Memoirs of Museum Victoria 63(2): 107-128 (2006)
ISSN 1447-2546 (Print) 1447-2554 (On-line)
http://www.museum.vic.gov.au/memoirs/index.asp
A review of Australian long-horned caddisflies in the Oecetis pechana - group
(Trichoptera: Leptoceridae), with descriptions of thirteen new species
Alice Wells
Australian Biological Resources Study, Department of the Environment and Heritage, GPO Box 787, Canberra ACT 2601,
Australia (alice.wells@deh.gov.au)
Abstract Wells, A. 2006. A review of Australian long-horned caddisflies in the Oecetis pechana- group (Trichoptera: Leptoceridae),
with descriptions of thirteen new species. Memoirs of Museum Victoria 63(2): 107-128.
This final paper in a series reviewing Australian members of the long-horned caddisfly genus Oecetis McLachlan
(Trichoptera: Leptoceridae) deals with 21 species, 13 of them new, and all assigned here to the pechana-growp, so-named
for the most common and widespread of Australian Oecetis species. This group is diagnosed as having fork 1 in the
forewing sessile and, in the male, scale patches on the forewing and a single internal paramere in the phallus, although in
several species one or other of these male features is absent. Keys are given to the Oecetis species groups recognised in
Australia and to species in the pechana- group. Established pechana- group species are diagnosed and new species are
described. Male genitalic features are illustrated for all species save O. lurida Kimmins which was described from a female
and is probably a doubtful name. A homonym created in the name O. dilata Wells, 2004 is replaced with the name Oecetis
kimberleyensis nom. nov., and Oecetis cepaforma Wells, 2004 is synonymised with the widespread Oriental-Australian
O. hemerobioides McLachlan, 1866. Seventy-four Oecetis species are now recognised in the Australian fauna.
Keywords Taxonomy, Trichoptera, Leptoceridae, Oecetis pechana- group, new species, Australia
Contents
Introduction 107
Methods 108
Key to males of Oecetis species groups in Australia 108
Key to males of the Oecetis pechana species group in
Australia 109
Australian pechana - group species
Oecetis pechana Mosely, 1953 110
Oecetis walpolica Neboiss, 1982 110
Oecetis umbra Neboiss, 1977 110
Oecetis magelensis sp. nov. 110
Oecetis suteri sp. nov. Ill
Oecetis radonensis sp. nov. 116
Oecetis humphreyi sp. nov. 116
Oecetis kateae sp. nov. 116
Oecetis geniculata sp. nov. 117
Oecetis litua sp. nov. 117
Oecetis jenniae sp. nov. 120
Oecetis mouldsi sp. nov. 120
Oecetis theischingeri sp. nov. 121
Oecetis australis Banks, 1920 121
Oecetis quadrula sp. nov. 121
Oecetis buitenzorgensis Ulmer, 1951 ‘ -122
Oecetis burtoni Neboiss, 1979 123
Oecetis searica sp. nov. 123
Oecetis ada sp. nov. 123
Oecetis gilva Neboiss, 1977 127
Oecetis lurida Kimmins, 1953 128
Nomen novum and new synonymy 128
Acknowledgements 128
References 128
Introduction
Among Australian representatives of the long-horned caddisfly
genus Oecetis McLachlan (Trichoptera: Leptoceridae), a
persistent taxonomic problem has been the precise
determination of the scope of the species known as
Oecetis pechana Mosely, 1953. The name has been applied to
almost all Oecetis specimens having patches of scales on the
male forewing, collected Australia-wide, including the water
bodies and even stock watering troughs of the arid inland.
However, close inspection of males of many of the putative
O. pechana specimens reveals a range of forms sharing all or
some of a suite of character states. Here the Oecetis pechana-
growp is recognised for a set of 21 species, 13 of which are
newly described, and is defined to include a number of species
that lack scales on the male forewing, but share other
synapomorphies.
108
Alice Wells
Oecetis pechana was described originally from the south-
west of Western Australia, and subsequently was redescribed
and figured by Neboiss, 1977. Neboiss illustrated his work
with new figures of O. pechana from Tasmania, showing a key
feature of the species: the very elongate phallus. This feature
is also seen in a newly recognised species, O. suteri sp. nov.,
but is absent in superficially similar specimens that are
recognised here as representing several distinct species. O.
pechana as defined here is the most common and widespread
Australian species in the genus Oecetis. Adults vary greatly in
size, forewing length of those in the northern regions being as
little as half that of southern forms. Such a difference in size
range probably reflects differences in growth rates, the life
cycles being completed far more rapidly in the warm northern
waters than in the cooler southern waters. In support of this
contention, larger-sized adults were collected in the southern
Mt Lofty Ranges of South Australia, where a main late-spring
emergence of adults is followed by low numbers of specimens
throughout summer, and none in the cool winter months. In
contrast, samples from the north of the Northern Territory
contain only small-sized adults and, throughout 14 consecutive
months of light trap sampling on the Magela Creek at Jabiru
(1991-1992), Oecetis pechana adults were collected
regularly.
Oecetis pechana is often abundant in lentic water bodies
such as farm dams and lakes, the larvae in their tubular sand-
grain cases probably scavenging on organic material on the
substrate. Similarly, all other pechana -group species for which
larvae have been associated, have cases built of sand grains,
including Oecetis gilva Neboiss, 1977. This species also occurs
in large populations in some farm dams in the Adelaide Hills,
South Australia, and has been taken in large numbers from
several lakes in Victoria. Oecetis gilva as described by Neboiss
lacks scales on the male forewing, although conforming with
members of the pechana- group in other respects. Indeed,
membership of this group is confirmed by the fact that light
trap collections referable to O. gilva taken at two Victorian
lakes include males that have scales on the forewing. Quite
conceivably, these are simply variants of O. gilva, or even
hybrids between O. pechana and O. gilva. For the present,
they are accepted as O. gilva.
Thus, the Oecetis pechana- group is defined by a suite of
character states, not all shared by all members: forewing fork
1 without a footstalk, usually with cross veins, forks and
anastomoses marked by darkened membrane and hairs; tibial
spur formula 1,2,2; in the male, forewing usually with scale
patches, phallus usually with a single internal spine or
paramere, never with two, abdomen usually with tergites II-IV
darkly sclerotised, and genitalia usually with a small setae -
lined pouch on inner dorsal side of inferior appendages; and
larvae with tubular to conical or cornucopia-shaped cases
constructed of sand grains held together by silk secretion.
Species of Oecetis in the fauna of New Zealand, New Guinea
and Java also share the synapomorphies of these pechana-
group members.
This is the final work in a series of papers (Neboiss, 1989;
Wells, 2000, 2004) revising Australian species of Oecetis, and
raises to 74 the number of species recorded in this genus in
Australia. The key provided by Wells, 2004 to distinguish
males of the five Oecetis groups recognised for Australia is
reproduced here with slight modifications, and a key to males
of pecliana-growp species is given. The name dilata applied to
an Australian Oecetis species by Wells, 2004, thus creating a
homonym, is replaced; and the name Oecetis cepaforma
Wells, made available in the same work is here suppressed
in synonymy.
Methods
Material examined is lodged in the collections of Museum
Victoria, Melbourne (NMV), the Australian National Insect
Collection, Canberra (ANIC), the Queensland Museum,
Brisbane (QM) and the Northern Territory Museum of Arts
and Sciences, Darwin (NTM). Other abbreviations used here
are BMNH (The Natural History Museum, London, UK);
MCZ (Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts, USA); ARR is used for the
Alligator Rivers region, NT; OSS for the Office of the
Supervising Scientist, Darwin (previously at Jabiru); and WTH
is an NMV code for the Wet Tropics Heritage area of north-
eastern Queensland. In the locality data included with the lists
of material examined, the Australian political states are
abbreviated as follows: NSW (New South Wales); NT
(Northern Territory); Qld (Queensland); SA (South Australia);
Tas. (Tasmania) ; Vic. (Victoria); and WA (Western Australia).
Many of the scaly-winged species in this group are very
difficult to distinguish without dissection of the male abdomen
or clearing of the male genitalia to determine the length of the
phallus and size and shape of the single internal paramere.
Key to males of Oecetis species groups in Australia
[Modified from Wells, 2004]
1. Abdominal tergite VIII sculptured, expanded and
extended distally, forming shield over terminal abdominal
segments and genitalia reticulata- group (see Neboiss,
1989: fig. 2)
— Abdominal tergite VIII unmodified 2
2. Wings with veins strongly pronounced, fork 1 without a
footstalk (sessile)
longiterga- group (see Wells, 2004: figs 91, 94)
— Wings with veins not strongly pronounced, fork 1 with or
without footstalk 3
3. Phallus simple, lacking parameres
laustra- group (see Wells, 2004: figs 18, 46, 59)
— Phallus with 1 or more internal or external parameres or
spines 4
4. Forewing fork 1 sessile, wing lamina often bearing patches
of scales (androconia) (fig. 1); phallus usually with a single
internal paramere or spine, never with more than 1 spine
associated with the phallus (e.g. figs 3, 7, 8)
pechana- group (see below)
— Forewing fork 1 stalked, wing never bearing patches of
scales; phallus with 1 or more external parameres (spines
of the phallotheca)
complexa-growp (see Wells, 2000: figs 25, 26-33)
Australian Oecetis pechana - group
109
Key to males of the Oecetis pechana - species group in
Australia
[excluding O. lurida for which males are not associated]
1. Forewing with 1 or more patches of scales (androconia)
on lamina (figs 1, 6) 2
— Forewing without scales on lamina (figs 38, 39) 17
2. Inferior appendages in ventral view with length less than
twice width at base (figs 13, 31) 3
— Inferior appendages in ventral view with length about
twice width at base, or longer (figs 3, 7, 17), 4
3. Inferior appendages in ventral view more or less quadrate;
phallus in lateral view with paramere about 2-3 times
length of abdominal segment VIII, strongly arched
dorsally (figs 31, 32) O.jenniae
— Inferior appendages in ventral view more or less quadrate;
phallus about 3-4 times length of abdominal segment
VIII, almost straight (figs 13, 15) suteri
4. Inferior appendages in ventral view abruptly angled meso-
dorsally at about two-thirds length, rather than smoothly
curved, apices truncate (fig. 25) O. geniculata
— Inferior appendages not as above 5
5. Pre-anal appendages with length about 3 times width (figs
52, 53) O. ada
— Pre-anal appendages with length less than 3 times width
(figs 34, 36, 44, 45) 6
6. Inferior appendages in ventral view with a triangular
meso-basal lobe dorsally (figs 33, 35) 7
— Inferior appendages without a lobe as above 8
7. Inferior appendages in ventral view with the triangular
meso-basal dorsal lobe prominent and proximal to a sharply
triangular apico-mesial angle (fig. 35) O. theischingeri
Inferior appendages in ventral view with the triangular
meso-basal dorsal lobe less prominent and without a sharply
triangular apico-mesial angle (fig. 33) O. mouldsi
8. Inferior appendages in ventral view with length more than
3 times basal width; phallus short, about length of 1
abdominal segment, sclerotised paramere slender (figs 43,
45) O. buitenzorgensis
— Not as above 9
9. Inferior appendages in ventral view fused mesially in
basal third; in lateral view with length about 4 times width
(figs 49, 50) O. searica
— Not as above 10
10. Inferior appendages in ventral view stout, mesial margin
excavated in distal half; phallus greater than 4 times length
of abdominal segment VIII (fig. 3) O. pechana
— Not as above 11
1 1 . Inferior appendages in ventral view stout at base, gradually
excavated towards apex, slender distally; phallus in lateral
view with sclerotised paramere arched ventrally (figs 22-
24) O. kateae
— Not as above 12
12. Inferior appendages in ventral view stout throughout
length, mesially gradually excavated from base, apices
rounded (figs 7, 54) 13
— Not as above 15
13. Phallus with sclerotised paramere equal to about 2-3
times length of abdominal segment VIIF,^ 14
Phallus without sclerotised paramere (fig. 54)
O. gilva (part)
14. Inferior appendages in ventral view dilated lateral at about
half length, somewhat sinuous in appearance (fig. 7)
O. walpolica
— Inferior appendages in ventral view swollen basally,
gradually tapered towards apices (Neboiss, 1977: fig.
771) O. umbra
15. Inferior appendages in ventral view stout basally, sharply
excavated on mesial margin at about two-thirds length,
apices rounded; sclerotised paramere of phallus about
length of 2 abdominal segments (figs 9, 10) . O. magelensis
— Not as above 16
16. Inferior appendages about equal width throughout length,
sinuous in lateral view, apices rounded to truncate (figs
16, 17) , , O. radonensis
Inferior appendages about equal width for most of length,
distally tapered to acute convergent apices (figs 19, 20)
O. humphreyi
17. Forewing with distinctly spotty pattern; inferior
appendages close-pressed meso-ventrally for proximal
half, mesial margin excavated in distal half; phallus about
length of 2 abdominal segments, with sclerotised paramere
strongly arched dorsally, down-turned in distal section,
apical ly acute (figs 27, 28) O. litua
— Forewing without distinctly spotty pattern, although
usually with crossveins and vein forks and anastomoses
marked by dark membrane and hair; inferior appendages
not close-pressed for more than one-fifth length; phallus
usually shorter than length of 2 abdominal segments,
sclerotised paramere strongly reduced, absent or very
slender 18
18. Inferior appendages with length longer than basal width
(fig. 54) O. gilva (part)
— Inferior appendages with basal width greater than length
(figs 37, 40, 46) 19
19. Inferior appendages in ventral view with apico-mesial
angles acute; phallus with sclerotised paramere finely
whip-like distally; a pair of membranous lobes flanking
abdominal segment X (fig. 37) O. australis
— Not as above 20
20. Abdominal segment X in lateral view with an apically
acute ventral process at about half length; inferior
appendages in lateral view tapered toward apex (figs 40,
42); antennal segment with brush of long setae on leading
edges O. quadrula
— Abdominal segment X in lateral view expanded towards
apex, lacking a ventral process; inferior appendages in
lateral view rod-shaped (fig. 48); each antennal segment
with setae all about equal length O. burtoni
110
Alice Wells
Oecetis pechana Mosely
Figures 1-6, 56
Oecetis pechana Mosely in Mosely and Kimmins, 1953: 302, figs
213-215. -Neboiss, 1977: 143, figs 764-769. -Neboiss, 1986: 267.
Material examined. Holotype. Male, Yanchep, WA, 31°32.9'S,
115°41.2'E (BMNH).
Other material: Some 240 samples from all states of Australia
were examined, in collections of NMV, ANIC, QM, NTM.
Diagnosis. Male. Recognised by a combination of wing
features, including scale patches, and the general form of the
genitalia with inferior appendages in ventral view stoutly
clasper-shaped, smoothly curved mesially, and broadly rounded
apically; abdominal segment X a simple membranous plate; the
phallus long and straight, extending the length of 3-4 abdominal
segments, and having the paramere stout and darkly sclerotised.
Male forewing (fig. 1) length 5. 5-9.6 mm.
Males of closely similar species differ from O. pechana as
follows: O. walpolica has the inferior appendages in lateral view
tapered to narrowly rounded apex, phallus shorter, its length
equalling about 2 abdominal segments, and the paramere
strongly arched dorsally; O. katae has inferior appendages in
lateral view even more narrowly tapered and curved upwards,
phallus shorter, paramere arched ventrally; O. magelensis has
the inferior appendages shallowly excavated mesially, phallus
shorter, about 2-3 abdominal segments in length; O. gilva
usually has no scales on the forewing, inferior appendages in
lateral view attenuate apically, and phallus very short, about
length of 1.5 abdominal segments, without paramere.
Distribution. Australia-wide.
Remarks. As defined here, O. pechana is difficult to distinguish
from closely similar species, since the key diagnostic feature is
the length of the male phallus. This feature is surely biologically
significant, acting as a reproductive isolating mechanism,
determining mating success. Nevertheless, it does appear likely
that hybridisation between O. pechana and other species occurs,
and this may be confirmed if molecular studies are carried out
in the future. Oecetis suteri similarly has the phallus elongate,
but can easily be distinguished by the much reduced inferior
appendages. Oecetis pechana is often common in both lentic
and lotic systems, where its larvae build tube cases of sand.
Oecetis walpolica Neboiss
Figures 7, 8, 57
Oecetis walpolica Neboiss, 1982: 321, figs 123-125. —Neboiss,
1986: 267.
Holotype. Male, Frankland R., Circular pool, 6 km NE of Walpole,
WA (NMV: T-6596).
Material examined. Forty-five samples, most from WA, several from
NT.
Diagnosis. Male. Closely resembling O. pechana, especially in
form of the wings, shape of inferior appendages in ventral view,
and presence of the basi-dorsal pouch on the inferior appendages.
Distinguished from that species by the male genitalia with inferior
appendages attenuate apically in lateral view, and phallus about
half length of that of O. pechana and strongly arched dorsally.
Also resembling O. kateae sp. nov. in general form, but the
phallus of kateae is less strongly curved and arches ventrally, and
the inferior appendages in ventral view are more widely curved
and attenuate apically. Male forewing length 5.7-10 mm.
Distribution. Distributed from south-western WA inland and
northwards to the far north of NT.
Remarks. Oecetis walpolica is easily mistaken for O. pechana
and, similarly, is found in waterholes of the arid inland.
Oecetis umbra Neboiss
Figure 58
Oecetis umbra Neboiss, 1977: 144, figs 770-773. —Neboiss,
1986: 268.
Holotype. Male, Tas., Waldheim, Cradle Mountain National Park
(NMV T-5489).
Material examined. Tas.: 1 male. Lake Dobson, 20 Feb 1967,
E.F. Riek (ANIC); numerous males, females, Franklin R., Roaring
Creek junction, 1 km above Gordon R., 8 Jan 1977, Coleman, Neboiss,
Allbrook (NMV); 1 male, Gordon R., 1 km above First Split, 11 Jan
1977, Coleman, Neboiss, Allbrook, Swain (NMV); 2 males, 1 female,
swamp near Olga R., 19 km above Gordon R. Junction, 13 Jan 1977,
Neboiss and Swain (NMV); 2 males, 2 females. Caves Camp, NE of
New R. Lagoon, 2 Jan 1979, S.F. McEvey (NMV); numerous males,
females, south coast. Cavers Camp, New R. Lagoon, 3 Jan 1979, S.F.
McEvey (NMV).
Diagnosis. Male. Closely resembling O. pechana and O.
walpolica in general features such as scales on the wing and
clasper-shaped inferior appendages, but distinguished from
both these species by shorter phallus with the paramere only
slightly curved, and inferior appendages divergent apically,
rather than convergent.
Distribution. Known only from Tas.
Remarks. Neboiss, 1977 compared this species to the New
Zealand Oecetis unicolor McLachlan and Chatham I.
O. chathamensis Tillyard and it certainly resembles those
species more closely than it does Australian species.
Oecetis magelensis sp. nov.
Figures 9-11, 59
Material examined. Holotype. Male, NT, Alligator R. region, Magela
Creek, OSS Site 009, 12°42'S, 132°57'E, 17 Jan 1992, Wells (ANIC).
Paratypes. NT: 24 males, ARR, Magela Creek site 009, 15 Feb
1991, Wells (ANIC); 5 males, 5 females, ARR, Magela Creek at Rum
pipeline, 18-19 Feb 1991, P. Dostine (NMV).
Other material. NT: 1 male, Mataranka HS, Roper R., 25 Jan 1977,
M.S. and B.J. Moulds (NMV); 1 male, 1 female, 12°42'S, 132°57'E,
Kakadu National Park, Magela Creek, OSS Site 009, 28-29 Jan 1991,
P. Dostine (NTM); 2 males, 12°42'S, 132°57'E, Kakadu National Park,
Magela Creek, OSS Site 009, 15 Feb 1991, Wells (NTM); 3 males,
ARR, Magela Creek at Rum pipeline, 1-2 Apr 1991, P. Dostine (NMV);
3 males, 12°42'S, 132°57'E, Kakadu National Park, Magela Creek, OSS
Site 009, 1-2 Apr 1991, P. Dostine (NMV); 3 males, 1 female, Jabiru
Town Lake, 12°40'S, 132°53'E, 5 Apr 1991, Wells and Webber (NTM);
6 males, 12°42'S, 132°57'E, Kakadu National Park, Magela Creek, OSS
Australian Oecetis pechana - group
111
Site 009, 24 Apr 1991, Wells and Webber (NTM); 1 male, 1 female,
12°42'S, 132°57'E, Alligator R. Region, Magela Creek, OSS Site 009,
28-29 Apr 1991, R Dostine (NTM); 1 male, 1 female, Gregory National
Park, Limestone Gorge, 25-26 Oct 1991, J. Webber (NTM); 1 male,
1 female, 12°42'S, 132°57'E, Kakadu National Park, Magela Creek,
OSS Site 009, 3^1 Feb 1992, P. Dostine (NTM); 1 male, 12°42'S,
132°57'E, Kakadu National Park, Magela Creek, OSS Site 009,
21-22 Mar 1992, P. Dostine (NTM). WA: 2 males, 2 females, Maggie
Creek, 3 Feb 1978, J.E. Bishop (NMV); 3 males, Maggie Creek, 90 km
Kununurra-Wyndham, 3 Feb 1978, J.E. Bishop (NMV); 1 male, stream
opposite Dead Horse Gap, Lake Argyle, 19 Feb 1977, J.E. Bishop
(NMV); 1 male, 4 females, Lily Creek Waterhole, 15 km W of
Kununurra, 22 Feb 1977, J.E. Bishop (NMV); 1 male, 1 female, Dunham
R. W of Kununurra, J.E. Bishop, 22 Feb 1977 (NMV); 1 male, Ord R.
at Kununurra Dam, 22 Feb 1977, J.E. Bishop (NMV).
