolution 
of several major pectiniform clades like prioniodontids and 
balognothids (Stouge & Bagnoli, 1999). We retain Acodus 
as a valid genus, although Pander’s original specimens of 
the genotype, A. erectus, are irretrievably lost. 

The type species of Tripodus, T. laevis Bradshaw, was 
originally defined as a form species on the basis of three 
specimens which show sharp anterior and posterior margins, 
a sharp blade-like costa on the outer lateral face situated 
more towards anterior margin, and a broad carina on the 
inner lateral face (Bradshaw, 1969). Based on material from 
the Ibex area of Utah, Ethington & Clark (1982) revised 
the type species of Tripodus as consisting of a quinqui- 
membrate apparatus. This is interpreted herein as including 
geniculate M (oistodiform element, see Ethington & Clark, 
1982, fig. 33A,B), alate triform Sa (trichonodelliform 
element, their fig. 33C), asymmetrical tetra-costate Sb 
(distacodiform element, their fig. 33F,G), asymmetrical 
laterally compressed Sc (drepanodiform element, their fig. 
33H), and asymmetrical multi-costate Sd (paltodiform 
element, their fig. 33D,E) elements. Apparently the type 


material of T. laevis would fall into the definition of the Sb 
element of the multi-element concept of the species, and 
the types of Scolopodus alatus Bradshaw, 1969, which 
Ethington & Clark (1982) regarded as part of the T. laevis 
apparatus, include both Sa (Bradshaw, 1969, pi. 132, fig. 4, 
paratype) and Sd (Bradshaw, 1969, pi. 132, figs 1-3, 
holotype) elements. Ethington & Clark (1982) also regarded 
Triangulodus van Wamel, 1974 as a junior synonym of their 
revised Tripodus. This raised the question as to whether 
Tripodus is likely a senior synonym of Triangulodus or a 
junior synonym of Acodus. 

Based on material from western Newfoundland, Stouge 
(1984) regarded Tripodus as a junior synonym of Acodus, 
and suggested that Acodus combsi Bradshaw, 1969, T. laevis 
Bradshaw, and S. alatus Bradshaw represented different 
elements of one species apparatus which he called A. combsi. 
By assigning the prioniodiform P element represented by 
the form species A. combsi to the species apparatus, Stouge 
(1984) expanded the definition of T. laevis given by 
Ethington & Clark (1982), although at the time he was 
unaware of their work. Further differentiation of the 
prioniodiform Pa and Pb elements has completed the species 
apparatus as septimembrate (Albanesi in Albanesi et al., 
1998). Acodiform specimens assignable to the form species, 
A. combsi Bradshaw, 1969 were also reported from the Ibex 
area of Utah under the name “ Scandodus ” robustus Serpagli, 
1974 (Ethington & Clark, 1982, pi. 10, fig. 25). This species 
definition for T. laevis has been more or less accepted by 
many Ordovician conodont workers (Sweet, 1988, Stouge 
& Bagnoli, 1988; Albanesi in Albanesi et al., 1998; 
Bergstrom & Albanesi, 2001; Pyle et al., 2003), although 
others (Johnston & Barnes, 2000; X.H. Chen et al., 2003) 
retained the original concept of oistodiform M and costate 
S elements only as defined by Ethington & Clark (1982). 

By taking A. combsi as the name bearer of the species, 
Stouge’s work (1984) raised two questions: firstly, should 
T. laevis or A. combsi be the name bearer of the revised 
multi-element species, and secondly, to which genus should 
this species be assigned? In a recent documentation, 
Bergstrom & Albanesi (2001) reviewed the validity of the 
species name Tripodus laevis, and concluded that based on 
the most recent edition of ICZN rules, T. laevis is the valid 
name and A. combsi is a junior synonym of T. laevis. As for 


Zhen et al Early Ordovician prioniodontid species 307 


the second question, some conodont workers follow Sweet 
(1988) in maintaining Tripodus as a valid genus and 
assigning Acodus as a nomina dubia, whereas others (e.g., 
Stouge & Bagnoli, 1988; Johnston & Barnes, 2000) utilize 
both Acodus and Tripodus by consigning species to the 
respective genus on historical reasons or on the details of 
the P and S elements. Based on the definitions of Acodus 
given by Lindstrom (in Ziegler, 1977) and of Triangulodus 
given by van Wamel (1974), both bear costate S and 
geniculate M elements, but Acodus has prioniodiform 
(acodiform) P elements instead of scandodiform P elements 
without costa on lateral faces as in Triangulodus. However, 
as discussed above, the form species of T. laevis may 
represent the Sb element of the multi-element species 
apparatus adopted herein. If Stouge’s (1984) revision of T. 
laevis is accepted, Tripodus is likely a junior synonym of 
Acodus. Ethington (pers. comm., 2004) suggested that the 
S elements of Tripodus were much more ornate in their 
surface morphologies than typical Acodus as demonstrated 
by A. deltatus Lindstrom, 1955 and A. triangularis Ding 
described herein, and the P elements of T. laevis (A. combsi 
s.f. of Bradshaw) were deeply albid with very shallow basal 
cavities and without prominent carinae on the cusps. Until the 
type species of Tripodus can be revised in detail the taxonomic 
relationship of these two genera remains uncertain. 

In the type sections of the Marathon Basin of Texas, all 
three form species (T. laevis, A. combsi and S. alatus ) were 
established based only on a few specimens, and T. laevis 
and A. combsi were not found in association (Bradshaw, 
1969). Furthermore, co-occurrence of Histiodella sinuosa 
(Graves & Ellison) and Periodon aculeatus Hadding, 1913 
in the Fort Pena fauna also suggests a younger age (Middle 
Ordovician, Yapeenian equivalent). Considering that the 
FAD (first-appearance datum) of T. laevis is a potential 
candidate for defining the base of the Middle Ordovician, 
detailed revision of this species at the type locality is 
urgently needed. 

Acodus triangularis (Ding in Wang, 1993) 

Figs 3-5 

?Triangulodus cf. alatus Dzik.-An et al.: 1985: pi. 8, figs 1, 4, 8. 
ITripodus alatus (Dzik).-An, 1987: p. 193, pi. 16, figs 3-6. 
Oistodus triangularis Ding in Wang 1993: p. 185, pi. 19, figs 1, 2. 

Material. 222 specimens from ten samples (Table 1). 

