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ISSN 0067-1975
RECORDS OF THE
AUSTRALIAN
MUSEUM
VOLUME 57
NUMBER 2
8 June 2005
RECORDS OF THE AUSTRALIAN MUSEUM
Director: Frank Howarth
Editor: Shane F. McEvey
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Chair: G.D.F. Wilson (Invertebrate Zoology)
M.S. Moulds (Invertebrate Zoology)
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J.M. Leis (Vertebrate Zoology)
S. Ingleby (Vertebrate Zoology)
I.T. Graham (Geology)
D.J. Bickel (Invertebrate Zoology)
V.J. Attenbrow (Anthropology)
S.T. Ahyong (Invertebrate Zoology)
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ISSN 0067-1975
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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.
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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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© 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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© 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).
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entomologica Italiana, Bollettino 42: 3-38.
Giglio-Tos, E., 1917. Mantidi Esotici. Generi e specie nouve.
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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.
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1. London: British Museum (Natural History).
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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.
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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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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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Barnard, J.L., 1970. Sublittoral gammaridea (Amphipoda) of the
Hawaiian Islands. Smithsonian Contributions to Zoology 34:
1-286.
Barnard, J.L., 1972. Gammaridean Amphipoda of Australia, part
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Barnard, J.L., 1974. Gammaridean Amphipoda of Australia, part
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tropics: a review. Micronesica 12(1): 169-176.
Bate, C.S., 1862. Catalogue of the Specimens of Amphipodous
Crustacea in the Collections of the British Museum London,
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of New England, 312 pp. Ithaca and London: Cornell University
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biotopes sableux et sablo-vaseux de la region de Tulear et de
Nosy-Be (Madagascar). Tethys Supplement 5: 51-94.
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Memorie del Museo Civico di Storia Naturale di Verona (II
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Ledoyer, M., 1982. Crustaces Amphipodes Gammaridiens famille
des Acanthonotozomatidae a Gammaridae. Faune de
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group, with a new species from the Andaman Sea (Crustacea,
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Special Publication 23: 197-212.
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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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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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Stuttgart: E. Schweizerbart’sche.
Manuscript received 6 September 2004, revised 8 December 2004 and
accepted 21 December 2004.
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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