Diagnosis. Males resemble O. pechana and O. walpolica, the
male genitalia appearing closely similar in ventral view, but are
distinguished from the former by the shorter phallus with a
strongly arched paramere, and from both O. pechana and
O. walpolica by features of the inferior appendages: in ventral
view, the mesial margin is lined proximally with a row of small
setae, then excavated to form a small concavity; apically a
cluster of short setae; in lateral view the narrower distal section
of the inferior appendages is shorter and more closely resembles
that of O. suteri sp. nov.
Description. Male. Forewing slender, length 5. 0-9.0 mm, with
short hair on veins, dark spots at vein junctions formed by
pigmented membrane and darker vestiture; and a single broad
patch of scales. Abdominal segments II-IV with tergites darkly
scerotised, IX short, pale, slightly produced mid-dorsally, X a
simple, membranous plate, triangular distally. Genitalia (figs
9-11). Pre-anal lobes length about 1.5 times width, apically
rounded. Phallus membranous, length equal to 2-3 abdominal
segments, apico- ventral portion of phallotheca sharply beak-like
and down-turned, paramere darkly sclerotised, in lateral view
strongly arched dorsally. Inferior appendages stout throughout
length in ventral view, apically rounded, mesial margin lined
proximally with short setae, abruptly excavated at about two-
thirds length, with a pouch meso-dorsally on basal section; in
lateral view stout, with a short, rounded lobe dorsally.
Distribution. Found in the north of WA and the NT.
Remarks. Oecetis magelensis is difficult to distinguish from
O. walpolica , but has the inferior appendages in lateral view
with distal narrower section shorter and in ventral view tending
to converge, whereas in O. walpolica the inferior appendages
tend to turn outward toward apices.
Oecetis suteri sp. nov.
Figures 12-15, 60
Material examined. Holotype. Male, Holmes Jungle NT, 7 Jun 1991,
Horak and Wells (ANIC, on slide).
Paratypes. NT: 2 males. Kakadu National Park, Baroalba Springs,
25 Apr 1991, Wells and Webber (ANIC); 2 males, 12°42'S, 132°57'E,
Kakadu National Park, Magela Creek, OSS Site 009, 28-29 Jan 1992,
P. Dostine (NTM); 2 males, same locality, 3-4 Feb 1992, P. Dostine
(NTM); 2 males, Litchfield National Park, Walker Creek, 18-19 Apr
1992, Wells (NMV).
Other material. NT: 2 males, 12°52’S, 132°50’E, Koongarra, 15 km
E of Mt Cahill, 15 Nov 1972, J.C. Cardale (ANIC); numerous males,
females, 16°40’S, 135°51’E, Bessie Spring, 8 km ESE of Cape Crawford,
25 Oct 1975, J.C. Cardale (ANIC); 1 male, 1 female, 16°40'S, 135°51'E,
8 km ESE of Cape Crawford, 26 Oct 1975, J.C. Cardale (ANIC);
2 males, 1 female, 16°32'S, 136°10'E, Cattle Creek, 54 km S by W of
Borroloola, 27 Oct 1975, J.C. Cardale (ANIC); 1 male, 1 female, SAR,
14 Jun 1988, P. Dostine, Site 1 (NTM); 1 male (headless), ARRS, Radon
Springs, 13-14 Apr 1989, Suter and Wells (NMV); 2 males. Kakadu
National Park, Baroalba Springs, 25 Apr 1991, Wells, Webber and
Bickel (NTM); 3 males, 2 females. Berry Springs, 30 Oct 1991, Wells
and Webber (NTM); 2 males, 12°42'S, 132°57'E, Kakadu National
Park, Magela Creek, OSS Site 009, 28-29 Jan 1992, P. Dostine (NTM);
2 males, same locality and collector, 3^1 Feb 1992 (NTM); 2 males,
Litchfield National Park, Walker Creek, 18-19 Apr 1992, Wells (NTM).
Qld: 1 male, Lockerbie Scrub, Cape York Peninsula, 15 Apr 1975, M.S.
Moulds (NMV); 1 male. Cape York Peninsula, upper Jardine R.,
11° ITS, 142°35'E, 13 Oct 1979, M.S. and B.J. Moulds (NMV); 2 males,
same locality and collectors, 17 Oct 1979 (NMV); 14 males, 6 females,
Bertie Creek, 1 km SE Heathlands HS, 4 Feb 1992, D. Cartwright and
A. Wells (ANIC); 2 males, 1 female, tributary of Bertie Creek, 250 m
SW Heathlands HS, 4 Feb 1992, D. Cartwright and A. Wells (NMV); 1
male, Eliot Creek upstream junction Canal Creek, 6Feb 1992, D.
Cartwright and A. Wells (NTM); 4 males, 7 females, Dulhunty R., at
Telegraph Crossing, 10 Feb 1992, D. Cartwright and A. Wells (NMV);
17 males, 7 females, tributary of Bertie Creek, 250 m SW Heathlands
HS, 11 Feb 1992, D. Cartwright and A. Wells (ANIC, 2 males on slides);
5 males, 3 females. Gunshot Creek at Telegraph Crossing, 14-15 Feb
1992, D. Cartwright and A. Wells (QM); 1 male, 1 female, same locality
and collectors, 17 Feb 1992 (QM); numerous males, females. Gunshot
Creek at Telegraph Crossing, 4-5 Apr 1 992, M. Crossland (ANIC). WA:
1 male, 1 female, Millstream Crossing Pool, 2 Oct 1970, J.C. Cardale
(ANIC); 4 males, Drysdale R., upper reaches, 16°09’S, 125°58'E, 7 Sep
1996, 1. Edwards (NMV).
Diagnosis. Alone in the pechana-growp, O. suteri lacks fork
1 in the posterior wing; the forewing is more rounded apically
than most species in this group and the scale patches narrower.
Otherwise it resembles O. pechana and O. magelensis
superficially, but in the male is distinguished from both by the
inferior appendages broader than long in ventral view. Like
O. pechana , O. suteri has the phallus about the length of 4
abdominal segments.
Description. Male. Wings narrower than in O. pechana, with
dark markings at crossveins, anastomoses and at marginal ends
of veins; vestiture short; forewing with a narrow, elongate patch
of scales, discoidal and thyridial cells long, narrow, discoidal
cell extending distally beyond M-R, M-R distal to thyridial
cell; hind wing without fork 1. Forewing length, 4.3-4.6 mm.
Abdominal tergites II-IV darkly sclerotised. Genitalia, see figs
12-15. Abdominal segment IX of moderate length, dorsally
with paired papillae apico-mesially. Abdominal segment X
sub -triangular, short setae scattered apically. Inferior
appendages broad basally, slightly extended laterally, in lateral
view triangular; a rounded dorsal pouch present. Phallus
elongate, about 4 times length of abdominal segment IX, with
a long, almost straight paramere.
Distribution. Collected in WA from the Millstream and
Kimberley regions, in NT from the north, and in Qld, from the
tip of Cape York.
Etymology. Named for Phillip Suter.
112
Alice Wells
Figures 1-6, Oecetis pechana Mosely, male: 1, forewings and hind wings; 2-5, genitalia in lateral, ventral, dorsal and lateral views, respectively
(4, after Mosely and Kimmins, 1953; 5, image taken from slide preparation); 6, wing scales (after CSIRO 1991).
Australian Oecetis pechana - group
113
Figures 7, 8, Oecetis walpolica Neboiss, male genitalia in ventral and lateral views (8, image taken from slide preparation).
Figures 9-11, Oecetis magelensis sp. nov., male genitalia: 9, 10, ventral view and lateral view; 11, dorsal view of right inferior appendage.
114
Alice Wells
Figures 12-15, Oecetis suteri sp. nov, male genitalia: 12, dorsal view; 13, ventral view, 14, 15 lateral views (14, image taken from slide
preparation).
Figures 16, 17, Oecetis radonensis sp. nov., male genitalia in lateral and ventral views (16, image taken from slide preparation).
Australian Oecetis pechana - group
115
Figures 18-20, Oecetis humphreyi sp. nov., male genitalia in dorsal, lateral and ventral views.
Figures 21-24, Oecetis kateae sp. nov, male genitalia in dorsal, ventral and lateral views (24, image taken from slide preparation).
116
Alice Wells
Oecetis radonensis sp. nov.
Figures 16, 17, 61
Material examined. Holotype. Male, NT, Kakadu National Park,
Radon Springs, Suter and Wells, 13-14 Apr 1989 (NMV T-18840).
Paratypes. NT: 17 males, 14 females, same data as for holotype
(NMV and NTM: 1 male on slide); 4 males, 3 females. Little Baroalba
Creek, 25-26 Apr 1991, Wells and Webber (ANIC); 1 male. Kakadu
National Park, Baroalba Springs, 12°48'S, 132°49'E, 4 Mar 1992,
Wells (NTM).
Other material. NT: 1 male. Radon Creek, Kakadu National Park,
3 Sep 1979, J. Blyth (NMV). WA: 1 male, Mitchell Plateau, 30 Jan
1978, J.E. Bishop (NMV).
Diagnosis. Closely similar to O. humphreyi sp. nov. but male
distinguished by having inferior appendages in ventral view,
stout throughout length, rounded apically, and with 2 small
patches of setae at about two-thirds length; phallus longer with
paramere only very slightly curved.
Description. Male. Wings narrow, dark markings at crossveins,
anastomoses and at marginal ends of veins; vestiture short;
with a single elongate patch of scales; forewing length 4.8-5.5
mm. Abdominal segments III and IV with tergites sclerotised;
segment IX excavated mid-ventrally and thus appearing to have
well-developed lateral lobes; X spatulate in dorsal view, slender
and elongate in lateral view. Genitalia, see figs 16, 17. Pre-anal
appendages sub -triangular. Phallus about 1.5 segments long;
phallotheca with membranous apex sharply down-turned;
paramere in lateral view slightly curved. Inferior appendages
elongate and in ventral and lateral views almost uniformly
stout throughout length; in ventral view with apices in-turned,
without a basi-dorsal pouch; dorsally at about two-thirds length
a small group of short setae on a raised papilla, another cluster
on mesial margin, these possibly homologous with the sets of
setae on papillae in the basi-dorsal pouches seen in O. pechana
and other species.
Distribution. From the Kimberley region of northern WA and
the north of the NT, but not a commonly collected species.
Etymology. Named for the type locality, on the edge of Mt
Brockman in Kakadu National Park.
Oecetis humphreyi sp. nov.
Figures 18-20, 62
Material examined. Holotype. Male, NT, Radon Springs, 13-14 Apr
1989, Suter and Wells (NMV T-18841).
Paratypes. NT: 2 males, same data as for holotype (NMV);
1 male, 12°48'S, 132°49'E, Kakadu National Park, Baroalba Springs,
25 Apr 1991, Wells and Webber (NTM); 4 males, females, same
locality and collectors, 29 May 1991 (NTM); 2 males, same locality
and collectors, 4 Oct 1991 (NTM); 1 male. Berry Springs, edge of
Melaleuca nr Billabong, 8-9 Apr 1991, Wells and Horak (NTM);
1 male. Berry Springs, 4 Oct 1991, Wells (NTM).
Other material. NT: 3 males, 1 female. Butterfly Gorge, Katherine
Gorge National Park, 27 Jan 1977, M.S. Moulds (NMV); numerous
males, females, ARR Magela Creek OSS site 009, 15 Feb 1991,
A. Wells (NMV); 2 males, ARR Magela Creek 009, 15 Mar 1991,
Wells and Webber (NTM); numerous males, females, 12°40'S,
132°53'E, Jabiru, Town Lake, 2 Apr 1991, Wells and Webber (NTM); 1
male, 12°42'S, 132°57'E, Kakadu National Park, Magela Creek, at Rum
Pipeline, 6-7 May 1991, P. Dostine (NMV); 1 male, 1 female, Gregory
National Park, Limestone Gorge, 25-26 Oct 1991, J. Webber (NTM);
1 male, 12°42'S, 132°57'E, Kakadu National Park, Magela Creek, OSS
Site 009, 14 Apr 1992, Wells and Webber (NTM); 1 male, same locality,
21-22 Apr 1992, P. Dostine (NTM). WA: 1 male, 1 female. Dead Horse
Springs, Lake Argyle, 19 Feb 1977, J.E. Bishop (NMV); 1 male. Fine
Springs Creek, on road between Lake Argyle tourist village and
Duncan Highway, 23 Feb 1977, J.E. Bishop (NMV).
Diagnosis. Closely similar to Oecetis radonensis, but male
with inferior appendages distally more strongly tapered to
sharper apices, a row of setae on the inner margin sub-apically;
abdominal segment X triangular; and phallus of medium length
with paramere, in lateral view, strongly arched dorsally.
Description. Male. Wings with vestiture short; forewing with a
broad scale patch. Anterior wing length, 4. 6^1.7 mm. Abdominal
tergites II-IV darkly sclerotised; segment IX strongly excavated
ventrally, with a pair of small apico-dorsal papillae; segment X
broad-based, triangular in dorsal view, elongate and slender in
lateral view. Genitalia, see figs 18-20. Pre-anal appendages well
separated, in dorsal view, triangular. Phallus of moderate length,
apically beak-like in lateral view; paramere strongly twisted at
base in ventral view, in lateral view strongly arched dorsally.
Inferior appendages in lateral view tapered towards apex, in
ventral view stout with apices convergent, short setae on inner
subapical margin, basi-dorsal pouch small and rounded.
Distribution. Collected from the north of the NT and the
Kimberley Region of northern WA.
Etymology. Named for Chris Humphrey.
Oecetis kateae sp. nov.
Figures 21-24, 63
Material examined. Holotype. Male, NT, 12°40’S, 132°53’E, Jabiru,
Town Lake, 16 Feb 1991, Wells (ANIC).
Paratypes. NT: 2 males, 14 females, same data as for holotype
(NTM, ANIC); 22 males, 38 females, 12°42'S, 130°58'E, Berry
Springs, 9 Apr 1991, A. Wells and M. Horak (NTM, NMV);
14 males, 23 females, 13°03'S, 130°47'E, Litchfield National Park,
Florence Falls, 6 Jun 1991, Wells and Webber (ANIC).
Other material. Ninety-seven samples from NT, Qld, NW WA and
NE NSW.
Diagnosis. Recognised by inferior appendages more slender
and smoothly curved than those of O. pechana and O. walpolica,
and by the ventrally arched paramere of the phallus.
Description. Male. Wings with short hair, darkly pigmented
hair and membrane at anastomoses of veins and crossveins;
forewing with a single large patch of scales. Forewing length,
5. 1-5.9 mm. Abdominal tergites II-IV darkly sclerotised,
segment IX narrow, almost a regular annulus, dorsally slightly
pointed apico-mesially, segment X narrow, tapered towards
apex. Genitalia, see figs 21-24. Pre-anal appendages short,
wider than long. Inferior appendages stouter basally than distally
but almost C-shaped in ventral view, in lateral view broad-based,
slender, upturned distally, basi-dorsal pouch present. Phallus
about length of 3 abdominal segments, membranous ventral
phallotheca small, not down-turned, paramere arched ventrally.
Australian Oecetis pechana - group
117
Distribution. A widely distributed species that is often abundant
in light trap samples from the Kimberley of WA, through the
north of the NT to northern Qld, and south east to the border of
Qld and NSW.
Remarks. Superficially males of this species resemble those of
O. walpolica save that the inferior appendages are distally
more slender, but close examination of the male genitalia
confirms the unique state of the paramere of the phallus,
arching in the opposite direction from that of O. walpolica, and
far less strongly curved.
Etymology. Named for Kate Humphrey.
Oecetis geniculata sp. nov.
Figures 25, 26, 64
Material examined. Holotype. Male, NT, Litchfield National Park,
Billabong, 18-19 Apr 1992, A. Wells (ANIC).
Paratypes. NT: 21 males, 33 females, same data as for holotype
(NTM and ANIC); 17 males, 15 females, Humpty Doo, Solar Village,
9 Apr 1991, Horak and Upton (NMV).
Other material. Forty-five samples from the north of NT.
Diagnosis. Readily distinguished by the almost jointed
appearance of the inferior appendages in ventral view, the
unusually slender wings, and markings on the forewings.
Description. Male. Wings with vestiture short; forewing with a
patch of scales, but scales appear to be readily dehiscent so that
often specimens are collected without scales. Forewing length
4.6-5.0 mm. Abdominal segments II-IV darkly sclerotised,
segment IX of moderate length, segment X sub -triangular.
Genitalia, see figs 25-26. Pre-anal appendages almond-shaped.
Inferior appendages stout, elongate, ridged and turned inward
at about two-thirds length, giving a jointed appearance; without
a basi-dorsal pouch. Phallus about 1.5 abdominal segments
long, paramere straight.
Distribution. Often abundant in samples from the north-west
of NT.
Etymology. The name derives from the Latin geniculatus —
knotty, for the somewhat jointed appearance of the inferior
appendages.
Remarks. This species is small and pale, its narrow wings
lacking the usual dark markings of the pechana-gvowp. It is
commonly collected at billabongs or other Stillwater bodies.
Oecetis litua sp. nov.
Figures 27-29, 65
Material examined : Holotype. Male, 12°40'S, 132°53'E, Jabiru, Town
Lake, 30 May 1991, Wells and Webber (ANIC).
Paratypes. NT: 46 males, 9 females. East Alligator R. at Cahill's
Crossing, Wells and Suter, 27 May 1988 (NMV).
Other material. NT: 2 males, 12°17'S, 133°20'E, Cooper Creek,
11 km SW of Nimbuwah Rock, 3^1 Jun 1973, J.C. Cardale (ANIC);
3 male, 3 females, 12°06'S, 133°04'E, Cooper Creek, 19 km SE of Mt
Borrodaile, 5-6 Jun 1973, J.C. Cardale (ANIC); 1 male, 12°25’S,
132°58'E, 1 km of Cahills Crossing (East Alligator R.), 7-8 Jun 1973,
J.C. Cardale (ANIC); 3 males, 12°52’S, 132°50’E, Koongarra,
15 km E of Mt Cahill, 12-13 Jun 1973, J.C. Cardale (ANIC); 1 male,
12°52'S, 132°47'E, Nourlangie Creek, 8 km E of Mt Cahill, 14-15 Jun
1973, J.C. Cardale (ANIC); 2 males, 12°57’S, 132°33'E, Jim Jim Creek,
19 km WSW of Mt Cahill, 17 Jun 1973, J.C. Cardale (ANIC); 1 male,
1 female, 15°58'S, 136°21'E, 12 km NNE of Borroloola, 1 Nov 1973,
J.C. Cardale (ANIC); numerous males, females, Katherine R. Gorge
Nat. Pk, 13 Aug 1979, J. Blyth (NMV); numerous males, females,
Adelaide R„ 15 km E of Stuart Highway, 15 Aug 1979, J. Blyth (NMV);
5 males, 1 female. South Alligator R., UDP Falls [Gunlom], 7 Sep
1979, J. Blyth (NMV); 5 males, junction of Arnhem Highway and
Oenpelli Rd, 26-27 Jun, 1980, M.B. Malipatil (NMV); 2 males, UDP
Falls [Gunlom], 18-19 Jul 1980, M.B. Malipatil (NMV); 2 males,
ARRS, South Alligator R. at Gimbat OSS Station, 28 Apr 1988, P.