Diagnosis. A species of Acodus consisting of a septimem- 
brate apparatus including pastinate Pa and Pb, makellate 
M, triform alate Sa, triform asymmetrical Sb, modified 
cordylodiform Sc, and tetra-costate Sd elements; elements 
mostly albid and small in size, all bearing short adenticulate 
processes; P elements with a triangular base bearing a large, 
open basal cavity; S elements bearing a longer posterior 
process typically with a thin, blade-like crest along the upper 
edge. 

Description. The pastinate Pa element has a reclined to sub¬ 
erect cusp and a large, triangular base (Fig. 3A-0). The 
cusp is triangular in cross section (Fig. 3F) with a sharp 
costa along the anterior and posterior margins, a mid-costa 
on the convex outer lateral face, and a smooth, more or less 
flattened inner lateral face. Three costae extend basally to 
merge with the upper margin of the anterior, posterior and 


outer lateral processes respectively (Figs 3A,G,K,I,M, 4 O). 
Fine striations are best developed in the area posterior to 
the outer lateral costa on the outer lateral face (Fig. 3L). All 
three processes have a blade-like upper edge (Fig. 3E,G) 
and an expanded base with posterior and anterior processes 
extending in the same plane (Fig. 3H) or with anterior 
process slightly turned inner laterally (Fig. 3M-0). The 
outer lateral process extends basally and outer laterally in a 
direction nearly normal to the anteroposterior plane in upper 
or basal view (Fig. 3B,F), and is triangular in outline in 
anterior view (Fig. 3C,I) or posterior view (Fig. 3E). 
Posterior process extends posteriorly and typically longer 
than the other two (Fig. 3G,K). The anterior process is 
anticusp-like, triangular in outline in the lateral view (Fig. 3D,I- 
K). Basal cavity is pyramidal, large and moderately deep, with 
more or less straight basal margin or with both sides of the 
outer lateral process slightly restricted inward (Fig. 3B,H). 

The Pb element is similar to the Pa element, but with a 
slightly proclined cusp (Fig. 4C,F,G,J,M), a shorter and less 
basally extended anterior process (Fig. 4A,B,G), a shorter 
and anterolaterally located outer lateral process (Fig. 
4A,E,F,N), and a more open basal cavity (Fig. 4D,H). The 
side between the anterior process and the outer lateral 
process is less inwardly restricted, and the basal margins 
on both sides of the outer lateral process are less restricted 
inward in upper or basal view (Fig. 4B,D,E). 

The M element is geniculate, anteroposteriorly 
compressed, with a low and well-developed outer lateral 
process and a short, anticusp-like inner lateral process, 
which is triangular in outline in anterior (Fig. 4K) or 
posterior view (Figs 3P, 4L). The anterior face is broadly 
convex and smooth, while the posterior face bears a broad 
carina and a weakly developed basal buttress. The basal 
margin is gently arched (Figs 3P, 4K,L). 

The Sa element is triform and symmetrical with a robust 
cusp, a short adenticulate posterior process, and a blade¬ 
like adenticulate lateral process on the anterolateral corner 
of each side (Figs 4P-U, 5A). The cusp is isosceles 
triangular in cross section with a broad anterior face (Fig. 
5A), a sharp costa along the posterior margin, and a sharp 
costa on each lateral side (Figs 4R,T, 5A). The three costae 
extend basally to form short, blade-like posterior and lateral 
processes (Fig. 4R,Q). Basal cavity is triangular in outline 
with moderate depth (Fig. 4R,U). Some specimens show a 
few small, rudimentary denticles along the thin upper edge 
of the posterior process (Fig. 4S). 

The Sb element resembles the Sa element in being 
tricostate, but is distinguished by being markedly 
asymmetrical (Fig. 5B-J). The cusp is proclined, scalene 
triangular in cross section with a sharp costa along the 
anterior and posterior margins and a costa on the outer lateral 
face. The inner lateral face is concave and smooth or bearing 
a broad carina, with anterior margin prominently curved 
inwards (Fig. 5B,C,G). The outer lateral face is divided into 
a broadly convex area defined by the anterior costa and the 
outer lateral costa, and a gently concave area between the 
outer lateral costa and the posterior costa (Fig. 5D,F,H,I). 
The base is triangular in outline when posterior process is 
shorter (Fig. 5J), but in more advanced forms, it is strongly 
laterally compressed (Fig. 5E) as a long, adenticulate blade 
with a straight basal margin and a straight or weakly arched 
upper margin. The anterior process is anticusp-like, 
triangular in outline in the lateral view (Fig. 51). The outer 


308 Records of the Australian Museum (2005) Vol. 57 


Fig. 3. Acodus triangularis (Ding in Wang, 1993): ( A-D ) Pa element, AMF126732, AFI993, (A) outer lateral view (IY52005), (B) 
basal view (IY64002), (C) antero-outer lateral view (IY52007), (D) inner lateral view (IY64001); (E-G) Pa element, AMF126733, 
AFI993, ( E ) upper-posterior view (IY52003), (F) outer lateral view (IY52004), (G) upper view (IY52001); ( H-K) Pa element, 
AMF126734, AFI993, (H) basal view (IY52009), (7) antero-outer lateral view (IY64005), (7) inner lateral view (IY52008), (K) outer 
lateral view (IY64003); ( L-0 ) Pa element, AMF126735, THH8, (L) outer lateral view showing the fine striae in the area posterior to 
the outer lateral costa (IY64027), (M) antero-outer lateral view (IY64026), (AO outer lateral view (IY64025), ( O ) inner lateral view 
(IY59016). (P) M element, AMF126736, AFI992, posterior view (IY65034). Scale bars 100 pm unless otherwise indicated. 


[Fig.4 caption continued ] ... (K,L) M element, AMF126740, AFI993, (K) anterior view (IY64009), (L) posterior view 
(IY52017). (M,N) Pb element, AMF126741, THH3, (M) outer lateral view (IY66032), (AO antero-outer lateral view 
(IY66034). ip) Pa element, AMF126742, THH10, outer lateral view (IY66037). (P-R) Sa element, AMF126743, AH993, 
(. P,Q ) posterolateral views (IY66013, IY66014), (R) basal-posterior view (IY66012). (S-U) Sa element, AMF126744, 
AFI997, (5) lateral view (IY66010), (7) posterior view (IY66009), (U) basal view (IY66011). Scale bars 100 pm. 


Zhen et al Early Ordovician prioniodontid species 309 


Fig. 4. Acodus triangularis (Ding in Wang, 1993): ( A-E ) Pb element, AMF126737, AFI992, (A) outer lateral view (IY53026), ( B ) 
upper view (IY53025), (C) inner lateral view (IY65016), (D) basal view (IY65017), (E) antero-outer lateral view (IY53027); (F,G) 
Pb element, AMF126738, AFI997, (F) outer lateral view (IY64010), (G) inner lateral view (IY52050); (H-J) Pb element, AMF126739, 
AFI993, ( H) basal-inner lateral view (IY52011), (/) outer lateral view (IY64006), (7) inner lateral view (IY52010)... [continued p. 308 ] 


310 Records of the Australian Museum (2005) Vol. 57 


Fig. 5. Acodus triangularis (Ding in Wang, 1993): (A) Sa element, AMF126745, AFI992, upper view (IY66022). ( B-D ) Sb element, 
AMF126746, AFI992, (B) basal-inner lateral view (IY65013), (C) inner lateral view (IY65012), (D) outer lateral view (IY53036); 
(E-H) Sb element, AMF126747, AFI992, (E) posterior view (IY53037), (F) antero-outer lateral view (IY65015), (G) basal-inner 
lateral view (IY53038), ( H) outer lateral view (IY65014); (7) Sb element, AMF126748, AFI992, outer lateral view (IY65032); ( J) Sb 
element, AMF 126749, AFI992, outer lateral view (IY66029). (K,L) Sc element, AMF126750, THH10, (K) outer lateral view (IY66040), 
(L) inner lateral view (IY66041); (M) Sc element AMF 126751, AFI992, inner lateral view (IY66027); (AO Sc element, AMF126752, 
THH3, upper view (IY66031); (O) Sc element, AMF126753, AFI992, inner lateral view (IY66026). ... [ continuedp. 311] 


Zhen et al Early Ordovician prioniodontid species 311 


lateral process is also anticusp-like extending basally and 
outer-laterally with blade-like upper margin curved 
posteriorly. Basal cavity is moderately deep and triangular 
in outline and extends as narrow grooves underneath the 
posterior and anterior processes (Fig. 5B). 

The Sc element is asymmetrical, strongly compressed 
laterally with a robust, proclined to suberect cusp, a posterior 
process varying from short (Fig. 5K,L) to relatively long 
(Fig. 5 O), and a short, anticusp-like anterior process (Fig. 
5K-0). The cusp is convex lens-like in cross section, slightly 
bent inward with a sharp costa along its anterior and 
posterior margins, and with smooth outer and inner lateral 
faces, or with a weakly developed broad carina on the inner 
lateral face (Fig. 5 O). The inner lateral face is less convex 
with anterior margin gently curved inward. Basal cavity is 
convex lens-like in outline with gently arched basal margin 
in lateral view (Fig. 5M). 

The Sd element is tetra-costate and asymmetrical, with 
a sharp costa along the anterior and posterior margins, and 
a sharp costa on each lateral side (Fig. 5P-AA). The outer 
lateral side is more convex with a broad convex area between 
the anterior process and the outer lateral process, and a less 
convex to concave area defined by the outer lateral process 
and the posterior process (Fig. 5P,U). The inner lateral side 
is less convex to concave with an inner laterally curved 
anterior margin (Fig. 5R,T,Y). Costae extend basally to 
merge with the upper margin of four corresponding short, 
blade-like processes. Small rudimentary denticles may occur 
along the upper edge of the posterior process of some 
specimens (Fig. 5P). However, the broken upper edge in 
some specimens superficially exhibits a denticulate 
appearance (Fig. 5U). 

Remarks. Oistodus triangularis was originally proposed 
as a form species with the type material from the 
Honghuayuan Formation of Anhui Province (Ding in Wang, 

1993) . The two figured specimens are both pastinate with a 
large, triangular base and a robust cusp which bears a sharp 
costa along its anterior and posterior margins, and a third 
costa on the outer lateral face (Ding in Wang 1993, p. 185). 
The holotype has a reclined cusp and less downwardly 
extending anterior process (Wang, 1993, pi. 19, fig. 1), 
whereas the other figured specimen (Wang, 1993, pi. 19, 
fig. 2), with a suberect cusp and a longer and more 
downwardly extending anterior process, is identical with 
the Pa element described herein from Guizhou. 

Specimens previously referred to Triangulodus alatus 
Dzik, 1976 from the lower Dawan Formation of South China 
(An et al., 1985; An, 1987) possibly represent the S elements 
of Acodus triangularis. The type material of T. alatus, 
consisting of large-sized hyaline elements, occurs in late 
Darriwilian erratic boulders of NW Poland. The holotype 
(Dzik, 1976, p. 433, fig. 20k) is a scandodiform P element 
with an adenticulate posterior process and a short 
adenticulate anticusp-like anterior process. Dzik (1983, 

1994) re-assigned this Baltic species to Eoneoprioniodus 


Mound, 1965, which is likely a senior synonym of 
Triangulodus van Wamel, 1974 (see Clark et al., 1981). 
Triangulodus alatus was also reported from the upper 
Darriwilian ( serra Zone) of the Tarim Basin (Zhao et al., 
2000), and from the upper Darriwilian ( serra Zone) of 
Oklahoma (Bauer, 1987). 

Specimens referred to Acodus deltatus Findstrom, 1955 
from the Emanuel Formation of Western Australia 
(McTavish, 1973) are reassigned herein to A. longibasis 
which differs from A. deltatus mainly in having a more 
extended posterior process. This species from the Canning 
Basin shows some resemblance to A. triangularis from 
South China, in particular the Pa and S elements. Based 
mainly on the length of posterior process of the S elements 
in the Emanuel Formation, McTavish (1973) split this 
species into several subspecies of A. deltatus including A. 
deltatus longibasis McTavish with long posterior process, 
A. deltatus deltatus with a shortest posterior process, and 
A. deltatus tortus McTavish with an inwardly twisted cusp 
of the S elements. Similar variations are also observed in 
the species from South China with the posterior process of 
the S elements varying from short (Fig. 5 J,L,K) to relatively 
long (Fig. 5B-H,M,0). 

Prioniodus Pander, 1856 

Type species. Prioniodus elegans Pander, 1856. 