Dostine (NMV); 1 male, Magela Creek, Stoned Billabong, 15 May
1988, Wells and Suter (NMV); 2 males 3 females, ARRS, Radon
Springs, 18 May 1988, A. Wells and P. Suter (NMV); 3 males,
Coonjimba Billabong, 19 May 1988, P. Suter and A. Wells (NMV); 1
male, Magela Creek at Ranger pipe outlet, 20 May 1988, A. Wells and
P. Suter (NMV); 1 male, ARRS, Ranger Mine RP1, 20 May 1988, A.
Wells and P. Suter (NMV); 2 males, Magela Creek at Ranger pipe
outlet, 23 May 1988, A. Wells and P. Suter (NMV); 1 male, 1 female,
ARRS, South Alligator R. below Fisher Creek junction, 24 May 1988,
A. Wells and P. Suter (NMV); 2 males, ARRS, South Alligator R. at
Gimbat OSS Station, 24 May 1988, A. Wells and P. Suter (NMV); 2
males. South Alligator R. below BHP camp, 25 May 1988, P. Suter and
A. Wells (NMV); 1 male, 1 female. South Alligator R., 14 Jun 1988, P.
Dostine, site 1 (NMV); 2 males, 12°36'S, 132°53'E, ARRS Gulungul
Creek, inlet to Gulungul Billabong, 20 Apr 1989, A. Wells and P. Suter
(NMV); 1 male, Litchfield National Park, Florence Falls, 9 Apr 1991,
Horak, Wells and Upton (NTM); 1 male, 12°42'S, 130°58'E, Berry
Springs, 9 Apr 1991, Wells and Horak (NTM); numerous males,
females, 12°42’S, 132°57’E, Kakadu National Park, Magela Creek,
OSS Site 009, 24 Apr 1991, Wells and Webber (NTM); 1 male. Little
Baroalba Creek, 25-26 Apr 1991, Wells and Webber (NTM); 4 males,
1 female, Jabiru Town Lake, 13 May 1991, C. Humphrey (ANIC); 1
male, Jabiru Town Lake, 30 May 1991, Wells and Webber (NMV); 1
male, 1 female, 12°48’S, 132°49’E, Kakadu National Park, Baroalba
Springs, 16 Aug 1992, Wells and Webber (NTM); 1 male, 13°16’S,
132°49’E, Kakadu National Park, Jim Jim Falls, camp site, 13 Aug
1996, 1. Edwards (NMV). WA: 2 males, Drysdale R. National Park via
Carson R. Station, 14°37'S, 125°56'E, 31 Aug 1996, I. Edwards
(NMV); 2 males, 1 female, Drysdale R. Crossing, Kalumbaru Rd,
15°42'S, 126°22'E, 28 Aug 1996, 1. Edwards (NMV). Qld: numerous
males, females. Palmer R., 20 Jun 1971, E.F. Riek (ANIC); 1 male,
Mulgrave R„ W of Gordonvale, 29 Apr 1979, A. Wells (NMV).
Diagnosis. 1 of several Australian species with distinctively
spotted wings that give a somewhat moth-like appearance. In
general appearance resembling O. hemerobioides McLachlan,
1866 and O. dostinei Wells, 2004, but distinguished from those
2 laustra- group species by having males with stout inferior
appendages, and the phallus with a slender internal paramere.
Description. Male. Wings of typical shape, without scales, but
with pattern of spots, and long downy hair on veins. Anterior
wing length 5.7-6. 0 mm. Abdominal segment IX narrow mid-
ventrally, with pronounced lateral lobes, segment X broad-
based, triangular in dorsal view, slender, tapered in lateral
view. Genitalia, see figs 27-29. Pre-anal appendages rounded
apically. Phallus stout, short; aedeagus membranous with a
sclerotised band apically forming a sharp ‘beak’ in lateral view;
paramere slender, in lateral view strongly curved, slightly
sinuous in dorsal and ventral view. Inferior appendages stout at
118
Alice Wells
Figures 25, 26, Oecetis geniculata sp. nov., male genitalia in ventral and dorsal views.
Figures 27-29, Oecetis litua sp. nov., male genitalia in ventral, lateral and dorsal views.
Figures 30-32, Oecetis jenniae sp. nov., male genitalia in dorsal, ventral and lateral views.
Australian Oecetis pechana - group
119
Figures 33, 34, Oecetis mouldsi sp. nov., male genitalia in ventral and lateral views.
Figures 35, 36, Oecetis theischingeri sp. nov., male genitalia in ventral and lateral views.
Figures 37-39, Oecetis australis (Banks): 37, male genitalia in ventral view; 38, forewing; 39, downy hair on wing veins.
120
Alice Wells
base, the mesial ventral margins closely adpressed for 1st half
length, then abruptly excavated such that the appendages taper
to conical lobes distally ; in lateral view quite irregular in shape,
broadest medially; dorsal pouch reduced to an angled row of
stout setae in basal part of appendage.
Distribution. Known from the north of NT and northern Qld,
and from 1 sample from northern WA.
Remarks. Within the pechana- group, this species is distinctive
in its wing pattern and colouration. Yet the paramere and
vestiges of the basi-dorsal pocket on the inferior appendages,
clearly place it with other pechana-growp species.
Etymology. The name is derived from the Latin — lituus, a
curved staff, being descriptive of the paramere of the phallus in
lateral view.
Oecetis jenniae sp. nov.
Figures 30-32, 66
Material examined. Holotype. Male, NT, 12°42'S, 130°58'E, Berry
Springs, 31 Mar 1992, Wells (ANIC).
Paratypes. NT: 18 males, 17 females, same data as for holotype
(ANIC, NMV); 42 males, 22 females, same locality, 17 May 1992,
Wells (NTM).
Other material. 105 samples, some comprising numerous specimens,
from NT, WA, Qld and NSW (one sample only).
Diagnosis. Oecetis jenniae resembles O. mouldsi, having
similar male genitalia; however, it lacks a prominent apico-
dorsal lobe, and has wing venation of a more normal pechana
type, the veins not particularly emphasised. In having wing
scales and in the pattern of wing markings, O. jenniae resembles
O. pechana, but the male inferior appendages are wider
than long and not clasper-shaped, features shared with
O. theischingeri from which it is distinguished by lacking an
apico-mesial lobe on the inferior appendages. The inferior
appendages of this species resemble most closely those of
O. burtoni and O. quadrula, but both of those species have long
downy vestiture on the male wing and no scales.
Description. Male. Wings with vestiture short, pattern of dark
areas at vein anastomoses and at margin; forewing with a patch
of scales, which may be narrow or broad. Anterior wing length,
3.9-4.7 mm. Abdominal tergites II-IV darkly sclerotised,
segment IX annulate, segment X short, rounded in ventral view,
slender in lateral view. Genitalia, see figs 30-32. Pre-anal
appendages rounded apically. Phallus about length of 2.5 to 3
abdominal segments; paramere S-shaped in ventral view, in
lateral view arched dorsally. Inferior appendages stout,
subquadrate in ventral view, with a short dorso-lateral lobe,
basi-dorsal pouch present.
Distribution. Widespread in the north of the continent, but also
occurring in the south-east of Qld, west of the divide in central
NSW, and in the Pilbara region of WA.
Etymology. Named for Jenni Webber.
Remarks. Oecetis jenniae as recognised here is rather variable
in features of male genitalia and in the breadth of the scale
patches on the male wing. Most specimens from across northern
Australia conform to the type specimen from the north of the
Northern Territory in having a broad scale patch on the
forewing, and the apex of the dorsal lobe of the inferior
appendages rounded, rather than acuminate. Some specimens
from north-western Western Australia have only a narrow scale
patch on the forewing, and the apex of the dorsal lobe of the
inferior appendages acuminate, rather than more rounded. For
the present a single species is recognised; it is often abundant
in light trap collections.
Oecetis mouldsi sp. nov.
Figures 33, 34, 67
Material examined. Holotype. Male, WA, Midstream, Fortescue R.,
S of Roeborne, 12 Nov 1978, M.B. and B.J. Moulds (NMV T-18842).
Paratypes. WA: 1 male, Drysdale R. headwaters, 30 km NW
Mt Elizabeth HS, 30 Sep 1979, J. Blyth (NMV); 1 male, 20°18.48'S,
119°15.16’E, N Pilbara, de Grey R., 25 Apr 1992, P.S. Cranston and
PJ. Gullan (ANIC); 5 males, 9 females, 18°06'S, 125°42'E, Geikie
Gorge, 5 Oct 1996, 1. Edwards (NMV). NT: 1 male, 2 females, 12°40'S,
132°53’E, Jabiru, Town Lake, 17 May 1991 Wells and Webber
(ANIC).
Other material. WA: 15°12'S, 128°12'E, Spillway Creek, Ord R.
Dam, 20 Feb 1977, J.E. Bishop (NMV); 1 male, 1 female. Spillway
Creek, 3 Feb 1978, J.E. Bishop (NMV); 2 females. Limestone Gorge,
16°02’S, 130°23'E, 23-26 Jun 1986, M.B. Malipatil, Operation Raleigh
1986 (NMV); 1 male, Wickham R., 23 km downstream Humbert R.
Station, 16°49'S, 131°28'E, 11 Jul 1986, I. Archibald, Operation
Raleigh 1986 (NMV); 1 male, 17°01'S, 126°14'E, Bell Gorge,
Melaleuca Hole, 13 Sep 1996, I. Edwards (NMV); 1 male, 16°58'S,
125°59'E, Moll Gorge, Hann R„ 15 Sep 1996, I. Edwards (NMV).
NT: 1 male, 5 km W of King R., SW of Katherine,
5 Feb 1977, M.S. and B.J. Moulds (NMV); 1 male, 1 female, Katherine
R. Gorge National Park, 13 Aug 1979, J.E. Blyth (NMV); 1 male,
3 km E of Howard Springs, 17 Aug 1979, J. Blyth (NMV); 3 males,
5 females, Jasper Creek, Victoria R. Downs Rd., 45 km SSE Timber
Creek, 17 Sep 1979, J. Blyth (NMV); 4 males, 1 female, UDP Falls
[Gunlom], 18-19 Jul 1980, M.B. Malipatil (NMV); 1 male, 4 females,
Roderick Creek, Gregory National Park, 15°38’S, 131°22'E, 4-6 Jul
1986, I. Archibald, Operation Raleigh 1986 (NMV); 3 males, ARR,
Magela Creek, 15 Feb 1991, A. Wells (NMV). Qld: 9 males, 5 females.
Upper Ross R. below weir, S of Townsville, 8 May 1979, A. Wells
(NMV); 1 male, 21°07'S, 148°38'E, Finch Hatton Gorge, 14 Nov 1982,
T. Hinger (NMV).
Diagnosis. Closely resembling O. jenniae, but distinguished
from that species by the more pronounced venation of the
forewing, and in the male genitalia, by the inferior appendages
in ventral view well separated mesially by a triangular apico-
mesial process.
Description. Male. Wings with veins more strongly pronounced
than usual for pechana-growp species, with vestiture short and
pattern of dark areas at vein anastomoses and at margin;
forewing with a large patch of scales, length 6. 0-6.3 mm.
Abdominal tergites II-IV darkly sclerotised; segment IX
annulate, postero-lateral margin produced slightly, forming a
short lateral lobe; segment X membranous, slender in lateral
view. Genitalia, see figs 33, 34. Pre-anal appendages rounded
apically. Phallus about length of 2 abdominal segments, in
Australian Oecetis pechana - group
121
lateral view with paramere arched dorsally. Inferior appendages
with a basi-dorsal pouch; in ventral view broad-based, with
slender dorso-lateral lobe, and a triangular process near base
on mesial margin; in lateral view, stout.
Distribution. From north-western WA to the north of the NT,
and from 2 localities in north-eastern in Qld.
Remarks. Superficially males of this species resemble those of
O. jenniae, with which they are often collected, but they are
clearly differentiated from that species by the more pronounced
wing venation, and narrower inferior appendages.
Etymology. Named for Max Moulds.
Oecetis theischingeri sp. nov.
Figures 35, 36, 68
Material examined. Holotype. Male, Qld, Booloumba Creek, 8 km
SW of Kenilworth, 26°39'S, 152°39'E, 12 Dec 1984, G. Theischinger
(NMV T-18843, on slide).
Paratypes. Qld: 4 males, same data as for holotype (NMV, 1 on
slide); 1 male. Finch Hatton Gorge, 21°07'S, 148°38'E, 14 Nov 1982,
T. Hinger (NMV, on slide); 13 males, Strathpine, nr Brisbane, 27°19’S,
153°00’E, 3 Dec 1984, G. Theischinger (NMV, 1 male on slide).
Other material. Qld: 1 male, 1 female. Upper Brookfield nr
Brisbane, 4 Apr 1967, N. Dobrotworsky (NMV); 1 male. West Claudie
R., 17 Sep 1974, M.S. Moulds (NMV); 1 male, same locality and
collector, 30 Sep 1974 (NMV); 1 male. Middle Claudie R., Iron Range,
29 Sep 1974, M.S. Moulds (NMV); 12 males, 1 female, same locality
and collector, 2-9 Oct 1974 (NMV); 2 males, same locality and
collector, 23 Oct 1974 (NMV); 1 male. Funnel Creek, 53 km SW of
Sarina, 4 Jan 1975, M.S. Moulds (NMV); 1 male, 1 female, Bulimba
Creek, Site 5, Boorabbin Park, 9 Oct 1979, J.C. Cardale (ANIC);
2 males, Bulimba Creek, near Brisbane, Site Rl, Kimmax Street riffle,
23 Oct 1979, [collector not given], (NMV); 4 males, Bulimba Creek,
near Brisbane, Site R3, Willowbend lawns, 13 Mar 1980, A. Neboiss
(NMV); 1 male. Obi Obi Creek, 8 km SW of Mapleton, 23 Oct 1980,
A. Neboiss (NMV); 1 male, 7 females, Coondoo Creek, 30 km NE
Gympie, Toolara State Forest, 28 Oct 1980, A. Neboiss (NMV);
1 male, 28°03’S, 152°24’E, Cunninghams Gap, Western Fall,
1 Sep 1984, G. Theischinger (NMV). NSW: 1 male. Kings Gap, S
of Sandy Hollow, 5 Dec 1976, G. Daniels (NMV). Vic.: 1 male,
Dartmouth R. Survey, Stoney Creek, 14 Feb 1973, Foe VB (NMV)
[doubtful record].
Diagnosis. Resembling O. jenniae and O. mouldsi in having
short stout inferior appendages, but distinguished by having the
apico -mesial angle of the inferior appendages triangular, and
triangular apico-mesial lobes on the inferior appendages. In the
latter feature, O. theischingeri resembles O. australis-, but
O. australis is clearly distinct in having the apico-mesial lobes
on the inferior appendages with apices acute and wings with
vestiture downy.
Description. Male. Wings with vestiture short, pattern of dark
areas at vein anastomoses and at margin; forewing with a large
patch of scales. Anterior wing length, 5.0-5.2 mm. Abdominal
tergites II-IV darkly sclerotised; abdominal segment IX
annulate; segment X elongate -triangular in dorsal view, stout
in lateral view, apically capped by a cluster of short setae.
Genitalia, see figs 35, 36. Pre-anal appendages almond-shaped.
Phallus about length of 2-3 abdominal segments, paramere
more sinuous than arched. Inferior appendages wide basally in
ventral view with a triangular apico-mesial lobe; in lateral view
stoutly rounded dorsally; a basi-dorsal pouch present.
Distribution. Known from north-, central- and south-eastern
Qld and from 1 site in NSW. A single specimen bearing locality
data of Vic. is undoubtedly this species, but as it is the only
specimen from as far south, despite intensive collecting in that
area, it is considered a doubtful record until verified by further
collecting.
Etymology. Named for Gunther Theischinger.
Oecetis australis Banks
Figures 37-39, 69
Oecetina australis Banks, 1920: 350, pis 1-7. — Mosely and
Kimmins, 1953: 305. Neboiss, 1977: 145, figs 777-783. Neboiss, 1979:
840.
Holotype. Male, Vic., Melbourne (‘Thorey’) (ANIC, ex MCZ),
examined.
Oecetis situlus Korboot, 1964: 32, figs 1-7 (synonymised by
Neboiss, 1987: 139).
Holotype. Male, Qld, Cedar Creek, Mt Tamborine (QM).
Material examined. Ninety-two samples from Qld, Vic., NSW, Tas.,
SA and the NT.
Diagnosis. In males closely resembling O. burtoni in wing
form, and most similar to O. theischingeri in form of the
inferior appendages, but distinguished from both species by the
acute ventro-mesial processes on the inferior appendages and
from the latter by the downy vestiture on the inferior appendages,
the acute apices on the ventro-mesial processes and the whip-
like flagellum into which the paramere of the phallus extends.
Forewing length, 6.4-7.0 mm.
Distribution. Widespread in eastern Australia, from Tas. to
northern Qld and also collected from Kangaroo I., SA, and the
north-west of the NT.
Oecetis quadrula sp. nov.
Figures 40-42, 70
Material examined. Holotype. Male, NT, Kakadu National Park, UDP
Falls, 7 Sep 1979, J. Blyth (NMV T-18844, on slide).
Paratypes. NT: 1 male, Fitchfield National Park, Florence Falls,
9 Apr 1991, Horak, Upton and Wells (ANIC); 1 male, Jabiru Town
Fake, 13 May 1991, C. Humphrey (ANIC);1 male. Kakadu National
Park, Baroalba Springs, 4 Oct 1991, Wells and Webber (NTM);
1 male, same locality, 16 Jan 1992, Wells, Webber and Bickel (NTM);
1 male, Magela Creek Site 009, 3 Mar 1992, A. Wells (NTM).
Other material. NT: 1 male, Muirella Park, 12 Oct 1972, J.C.
Cardale (ANIC); 1 male, 12°17'S, 133°13'E, Birraduk Ck, 18 km E by
N of Oenpelli, 4-5 Jun 1973, J.C. Cardale (ANIC); 1 male, 12°31'S,
132°54'E, 9 km N by E of Mudginberri HS, 10-11 Jun 1973, J.C.
Cardale (ANIC); 1 male, 12°50'S, 132°51'E, 16 km E by N of Mt
Cahill, 16 Jun 1973, J.C. Cardale (ANIC); 1 male, 16°25’S, 136°05'E,
45 km SW by S of Borroloola, 5 Nov 1975, J.C. Cardale (ANIC);
male, Jim Jim Creek, 3 km below falls. Kakadu National Park,
1 Sep 1979, J. Blyth (NMV, on slide); male. South Alligator R. at
122
Alice Wells
UDP Falls [Gunlom], 7 Sep 1979, J. Blyth (NMV, on slide); 1 male,
ARR, creek 5 km W of OSS South Alligator Field Station, 19 Apr
1989, Wells and Suter (NMV); 1 male, Melville I., Andranangoo
Creek, ll 0 31'S, 130°54'E, 8 Oct 1996, G.R. Brown (NTM); 1 male,
Melville I., Maxwell Creek, 11°33'S, 130°35'E, 14 Oct 1996, G.R.
Brown (NTM); 1 male, same locality and collector, 15 Oct 1996
(NTM). WA: numerous males, females, 15°02’S, 126°55'E, Drysdale
R., 3-5 Jul 1975, 1.B.F. Common (ANIC); numerous males, females,
15°02'S, 126°55'E, Drysdale R„ 3-8 Aug 1975, I.B.F. Common and
M.S. Upton (ANIC); male, Mitchell Plateau, Camp Creek,
31 Jan 1978, J.E. Bishop (NMV, on slide); male, Mitchell Plateau,
Camp Creek at Crusher, 15 Feb 1979, J.E. Bishop (NMV, on slide);
3 males, Mitchell Plateau, at Mining Camp, 14°49'S, 125°50'E, 9-19
May 1983, J.C. Cardale (ANIC); 1 male, Kimberley, Mt Barnett
Station, Manning Gorge, 30 Apr 1992, P.S. Cranston (ANIC);
1 male, 15°55'S, 127°56'E, El Questro Station, Chamberlain R. pool,
26 Aug 1996, 1. Edwards, NMV); numerous males, females, 15°42'S,
126°22'E, Drysdale R. Crossing, Kalumburu Road, 28 Aug 1996,
I. Edwards (NMV); 2 males, 14°54'S, 126°12’E, King Edward R„
3 Sep 1996, 1. Edwards (NMV); 1 male, 16°40'S, 125°56’E, Manning
R. near Mt Barnett, 8 Sep 1996, 1. Edwards (NMV). Qld: 1 male. The
Crater, Atherton Tableland, 25 Apr 1970, S.R. Curtis (ANIC);
numerous males, females, Tewah Creek, Tin Can Bay, 17-18 Oct
1971, S.R. Monteith (ANIC); 1 male. Upper Freshwater Creek,
Whitfield Range near Cairns, 15 Dec 1974, Moulds (NMV, on slide);
1 male, Mulgrave R. W of Gordonvale, 29 Apr 1979, Wells (NMV); 1
male, 15°12'S, 143°52'E, Hann R. NW by W of Laura, 27 Jun 1986,
J. C. Cardale (ANIC). NSW: 4 males, 8 females, Wallagarough R.,
Princes Highway bridge, 29 Jan 1975, A. Neboiss (NMV). Vic.:
1 male, Wingan R., 8 km S Princes Hwy, 30 Jan 1975, A. Neboiss
(NMV, slide); 1 male, Tanjil R. near Old Tanjil, S34, Latrobe C.