Remarks. The type species of Oepikodus Findstrom, 1955, 
O. smithensis Findstrom, 1955, was originally proposed as 
a form species based on the oepikodiform elements 
(Findstrom, 1955, pi. 6, figs 1-3). Subsequently, Findstrom 
(1971) considered Oepikodus as a junior synonym of 
Prioniodus, 1856, and its type species, O. smithensis, along 
with Oistodus longiramis Findstrom, 1955 to be parts of 
the revised multi-element species Prioniodus evae 
Findstrom, which was also originally proposed as a form 
species based on the prioniodiform elements (Findstrom, 
1955, pi. 6, figs 4-10). Hence Findstrom (1971) defined P. 
evae as consisting of a trimembrate apparatus including 
prioniodiform, oepikodiform and oistodiform elements. 
However, Bergstrom & Cooper (1973, pp. 323-324) pointed 
out that “the apparatus of P. evae differs from that of P. 
elegans ”, the type species of Prioniodus, and “no element 
clearly homologous to the hibbardelliform element” existed 
in the latter. They further considered Oepikodus to be a 
subgenus of Prioniodus. This suggestion was formalized 
by Serpagli (1974). However, van Wamel (1974) and 
Findstrom (in Ziegler, 1975) restored Oepikodus as a 
separate multi-element genus consisting of a trimembrate 
apparatus. Bergstrom (in Clark et al., 1981) and Stouge & 
Bagnoli (1999) categorized Oepikodus and Prioniodus as 
belonging to different families. 

Serpagli (1974) and Findstrom (in Ziegler, 1975) 
redesignated the form species P. evae Findstrom, 1955 as 
the type species of Oepikodus, since the originally 


[Fig. 5 caption continued] ... (P-R ) Sd element, AMF126754, AFI997, (P) outer lateral view (IY53002), (Q) posterior view 
(IY53001), (R) inner lateral view (IY53003); ( S ) Sd element, AMF126755, AFI997, basal-posterior view (IY66006); ( T) Sd 
element, AMF126756, AFI997, inner lateral view (IY66001); (U,V) Sd element, AMF126757, AFI993, (U) outer lateral 
view (IY66017), (V) anterior view (IY66016); (W-Y) Sd element, AMF126758, THH3, (W) upper view (IY58017), (Y) outer 
lateral view (IY58019), (Y) inner lateral view (IY58018); (Z,AA) Sd element, AMF126759, AFI993, (Z) inner lateral view 
(IY66018), (AA) posterior view (IY66020). Scale bars 100 pm. 


312 Records of the Australian Museum (2005) Vol. 57 


designated type species O. smithensis became a synonym 
of Prioniodus evae (see Lindstrom in Ziegler, 1975, p. 237). 
As correctly pointed out by van Wamel (1974, p. 74), the 
form species O. smithensis is not a junior synonym of the 
form species P. evae, and the form species O. smithensis 
must therefore remain as the type species of Oepikodus. 
Although O. smithensis has page priority among the three 
form species erected by Lindstrom in the same publication 
for three different genera, they ( O. smithensis Lindstrom, 
1955, P. evae Lindstrom, 1955, and Oistodus longiramis 
Lindstrom, 1955) form the apparatus of a multi-element 
species, which was named as P. evae by Lindstrom (1971). 
With Lindstrom (1971) as the first reviser (see Article 24.2.2, 
ICZN of 1999 edition), O. evae (Lindstrom, 1955) emend. 
Lindstrom (1971) is recognized as a valid and proper name 
of the species with this multi-element apparatus. 

Following the definition of Oepikodus as trimembrate 
(van Wamel, 1974) and then quadrimembrate (Bergstrom 
in Clark et al., 1981), Stouge & Bagnoli (1988) suggested 
that Oepikodus consisted of a seximembrate apparatus that 
could be distinguished from Prioniodus by lack of the Sa 
element. Albanesi (in Albanesi et al., 1998) concurred with 
this concept. 

Oepikodus, as most recently revised by Nicoll & 
Ethington (2004), was defined as consisting of a septimem- 
brate apparatus including pastinate Pa and Pb, makellate M 
and quadriramate S (Sa, Sb, Sc and Sd) elements. Prioniodus 
consists of a similar septimembrate apparatus that includes 
a triform alate Sa element, rather than a quadriform alate 
Sa element as in Oepikodus. Therefore, occurrence of the 
triform alate Sa element in the new species from the 
Honghuayuan Formation confirms its generic assignment 
to Prioniodus. 


Prioniodus honghuayuanensis n.sp. 

Figs 6-8 

Baltoniodus communis (Ethington & Clark).-An, 1981: pi. 4, figs 
20-23, 25, 27-29, ?24. 

Baltoniodus communis (Ethington & Clark).-An & Ding, 1985: 
p.8,9, pi. 1, figs 1-5,8,79. 

Baltoniodus? communis (Ethington & Clark).-An, 1987: p. 125, 
126, pi. 19, figs 1-11. 

Baltoniodus communis (Ethington & Clark).-Ding et al. in Wang, 
1993: p. 161, pi. 23, figs 1-12. 

Etymology. After Honghuayuan Village, where the type 
section of the Honghuayuan Formation is exposed on the 
hill slope located to the southeast of the village. 

Material. 435 specimens, including holotype (AMF126760, 
from sample AFI993) and 19 paratypes (AMF126761- 
AMF126779), from eight samples (Table 1). 

Diagnosis. Species of Prioniodus consisting of a 
septimembrate apparatus, including pastinate Pa and Pb 
elements with denticulate posterior and outer lateral process 
and adenticulate (or with rudimentary denticles) anterior 


process which curves distally outer laterally; makellate M 
element with low but long inner lateral and outer lateral 
processes; alate triform Sa, triform asymmetrical (modified 
quadriramate) Sb, bipennate (modified quadriramate) Sc, and 
quadriramate Sd elements; all S elements with a proclined cusp, 
a long denticulate posterior process, and adenticulate (or 
mdimentarily denticulated) lateral and anterior processes. 

Description. All elements are small, mostly albid, thin and 
fragile; the long denticulate posterior process of the P and 
S elements is broken in most specimens. Pastinate P 
elements bear a robust, laterally compressed cusp, with a 
sharp costa along its anterior and posterior margins, and 
with a prominent costa on the outer lateral face (Fig. 6). 
Surface of the cusp is ornamented with fine striation, which 
are best developed posterior to the outer lateral process on 
the outer lateral face (Fig. 6P). The posterior process is 
laterally compressed as a long blade with closely spaced, 
basally confluent, and more or less equal-sized denticles 
along its upper margin. The anticusp-like anterior process 
extends downward with its anterior margin strongly curved 
outwards and occasionally with confluent and small 
rudimentary denticles along its edge (Fig. 6F). The outer 
lateral costa extends downward to merge with the upper 
margin of the outer lateral process, which is long and bearing 
small, closely spaced denticles (Fig. 6B,M,N). The outer 
lateral process that extends downward laterally outward with 
an angle slightly >90° (Pb, Fig. 60) or nearly normal (Pa, 
Fig. 6D,G) to the posterior process is often broken in the 
present collection. Basal cavity is triangular in shape 
extending underneath the three processes as narrow grooves 
(Fig. 6E,F,Q,0). The Pa element has a suberect cusp with 
anterior process curved laterally outward and with the outer 
lateral process extending nearly normal to the posterior 
process in the upper view (Fig. 6D,G). The Pb element 
resembles the Pa, but with a reclined cusp and an inner- 
laterally bent posterior process (Fig. 6K-S). 

The M element has a robust and strongly outer laterally 
reclined cusp, and adenticulate, low and long outer lateral and 
inner lateral processes (Fig. 7A-D). The cusp is sightly curved 
posteriorly and anteroposteriorly compressed, with a weak 
costa on the broad anterior face, and a broad carina on the 
posterior face. The outer lateral process has gently arched upper 
and basal margins, while the inner lateral process has nearly 
straight upper and basal margins. Basal cavity is relatively 
narrow, without prominent basal buttress (Fig. 7B,C). 

The Sa element is triform and symmetrical with a proclined 
cusp, a long denticulate posterior process, and an anticusp¬ 
like lateral process on each side (Fig. 7E-S). The cusp is 
triangular in cross section with a broad anterior face (Fig. 7M, 
O), a sharp blade-like costa along the posterior margin, and a 
sharp costa along the anterolateral comer of each side (Fig. 7J, 
S). The posterior process is long, straight and strongly 
compressed laterally bearing small, closely spaced denticles 
(Fig. 7P-R). The blade-like costa on each side is gently curved 
posteriorly and extends basally to merge with the upper margin 
of the lateral process, which often bears rudimentary denticles 
(Fig. 7J,R). The basal cavity is an isosceles triangle in outline 
with moderate depth (Fig. 7E,J). 


[Fig.6 caption continued ] ... AMF126762, (K) inner lateral view (IY63005), (L) outer lateral view (IY63006), (M) upper view 
(IY63007), (AO anterior view (IY63004), (O) basal view (IY50035), (P) close up of outer lateral view, showing the fine striae in 
the area posterior to the outer lateral costa (IY50036); (Q-S) Pb element, paratype, AMF126763, (Q) basal view (IY50027), (R) inner 
lateral view (IY63011), (5) outer lateral view (IY50028). All from sample AFI993; scale bars 100 pm unless otherwise indicated. 


Zhen et al Early Ordovician prioniodontid species 313 


Fig. 6. Prioniodus honghuayuanensis n.sp.: (A-E) Pa element, paratype, AMF126761, (A) antero-outer lateral view (IY50030), ( B ) 
basal-outer lateral view (IY50032), (C) outer lateral view (IY50033), (D) upper view (IY63010), (E) inner lateral view (IY63009); 
(F-J) Pa element, holotype, AMF126760, (F) basal view (IY50039), (G) upper view (IY63003), ( H) outer lateral view (IY50038), 
(I) inner lateral view (IY63001), (7) postero-outer lateral view (IY50040). (K-P) Pb element, paratype, ... [continuedp. 312 ] 


314 Records of the Australian Museum (2005) Vol. 57 


Fig. 7. Prioniodus honghuayuanensis n.sp.: (A,B) M element, paratype, AMF126764, AFI993, (A) anterior view (IY51011), ( B ) 
posterior view (IY63016); (C,D) M element, paratype, AMF126765, AFI993, (C) basal-posterior view (IY51015), (D) anterior view 
(IY63018). (E-G) Sa element, paratype, AMF126766, AFI993, ( E ) basal view (IY51003), (F,G) lateral views (IY51002, IY63012); 
(H,I) Sa element, paratype, AMF126767, AFI993, lateral views (IY63013, IY51005); U-N) Sa element, paratype, AMF126768, 
AFI993, (7) basal-posterior view (IY51009); ( K,L ) lateral views (IY51010, IY51008), ( M) anterior view (IY63014); (AO basal view 
(IY63015); (O) Sa element paratype, AMF126769, AFI993, anterior view (IY51006); (P-S) Sa element, paratype, AMF126770, 
AFI997, (P, Q) lateral views (IY51046, IY51048), (R) upper view (IY51045), (S) upper view showing the cross section of the cusp 
(IY63026). Scale bars 100 pm. 


[Fig.8 caption continued ] ... AMF126774, AFI993, (7) inner lateral view (IY63021), (7) outer lateral view (IY51026), ( K ) 
basal view (IY51025); (L-P) Sc element, paratype, AMF126775, AFI997, (L) basal view (IY51050), (M) inner lateral view 
(IY51049), (AO outer lateral view (IY63028), (O) upper view (IY51051), (P) close up of the upper view showing the cross 
section of the cusp (IY51052); (0 Sc element, paratype, AMF126776, AFI993, outer lateral view (IY51022). (R-T) Sd 
element, paratype, AMF126777, AFI993, ( R ) outer lateral view (IY63025), ( S ) inner lateral view (IY51037), (T) posterior 
view (IY51036); (U-X) Sd element, paratype, AMF126778, AFI993, ( U) inner lateral view (IY63023), (V) outer lateral 
view (IY51033), (W) posterior view (IY51035), (A) postero-outer lateral view (IY51034); (Y-AA) Sd element, paratype, 
AMF126779, AFI993, (7) postero-inner lateral view (IY51032), (Z) outer lateral view (IY63022), (AA) inner lateral view 
(IY51031). Scale bars 100 pm unless otherwise indicated. 


Zhen et al Early Ordovician prioniodontid species 315 


Fig. 8. Prioniodus honghuayuanensis n.sp.: (A-C) Sb element, paratype, AMF126771, AFI993, (A) inner lateral view (IY51013), (. B) 
basal-inner lateral view (IY51014), (C) outer lateral view (IY63017); ( D-F) Sb element, paratype, AMF126772, AFI993, (D) basal 
view (IY51020), (E) inner lateral view (IY51021), ( F) outer lateral view (IY63020); (G,H) Sb element, paratype, AMF126773, 
AFI993, (G) inner lateral view (IY63019), ( H) outer lateral view (IY51017). (I-K) Sc element, paratype, ... [continuedp. 314 ] 


316 Records of the Australian Museum (2005) Vol. 57 


The Sb element is strongly asymmetrical with a proclined 
and laterally compressed cusp, which bears a sharp costa 
along the anterior and posterior margins, and the third costa 
on the outer lateral face (Fig. 8A-H). The anterior costa is 
inner laterally curved and extends basally to merge with 
the upper margin of the anticusp-like downwardly extending 
adenticulate anterior process (Fig. 8A,E,G). The long 
posterior process is laterally compressed as a high blade 
bearing small, closely spaced denticles of similar sizes along 
its upper margin. The costa on the outer lateral face extends 
downward to merge with the upper margin of the downward 
extended, shorter, adenticulate outer lateral process. The 
inner lateral face is less convex, and bears a broad carina 
(Fig. 8A,B,E,G). 