Survey, 5 Feb 1980 (NMV).
Diagnosis. In the male, resembling O. australis and O. burtoni
in having long downy vestiture on the wings and no scales, but
distinguished by absence of acute ventro-mesial processes on
the inferior appendages, and inferior appendages short and
stout in ventral view, tapered distally in lateral view, phallus
short and beak-like, with a short slightly curved paramere.
The form of the inferior appendages of this species is similar
to O. jenniae, but the phallus is far shorter, its paramere short
and slender.
Description. Male. Wings rounded apically, with long hair on
veins; forewing without scales, length 4.6-7.0 mm. Antennae
with a brush of long setae on leading edge of segments.
Abdominal segments 3-5 with tergites sclerotised; segment IX
narrow, lateral angles produced, segment X elongate cone-
shaped in ventral view, in lateral view elongate, rounded
apically and with a small triangular ventral process. Genitalia,
see figs 40-42. Pre-anal appendages short, rounded, phallus
scarcely longer than wide, with membranous apex of aedeagus
folded and down-turned; a short, slightly curved paramere
present. Inferior appendages in ventral view subquadrate, with
apico-lateral lobes short, convergent; in lateral view, simple,
straight, tapered at apex, without basi-dorsal pouch.
Distribution. Found across northern Australia, from WA, NT
and Qld, and from south-eastern Vic.
Remarks. With a dense brush of setae on the leading edge of
the antennae, antennae erect (at least in spirit material), and
long maxillary palpi, males of this species look particularly
moth-like. Material here referred to O. quadrula shows
variability in genitalic form, but a conservative approach is
taken in assigning all to a single species as examination of
more material from the south-east of the continent is required
to justify recognition of two distinct morpho-species.
Etymology: Derived from the Latin, quadra — square, for the
shape of the inferior appendages in ventral view.
Oecetis buitenzorgensis Ulmer
Figures 43-45, 71
Oecetis buitenzorgensis Ulmer, 1951: 459, figs 708-712.
Holotype. Java (ZMB).
Material examined. Qld: 7 males. Little Mulgrave R., 28 Jun 1971,
E.F. Riek (ANIC); 3 males. West Claudie R., Iron Range,
17 Sep 1974, M.S. Moulds (NMV, 1 male on slide); 1 male. Middle
Claudie R., Iron Range, 2-9 Oct 1974, M.S. Moulds (NMV); 1 male,
Mcleod R., 15 km W of Mt Carbine, 22-23 Jun 1975, S.R. Monteith
(ANIC); 3 males, 1 female, Mulgrave R., W of Gordonvale,
29 Apr 1979, A. Wells (NMV: WTH 1387, 1 male on slide); 2 males,
5 km W by N Rounded Hill nr Hope Vale Mission, 15°17'S, 145°10'E,
7 Oct 1980, J.C. Cardale (ANIC); 1 male, 3 km ENE Mt Tozer,
12°44'S, 143°14'E, 2 Jul 1986, J.C. Cardale (ANIC); 4 males, 2
females, 9 km ENE Mt Tozer, 12°43'S, 143°17'E, 5-10 Jul 1986, J.C.
Cardale (ANIC); 3 males, 12°44’S, 143°16’E, Claudie R., Iron Range
National Park, 25 km NW Lockhart R., 10 Nov 1988, K. Walker
(NMV); 3 males, 21 females, Bertie Creek, 1 km SE Heathlands HS,
4 Feb 1992, D. Cartwright and A. Wells (QM); 1 male, 19 females.
Gunshot Creek at Telegraph Crossing, 18 Feb 1992, D. Cartwright
and A. Wells (QM); 4 males, 2 females, Kearney Falls, Goldsborough
Valley, 26-27 Mar 1997, G. Theischinger and L. Muller (ANIC,
1 male on slide); 7 males. Cape York Peninsula, Iron Range, Rainforest
Camping Ground, 6 Oct 2002, G. Theischinger (ANIC).
Diagnosis. Wings with long downy hair and, as in O. burtoni
and O. australis, male forewings lack scale patches. Males are
distinguished by genitalic features (figs 43-45): shape of the
inferior appendages which are stout and elongate in ventral
aspect, in lateral view narrowly tapered apically and with a
dorso-mesial expansion; and pre-anal lobes that are broadly
almond-shaped. Male forewing length, 3.9-5.5 mm.
Distribution. Known in Australia only from far north-eastern
Qld. Described from Bogor (= Buitenzorg), Java, and recorded
subsequently from New Guinea (Kimmins, 1962, from Kokoda,
Papua New Guinea).
Remarks. Oecetis buitenzorgensis is one of several
Oriental/New Guinean species recorded from far northern
Queensland. Chen, 1992 placing O. buitenzorgensis with a
group of Palaearctic and Oriental species (African, Indian
and Thai), commented that it is a nomen dubium, not
recognisable from Ulmer’s illustrations. However, the north-
eastern Australian specimens agree closely with Ulmer’s
figures, and conform well to other members of the pechana-
group as recognised here.
Australian Oecetis pechana - group
123
Oecetis burtoni Neboiss
Figures 46-48, 72
Oecetis burtoni Neboiss, 1979: 841, figs 37-40.
Holotype. Male, Qld, Fraser I. (NMV T-5749).
Material examined. Qld: 1 male, Mossman Gorge, 16 Jun 1971, E.F.
Riek (ANIC); numerous males, females, Tewah Creek, Tin Can Bay,
17-18 Oct 1971, S.R. Monteith (ANIC); 4 males. Iron Range, West
Claudie R., 17 Sep 1974, M. Moulds (NMV); 1 male. Iron Range,
Middle Claudie R„ 19 Sep 1974, M. Moulds (NMV); 1 male.
Cap Creek, Mt Finlayson Range, S of Cooktown, 23 Nov 1974,
M.S. Moulds (NMV); numerous males, females. Lake Wabby, S end
Fraser I., 14 Dec 1975, H. Burton (NMV); 1 male, Bluewater State
Forest, S end Paluma Range WNW of Townsville, 31 Jan 1981, M.S.
and B.J. Moulds (NMV); 4 males, 5 females, Seary’s Creek Rainbow
Beach, 25°58'S, 153°04'E, 6 Dec 1984, G. Theischinger (ANIC);
5 males, 6 females, 12°44’S, 143°35’E, Claudie R., Iron Range
National Park, 25 km NW Lockhart, 10 Nov 1988, K. Walker (NMV);
3 males, 5 females, 11°45'S, 142°35'E, Heathlands, 15-26 Jan 1992, 1.
Naumann and T. Weir (ANIC); 6 males, 2 females, Bertie Creek, 1
km SE Heathlands HS, 4 Feb 1992, D. Cartwright and A. Wells
(NMV); 3 males, 6 females. Tributary of Bertie Creek, 250 m SW
Heathlands HS, 4 Feb 1992, D. Cartwright and A. Wells (NMV); 1
male, Eliot Creek upstream junction Canal Creek, 6 Feb 1992, D.
Cartwright and A. Wells (NMV); 1 male, Dulhunty R. at Telegraph
Crossing, 11°50’S, 142°30'E, 8-9 Feb 1992, D. Cartwright and A.
Wells (NMV); 29 males, 4 females, Dulhunty R., at Telegraph
Crossing, 10 Feb 1992, D. Cartwright and A. Wells (NTM); 8 males,
13 females. Tributary of Bertie Creek, 250 m SW Heathlands HS, 11
Feb 1992, D. Cartwright and A. Wells (NMV); 5 males, 7 females.
Tributary of Bertie Creek, 250 m SW Heathlands HS, 11 Feb 1992, D.
Cartwright and A. Wells (NMV); numerous males, females. Cockatoo
Creek and McDonnell Creek junction, 11°39'S, 142°28'E, 13 Feb
1992, D. Cartwright and A. Wells (NMV); 1 male, 1 female. Gunshot
Creek at Telegraph Crossing, 18 Feb 1992, D. Cartwright and A.
Wells (NMV); 1 male. Cape York Peninsula, Heathlands, Bertie
Creek, 23 Mar 1993, M. Crossland (ANIC); 2 males, 3 females.
Gunshot Creek at Telegraph Crossing, M. Crossland, 4-5 Apr 1993
(ANIC); numerous males, females. Gunshot Creek at Telegraph
Crossing, M. Crossland, 10-11 Apr 1993 (ANIC). WA: 3 males, 1
female, Drysdale R. at Kalumburu Road crossing, Kimberley, 28 Sep
1979, J. Blyth (NMV). NT: 2 males, 2 females. Devil Devil Creek, 70
km SW of Daly R. Mission, 23 Aug 1979, J. Blyth (NMV).
Diagnosis. In lacking wing scales in the male, but having long
downy vestiture on the wings, this species resembles
O. australis, O. quadrula and O. buitenzorgensis, but in
features of male genitalia it more closely resembles
O. theischingeri from which it is distinguished by the very
short phallus with an equally short paramere, absence of the
triangular process apico-mesially on inferior appendages and
of the dorso-basal pouch on inferior appendages. Male
forewing length, 4.0-4.6 mm.
Distribution. Quite widespread around coastal areas of northern
Australia, although not very commonly collected. Possibly
records represent more than 1 species, as some variability is
recognised in form of the inferior appendages and tergite X. At
present this is considered too slight to warrant designation of
separate morpho-species.
Oecetis searica sp. nov.
Figures 49, 50, 73
Material examined. Holotype. Male, Qld, Seary’s Creek Rainbow
Beach, 25°58'S, 153°04'E, 6 Jan 1986, G. Theischinger (NMV T-
18845, on slide).
Paratypes. 3 males, same locality and collector as for holotype,
6 Dec 1984 (NMV, 1 on slide).
Diagnosis. One of the species with scales on the male forewing,
but distinguished from others by the medium length paramere
in the phallus strongly arched dorsally and elongate inferior
appendages which, in lateral view, are rod-like, taper gradually
toward the apex and lack a dorsal lobe.
Description. Male. Wings with short hair on veins, forewing
with a patch of scales. Anterior wing length, 4.3 mm. Abdominal
segment IX broader dorsally, with pronounced lateral lobes,
segment X triangular in dorsal view. Genitalia, see figs 49, 50.
Pre-anal appendages longer than wide. Phallus about length of
1.5 abdominal segments; paramere strongly arched dorsally.
Inferior appendages clasper-shaped in ventral view, stout
basally, narrow distally, rounded apically, in lateral view
elongate, with length almost 4 times as long as basal width,
tapered distally, without basi-dorsal pocket.
Distribution. Known only from the type locality in north-
eastern Qld.
Etymology. Named for the type locality.
Oecetis ada sp. nov.
Figures 51-53, 74
Material examined. Holotype. Male, NT, Litchfield National Park,
Ada Creek, at jump up, 24-25 Jun 1992, Wells and Webber (NTM).
Diagnosis. The patches of wing scales clearly place this species
in the pechana- group, but, like O. gilva, it lacks a paramere in
the very short phallus; and the pre-anal appendages are
unusually long for a pechana-growp species, being over 3 times
longer than wide.
Description. Wings with short hair on veins, fore wing with a
patch of scales. Male anterior wing length, 6.6 mm. Abdominal
segment IX narrow, lateral angles produced; segment X about as
long as pre-anal appendages, slender in dorsal view. Genitalia, see
figs 51-53. Pre-anal appendages elongate-ovoid. Inferior
appendages stout at base, in ventral view tapered to rounded
apices; in lateral view, with a triangular basi-dorsal lobe, and basi-
dorsal pouch. Phallus about length of 1 abdominal segment, with
a sharply returned, pointed apex; paramere absent.
Distribution. Known only from the holotype male from the
north of the NT.
Etymology. Named for the type locality.
Remarks. Although this species lacks a paramere it clearly
groups with pechana- group species on the basis of the general
form of the phallus, wing venation and presence of scale
patches.
124
Alice Wells
Figures 40-42, Oecetis quadrula sp. nov., male genitalia in ventral, dorsal and lateral views.
Figures 43-45, Oecetis buitenzorgensis Ulmer, male genitalia in ventral, dorsal and lateral views.
Figures 46-48, Oecetis burtoni Neboiss, male genitalia in ventral, dorsal and lateral views.
Australian Oecetis pechana - group
125
Figures 49, 50, Oecetis searica sp. nov., male genitalia in ventral and lateral views.
Figures 51-53, Oecetis ada sp. nov., male genitalia in ventral, dorsal and lateral views.
Figures 54, 55, Oecetis gilva Neboiss, male genitalia in ventral and lateral views.
126
Alice Wells
Figures 56-67, Distribution of species within Australia: 56, O. pechana Mosely; 57, O. walpolica Neboiss; 58, O. umbra Neboiss; 59, O.
magelensis sp. nov.; 60, O. suteri sp. nov.; 61, O. radonensis sp. nov.; 62, O. humphreyi sp. nov; 63, O. kateae sp. nov.; 64, O. geniculata sp. nov.;
65, O. litua sp. nov.; 66, O.jenniae sp. nov.; 67, O. mouldsi sp. nov.
Australian Oecetis pechana - group 1 27
Figures 68-75, Distribution of species within Australia: 68, O. theischingeri sp. nov.; 69, O. australis (Banks); 70, O. quadrula sp. nov.; 71, O.
buitenzorgensis Ulmer; 72, O. burtoni Neboiss; 73, O. searica sp. nov.; 74, O. ada sp. nov.; 75, O. gilva Neboiss.
Oecetis gilva Neboiss
Figures 54, 55, 75
Oecetis gilva Neboiss, 1977: 145, figs 774-776. —Neboiss, 1979:
840, figs 35, 36; Neboiss, 1986: 268.
Holotype. Male, Tas., South Esk R. near Evandale (NMV
T-5491).
Material examined. Eighty-seven samples examined, from localities
in south-eastern Australia; 2 samples examined from far north-
eastern SA.
Diagnosis. In features of wing venation, pattern of markings,
and male genitalia resembling O. pechana superficially, but
wings broader and more rounded apically, and male generally
without scales on anterior wing, inferior appendages tapered
distally. This species lacks a paramere in the phallus, a feature
shared with O. ada sp. nov., but unlike that species the
phallotheca forms a pale dorsal spine.
Revised description. Wings relatively broad, forewing length
9.1-9.6 mm, rounded apically, male forewing with or without
128
Alice Wells
scales. Abdominal segments III-V withtergites well sclerotised;
IX short; segment X narrow, membranous, rounded apically.
Male genitalia; figs 54, 55. Pre-anal appendages short, rounded
apically. Phallus about twice length of segment IX, beak-like
apico-ventrally, without a paramere, but in lateral view extended
in a short, pale apico-dorsal spine. Inferior appendages clasper-
shaped in ventral view, with basi-dorsal dorsal pouch; in lateral
view, expanded and rounded dorsally, tapered to acute apex.
Distribution. A common species in SE Australia, O. gilva
occurs from northern NSW, throughout Vic., to the Flinders
Ranges in SA and south to Tas.
Remarks. The diagnosis of O. gilva is expanded here to include
the curious scaly winged form that has been collected together
with O. gilva, s. str. from several lakes in Victoria. In all
respects other than presence of patches on the wings, these
animals appear to be identical. The winged forms could
conceivably be hybrids between O. gilva and O. pechana.
O. gilva is often abundant in still to slow waters; the larvae
build tube cases of sand.
Oecetis lurida Kimmins
Oecetis lurida Kimmins in Mosely and Kimmins, 1953 : 298, fig.
210. -Neboiss, 1986: 268.
Material examined. Holotype. Female, Broken Hill, NSW (BMNH).
Remarks. This species was described from a female with
uniformly pale, unspotted wings and while it may eventually
be found to be a senior synonym of O. gilva, which has been
found in inland waters, O. lurida is probably best considered a
doubtful name.
Nomen novum and new synonymy
The name Oecetis kimberleyensis nom. nov. is proposed to
replace the name, Oecetis dilata Wells, 2004: 98, preoccupied
by O. dilata Yang and Morse, 2000 (J. Morse, in litt.); and,
following advice from H. Malicky (in litt.), Oecetis cepaforma
Wells, 2004 is suppressed in synonymy with Oecetis
hemerobioides McLachlan, 1866. This action extends into
northern Australia the range of O. hemerobioides, described
from Sulawesi (as Celebes) and recognised previously Chen,
1992 from Thailand and Malaysia through Indonesia and the
Philippines to New Guinea. It is probable, too, that
O. oecetinellae Mey, 1990 from the Philippines is also referable
to this species.
Acknowledgements
Facilities for laboratory work were provided to me as a visiting
scientist at the Australian National Insect Collection, CSIRO
Entomology, Canberra, and facilities at the Australian
Biological Resources Study were used for preparation of the
manuscript and manipulation of images. The study is based on
collections in Museum Victoria, the Museum and Art Galleries
of the Northern Territory, Darwin, and the Australian National
Insect Collection, Canberra. CSIRO Entomology is thanked
for permission to reproduce the photograph of O. pechana
scales.
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People’s Republic of China. Memoirs of the American
Entomological Institute 64: viii + 309 pp.
Memoirs of Museum Victoria 63(2): 129-173 (2006)
ISSN 1447-2546 (Print) 1447-2554 (On-line)
http://www.museum.vic.gov.au/memoirs/index.asp
Nannastacidae (Crustacea: Cumacea) from eastern Bass Strait, the south-eastern
Australian slope, and Antarctica in the collections of Museum Victoria
IORGU PETRESCU
“Grigore Antipa” National Museum of Natural History, Kiseleff 1, Bucharest 011341, Romania (iorgup@antipa.ro)
Abstract Petrescu, I. 2006. Nannastacidae (Crustacea: Cumacea) from eastern Bass Strait, the south-eastern Australian slope, and
Antarctica in the collections of Museum Victoria. Memoirs of Museum Victoria 63(2): 129-173.
Forty-five species of Nannastacidae were identified in collections in Museum Victoria from eastern Bass Strait and the
south-eastern Australian slope. Twenty-eight are described as new: Campylaspis anae, C. angelae, C. australiensis, C.
edenensis, C. grossui, C. halei, C. hirsuta, C. latimera, C. longidentata, C. lynseyae, C. nowrae, C. poorei, C. rectangulata,
C. sculpta, C. serrata , C. setifera, C. spinifera, C. tasmaniensis, C. trisulcata, Procampylaspis australiensis, P. spinifera,
P. tasmaniensis, Styloptocuma australiense, S. granulosum, S. nodosum, S. poorei, S. spinosum, Styloptocumoides
australiensis and Vemacumella bacescui. The new genus Styloptocumoides is diagnosed and Styloptocuma Bacescu and
Muradian, 1974 and Vemacumella Petrescu, 2001 are recorded for the first time from Australia. The new species
Procampylaspis poorei is described from Antarctic waters.
Keywords Crustacea, Cumacea, Nannastacidae, new taxa, new records
Contents
Introduction 130
Campylaspis Sars, 1865 130
Campylaspis anae sp. nov. 131
Campylaspis angelae sp. nov. 132
Campylaspis aspera Hale, 1945 133
Campylaspis australiensis sp. nov. 133
Campylaspis echinata Hale, 1945 135
Campylaspis edenensis sp. nov. 135
Campylaspis grossui sp. nov. 136
Campylaspis halei sp. nov. 138
Campylaspis hirsuta sp. nov. 139
Campylaspis johnstoni Hale, 1937 140
Campylaspis latidactyla Hale, 1945 140
Campylaspis latimera sp. nov. 140
Campylaspis longidentata sp. nov. 142
Campylaspis lynseyae sp. nov. — 144
Campylaspis nowrae sp. nov. 145
Campylaspis pileus Foxon, 1932 146
Campylaspis poorei sp. nov. 146
Campylaspis pustulosa Hale, 1945 147
Campylaspis rectangulata sp. nov. 147
Campylaspis sculpta sp. nov. 148
Campylaspis serrata sp. nov. 149
Campylaspis setifera sp. nov. 151
Campylaspis spinifera sp. nov. 152
Campylaspis tasmaniensis sp. nov. 153
Campylaspis thetidis Hale, 1945 155
Campylaspis thompsoni Hale, 1945 155
Campylaspis triplicata Hale, 1945 155
Campylaspis trisulcata sp. nov. 155
Campylaspis uniplicata Hale, 1945 156
Campylaspis unisulcata Hale, 1945 156
Procampylaspis Bonnier, 1896 156
Procampylaspis australiensis sp. nov. 156
Procampylaspis poorei sp. nov. 158
Procampylaspis sordida Hale, 1945 159
Procampylaspis spinifera sp. nov. 159
Procampylaspis tasmaniensis sp. nov. 161
Scherocumella Watling, 1991 162
Scherocumella nichollsi (Hale, 1945) 162
Scherocumella sheardi (Hale, 1945) 162
Schizotrema Caiman. 191 1 162
Schizotrema aculeatum Hale, 1936 163
Styloptocuma Bacescu and Muradian, 1974 163
Styloptocuma australiense sp. nov. 163
Styloptocuma granulosum sp. nov. 164
Styloptocuma nodosum sp. nov. 165
Styloptocuma poorei sp. nov 167
Styloptocuma spinosum sp. nov. 168
Styloptocumoides gen. nov. 169
Styloptocumoides australiensis sp. nov. 169
Vemacumella Petrescu, 2001 170
Vemacumella bacescui sp. nov. 170
Acknowledgements 171
References 171
130
lorgu Petrescu
Introduction
The most important studies on Australian Cumacea were made
by the famous specialist, Herbert M. Hale, who dealt with 90%
of Australian fauna between 1936 and 1949. He added 32
species of Nannastacidae to the Australian list, 29 being new to
science. However, the first species known from this family from
Australia were Cumella cyclaspoides, Cumella gibba, and
Scherocumella nasuta, described by Zimmer, 1914 from Shark
Bay, Western Australia. Foxon (1932) described Campylaspis
pileus from the Great Barrier Reef. Two other papers have since
been published: Bacescu (1991) describing Campylaspis wardi
and Tafe and Greenwood (1996) adding a new species of
Schizotrema. Up to now, 40 species of Nannastacidae are known
from Australia (Stoddart and Lowry, 2003).
The material in Museum Victoria selected for study was 27
samples collected during two sampling campaigns. One was in
eastern Bass Strait at shallow shelf depths (see Coleman et al.,
1997 for a discussion of the fauna of this sampling program).
The second was on the south-eastern Australian continental
slope between Sydney, New South Wales and eastern Tasmania
(see Poore et al., 1994, for discussion of the diversity of isopods
from these deep-water samples). Only nine of numerous
samples were searched for nannastacids. Forty-five species of
Nannastacidae including 28 new species, one in a new genus,
were identified, bringing the total for Australia to 68. One
additonal new species was described from Antarctica.
Specimens are now registered in Museum Victoria, Melbourne
(NMV) and representatives are in the “Grigore Antipa” National
Museum of Natural History, Bucharest, Rumania (MGAB). Type
localities are given in full but other records are given only by
station number, and are expanded in the appendix.
Campylaspis Sars, 1865
Campylaspis Sars, 1865: 200.— Bacescu, 1992: 178-179.—
Stoddart and Lowry, 2003: 373-418.
Bacescua Muradian, 1979: 105.
Sarsicuma Muradian, 1979: 104.
Type species. Cuma rubicunda Liljeborg, 1855.
Remarks. Species of Campylaspis are cumaceans with a bulky
carapace, longer than 0.4 of total length, having a mandible
with large pars incisiva and lacinia mobilis, acute pars molaris,
and maxilliped 2 with a 3 -toothed dactylus. Fifteen species
are previously known from Australia; another 20 taxa are
added from this study (19 of them being new to science).
Key to species of Campylaspis from Australian waters
1. Eye lenses present 2
— Eye lenses absent 14
2. Carapace smooth without tubercles, spines, carinae or
lateral sulcus 3
— Carapace with tubercles, spines, carinae or a lateral
furrow or sulcus 4
3. Uropodal peduncle 3 times as long as its endopod
C. thompsoni Hale, 1945
— Uropodal peduncle 2.3 times as long as its endopod
C. similis Hale, 1945
4. Carapace with lateral sulcus, without ridges 5
— Carapace without lateral sulcus, with ridges and carinae
...7
5. Dactylus of pereopod 2 dilated distally
C. latidactyla Hale. 1945
— Dactylus of pereopod 2 not dilated distally 6
6. Dactylus of pereopod 2 as long as carpus plus propodus ....
C. unisulcata Hale, 1945
— Dactylus of pereopod 2 shorter than carpus plus
propodus C. minor Hale, 1945
7. Carapace with single, lateral rounded carina , w . K
C. uniplicata Hale, 1945
— Carapace with tubercles, spines, depressions and ridges
8
8. Carapace with ridges and lateral depressions 9
— Carapace with tubercles, spines and ridges 10
9. One ridge delimiting 1 large lateral excavation and 2 in
posterior half C. nipt a Hale, 1945
— Three lateral ridges delimiting 2 curved depressions
C. triplicata Hale, 1945
10. Carapace covered with spiniform projections, spines on
rest of body C. echinata Hale, 1945
— Carapace coverd with small tubercles 11
11. Carapace with very small tubercles anteriorly and
dorsolaterally C. roscida Hale, 1945
— Carapace with conical tubercles 12
12. Maxilliped 3, 3 times as long as wide
C. pustulosa Hale, 1945
— Maxilliped 3, 2 times as long as wide
C. aspera Hale. 1945
13. Dactylus of maxilliped 2 with 3 long teeth, twice as long
as propodus C. longidentata sp. nov.
— Dactylus of maxilliped 2 with teeth shorter or as long as
propodus 14
14. Carapace with lateral depressions .......... 15
— Carapace without lateral depressions 34
15. Carapace without ridges or carinae 16
— Carapace with ridges or carinae 29
16. Carapace with sulcus open on anterior extremity of
carapace „ ;V , 17
— Carapace with sulcus not reaching the anterior extremity
of carapace 25
17. Carapace smooth, excepting lateral depression 18
— Carapace with tubercles 19
18. Uropodal peduncle robust, with median crest, twice as
long as exopod C. anae sp. nov.
— Uropodal peduncle slender, no median crest, 1.4 times as
long as exopod C. rectangulata sp. nov.
19. Uropods long, elongated 20
— Uropods short, robust 22
20. Carapace with several rows of tubercles
C. tasmaniensis sp.nov.
— Carapace with 1-2 tubercles 21
21. Carapace with 1 pair of tubercles at basis of frontal lobe ..
ujgm C. lynseyae sp. nov.
— Carapace with 1 pair of tubercles at basis of frontal lobe,
2nd under sulcus C. spinifera sp. nov.
New nannastacid cumaceans from Museum Victoria
131
22. Dactylus of pereopod 2 long 23
— Dactylus of pereopod 2 short 24
23. Pseudorostral lobes meeting for distance of 3 times length
of ocular lobe C. thetidis Hale, 1945
— Pseudorostral lobes meeting for distance of half ocular
lobe C. sculpta sp. nov.
24. Uropods with terminal short setae with oval structures
C. wardi Bacescu, 1991
— Uropods with terminal short setae without oval
structures C. grossui sp. nov.
25. Carapace with tubercles 26
— Carapace without tubercles 27
26. Integument with acute tubercles
C. australiensis sp. nov.
— Integument with spiny tubercles C. hirsuta sp. nov.
27. Maxilliped 3 and pereopod 1 with large and concave
merus C. latimera sp. nov.
— Maxilliped 3 and pereopod 1 without large and concave
merus 28
28. Pereopod 2 with digitiform dactylus tip
C. halei sp. nov.
— Pereopod 2 with straight dactylus tip
C. poorei sp. nov.
29. Carapace with transversal dorsal carinae 30
— Carapace without transversal dorsal carinae 31
30. Carapace with 1 transversal dorsal carina
C. johnstoni Hale. 1937
— Carapace with 2 transversal dorsal carinae
C. nowrae sp. nov
31. Carapace with dorsal tubercles C. angelae sp. nov.
— Carapace without tubercles 32
32. Pseudorostrum long, lobes meeting for a distance of sixth
carapace length C. pileus (Foxon, 1932)
— Pseudorostrum short, lobes meeting for a distance shorter
than eighth carapace length 33
33. Carapace with 2 lateral ridges delimiting 1 depression
C. edenensis sp. nov.
— Carapace with 4 lateral ridges delimiting 3 depressions
C. trisulcata sp. nov.
34. Carapace integument smooth C. setifera sp. nov.
— Carapace with long club-like setae, body and appendices
highly serrated C. serrata sp. nov.
Campylaspis anae sp. nov.
Figures 1, 2
Material examined. Holotype female, NSW, off Nowra, 34°59.52'S,
151°05.94'E, 204 m, coarse shell, WHOI epibenthic sled, G.C.B.
Poore et al„ RV Franklin , 14 Jul 1986 (stn SLOPE 1), NMV J52946.
Diagnosis. Carapace with long lateral sulcus, rectangular in
dorsal view. Maxilliped 3 with large merus and carpus. Uropod
peduncle more than 2.5 times as long as pleonite 6; exopod a
little shorter than endopod.
Description. Body with smooth integument. Length: 3.5 mm.
Antenna 1, basal article of peduncle longest, median and
apical articles equal; main flagellum 3 -articulate, accessory
flagellum minute, uniarticulate.
Figure 1. Campylaspis anae sp. nov. female holotype: a, body, lateral
view; b, body, dorsal view; c, antenna 1; d, maxilliped 2; e, maxilliped
3. Scale (in mm): a, b, 1; c, e, 0.25; d, 01.
Figure 2. Campylaspis anae sp. nov. female holotype: a, pereopod 1;
b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod 5; f, pleonite
6 and left uropod. Scale (in mm): a-f, 0.25.
132
lorgu Petrescu
Maxilliped 2 basis (fused with ischium) and merus with
long plumose seta on distal inner corner, carpus produced
bluntly between 2 simple setae on inner margin, propodus
longer than carpus, as long as its distal outer strong seta,
dactylus with 3 teeth shorter than propodal seta. Maxilliped 3
basis less than half as long as appendage, with serrated inner
margin, with plumose seta on inner and outer distal corners,
without outer process; ischium small; merus much bigger
(second longest article), with serrated margins and plumose
seta on outer distal corner; carpus as long as its distal outer
strong seta, dactylus with 3 teeth shorter than propodal seta.
Maxilliped 3 basis shorter than half length of appendage, with
serrated inner margin, with plumose seta on inner and outer
distal corners, without outer process; ischium small; merus
larger (second longest article), with serrated margins and
plumose seta on outer distal corner; carpus 0.65 of merus
length, large, with strong serration on outer margin, 1 plumose
seta on outer distal corner; propodus little shorter than carpus,
much thinner; dactylus less than half of propodus, with 2
microserrate terminal setae twice dactylus length.
Pereopod 1 basis less than half as long as appendage, with
serrated margins; merus 2.6 times as long as ischium; carpus
to dactylus progressively shorter, with numerous simple setae
on both margins, dactylus with long terminal simple setae.
Pereopod 2 basis less than half as long as appendage, with a
plumose seta on inner distal corner; merus 4 times as long as
ischium, with long plumose seta on inner margin; carpus 1.4
times as long as ischium and merus combined, 2 simple setae
on its inner distal corner; dactylus with broken tip, 4 times as
long as propodus, with several simple setae marginally.
Pereopod 3 basis longer than rest of appendage, ischium as
long as merus, ischium and merus with simple seta on inner
distal corner; carpus twice as long as merus; dactylus with
long stout terminal seta. Pereopods 4 and 5 with shorter basis
and longer carpus than pereopod 3. Exopods on maxilliped 3
and pereopods 1, 2.
Uropod peduncle more than 2.5 times as long as pleonite
6, with strong serrated inner margin and median crest, twice
as long as exopod; exopod shorter than endopod, with
subterminal and terminal (much longer) stout setae; endopod
with median crest, with 4 microserrate setae on inner margin
and a long stout terminal seta.
Etymology. The species is dedicated to my lovely daughter,
Ana-Maria.
Distribution. Off Nowra, NSW; 204 m depth.
Remarks. Campylaspis anae has a large lateral sulcus that does
not reach the posterior extremity of the carapace. No other
species previously known from Australia or from the western
Pacific has such a sulcus. The shape of the carapace is similar
to that of C. ovalis Stebbing, 1912 from South Africa which has
an even larger sulcus that is confluent with the posterior
extremity of the carapace. Of the 16 new species with a lateral
sulcus described herein, C. anae is closest to the immature
male holotype of C. rectangulata. It differs from C. rectangulata
in possessing longer teeth on maxilliped 2, the merus of
maxilliped 3 without a concavity and larger, the dactylus of
maxilliped 3 with shorter setae, the uropod with a longer and
wider peduncle with a longitudinal median crest, shorter rami,
and an endopod with four microserrate setae on its inner margin
(3 similar setae in C. rectangulata ).
Campylaspis angelae sp. nov.
Figures 3, 4
Material examined. Holotype female, NSW, off Nowra, 34°59.52'S,
151°05.94'E, 204 m, coarse shell, WHOI epibenthic sled, G.C.B. Poore
et al„ RV Franklin , 14 Jul 1986 (stn SLOPE 1), NMV J52960.
Paratype: 1 female, dissected, type locality, NMV J52961.
Diagnosis. Carapace with a lateral sulcus, 2 dorsal parallel rows
of tubercles on carapace, without eyes. Lateral keel on pleonites
1-5 and 2 parallel dorsal keels on pereonite 5 and pleonites 1^1,
unique on last 2 pleonites. Antenna 1 short, robust. Maxilliped
3 with enlarged merus to propodus. Pereopod 1 with long
ischium. Dactylus of pereopod 2 with finger-like tip. Serrated
uropod with a median keel on peduncle.
Description. Body with smooth, highly calcified integument.
Length: 2.55 mm.
Carapace almost half of body length, 1.8 times as long as
high, with a large lateral sulcus delimited by a transverse crest
on anterior part, meeting ventral undulated crest that unites
anteroventral corner and oblique posterior crest on each side
of carapace; 2 dorsal parallel rows of small tubercles, 1
tubercle on top of ocular lobe, another on pseudorostrum;
antennal notch marked, anteroventral corner serrated, ventral
margin smooth; prominent ocular lobe without eyes. Parallel
dorsal crests on pereonite 6, pleonite 1-4, and single dorsal
serrated crest on pleonites 5-6. Pleonites 1-4 produced
laterally, pleonite 5 weakly produced laterally.
Antenna 1 short, robust, articles of peduncle decreasing in
length distally, main flagellum 3-articulate ending with short
aesthetascs; accessory flagellum tiny, uniarticulate.
Maxilliped 2 basis fused with ischium, with stout inner
plumose seta, similar seta on merus, carpus with tooth and 2
simple setae on inner margin, propodus with strong robust seta
on outer distal corner near articulation with short 3 -toothed
dactylus. Maxilliped 3 basis less than half as long as appendage,
without outer process, with 2 long plumose setae on outer
distal corner, ischium to propodus with serrated inner margins,
merus second longest article, with 3 teeth and plumose seta on
outer margin, carpus 0.66 merus length, with 3 long teeth and
long plumose seta on outer margin, propodus as long as carpus,
dactylus less than half as long as propodus, with terminal setae
longer than dactylus.
Pereopod 1 basis longer than rest of articles combined,
basis to propodus with serrated margins, short ischium with
inner plumose seta, merus, carpus and propodus enlarged,
merus second largest article, longer than carpus and propodus
combined, dactylus and propodus subequal. Margins of
merus-propodus serrated on both margins. Pereopod 2 basis
less than half as long as appendage, with short simple seta on
distal inner corner, merus 5 times as long as ischium, with
simple seta on inner margin, carpus 1.66 times as long as
merus, with short simple setae on distal inner corner, dactylus
New nannastacid cumaceans from Museum Victoria
133
Figure 3. Campylaspis angelae sp. nov. female holotype: a, body,
lateral view; b, body, dorsal view; c, antenna 1; d, maxilliped 2; e,
maxilliped 3. Scale (in mm): a, b, 0.5; c, d, 0.1; e, 0.2.
Figure 4. Campylaspis angelae sp. nov. female holotype :
A, pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4;
E, pereopod 5; f, pleonite 6 and left uropod. Scale (in mm): a, 0.2;
B-f, 0.25.
3 times as long as propodus, bluntly produced terminally, with
2 subterminal short simple setae, no evidence of terminal
setae. Pereopod 3 basis longer than rest of articles combined,
carpus little longer than merus, with only hair-like distal seta,
dactylus fused with its terminal short stout seta. Pereopods 4
and 5 with progressively shorter basis than in pereopod 3,
carpus twice as long as merus. Exopods on maxilliped 3 and
pereopods 1, 2.
Uropod with robust peduncle, 1.6 times as long as pleonite
6, with median serrated crest and serrated margins, 1.3 times
as long as endopod; exopod little shorter than endopod, with
strong serrated margins, 2 terminal robust setae, 1 long,
endopod 1.15 times as long as exopod, with serrated margins
and serrated median crest, 3 stout setae on inner margin and
terminal robust microserrate seta.
Etymology. This species is dedicated to my wonderful, lovely
and highly devoted wife, Angela.
Distribution. NSW, off Nowra, shelf break, 204 m.
Remarks. This new species has a carapace with a lateral sulcus
posteriorly bordered by an oblique crest as in C. pileus Foxon,
1932. It differs mainly by a shorter pseudorostrum, small
tubercles on the carapace, and shorter and more robust uropods
with serrated margins. C. angelae is also more similar than other
species of the genus to C. calmani Petrescu, 1995 from Indonesia,
which also has a lateral sulcus with a transverse crest on the
anterior part (close to the ocular lobe) and small tubercles.
However, tubercles just border the sulcus in C. calmani and are
otherwise disposed in C. angelae, the dactylus of pereopod 2 is
with a digitiform extremity only in C. angelae, and there is a
longitudinal serrated crest on the uropodal peduncle in C.
angelae, missing in C. calmani. The new species is also related
to C. edenensis sp. nov., which has an almost similar lateral
sulcus but without tubercles, similar to pereopod 2 but with
longer dactylar teeth than maxilliped 2, and uropodal peduncle
without a longitudinal crest.
Campylaspis aspera Hale, 1945
Campylaspis aspera Hale, 1945: 209, figs 45, 46.
Material examined. 8 females, 1 male (stn SLOPE 1), NMV J52964;
2 females, 1 male (stn SLOPE 1), MGAB CUM 1613; 11 females, 7
males (stn SLOPE 45), NMV J52965; 2 females, 1 male (stn SLOPE
46), NMV J52966; 1 female (stn MSL-EG 20), NMV J52967.
Distribution. NSW, 11m depth.
Remarks. The species was previously recorded by Hale (1945)
from Eden, New South Wales, at 70 m depth. The geographic
and depth ranges are considerably extended to South Australia,
Victoria and Tasmania and from 800 m depth.
Campylaspis australiensis sp. nov.
Figures 5, 6
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°00.20'S, 148°37.70'E, 720 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn SLOPE 46),
NMV J52415.
134
lorgu Petrescu
Figure 5. Campylaspis australiensis sp. nov. female holotype:
a, body, lateral view; b, body, dorsal view; c, antenna 1;
d, maxilliped 2; e, maxilliped 3; f, pereopod 1. Scale (in mm): a, b,
1; c, e, f, 0.25; d, 0.2.
Figure 6. Campylaspis australiensis sp. nov. female holotype: a,
pereopod 2; b, pereopod 3; c, pereopod 4; d, pereopod 5; e, pleonite
6 and left uropod; f, pleonite 6 and right uropod of other specimen.
Scale (in mm): a-f, 0.5.
Paratypes: 5 females, 11 males, type locality, NMV J52933; 1
female, 2 males (stn SLOPE 46), MGAB CUM 1606; 1 manca, 1
female (stn SLOPE 47), NMV J52934; 2 males, 1 manca (stn
SLOPE 48), NMV J52419; 1 manca (stn SLOPE 1), NMV J52413;
1 female, 1 male (stn SLOPE 45), NMV J52414; 1 immature male,
dissected, NMV J52416.
Diagnosis. Carapace with acute tubercles, with a lateral sulcus.
A pair of small dorsal tubercles on pereonite 5 and all pleonites,
small lateral tubercle on pereonites 3-5 and pleonites 1-5.
Antenna 1 accessory flagellum of 2 articles. Propodus of
maxilliped 2 with long robust distal seta, longer than dactylar
teeth. Merus and carpus of maxilliped 3 with teeth on outer
margin. Dactylus of pereopod 2 3.7 times as long as propodus,
with tapering extremity. Dactylus of pereopods 3-5 not fused
with terminal seta. Uropod peduncle 1.8 times as long as
pleonite 6, 1.4 times as long as endopod, with strongly serrated
margins and longitudinal serrated crest, endopod slightly longer
than exopod, with 3 microserrate setae on inner margin.
Description. Body with well calcified and hardened reticulated
integument. Length: 3.06 mm.
Carapace 0.5 body length, 1.7 times as long as high, with a
lateral sulcus, 0.5 of carapace length, open towards anterior
part, bordered with acute tubercles, bigger dorsal tubercles,
parallel row of smaller dorsal ones, 2 small tubercles on top of
eyeless ocular lobe, antennal notch marked, pseudorostral
lobes meeting a short distance in front of ocular lobe.