The Sc element is bipennate with a broad carina on each 
lateral face (more prominent on the inner lateral face), a 
short, adenticulate anterior process, and a long, laterally 
compressed, denticulate posterior process (Fig. 8I-Q). The 
cusp is proclined (Fig. 8Q), and laterally compressed with 
a sharp blade-like costa along its posterior and anterior 
margins (Fig. 8 0,P), which extends basally to merge 
respectively into the upper margin of the anterior process 
and the posterior process (Fig. 81,M). The anterior process 
is anticusp-like, and curved laterally inward (Fig. 81,M). 
The posterior process bears small, closely spaced denticles 
of more or less similar sizes (Fig. 8Q). The basal cavity is 
biconvex in outline, and extends as a narrow groove 
underneath the anterior and posterior processes (Fig. 8K,L). 

The Sd element is quadriramate and asymmetrical, with 
an anterior process, a posterior process, and a lateral process 
on each side (Fig. 8R-AA). The cusp is proclined with a 
sharp blade-like costa along the anterior and posterior 
margins and on each lateral side (Fig. 8T,W,Y,AA). Anterior 
costa is inner laterally curved (Fig. 8S,U,Y), and extends 
basally to merge into the upper margin of the anticusp-like 
anterior process which is adenticulate or has small, fused 
rudimentary denticles along its distal edge (Fig. 8Y). As in 
the other S elements, the posterior process is long, laterally 
compressed, and blade-like bearing small, closely spaced 
denticles of similar sizes. The area defined by the anterior 
costa and the outer lateral costa is more broadly convex 
(Fig. 8R,V). The lateral costa on each side extends basally 
to merge respectively with the upper margin of the short 
inner and outer lateral processes. Both lateral processes 
extend downward and often bear small rudimentary 
denticles along their distal edges (Fig. 8S-U,W,X,AA). 

Remarks. Although P elements of the new species show 
some resemblance to those of Oepikodus communis 
(Ethington & Clark, 1964), they differ in having denticulate 
outer lateral processes, and an anterior process which bears 
rudimentary denticles along its distal edge. The anterior 
process of P. honghuayuanensis extends strongly downward 
as an anticusp, and curves strongly outer laterally (Fig. 
6F,J,Q). The Pb element of O. pincallyensis Zhen in Zhen 
at al., 2003 has denticulate anterior, outer lateral and 
posterior processes, but the anterior process of the Pa 
element is adenticulate. However, the P elements of O. 
pincallyensis are distinctively curved inner laterally (Zhen 
et al., 2003, fig. 19E,K) rather than outer laterally as in P. 
honghuayuanensis (Fig. 6D,M). Furthermore, the S 
elements of the new species can be easily distinguished from 
those of O. communis, O. pincallyensis and other species 
of Oepikodus. The Sa element is triform and alate with a 


wide anterior face (Fig. 7M,0) and the open basal cavity is 
an isosceles triangle in outline (Fig. 7E,J). The Sb element 
can be described as asymmetrical triform (or modified 
quadriramate of some authors) with a strongly developed 
costa on the outer lateral face, which extends basally as a 
short but prominent adenticulate outer lateral process. The 
quadriramate Sd element of P. honghuayuanensis differs 
from that of O. communis in having more strongly developed 
costae, in having more prominent lateral processes, and in 
tending to develop rudimentary denticles along the distal edge 
of the anterior and lateral processes (Fig. 8S-U,W-Y,AA). 

Oepikodus communis was originally proposed as a form 
species based on pastinate elements (Ethington & Clark, 
1964). The type material shows a long denticulate posterior 
process with small, closely spaced denticles, and 
adenticulate anterior and outer lateral processes (Ethington 
& Clark, 1964, pi. 114, figs 6,14, text-fig. 2F). Subsequent¬ 
ly, Ethington & Clark (1982) revised O. communis in 
multielement taxonomy by accommodating another three 
form species which were reported in association with the 
type material of the form species Gothodus communis 
Ethington & Clark, 1964 in the El Paso Formation of Texas. 
These include Oepikodus equidentatus Ethington & Clark, 
1964 with typical quadriramate ramiform elements 
(including symmetrical and asymmetrical), Subcordylodus 
sp. aff. S. delicatus (Branson & Mehl) with modified 
quadriramate ramiform elements (bearing only a weakly 
developed broad carina on the lateral faces), and Oistodus 
longiramis Lindstrom, 1955 comprising a makellate element 
with a long and slender outer lateral process. Ethington & 
Clark (1982) recognized the symmetry transition among 
the ra mi form elements of four morphotypes, symmetrical 
quadriramate element, asymmetrical quadriramate element, 
modified quadriramate element with costa on one side, and 
modified quadriramate element without lateral costa. 
Therefore, they indicated a seximembrate apparatus for the 
species, although no formal notation was made. Repetski 
(1982) also suggested a seximembrate apparatus for O. 
communis, including prioniodiform, falodiform and four 
types of ramiform (belodiform, tetraprioniodiform, 
hibbardelliform, and cordylodiform) elements. Nicoll & 
Ethington (2004) defined Oepikodus as consisting of a 
septimembrate apparatus of a geniculate makellate M, four 
types of quadriramate or modified quadriramate S, and two 
types of pastinate P elements. 