Pereonites 3-5 with lateral small tubercle. Pleonites 1-5 with
pair of dorsal small tubercles, pleonite 5 with constriction at
about posterior one-third, armed with 2 or 3 small dorsal
tubercles, lateral tubercle on pleonites 1-5, pleonite 6 with 1 or
2 pairs of small dorsal tubercles.
Antenna 1 peduncle with robust articles, with serrated
margins, progressively shorter, main flagellum 3 -articulate,
longer than distal article of peduncle, accessory flagellum 2-
articulate, shorter than basal article of main flagellum.
Maxilliped 2 with long plumose setae on inner margin of
basis and merus, tooth and plumose seta on inner margin of
carpus, propodus second longest article, with long robust distal
seta, longer than dactylar teeth, with tooth on inner margin,
dactylus with 3 short, robust teeth. Maxilliped 3 basis longer than
rest of articles combined, with 2 long plumose setae on outer
margin, merus second longest article, with 2 teeth and plumose
seta on outer margin, carpus shorter than merus, with teeth on
both margins and plumose seta on outer margin, propodus shorter
than carpus, with 2 pappose setae on inner margin, dactylus half
of propodus length, with short simple setae.
Pereopod 1 with slender articles, basis shorter than rest of
articles combined, with plumose seta on inner distal corner,
merus second longest article, 2.5 times as long as ischium,
with small tooth and plumose seta on outer margin, carpus
shorter than merus, propodus shorter than carpus, dactylus 0.6
propodus length, with short simple setae. Pereopod 2 basis
less than one-third of entire pereopod length, with tooth and
plumose seta on inner margin, merus with tooth and 2 simple
setae on inner margin, carpus 1.6 times as long as merus, with
stout and simple seta on distal margin, dactylus 3.7 times as
long as propodus, with tapering tip and short setae. Pereopods
New nannastacid cumaceans from Museum Victoria
135
3-5 with progressively shorter basis and longer carpus,
propodus longer than merus, dactylus with long stout terminal
seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 1.8 times as long as pleonite 6, 1.4 times
as long as endopod, with strongly serrated margins and
longitudinal serrated crest, exopod shorter than endopod, with
subterminal stout seta and a terminal longer one, endopod with
3 microserrate setae on inner margin and more robust, longer
terminal seta.
Etymology. The species bears the name of type locality -
Australia.
Distribution. Tas., off Freycinet Peninsula, 600-720 m.
Remarks. The closest species to Campylaspis australiensis is
C. sagamiensis Gamo, 1967 from Japan. Both species have
similar tubercles on the carapace, pereon and pleon, and a
lateral sulcus, but in C. sagamiensis the pleon has lateral ridges
rather than tubercles; C. sagamiensis also differs in a longer
propodus seta, shorter and more robust teeth on the dactylus of
maxilliped 2, wider merus and dactylus with shorter terminal
setae on maxilliped 3, and shorter more robust uropods.
Similar carapace tubercle patterns around the lateral sulcus
also exist on C. clavata Lomakina, 1952 from the north-western
Pacific, C. horrida Sars, 1870 from the Atlantic Ocean, C.
horridoides Stephensen, 1915 from the Mediterranean and C.
antarctica Caiman, 1907 from Antarctica. C. clavata differs in
having a longer ocular lobe (reaching the extremity of the
carapace versus shorter in C. australiensis), shorter dactylus of
pereopod 2 and longer uropod peduncle. C. horrida differs in
having a sulcus not surrounded by acute tubercles, pleonite 6
without tubercles and a longer uropod peduncle. C. horridoides
differs in fewer dorsal tubercles on the carapace, maxilliped 3
with more teeth on its outer margin and a longer uropod
peduncle. C. antarctica differs in being without a lateral sulcus
on the carapace and its pleonite 6 being without spines.
Campylaspis echinata Hale, 1945
Campylaspis echinata Hale, 1945: 204, figs 41, 42.
Material examined. 1 female (stn SLOPE 1), NMV J52969; 1 immature
male (stn SLOPE 47), NMV J52970.
Distribution. NSW, 204 m depth.
Remarks. Hale, 1945 recorded the species from Eden, New
South Wales, at 70 m depth; these records represent new
geographic and depth extensions (Tasmania, from 600 depth).
Campylaspis edenensis sp. nov.
Figures 7-9
Material examined. Holotype female, NSW, off Eden, 37°07.30'S,
150°20.20'E, 520 m, grey coarse shell, WHOI epibenthic sled, G.C.B.
Poore et al„ RV Franklin, 20 Jul 1986 (stn SLOPE 19), NMV J52956.
Allotype: immature male, type locality, NMV J52957.
Paratypes: 3 females (stn SLOPE 19), NMV J52958; 2 females
(stn SLOPE 19), MGAB CUM 1612; 1 female, dissected (stn SLOPE
19), NMV J52959.
Figure 7. Campylaspis edenensis sp. nov. female paratype: a, body,
lateral view; b, body, dorsal view; c, antenna 1; d, maxilliped 2; e,
maxilliped 3. Scale (in mm): a, b, 0.5; c, d, 0.2; e, 0.25.
Figure 8. Campylaspis edenensis sp. nov. female paratype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and left uropod. Scale (in mm): a-f, 0.5.
136
lorgu Petrescu
Figure 9. Campylaspis edenensis sp. nov. male allotype: a, body,
lateral view; b, carapace, dorsal view; c, pleonite 6 and left uropod.
Scale (in mm): a, b, 1; c, 0.25.
Diagnosis. Carapace with large, long lateral sulcus not reaching
dorsal crest. Median dactylar tooth of maxilliped 2 longer than
other two. Maxilliped 3 basis more than one -third of entire
appendage. Pereopod 1 with numerous plumose setae on both
sides. Pereopod 2 with digitiform tip of dactylus, with long
simple subterminal seta. Pereopods 3-5 robust, dactylus fused
with terminal seta. Uropod peduncle 1.7 times as long as
pleonite 6, subequal rami.
Description of female. Body with well calcified glabrous
integument. Length: 3.3 mm.
Carapace half body length, 2.2 times as long as high, large
and long lateral sulcus not reaching dorsal crest, upper margin
ending on tip of pseudorostrum, transverse ridge on sulcus at
base of ocular lobe, large ocular lobe without eyes, numerous
dorsal clear lenses, antennal notch small, ventral margin
smooth, dorsal crest posteriorly. Pereon with 4 free thoracic
segments. Pleon 0.38 body length.
Antenna 1 peduncle with serrated margins, proximal
article longest, main flagellum 3 -articulate, longer than distal
article of peduncle, accessory flagellum tiny, uniarticulate.
Maxilliped 2 basis fused with ischium, with plumose seta
on inner distal corner, similar seta medially disposed on
merus, carpus with short tooth and 2 simple setae on inner
margin, propodus second longest article, with small tooth on
inner margin and outer stronger seta, longer than dactylar
teeth, dactylus fused with its 3 teeth, median one the longest.
Maxilliped 3 basis more than one -third of entire maxilliped,
with plumose seta on inner distal corner and 2 longer setae on
outer distal corner, merus second longest article, with serrated
margins, stronger on outer margin, with plumose seta on outer
distal corner, carpus as long as propodus, with 4 teeth on inner
margin and 2 on outer one, propodus with 2 teeth and 4
pappose setae on inner margin, 3 teeth on outer margin,
dactylus 0.35 of propodus, with simple short setae.
Pereopod 1 basis more than one -third of entire appendage,
with serrated inner margin, plumose setae on basis to propodus,
merus twice as long as ischium, carpus as long as propodus,
with strong inner serration, dactylus 0.6 of propodus length,
with simple short setae. Pereopod 2 basis little less than one-
third of appendage, basis and merus with plumose seta on
inner distal corner, merus 5 times as long as ischium, carpus
1.3 times as long as merus, with plumose seta and simple seta
on distal inner corner, dactylus 3.5 times as long as propodus,
with digitiform tip, few simple short setae on both margins, 3
subterminal simple setae, 1 much longer. Pereopods 3-5 with
progressively shorter basis and longer carpus, carpus twice as
long as merus in last pair, with simple robust seta on distal
outer corner, dactylus fused with its terminal robust seta.
Exopods on maxilliped 3 and pereopods 1, 2.
Uropod with robust peduncle 1.7 times as long as pleonite
6, twice as long as rami, with serrated inner margin, exopod
slightly shorter than endopod, with terminal longer robust
seta, endopod with 2 stout setae on serrated inner margin,
robust terminal seta shorter than endopod; terminal setae on
rami with single subterminal setule.
Description of male. Body length: 3.5 mm.
Carapace with stronger ridges than in female.
Uropod peduncle 1.5 times as long as pleonite 6, twice as
long as endopod, exopod little shorter than endopod, with 2
subterminal setae and a longer, more robust, terminal one,
endopod larger than exopod, with terminal robust seta and 5
stout short setae on inner margin.
Etymology. The new species bears the name of Eden, a port
town in south-eastern Australia.
Distribution. Off Eden, NSW; 520 m depth.
Remarks. The two species most similar to Campylaspis
edenensis are C. calmani Petrescu, 1995 and C. angelae.
Campylaspis edenensis is differentiated from C. calmani by
having the sulcus not bordered by small tubercles and the
dactylus of pereopod 2 with a digitiform tip. Campylaspis
edenensis differs from C. angelae by a lack of carapace
tubercles, dorsal serrated crests on the pereon and pleon, and
the longitudinal crest on the uropod peduncle.
Campylaspis grossui sp. nov.
Figures 10, 11
Material examined. Holotype female. Vic., eastern Bass Strait, 11.2
km E of eastern edge of Lake Tyers, 37°51.41'S, 148°13.16'E, 32 m,
sand-shell, Smith-Mclntyre grab. Marine Science Laboratories, RV
Sarda, 25 Sep 1990 (stn MSL-EG 27), NMV J27468.
Paratypes: 1 female (stn MSL-EG 27), NMV J27468; 1 female,
dissected (stn MSL-EG 27), NMV J52952; 1 female, 1 male (stn MSL-
EG 26), NMV J27467; 1 female (stn MSL-EG 26), MGAB CUM
1607; 1 immature male, NMV J 27469; 1 female, NMV J27470.
New nannastacid cumaceans from Museum Victoria
137
Figure 10. Campylaspis grossui sp. nov. female paratype: a, body,
lateral view; b, body, dorsal view; c, antenna 1; d, maxilliped 2; e,
maxilliped 3. Scale (in mm): a, b, 1; c, d, 0.3; e, 0.5.
Figure 11. Campylaspis grossui sp. nov. female paratype: a, pereopod
1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod 5; f,
pleonite 6 and left uropod. Scale (in mm): a-e, 0.5; f, 0.25.
Diagnosis. Carapace with large lateral sulcus (almost as long
as carapace), not meeting on posterior end, with dorsal
tubercles. Pleon with short segments. Maxilliped 2 with long
dactylar teeth. Pereopod 1 with large merus with an inner
concavity. Pereopod 2 with large articles, dactylus wide, with
short stout terminal seta. Pereopods 3-5 robust, dactylus fused
with short robust, terminal and curved seta. Uropod peduncle
1.4 times as long as pleonite 6, 1.5 times as long as its equal
rami, exopod and endopod with very short robust terminal seta,
2 short robust setae on inner margin of endopod.
Description. Body with highly calcified glabrous integument.
Length: 3.95 mm.
Carapace 0.5 body length, twice as long as high, with long
lateral sulcus, opened towards anterior, not meeting at
posterior end, large and small lateral and dorsal tubercles,
antennal notch wide, pseudorostrum meeting in front of
eyeless ocular lobe. Pereonites 1-4 covered almost completely
by carapace. Pleon with anteroposterior compressed segments,
shorter than carapace.
Antenna 1 short and robust, median article of peduncle
longest, main flagellum as long as distal article of peduncle,
3 -articulate, accessory flagellum tiny, uniarticulate.
Maxilliped 2, long plumose seta on inner margin of basis
and on distal article of merus, carpus with large tooth on inner
margin, propodus second longest article, a long robust distal
seta as long as dactylar teeth, dactylus with 3 teeth, 2 of them
longer than propodus, median one-half of other two. Maxilliped
3 with wide basis, shorter than rest of articles combined, with
6 long plumose setae on inner margin and 1 on outer margin,
ischium to dactylus slender, merus second longest article, with
plumose seta on outer distal margin, carpus with plumose seta
midway on outer margin, propodus 0.57 of carpus, dactylus
short, slender, half of propodus, with short simple terminal
setae.
Pereopod 1 basis less than half as long as appendage, with
2 plumose setae on inner margin, wide merus with inner
concavity with numerous setae, carpus more slender than
merus, second longest article, propodus shorter than merus,
dactylus 0.5 of propodus, with short simple terminal setae.
Pereopod 2 with short, very robust articles, basis less than half
as long as appendage, long plumose seta on inner margin of
basis, ischium and merus, carpus 1.2 times as long as merus,
with 2 simple setae on distal inner corner, dactylus broad,
twice propodus length, with short stout hooked terminal seta.
Pereopods 3-5 robust, with progressively shorter basis, and
longer carpus, dactylus fused with its robust terminal short
seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 1.4 times as long as pleonite 6, 1.5 times
as long as its equal rami, exopod and endopod with short
robust terminal seta, 2 short robust setae on inner margin of
endopod.
Etymology. The species is dedicated to the memory of the late
Prof. Dr Alexandra V. Grossu (1910-2004) as a sign of gratitude
for his moral and professional qualities as a brilliant scientist
and mentor. He was the most important Romanian specialist in
malacology, highly appreciated throughout the world, former
director of “Grigore Antipa” National Museum of Natural
138
lorgu Petrescu
History (1955-1957), founder of its scientific journal, professor
of zoology and Dean of the Faculty of Biology in Bucharest
(1957-1962).
Distribution. Eastern Vic.; 32 m depth.
Remarks. Campylaspis grossui is closely related to C. wardi
Bacescu, 1991 and C. sculpta. All three have a similar carapace,
and short and robust pereopods and uropods. Campylaspis
grossui differs from C. wardi in several characteristics: fewer
tubercles on the carapace; longer teeth on the dactylus of
maxilliped 2; more slender carpus of maxilliped 3; uropods
with smooth margins; two short stout setae on the inner margin
of endopod versus none; and terminal short setae without the
oval structures characteristic of C. wardi. Campylaspis grossui
differs from C. sculpta in the shape of the carapace in dorsal
view, more robust antenna 1, maxilliped 3 and pereopod 1 with
more slender articles, pereopod 2 with a much shorter dactylus
(twice as long as the propodus in C. grossui versus four times in
C. sculpta ) and without a digitiform extremity, and more slender
uropod rami with longer setae.
Campylaspis halei sp. nov.
Figures 12, 13
Material examined. Holotype female, Tas., off Freycinet Peninsula,
41°57.50'S, 148°37.90'E, 400 m, coarse shell, WHOI epibenthic sled,
M.F. Gomon et al., RV Franklin , 27 July 1986 (stn SLOPE 48),
NMV J52940.
Allotype male: (stn SLOPE 48), NMV J52941.
Paratypes: 2 females, 1 immature male, 2 mancas (stn SLOPE 1),
NMV J52943 ; 2 females (stn SLOPE 47), NMV J52416; 2 females
(stn SLOPE 47), MGAB CUM 1608; 3 females, 1 female, dissected
(stn SLOPE 48), NMV J52942; 1 female (stn SLOPE 19), NMV
J52420.
Diagnosis. Carapace with a small lateral sulcus. Accessory
flagellum of antenna 1 2-articulate. Maxilliped 2 with subequal
dactylar teeth. Merus of maxilliped 3 massive, merus, carpus
and propodus with teeth on inner margin. Dactylus of pereopod
2 2.8 times as long as propodus, with a digitiform extremity.
Uropod peduncle twice as long as pleonite 6 and its endopod,
exopod 0.8 length of endopod, endopod with 5 microserrate
setae on inner margin.
Description of female. Body with calcified smooth integument.
Length: 3.2 mm.
Carapace 0.45 length of entire body, with a small lateral
sulcus, large ocular lobe without lenses, antennal notch
distinct, anteroventral corner acute, ventral margin smooth.
Pleon 0.36 body length.
Antenna 1 peduncle with serrated margins, distal article
longer than median one, main flagellum 3 -articulate, longer
than last article of peduncle, basal article longer than rest of
articles combined, accessory flagellum tiny, 2-articulate.
Maxilliped 2 basis fused with ischium, with plumose seta
on inner distal corner, merus with similar seta medially, carpus
with tooth and 2 long simple setae on inner margin, propodus
second longest article, with robust outer seta longer than
dactylar teeth, dactylus with 3 subequal teeth.
Figure 12. Campylaspis halei sp. nov. female paratype: a, body,
lateral view; b, carapace, dorsal view; c, antenna 1; d, maxilliped 2;
e, maxilliped 3; f, pereopod 1. Scale (in mm): a, b, 0.5; c, 0.1; d, 0.1;
e, f, 0.25.
Figure 13. Campylaspis halei sp. nov. female paratype: a-e, male
f-h: a, pereopod 2; b, pereopod 3; c, pereopod 4; d, pereopod 5; e,
pleonite 6 and left uropod; f, body, lateral view; g, carapace, dorsal
view; i, pleonite 6 and right uropod. Scale (in mm): a, e, h, 0.25;
b-d, 0.5; f, g, 0.5.
New nannastacid cumaceans from Museum Victoria
139
Maxilliped 3 basis less than half as long as appendage,
with plumose seta on inner distal corner and longer 1 on outer
corner, inner distal corner of ischium with robust seta, merus
second longest article, longer than carpus, propodus and
dactylus combined, with row of teeth on inner margin and
plumose seta on outer distal corner, carpus shorter than
propodus, with serrated inner margin and plumose seta on
outer distal corner, propodus with teeth on both margins,
dactylus shorter than propodus, with terminal simple setae.
Pereopod 1 basis shorter than one -third of appendage
length, with 2 plumose setae on inner margin, numerous
plumose setae on inner margins of ischium to propodus, merus
second longest article, carpus longer than propodus, propodus
longer than dactylus, dactylus with simple setae. Pereopod 2
basis more than one-third length of appendage, plumose seta
on inner distal corner of basis, ischium and merus, carpus
twice as long as merus, with short stout seta, simple and
plumose setae on inner distal corner, dactylus 2.8 times as
long as propodus, digitiform dactylus tip with subterminal
short simple seta. Pereopods 3-5 with progressively shorter
basis and longer carpus, carpus and propodus with simple
stout seta on outer distal corner, dactylus with stout terminal
seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod with serrated margins, peduncle twice pleonite 6,
exopod shorter than endopod, with terminal robust seta shorter
than exopod, endopod half peduncle length, with robust
terminal seta and 5 microserrate setae on inner margin.
Description of immature male. Body length: 2.8 mm. Carapace
less vaulted than in female, with similar sulcus. Antenna 2,
pereopods 1-4 and exopods characteristic of an immature
stage.
Uropod peduncle with less serrated margins, more slender
and proportionally longer than in females, 2.3 times pleonite
6, rami longer than in females, endopod 0.77 peduncle length,
rami equal, exopod with smooth margins, endopod with 5
stout setae on inner serrated margin.
Etymology. The species is dedicated to the memory of Herbert
Mathew Hale (1895-1963), former director of the South
Australian Museum (1928-1960), one of the most important
specialists of Cumacea. He described most of the cumacean
taxa known from Australian waters.
Distribution. NSW, off Freycinet Peninsula, Tas.; 400-600 m
depth.
Remarks. Campylaspis halei has a smooth carapace surface
with a relatively small lateral sulcus, extending over about half
of the carapace. It resembles C. latidactyla Hale, 1945 but with
a different extremity of the dactylus of pereopod 2 (digitiform
tip in C. halei). Its sulcus is smaller than in C. minor Hale,
1945, where it is largely opened towards the anterior margin,
and has a marked antennal notch absent in Hale’s species
(which has a much smaller size). C. halei is also similar to C.
latimera, a species with a characteristic large merus on
pereopod 1 and to C. poorei which has a narrower sulcus,
slender pereopods 1 and 2 and a straight tip on the dactylus of
pereopod 2 (digitiform in C. halei).
Campylaspis hirsuta sp. nov.
Figures 14, 15
Material examined. Holotype male, Vic., south of Point Hicks,
38°25'S, 149°00'E, 1500 m, compacted clay, WHOI epibenthic sled,
G.C.B. Poore et al., RV Franklin , 22 Jul 1986 (stn SLOPE 27),
NMV J52411.
Paratype: 1 immature male (stn SLOPE 46), NMV J52951.
Diagnosis. Carapace with 2 rows of dorsal tubercles provided
with strong spines, spines on top of pseudorostrum, external
row of tubercles with stronger spines, large lateral sulcus.
Dorsal spines on pereon and pleon. Maxilliped 2 with dactylar
teeth. Maxilliped 3 with long terminal seta on dactylus.
Dactylus of pereopod 2 as long as propodus and carpus
combined. Uropod peduncle 1.7 times as long as pleonite 6, 1.6
times as long as endopod.
Description. Body with integument strongly calcified. Length:
4.7 mm.
Carapace 0.4 body length, 1.8 times as long as high, with
large lateral sulcus, pair of tubercles with strong spines at the
posterior corners of the frontal lobe, 3 pairs in the median part
and pair of smaller ones on posterior extremity, exterior row of
3 strong tubercles with spines on each side, ocular lobe acute,
without eyes, pseudorostral lobes large and rounded, little
upturned, with spines and long simple setae at the tip, antennal
notch marked, anteroventral corner acute, ventral serration
strong, long simple dorsal setae.
Antenna 1, median and distal article of peduncle subequal,
shorter than main flagellum 3 -articulate, accessory flagellum
tiny, uniarticulate.
Maxilliped 2 with strong plumose seta on distal inner
corner of basis and merus, ischium not visible, 2 simple setae
on inner margin of carpus, propodus second longest article,
with robust seta on distal outer corner, longer than dactylar
teeth, dactylus with 3 teeth, median one shorter. Maxilliped 3
basis half entire appendage, with 2 short plumose setae and
short distal serration on inner margin, 2 long plumose setae on
outer distal corner, merus second longest article, with serrated
inner margin, with tooth and plumose seta on outer margin,
short carpus with serrated outer margin, propodus 1.4 times as
long as carpus and twice as long as dactylus, dactylus with
terminal simple long seta, 3 times dactylus length.
Pereopod 1 basis little less than half as long as appendage,
with 2 long plumose setae on inner margin, serration on both
distal sides, ischium with serrated inner margin with long
plumose seta, merus second longest article, with numerous
setae on both margins, carpus little shorter than merus, densely
setose, propodus shorter than carpus, with numerous setae,
twice as long as dactylus, dactylus with short simple setae.
Pereopod 2 basis less than one -third rest of articles combined,
with plumose seta on inner distal corner, simple seta on inner
margin of merus, carpus 1.9 times as long as merus, short
simple seta and longer plumose seta on its distal inner corner,
dactylus 3.5 times as long as propodus, as long as propodus
and carpus combined, with simple setae on both margins,
subterminal and terminal simple long setae. Pereopod 3 basis
large, plumose seta on distal inner corner of ischium and
140
lorgu Petrescu
Figure 14. Campylaspis hirsuta sp. nov. female holotype: a, body,
lateral view; b, carapace, dorsal view; c, antenna 1; d, maxilliped 2
Scale (in mm): a, b, 0.5; c, 0.25; d, 0.3.
Figure 15. Campylaspis hirsuta sp. nov. female holotype: a,
maxilliped 3; b, pereopod 1; c, pereopod 2; d, pereopod 3; e,
pereopod 4; f, pereopod 5; g, pleonite 6 and left uropod. Scale (in
mm): a-f, 0.5; g, 0.25.
merus, carpus little longer than merus, with 2 plumose setae
on outer margin and robust seta on outer distal corner, small
dactylus with stout terminal seta. Pereopod 4 similar to
pereopod 3, with smaller basis and longer carpus. Pereopod 5
with 1 plumose seta on outer margin of carpus. 5 pairs of
exopods incompletely developed.
Uropod peduncle 1.7 times as long as pleonite 6, 1.6 times
as long as endopod, 3 proximal teeth and 4 simple setae on
inner margin, exopod little shorter than endopod, with short
simple subterminal seta, another seta on inner margin and
robust microserrate terminal seta, endopod with 4 microserrate
setae on inner margin, subterminal simple seta and terminal
robust microserrate seta.
Etymology. The species is named “ hirsuta ” because of its
integument with numerous setae and spines.
Distribution. South of Point Hicks, Vic., off Freycinet
Peninsula, Tas.; 720-1500 m depth.
Remarks. Such a combination of characters, tubercles with
spines on the carapace, carapace with a lateral sulcus, plus
dorsal spines on the pereon and pleon as in Campylaspis
hirsuta, is unique among species of Campylaspis. C. spinosa
Caiman, 1906 from the Mediterranean and the Atlantic Ocean
has acute tubercles without spines on the carapace and dorsal
spines on the pereon and pleon. The uropod endopod of C.
hirsuta has four microserrate setae on the inner margin instead
of three as in C. spinosa
Campylaspis johnstoni Hale, 1937
Campylaspis johnstoni Hale, 1937: 37-56, fig. 2.
Material examined. 4 females, 2 males (stn SLOPE 1), NMV J52975;
1 female, 1 immature male (stn SLOPE 2), NMV J54127; 1 female (stn
SLOPE 2), NMV J52963; 1 male (stn SLOPE 46), NMV J52974.
Distribution. Antarctica, Kerguelen Is., south-western Atlantic
(Argentina), 204 m depth (Bacescu, 1992); NSW, Tas., 720 m
depth.
Remarks. This is the first record of this widespread species
from Australian waters and from such a depth (720 m).
Campylaspis latidactyla Hale, 1945
Campylaspis latidactyla Hale, 1945: 194, figs 33, 34.
Material examined. 1 female, 1 male (stn SLOPE 22), NMV J52971.
Distribution. Moreton Bay, Qld, NSW; 200 m depth.
Remarks. The species was first recorded by Hale, 1945 from
Moreton Bay, Queensland; its distribution is now extended to
the southern New South Wales shelf.
Campylaspis latimera sp. nov.
Figures 16-19
Material examined. Holotype subadult female, Tas., off Freycinet
Peninsula, 42°00.20'S, 148°37.70'E, 720 m, coarse shelly sand, WHOI
epibenthic sled, M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn
SLOPE 46), NMV J52935.
New nannastacid cumaceans from Museum Victoria
141
Figure 16. Campylaspis latimera sp. nov. female paratype: a, body,
lateral view; b, carapace, dorsal view; c, fifth pleonite; d, antenna 1;
e, maxilliped 3. Scale (in mm): a, b, 1; c, 0.25; d, 0.25; e, 0.5.
Figure 18. Campylaspis latimera sp. nov. male paratype: a, body,
lateral view; b, pleonite, magnified; c, antenna 1; d, its flagella,
magnified; e, maxilliped 3; f, pereopod 1. Scale (in mm): a, 1; b, c,
0.25; d, 0.1; e, f, 0.5.
Figure 17. Campylaspis latimera sp. nov. female paratype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and left uropod. Scale (in mm): a-f, 0.5.
Figure 19. Campylaspis latimera sp. nov. male paratype: a, pereopod
2; b, pereopod 3; c, pereopod 4; d, pereopod 5; e, pleonite 6 and left
uropod; f, detail of uropodal peduncle, magnified. Scale (in mm): a-e,
0.5; f, 0.25.
142
lorgu Petrescu
Allotype: male (stn SLOPE 45), NMV J52936.
Paratypes: 2 females, 1 manca (stn SLOPE 1), NMV J52937; 1
female (stn SLOPE 1) MGAB CUM 1609; 1 immature male (stn
SLOPE 46), NMV J52938; 1 immature male, NMV J45784; 1 female,
1 immature male (stn SLOPE 48), NMV J52939.
Diagnosis. Carapace with a large lateral sulcus, small ocular
lobe without eyes. Maxilliped 3 with large concave merus.
Pereopod 1 with large merus. Pereopod 2 with long dactylus
(3.8 times as long as propodus). Pereopods 3-5 dactylus fused
with its terminal seta. Uropod peduncle 3 times as long as
pleonite 6 and 2.8 times as long as endopod.
Description of female. Body with calcified, smooth integument.
Length: 6.8 mm.
Carapace 0.44 body length, 1.7 times as long as high, large
lateral sulcus occupying most of lateral side of carapace (0.7 of
its length), ocular lobe small, acute, without eyes, antennal
notch small, ventral margin smooth. Pleonites each with
toothed posterior margin.
Antenna 1 basal article of peduncle the longest, distal one
the shortest, main flagellum shorter than last article of
peduncle, 3 -articulate, accessory flagellum tiny, uniarticulate.
Maxilliped 3 basis more than half length of appendage,
serrated inner margin with 2 plumose apical setae, 2 longer
plumose setae on outer corner (not produced), ischium with
strong tooth on inner margin, merus large and concave,
produced with small tooth, serrated outer margin with plumose
seta, carpus shorter than merus, with serrated outer margin
and plumose seta, propodus shorter than carpus, with numerous
inner plumose setae, dactylus half of propodus, with setae as
long as article.
Pereopod 1 basis more than half length of appendage, serrated
inner margin with 2 simple setae and long pappose seta on inner
distal comer, 2 plumose setae on outer margin, ischium with
pappose seta on serrated inner margin, merus large, second longest
article, 3 simple setae on serrated inner margin and 2 plumose
setae and spine on outer margin, carpus large with serrated
margins with simple setae on inner margin and plumose on outer
margin, shorter than merus, propodus shorter and thinner than
carpus, with numerous plumose setae on outer margin, dactylus
0.5 of propodus, with short simple setae. Pereopod 2 basis less
than half length of appendage, simple seta on inner margin of
merus, 2 unequal simple setae on outer distal comer, dactylus 3.8
times as long as propodus, with plumose setae on inner margin
and simple seta on outer margin, subterminal and terminal simple
setae. Pereopods 3-5 with progressively shorter basis and longer
carpus (twice as long as merus), dactylus fused with terminal
stout seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 3 times as long as pleonite 6 and 2.8
times as long as endopod, serrated inner margin without setae,
exopod slightly shorter than endopod, with subterminal simple
stout setae, endopod with 4 microserrate setae on inner margin,
terminal robust seta shorter than endopod.
Description of male. Body with numerous long simple setae on
posterior margin of pereonites and of pleonites 1-4. Length:
6.5 mm. Carapace less vaulted than in female, lateral sulcus
smaller, anteroventral corner serrated. Pleonites with lateral
groove for protecting the flagellum of antenna, provided with
stout setae on its margins keeping flagellum within groove.
Antenna 1 shorter than in female, main flagellum of 3
articles, 1 aesthetasc on articles 2-3. Accessory flagellum tiny,
uniarticulate. Maxilliped 3 with similar merus but thinner
propodus than in female.
Pereopod 1 as in female. Pereopod 2 basis larger, plumose
seta on inner margin of merus, carpus with plumose seta on
inner distal corner, dactylus 3.5 times as long as propodus, as
in female. Pereopods 3 and 4 basis broad, longer carpus in
pereopod 4 (twice as long as merus), dactylus with stout
terminal seta. Pereopod 5 carpus second longest article, little
longer than in pereopod 4.
Uropod peduncle 3.35 times as long as pleonite 6, longer
than in female, 2.65 times as long as endopod, rami little longer
than in female, numerous plumose setae on serrated inner
margin and a median groove for antenna provided with a lateral
row of stout setae and a parallel one with plumose setae, exopod
0.8 length of endopod, with short subterminal stout seta, inner
plumose seta and terminal stout long seta (longer than exopod),
endopod with 9 microserrate long setae on inner margin and a
terminal robust seta shorter than endopod.
Etymology. The species is named “ latimera ” because of the
large merus of its maxilliped 3 and pereopod 1.
Distribution. Off Nowra, NSW, off Freycinet Peninsula, Tas.;
204-720 m depth.
Remarks. Campylaspis latimera differs from other species from
this area with a lateral sulcus on the carapace by the characteristic
shape and large size of the merus of the first pereopod.
Maxilliped 3 also has a large merus with a concave inner margin
and a large carpus. Both features are present in males and
females. C. anae also has a large merus and carpus of maxilliped
3 but without the concavity of the merus; pereopod 1 of this
species has slender articles. Also, the merus and carpus of
maxilliped 3 of C. rectangulata are large, the merus with an
inner concavity but not as evident as in C. latimera and pereopod
1 has only moderately large articles. Campylaspis microdentata
Ledoyer, 1988 from off Comore Is. has a similar merus to
maxilliped 3 but the carapace has no lateral sulcus and pereopod
1 has slender articles, shaped normally.
Campylaspis longidentata sp. nov.
Figures 20, 21
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°00.20'S, 148°37.70'E, 720 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn SLOPE 46),
NMV J52944.
Diagnosis. Carapace with an oblique, concave ridge from dorsal
side toward the lower margin, covered with pits and spines.
Maxilliped 2 with long terminal teeth, longer than usual,
propodus with long robust distal seta and dactylus also with
long teeth. Maxilliped 3 with serrated margins. Pereopod 1 with
serrated margins on basis to propodus, with slender articles.
Pereopod 2 with dactylus 5 times as long as propodus, and with
long digitiform extremity. Slender uropod peduncle 2.48 times
as long as pleonite 6 and 1.9 times as long as equal rami.
New nannastacid cumaceans from Museum Victoria
143
Figure 20. Campylaspis longidentata sp. nov. female holotype: a,
body, lateral view; b, detail of its integument; c, carapace, dorsal
view; d, antenna 1; e, maxilliped 2; f, maxilliped 3; g, pereopod 1.
Scale (in mm): a, c, 1; d, 0.25; e-g, 0.25.
Figure 21. Campylaspis longidentata sp. nov. female holotype: a,
pereopod 2; b, pereopod 3; c, pereopod 4; d, pereopod 5; e, pleonite
6 and right uropod. Scale (in mm): a-e, 0.5.
Description. Body elongated, with strongly calcified integument.
Length: 3.7 mm.
Carapace about half of body, longer than high, covered
with numerous pits bordered by little spines, antennal notch
obvious, anterolateral corner acute, lower margin with tooth
on anterior half, pseudorostral lobes meeting in front of
eyeless ocular lobe at a distance longer than length of frontal
lobe, concave ridge from dorsal side toward lower margin. A
tooth on lateral plate of pereonite 5, a pair of dorsal teeth on
pleonites 1-3.
Antenna 1 peduncle articles progressively shorter, basal
and median articles with tubercle on inner margin, main
flagellum 3 -articulate, longer than distal article of peduncle,
basal article of flagellum much more than half flagellum
length, accessory flagellum tiny, uniarticulate.
Maxilliped 2 basis with plumose seta on distal margin,
ischium fused to basis, carpus second longest article, with 4
teeth (2 stronger) and axe -like tooth on inner margin, propodus
with long robust distal seta, dactylus with 3 long teeth, median
one shorter, twice as long as propodus. Maxilliped 2 visible
between pereopod 1 and maxilliped 3 in lateral view.
Maxilliped 3 basis shorter than 0.5 of appendage, with serrated
margins, 2 plumose setae on outer distal corner, merus second
longest article, with serrated margins and plumose seta on
outer margin, carpus longer than propodus, with plumose seta
on outer margin, propodus twice as long as dactylus, with 2
pappose setae on inner margin, dactylus with short simple
terminal setae.
Pereopod 1 with slender articles, basis shorter than rest of
articles combined, with serrated margins, plumose seta on
distal inner and outer corner, tooth on inner margin of ischium,
merus shorter than carpus, carpus shorter than propodus,
propodus 1.6 times as long as dactylus, with simple setae.
Pereopod 2 with slender articles, basis less than half length of
appendage, plumose seta on inner distal corner, simple and
plumose setae on inner margin of merus, carpus 2.8 times as
long as merus, with robust short seta and plumose seta on its
distal inner corner, dactylus 5 times as long as propodus, with
long digitiform extremity, and simple setae on both margins.
Pereopods 3-5, with progressively shorter basis and carpus,
dactylus with long stout terminal seta. Exopods on maxilliped
3 and pereopods 1, 2.
Uropod slender, peduncle 2.5 times as long as pleonite 6
and 1.9 times as long as equal rami, with serrated inner margin,
exopod with short stout terminal seta, endopod with 3 stout
setae on inner margin and robust longer terminal seta.
Etymology. The name of the species is derived from the Latin
“ longa ” - “long” and “dentata” - “with teeth”, due to the long
teeth of maxilliped 2.
Distribution. Off Freycinet Peninsula, Tas.; 720 m depth.
Remarks. The texture of the carapace integument with pits and
spines is as in C. sagamiensis Gamo, 1967 from Japan. It differs
mainly in maxilliped 2 and pereopod 2 with a digitiform tip to
the dactylus. The presence of such long dactylar teeth on
maxilliped 2 is a unique feature within the genus Campylaspis.
144
lorgu Petrescu
Campylaspis lynseyae sp. nov.
Figures 22, 23
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°02.20'S, 148°38.70'E, 800 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn SLOPE 45),
NMV J52962.
Paratypes: 2 immature males, 1 manca (stn SLOPE 45),
NMV J52963.
Diagnosis. Carapace with large lateral sulcus, 0.53 body length,
1.7 times as long as high, large lateral sulcus (0.8 of carapace
length). Small dorsal teeth on first 2 pleonites. Antenna 1
slender. Robust seta of propodus of maxilliped 2 as long as
dactyl ar teeth. Maxilliped 3 slender, dactylus with long terminal
simple setae. Pereopod 1 with merus twice as long as propodus,
few setae on both margins. Pereopod 2 with digitiform tip of
dactylus. Dactylus of pereopods 3-5 fused with its terminal
stout seta. Uropod peduncle little more than twice pleonite 6
length, 1.7 times as long as endopod.
Description. Body with calcified smooth integument. Length:
2.36 mm.
Carapace 0.53 body length, 1.7 times as long as high, large
lateral sulcus (0.8 of carapace length, 4.6 times as long as
high), not meeting the dorsal side, frontal lobe with large base,
pair of small tubercles at base of this lobe, ocular lobe with
acute top, without eyes, long pseudorostrum not upturned,
large siphon, small antennal notch, anteroventral corner acute,
short serration of ventral margin in anterior part. Pleonites 1
and 2 with a pair of dorsal teeth and a lateral tubercle. Pereonite
6 with small tooth on posteroventral corner.
Antenna 1 basal article of peduncle the longest, with 2
long simple setae, main flagellum, 3 -articulate, longer than
distal article of peduncle, accessory flagellum minute.
Maxilliped 2 with plumose seta on distal inner corner of
basis and median part of merus, tooth and 2 simple seta on
inner margin of carpus, propodus second longest article, with
robust seta on outer distal corner as long as dactylar teeth and
tooth on inner margin, dactylus fused with its forked-like
teeth, median one much shorter. Maxilliped 3 basis less than
half as long as appendage, with 2 plumose setae on inner and
outer margin, no outer process, long and slender merus, second
longest article, with small inner concavity, and plumose seta
on both margins, carpus half merus length, with 3 teeth on
outer margin, propodus longer than carpus, dactylus shorter
than propodus, with simple terminal setae much longer than
propodus. Pereopod 1 basis less than half as long as appendage,
merus 3 times as long as ischium, second longest article,
carpus 0.6 length of merus, 1.25 of propodus, propodus twice
as long as dactylus, few setae on ischium to dactylus, dactylus
with simple short setae. Pereopod 2 basis less than half as long
as appendage, plumose seta on inner margin of merus, carpus
1.4 times as long as merus, with simple and plumose seta on
inner distal corner, dactylus 5 times as long as propodus, with
few setae on both margins and a pronounced digitiform tip
with 2 simple subterminal setae. Pereopods 3-5 with
progressively shorter basis and longer carpus (carpus more
than twice merus), dactylus fused with its terminal stout seta.
Exopods in maxilliped 3 and pereopods 1, 2.
Figure 22. Campylaspis lynseyae sp. nov. female holotype: a, body,
lateral view; b, carapace, dorsal view; c, antenna 1; d, maxilliped 2;
e, maxilliped 3. Scale (in mm): a, b, 0.5; c, 0.2; d, 0.1; e, 0.25.
Figure 23. Campylaspis lynseyae sp. nov. female holotype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and left uropod. Scale (in mm): a-f, 0.25.
New nannastacid cumaceans from Museum Victoria
145
Uropod peduncle little more than twice length of pleonite
6, 1.7 times as long as endopod, serrated inner margin, without
any setae, exopod 0.8 of endopod length, with short stout
subterminal seta and much longer terminal one, endopod with
3 stout setae with subterminal setule on inner serrated margin
and terminal robust seta.
Etymology. The species is dedicated to Lynsey Poore as a sign
of gratitude for the kindness shown during my short visit to
Melbourne.
Distribution. Off Freycinet Peninsula, Tas.; 800 m depth.
Remarks. The new species has a large lateral sulcus like C. anae
and C. rectangulata. It differs from these species by presence of
a pair of small tubercles on the base of the ocular lobe, longer
antenna 1, slender maxilliped 3 and pereopod 1 and digitiform
extremity of the dactylus of pereopod 2 (extremity broken in C.
anae). C. spinifera also has a similar sulcus, a pair of tubercles
on the base of the ocular lobe but with a supplementary tubercle
with a spine on top on the lower margin of the sulcus near the
anterolateral corner, a shorter, larger merus of maxilliped 3, and
pereopod 2 with a tapering extremity to the dactylus.