We also interpret Oepikodus communis as consisting of 
a septimembrate apparatus. The pastinate Pa element is 
represented by the holotype (Ethington & Clark, 1964, pi. 
114, fig. 6) with a suberect cusp, and the pastinate Pb 
element by the other figured specimen of G. communis 
(Ethington & Clark, 1964, pi. 114, fig. 14) with a reclined 
cusp. The Sa and Sd elements are represented by the form 
species O. equidentatus Ethington & Clark (1964, pi. 113, 
figs 6, 8, 10, 11, 14). The Sa element is symmetrical or 
nearly symmetrical with a straight anterior margin, while 
the Sd element is markedly asymmetrical with a curved 
anterior margin (e. g. Ethington & Clark, 1964, pi. 113, fig. 
14). The Sc element of O. communis is represented by those 
described and illustrated by Ethington & Clark (1964, p. 
701, pi. 115, figs 1, 5, 7, 10) as Subcordylodus sp. aff. S. 
delicatus (Branson & Mehl, 1933). The makellate M 
element of O. communis is represented by those described 
and illustrated by Ethington & Clark (1964, p. 693, pi. 114, 
figs 2, 7) as Oistodus longiramis Lindstrom, 1955. The 


Zhen et al Early Ordovician prioniodontid species 317 


Fig. 9.A-P, Prioniodus sp.: (A,B) Pa element, AMF126781,100 feet above base of KanoshFm., Utah, (A) inner lateral view (IY66053), 
(. B ) upper view (IY66054). (C-£) Pb element, AMF126782, 83 feet above base of Kanosh Fm., Utah, (C) basal view (IY66051), ( D ) 
antero-outer lateral view (IY66052), (£) outer lateral view (IY66050). (F,G) Sd element, AMF126783, 78J-23, 150 feet above base of 
Juab Fm., Utah, (F) basal view (IY66061), (G) inner lateral view (IY66062). (H) Sa element, AMF126784, 78J-23, 150 feet above 
base of Juab Fm., Utah, lateral view (IY66060); ( I-K) Sa element, AMF126785, 78J-16, top ledge of Wah Wah Fm., Utah, (/) antero- 
upper view (IY66042), (J,K) lateral views (IY66045, IY66043). (L) M element, AMF126786, 78J-16, top ledge of Wah Wah Fm., 
Utah, posterior view (IY66049). (M,N) Sb element, AMF126787, 100 feet above base of Kanosh Fm., Utah, (A/) outer lateral view 
(IY66065), (N) inner lateral view (IY66066). (O) Sc element, AMF126788, 57 feet above base of Kanosh Fm., Utah, inner lateral 
view (IY66063); (P) Sc element, AMF126789, 78J-16, top ledge of Wah Wah Fm., Utah, outer lateral view (IY66047). (Q-U) 
Prioniodus ?honghuayuanensis n.sp.: ?Pc element, AMF126780, AFI999, ( Q ) inner lateral view (IY53012), (R) outer lateral view 
(IY65010), ( S ) upper view (IY53010), (7) outer-basal view (IY65008), (U) close up of outer lateral view showing fine striae in the 
area posterior to outer lateral costa (IY65009); Scale bars 100 pm unless otherwise indicated. 


318 Records of the Australian Museum (2005) Vol. 57 


asymmetrical Sb element is a modified quadriramate 
element with a more prominent costa on the outer lateral 
face (Albanesi in Albanesi et al., 1998, pi. 6, fig. 22). 

Oepikodus intermedius Serpagli, 1974, from the San Juan 
Formation of the Argentine Precordillera, is regarded herein 
as conspecific with O. communis ; it was originally described 
as consisting of a trimembrate apparatus (prioniodiform, 
oistodiform and oepikodiform). In more recent revisions of 
the San Juan faunas, both Lehnert (1995) and Albanesi (in 
Albanesi et al., 1998) did not recognize symmetrical Sa 
elements in any of the three species ( O. communis, O. 
intermedius and O. evae) they ascribed to Oepikodus. 

Lindstrom (in Ziegler, 1975) considered the form species 
Gothodus microdentatus van Wamel, 1974 from the 
Glauconitic Limestone of Sweden as part of O. communis 
species apparatus. However, not only have the well- 
recognized elements of O. communis apparatus not been 
recorded in association with this form species in Sweden, 
but also the triform, asymmetrical G. microdentatus shows 
rather different morphology in comparison with the 
ramiform elements of O. communis from North America. 
Based on the original description and illustrations (van 
Wamel, 1974), the form species G. microdentatus more 
likely represents the Sb element of a multi-element species 
of Prioniodus rather than of Oepikodus. 

Oepikodus communis is widely distributed in North 
America (Ethington & Clark, 1964, 1982; Repetski, 1982; 
Stouge, 1982; Stouge & Bagnoli, 1988; Johnston & Barnes, 
2000), Argentine Precordillera (Serpagli, 1974; Lehnert, 
1995; Albanesi in Albanesi et al., 1998), Australia 
(McTavish, 1973; Zhen et al., 2003), and Greenland (Smith, 
1991). However, as the result of inclusion of a triform alate 
Sa element in this species apparatus by An and other Chinese 
authors (An, 1981, 1987; An et al., 1985; Ding et al. in 
Wang, 1993), the occurrence of O. communis in South China 
needs to be re-examined. 

Specimens referable to the Pa, Pb, Sa, Sb, Sc and Sd 
elements of the present new species were first recorded from 
the top of the “Honghuayuan Formation” of Hexian, Anhui 
Province as Baltoniodus communis (An, 1981, 1987; An & 
Ding, 1985). An (1987) noted the abundant occurrence (up 
to 80% in some samples) of this species at the top of the 
Honghuayuan Formation in Guizhou, Hunan, Hubei and 
Anhui provinces, and correlated this interval at the top of 
the Honghuayuan Formation with the communis Zone of 
North America and with the elegans Zone of Baltoscandia. 
Although he recognized the occurrence of triform alate 
element as part of this species apparatus, An (1981, 1987) 
included this material into his rather broad interpretation 
of O. communis (Ethington & Clark, 1964). However, as the 
triform Sa element occurring in the material from South China 
had not been recognized at the type locality of O. communis in 
North America, An (1981, 1987) doubtfully assigned O. 
communis to Baltoniodus rather than Oepikodus. 

Specimens previously referred to as aff. Oepikodus? 
minutus (McTavish, 1973) from the upper Wah Wah 
Formation, and the overlying Juab and Kanosh formations 
in the Ibex area of Utah (Ethington & Clark, 1982) 
somewhat resemble P. honghuayuanensis. Prof. R. 
Ethington kindly provided sixty specimens of this Utah 
species from nine samples for comparative study. 
Stratigraphically the Ibex species (designated herein as 
Prioniodus sp.; Fig. 9A-P) occurs at a slightly younger level 


(basal Whiterockian, victoriae graptolite Zone) than the new 
species from South China. Similarities include the 
prioniodiform Pb element of Prioniodus sp. (Fig. 9C-E) 
that bears a denticulate outer lateral process with anterior 
margin curved outer laterally towards outer lateral process, 
and the ramiform S series that includes alate triform Sa (Fig. 
9H-K), asymmetrical triform Sb (Fig. 9M,N), bipennate 
Sc (Fig. 9 0,P), and quadriramate Sd (Fig. 9F,G) elements. 
However, the associated M element of the Ibex species 
shows a much shorter inner lateral process, and the Pa 
element has an adenticulate and weakly developed outer 
lateral process and a more or less straight anterior process. 
Bagnoli & Stouge (1997) referred this Ibex species to 
Gothodus Lindstrom, 1955. Based on the recent revision of 
Gothodus (Bagnoli & Stouge, 1997, and Stouge & Bagnoli, 
1999), it has prioniodiform P elements comparable with 
those of Oepikodus and Prioniodus, but the S elements are 
more related to Phragmodus Branson & Mehl, 1933. 
However, most other conodont authors regard it as a junior 
synonym of Prioniodus (e.g., Bergstrom in Clark et al., 
1981) or Baltoniodus (Bagnoli et al., 1988). 

The new species from the Honghuayuan Formation 
shows some resemblance to P. elegans documented from 
Sweden by van Wamel (1974, p. 87-89, pi. 6, figs 1-6) and 
by Lofgren (1978, pi. 9, figs 1-6), but Swedish material of 
P. elegans has the M element with a denticulate inner lateral 
process and the P and S elements with better developed 
denticles on the lateral and anterior processes. Prioniodus 
elegans described from the Leningrad region of Russia and 
Tulubacken of Sweden by Bergstrom (1988, pi. 3, figs 33- 
38), and from western Newfoundland by Stouge & Bagnoli 
(1988, pi. 13, figs 1-9) and Johnston & Barnes (2000, pi. 3, 
figs 1, 5-7, 11, pi. 16, fig. 17) has an even stronger 
development of denticulation with P elements bearing a 
longer, fully denticulated anterior process, and with S 
elements displaying well-denticulated anterior and lateral 
processes. The multi-element species definition of P. elegans 
followed herein conforms to the concept of Bergstrom 
(1971, 1981 in Clark et al., 1981), van Wamel (1974), and 
Stouge & Bagnoli (1988). Stouge & Bagnoli (1988, p. 134) 
noted that this species displayed a large degree of 
morphological variation and might be polymorphic. Some 
of the material of P. elegans documented by Bergstrom 
(1988, pi. 3, figs 33-37) came from Popowka to the south 
of the Leningrad region, from where Pander’s original type 
material (now lost) was collected. In comparison with those 
figured specimens of P. elegans, the P elements of P. 
honghuayuanensis display a more prominent cusp and 
smaller and less developed anterior and outer lateral 
processes (Fig. 6H,L), the P and S elements show less 
development of denticles on the anterior and lateral processes 
(Figs 6-8), and the M element lacks denticles on the inner 
lateral process (Fig. 7A-D). Also, the anterior process of the P 
elements in P. elegans is distinctively curved inner laterally 
(Bergstrom, 1988, pi. 3, fig. 33) rather than outer laterally (Fig. 
6D,F,G) as in P. honghuayuanensis. In South China, P. 
elegans has only been recorded from the Jinshan Formation 
(biostratigraphically coeval with the Honghuayuan Formation) 
in Zhejiang Province of southeast China (An, 1987). It was 
also recorded from sub-surface core samples of the Tarim 
Basin, northwest China (Zhao et al., 2000). 

One specimen from the very top of the Honghuayuan 
Formation at the Honghuayuan Section (AFI999, Fig. 2) is 


Zhen et al Early Ordovician prioniodontid species 319 


similar to the Pb element of P. honghuayuanensis, but is 
stellate in outline with an additional sharp, blade-like costa 
on the inner side of the cusp, which extends basally into a 
short process with a few small rudimentary denticles. It is 
tentatively referred to herein as a Pc element of P. 
honghuayuanensis (Fig. 9Q-U). 


Acknowledgments. Fieldwork by YYZ in Guizhou in late 2000 
was undertaken with the support of the Australian Academy of 
Sciences and the Academia Sinica (visiting grant). Professors 
Zhiyi Zhou and Jiayu Rong from Nanjing Institute of Geology 
and Palaeontology, Academia Sinica kindly provided financial 
assistance and made arrangement for the shipment of the conodont 
samples to Australia (NSFC projects 40272001, and Major Basic 
Research Projects of MST, China G2000077703). Further 
collecting in Tongzi was undertaken by JBL in the following years 
(2001 to 2002) with the support of the Special Funds for Major 
State Basic Research Project (G200077700) of PR. China, and 
these samples were processed at the Palaeontology Laboratory of 
the Geology Department, Peking University. Y.Y. Zhen’s study 
of the conodont fauna was partially funded by a Sydney Grammar 
School Science Fellowship. Prof. R. Ethington kindly provided 
material of a Prioniodus species from the Wah Wah, Juab and 
Kanosh formations of the Ibex area, Utah for comparative study. 
Gary Dargan (Geological Survey of New South Wales) assisted 
with acid leaching, residue separation and other laboratory work. 
Scanning electron microscope photographs were prepared in the 
Electron Microscope Unit of the Australian Museum. Reviews by 
R, Ethington and G. Albanesi assisted clarification of our ideas. IGP 
publishes with permission of the Director, Geological Survey of NSW. 

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Stock no. 005R57B 


CONTENTS 


Algal-tube dwelling amphipods in the genus Cerapus from Australia and Papua 

New Guinea (Crustacea: Amphipoda: Ischyroceridae). 

.J.K. Lowry & P.B. Berents 153 

A review of the Australian fossil storks of the genus Ciconia (Aves: Ciconiidae), 

with the description of a new species. Walter E. Boles 165 

A new flightless gallinule (Aves: Rallidae: Gallinula) from the Oligo-Miocene 
of Riversleigh, northwestern Queensland, Australia. Walter E. Boles 179 

Revision of the genera Sphodropoda, Trachymantis and Zopheromantis 
(Mantodea: Mantidae: Mantinae).G.A. Milledge 191 

A new Middle Devonian arthrodire (placoderm fish) from the Broken River area, 

Queensland. Gavin C. Young 211 

Amphipods of the genera Ceradocus, Dulichiella, Melita and Nuuanu (Crustacea: 

Melitidae) from Mauritius, Indian Ocean. 

.Chandani Appadoo & Alan A. Myers 221 

New and little-known melitid amphipods from Australian waters (Crustacea: 

Amphipoda: Melitidae).J.K. Lowry & R.T. Springthorpe 237 

Revision of two prioniodontid species (Conodonta) from the Early Ordovician 

Honghuayuan Formation of Guizhou, South China. 

.Yong Yi Zhen, Jianbo Liu & Ian G. Percival 303 


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RECORDS OF THE AUSTRALIAN MUSEUM 2005 Vol. 57 No. 2