Campylaspis nowrae sp. nov.
Figures 24, 25
Material examined. Holotype female, NSW, off Nowra, 34°51.90'S,
151°12.60'E, 770 m, crinoid dominated, WHOI epibenthic sled, G.C.B.
Poore and C.C. Lu, RV Franklin , 15 Jul 1986 (stn SLOPE 6), NMV J
J52945.
Diagnosis. Carapace with large lateral sulcus and 2 dorsal
transverse ridges. Robust antenna 1. Maxilliped 3 with slender
articles, dactylus with long terminal seta. Pereopod 1 with
merus, carpus and propodus subequal, dactylus with long
plumose subterminal seta. Pereopod 2 with digitiform extremity
of long dactylus. Dactylus fused with its terminal seta in
pereopods 3-5. Short uropods, peduncle 1.3 times as long as
pleonite 6 and 1.7 times as long as equal rami, endopod with 2
stout setae on inner margin.
Description. Body with highly calcified integument, smooth.
Length: 5.1 mm.
Carapace 0.49 body length, 2.3 times as long as high, with a
large lateral sulcus posteriorly continued by a large concavity up
to the median part, 2 dorsal transverse ridges in median part that
reach upper side of sulcus, 2 small tubercles on lateral side of
pseudorostrum, minute ocular lobe with a large base, without
eyes, antennal notch large, no lateral and lower serration. Pleonites
1^1 with lateral acute expansions, pleonite 6 the longest.
Antenna 1 with robust articles, articles of peduncle
progressively shorter toward the distal extremity, main
flagellum 3 -articulate, shorter than distal article of peduncle,
accessory flagellum minute, uniarticulate.
Maxilliped 3 basis little shorter than rest of articles
combined, with hairy margins, plumose setae on outer and inner
distal comers, mems with small inner concavity, with plumose
seta on outer margin, carpus shorter than merus, with plumose
seta on outer serrated margin, propodus second longest article,
dactylus half of propodus, with long simple terminal seta.
Figure 24. Campylaspis nowrae sp. nov. female holotype: a, body,
lateral view; b, body, dorsal view; c, antenna 1; d, maxilliped 3;
e, pereopod 1; f, pereopod 2. Scale (in mm): a, 1; b, 1; c, d, 0.25.
Figure 25. Campylaspis nowrae sp. nov. female holotype: a, pereopod
3; b, pereopod 4; c, pereopod 5; d, pleonite 6 and left uropod. Scale
(in mm): a-d, 0.5.
146
lorgu Petrescu
Pereopod 1 basis less than half as long as appendage, with
serrated outer margin and plumose seta on inner distal corner,
merus, carpus and propodus subequal with numerous plumose
setae on both margins, dactylus shorter than propodus, with
plumose terminal seta. Pereopod 2 basis less than half as long
as appendage, simple and plumose setae on inner margin of
merus, carpus 1.7 times as long as merus, with stout seta and
pappose seta on distal inner corner, dactylus 4.6 times as long
as propodus, as long as merus, carpus and propodus combined,
with lateral setae and digitiform short tip provided with short
simple seta. Pereopods 3-5 with progressively shorter basis
and longer carpus (carpus of 5th pair twice as long as merus),
simple stout long setae on outer margin of carpus, 1 seta on its
outer distal corner, dactylus fused with its terminal stout
microserrate seta. Exopods on maxilliped 3 and pereopods 1,
2; with 2nd article very long.
Uropod peduncle 1.3 times as long as pleonite 6 and 1.7
times as long as equal rami, with serrated inner margin,
exopod 2-articulate, with subterminal microserrate seta and a
terminal longer one, endopod with 2 stout microserrate setae
on inner margin, single subterminal setules and longer terminal
seta (shorter than endopod), and 3 pedunculate setae.
Etymology. The species bears the name of a town in NSW near
the type locality.
Distribution. Off Nowra, NSW; 770 m depth.
Remarks. The carapace of C. nowrae has two pairs of transverse
dorsal crests in the median region similar to the carapace of
form B of C. johnstoni Hale, 1937 found by Ledoyer (1993) in
the Weddell Sea and C. bulbosa Jones, 1974 from the Atlantic
coasts of south-western Africa. It differs in the posterior
extremity of the carapace with a different arrangement of
ridges, in much shorter uropods, and an endopod with two
setae on its inner margin.
Campylaspis pileus Foxon, 1932
Cumella pileus Foxon, 1932: 393, figs 9, 10.
Material examined. 6 females (stn SLOPE 45), NMV J52976; 1 female
(stn SLOPE 46), NMV J52977.
Distribution. Great Barrier Reef, Qld, Tas.; 22-800 m depth.
Remarks. These records extend the range of the species much
further south, to Tasmania, and to greater depth (800 m).
Campylaspis poorei sp. nov.
Figure 26
Material examined. Holotype female, NSW, off Nowra, 34°51.90'S,
151°12.60'E, 770 m, crinoid dominated, WHOI epibenthic sled, G.C.B.
Poore and C.C. Lu, RV Franklin , 15 Jul 1986 (stn SLOPE 6), NMV
J52417.
Diagnosis. Carapace with long, thin narrow sulcus. Antenna 1
long. Maxilliped 2 with 3 short dactylar teeth, median one
small. Merus of maxilliped 3 the 2nd longest article, dactylus
with microserrate terminal setae. Pereopod 1 with slender and
less setulated articles. Dactylus of pereopod 2 without digitiform
extremity. Uropod peduncle 1.6 times as long as pleonite 6 and
1.3 times longer than endopod, exopod shorter than endopod,
endopod with 4 short microserrate setae on inner margin.
Description. Body with smooth integument. Length: 3.1 mm.
Carapace 0.43 of entire body length, twice as long as high,
with long, thin lateral sulcus, big ocular lobe without eyes,
antennal notch small, ventral margin smooth.
Antenna 1 long, proximal article of peduncle longest
article, distal one longer than median article, main flagellum,
3 -articulate, as long as last 2 articles of peduncle, accessory
flagellum, minute, uniarticulate.
Maxilliped 2 basis with strong inner plumose seta, small
tooth and 2 simple setae on inner margin of carpus, propodus
second longest article, with robust seta on outer distal corner,
longer than dactylar teeth, dactylus fused with 3 short teeth,
median one minute. Maxilliped 3 basis half of appendage, 2
plumose setae on outer and inner distal corner, no outer
process, merus second longest article, with serrate inner
margin with plumose setae, long plumose seta on distal outer
corner, carpus less than half merus, with strong serration and
plumose seta on outer margin, propodus 1.5 times longer than
carpus, with 2 pappose setae on inner margin, dactylus half of
propodus, with microserrate setae longer than article.
Figure 26. Campylaspis poorei sp. nov. female holotype: a, body,
lateral view; b, antenna 1; c, maxilliped 2; d, maxilliped 3; e,
pereopod 1; f, pereopod 2; g, pereopod 3; h, pereopod 4; i, pereopod
5; j, pleonite 6 and right uropod. Scale (in mm): a, 0.5; b, c, 0.1; d-j,
0.25.
New nannastacid cumaceans from Museum Victoria
147
Pereopod 1 basis shorter than rest of articles combined,
margins smooth, merus 3 times as long as ischium, with simple
seta on outer margin, carpus second longest article, with 2
simple curved setae on inner margin, propodus 0.8 of carpus
length, with simple setae on inner margin, dactylus 0.7
propodus length, with simple short setae. Pereopod 2 basis
shorter than rest of articles combined, simple seta on inner
margin of merus, carpus 1.8 times as long as merus, with short
stout seta and longer simple one on distal inner corner, dactylus
2.6 times as long as propodus, shorter than carpus, with few
simple setae on both margins, subterminal and terminal
plumose setae. Pereopods 3-5 with progressively shorter basis
and longer carpus (carpus 3 times as long as merus in last
pair), carpus with simple stout long seta on outer distal corner,
dactylus with long stout terminal seta. Exopods on maxilliped
3 and pereopods 1, 2, with 2nd article long.
Uropod peduncle 1.6 times as long as pleonite 6 and 1.3
times as long as its endopod, with serrated inner margin,
exopod 0.8 length of endopod, with long terminal robust seta,
endopod with 4 short microserrate setae on inner margin and
short microserrate terminal seta.
Etymology. The species is dedicated to Gary Poore, Museum
Victoria, Melbourne, Australia, well-known specialist in
Peracarida, as a sign of gratitude for offering me the opportunity
to study this collection in his laboratory.
Distribution. Off Nowra, NSW; 770 m depth.
Remarks. The new species most resembles C. minor Hale,
1945. It differs in the lateral sulcus not reaching the anterior
margin of the carapace, pereopod 1 has slender articles, without
plumose setae, there are fewer plumose setae on pereopod 2,
and the uropod endopod has three setae on the inner margin
instead of two.
Campylaspis pustulosa Hale, 1945
Campylaspis pustulosa Hale, 1945: 207, figs 43, 44.
Material examined. 1 female, 1 male (stn SLOPE 1), NMV J52978; 1
female (stn SLOPE 2), NMV J52979; 2 females, 2 males (stn SLOPE
47), NMV J52980; 1 female, 1 male (stn SLOPE 47), MGAB CUM
1614; 2 females, 1 male (stn SLOPE 48), NMV J52981.
Distribution. Off Eden, NSW, Tas.; 70-600 m depth.
Remarks. New discoveries extend the range of this species
from New South Wales to Tasmania and to 600 m depth.
Campylaspis rectangulata sp. nov.
Figures 27, 28
Material examined. Holotype immature male, NSW, off Nowra,
34°57.90'S, 151°08'E, 503 m, bryozoa and shell, WHOI epibenthic
sled, G.C.B. Poore et al„ RV Franklin, 14 Jul 1986 (stn SLOPE 2),
NMV J54123.
Diagnosis. Carapace with large lateral sulcus, rectangular in
dorsal view, with short thoracic and abdominal segments.
Maxilliped 2 with short dactylar teeth, shorter than robust seta
of propodus. Merus of maxilliped 3 with inner concavity,
Figure 27. Campylaspis rectangulata sp. nov. female holotype: a, body,
lateral view; b, carapace and pereon, dorsal view; c, antenna 1; d,
maxilliped 2; e, maxilliped 3; f, pereopod 1. Scale (in mm): a, b, 0.5;
c-e, 0.25; f, 0.5.
a
Figure 28. Campylaspis rectangulata sp. nov. female holotype: a,
pereopod 2; b, pereopod 3; c, pereopod 4; d, pereopod 5; e, pleonite
6 and left uropod. Scale (in mm): a, 0.5; b-e, 0.25.
148
lorgu Petrescu
enlarged merus and carpus. Pereopod 1 with short articles
(except enlarged merus), with numerous plumose setae.
Dactylus of pereopod 2 without digitiform extremity. Uropod
peduncle twice as long as pleonite 6 and 1.4 times as long as its
equal rami, without setae, exopod with long robust terminal
seta, endopod with 3 microserrate setae on inner margin and
longer, more robust, terminal seta.
Description. Body with highly calcified smooth integument.
Length: 3.5 mm.
Carapace 0.51 body length, covering first 4 pereonites in
lateral view, 1.8 times as long as high, with large lateral sulcus
(0.6 of carapace length, 3 times as long as high), frontal lobe
with large base and small ocular lobe without eyes, antennal
notch almost absent, anterolateral corner with short serration.
Pleon elongated, 0.45 body length.
Antenna 1 peduncle with short and robust articles, with
serrated margins, main flagellum 3-articulate, longer than distal
article of peduncle, accessory flagellum tiny, uniarticulate.
Maxilliped 2 basis with strong plumose seta on inner
margin, similar seta medially on distal margin of merus,
carpus with 2 simple setae on inner margin, propodus second
longest article, with robust curved seta on outer distal corner,
longer than dactylar teeth, dactylus fused with 3 short, equal,
teeth. Maxilliped 3 basis less than half as long as appendage,
with serrated inner margin, with 2 plumose setae on inner
distal corner and another 2 longer on outer distal corner,
ischium with tooth and plumose seta on inner margin, merus
second longest article, with inner concavity with setae, tooth
and long plumose seta on outer margin, carpus enlarged,
shorter than merus, with 2 teeth and 2 plumose setae on outer
margin, propodus shorter than carpus, thinner, with 3 pappose
setae on inner margin, dactylus 0.58 propodus length, with 3
long simple terminal setae.
Pereopod 1 basis less than half as long as appendage, with
serrated margins (outer margin stronger), with 2 plumose setae
on inner distal corner, short and robust articles with serrated
margins and numerous plumose setae excepting dactylus with
short simple ones, merus second longest article, carpus longer
than propodus, propodus longer than dactylus. Pereopod 2
bulky, basis less than half as long as appendage, with serrated
margins, short plumose seta on inner distal corner, 1 simple and
1 plumose seta on inner distal corner of merus, short carpus (1.5
times as long as merus), with short plumose seta on distal inner
corner, dactylus 3 twice as long as propodus, with a right-angled
extremity, with simple and plumose setae on margins, long stout
terminal seta, shorter than dactylus. Pereopods 3-5: pair 3 and
4 with enlarged basis, progressively shorter, carpus progressively
longer (3 times as long as merus in 5th pair), dactylus with
robust long terminal seta. Exopods, 5 pairs incompletely
developed (characteristic of immature specimen).
Uropod peduncle twice as long as pleonite 6 and 1.4 times
as long as equal rami, without setae, with serrated inner
margin, exopod with long robust terminal seta, endopod with
3 microserrate setae on inner margin and longer, more robust,
terminal seta.
Etymology. The name “ rectangulata ” describes the carapace
in dorsal view, almost square.
Distribution. Off Nowra, NSW; 503 m depth.
Remarks. Campylaspis rectangulata is similar to C. anae (in
the form of the carapace and similar antenna 1). It differs in
having a maxilliped 2 with shorter teeth on the dactylus, a
shorter robust distal seta on the propodus, merus of maxilliped
3 with an inner concavity and a stout seta (versus none) and
shorter uropods with slender peduncle, longer subequal rami
(versus exopod shorter than endopod in C. anae), and three
microserrate setae on the inner margin of the endopod instead
of four. Pereopods are sexually dimorphic.
Campylaspis sculpta sp. nov.
Figures 29, 30
Material examined. Holotype immature male, dissected, Vic., eastern
Bass Strait, 7.3 km SSW of Cape Conran, 37°52.67'S, 148°42.06'E, 48
m, sand-shell, Smith-Mclntyre grab. Marine Science Laboratories,
RV Sarda, 28 Sep 1990 (stn MSL-EG 62), NMV J23259.
Paratypes: 1 immature male (stn MSL-EG 108), NMV J27471; 1
manca (stn MSL-EG 57), NMV J23258, 1 manca (stn MSL-EG 62),
NMV J52955.
Diagnosis. Carapace highly sculptured, with short thoracic and
abdominal segments. Maxilliped 3 and pereopod 1 with
enlarged articles, except dactylus, propodus 2nd longest article
after basis. Pereopod 2 with enlarged articles, with long
dactylus, longer than basis, with digitiform tip. Pereopods 3-5
with short and robust articles, dactylus fused with robust
terminal seta. Short and robust uropods, uropod peduncle 1.25
times as long as pleonite 6 and 1.3 times as long as equal rami.
Exopod and endopod (wider than exopod) with rounded tips,
with fine short setae.
Description. Body with strongly calcified and smooth
integument. Length: 6.08 mm.
Carapace 0.49 body length, 1.9 times as long as high, with
large, blunt tubercles around large lateral sulcus (0.8 of
carapace length), pseudorostrum short, ocular lobe large,
without lenses, pair of tubercles and transverse ridge at its
base, antennal notch small, anterolateral corner continued
towards ocular lobe by an oblique ridge.
Antenna 1 peduncle with short and robust articles, median
article as long as distal one, main flagellum with 3 short and
robust articles, shorter than last article of peduncle, median
and distal article with an aesthetasc, accessory flagellum tiny,
uniarticulate.
Maxilliped 3 basis half of appendage, with 2 plumose
setae on inner distal corner and another longer 2 on distal
outer corner, merus longer than carpus and ischium, shorter
than propodus (0.7), with long plumose seta on outer margin,
carpus about half propodus, with small tooth and plumose
seta on outer margin, propodus 2nd longest article, with short
plumose seta on outer margin and 2 fine setae on inner margin,
dactylus 0.2 length of propodus, enlarged, with fine terminal
seta.
Pereopod 1 basis half of appendage, with 3 long plumose
setae on inner margin, enlarged ischium to dactylus, ischium
with tooth and long plumose seta on inner margin, merus
widest article, with plumose setae on inner margin, carpus as
New nannastacid cumaceans from Museum Victoria
149
Figure 29. Campylaspis sculpta sp. nov. female holotype: a, body,
lateral view; b, body, dorsal view; c, antenna 1; d, maxilliped 3; e,
pereopod 1. Scale (in mm): a, b, 1; c, 0.5; d, e, 0.5.
Figure 30. Campylaspis sculpta sp. nov. female holotype: a,
pereopod 2; b, pereopod 3; c, pereopod 4; d, pereopod 5; e, pleonite
6 and right uropod. Scale (in mm): a-d, 0.5; e, 0.5.
long as merits, with plumose seta on outer margin, propodus,
2nd longest article, 1.1 times as long as carpus, with 3 plumose
setae on outer margin, oval dactylus 0.7 of propodus length,
with fine terminal seta. Pereopod 2 basis shorter, less than 3rd
of appendage length, ischium with 3 simple setae on inner
margin, robust merus, the widest article, with small tooth and
3 simple setae on inner margin, carpus little longer than merus,
with small tooth and short plumose seta on inner distal corner,
long, curved and with digitiform dactylus, 4.6 times as long as
propodus, longest article, with 3 fine subterminal setae.
Pereopod 3 basis more than half of appendage, merus twice as
long as ischium, carpus 0.7 of merus length, with stout long
seta on outer margin, propodus 0.6 of carpus length, with
robust distal seta, dactylus fused with robust terminal seta.
Pereopod 4 basis shorter than in previous pair, merus subequal
to carpus, propodus little longer than in previous pair. Pereopod
5 with shorter basis, carpus 1.8 times as long as merus.
Exopods, 5 pairs.
Pleonite 6 and uropod with hyaline fringes. Uropod short
and robust, uropod peduncle 1.25 times as long as pleonite
6 and 1.3 times as long as equal rami, serrated inner margin,
exopod and endopod (wider than exopod), with rounded
tips and serrated inner margins, exopod with 3 fine terminal
setae, endopod with 3 stout short setae, terminal one little
longer.
Etymology. The name “ sculpta ” refers to the sculptured
carapace.
Distribution. Eastern Vic.; 48-50 m depth.
Remarks. Campylaspis sculpta is similar to C. wardi Bacescu,
1991 and C. grossui. They have similar carapaces, short and
robust pereopods and uropods. C. sculpta differs from C. wardi
in: fewer tubercles on the carapace; larger sulcus; larger
propodus (second longest article) and rounded dactylus of
maxilliped 3; larger propodus and dactylus; rounded dactylus
of pereopod 1; much longer dactylus of pereopod 2; and the
uropod exopod with few terminal fine setae and the endopod
with three stout ones, versus the endopod with terminal short
setae with the oval structures characteristic of C. wardi. The
shape of the carapace in dorsal view is different from C.
grossui. The species also differ in: stronger antenna 1, tiny
rounded dactylus of maxilliped 3 and enlarged and rounded
dactylus of pereopod 1, versus slender antenna 1 and maxilliped
3 and pereopod 1 with larger articles and dactylus in C. wardi
and C. grossui. Pereopod 2 has a much longer dactylus, four
times as long as the propodus in C. sculpta, and with a
digitiform extremity, versus twice as long in C. wardi and C.
grossui, which have a normal extremity. The uropod rami are
stronger than in C. grossui and it also has finer setae.
Campylaspis s errata sp. nov.
Figures 31, 32
Material examined. Holotype female, NSW, off Nowra, 34°51.90'S,
151°12.60'E, 770 m, crinoid dominated, WHOI epibenthic sled, G.C.B.
Poore and C.C. Lu, RV Franklin, 15 Jul 1986 (stn SLOPE 6), NMV
J45786.
150
lorgu Petrescu
Figure 31. Campylaspis serrata sp. nov. female holotype: a, body,
dorsal view; b, detail of integument from carapace; c, detail of
integument of pereon and pleon; d, antenna 1; e, maxilliped 2. Scale
(in mm): a, 1; d, 0.25; e, 0.2; f, 0.25.
Figure 32. Campylaspis serrata sp. nov. female holotype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and right uropod. Scale (in mm): a, b, 0.5; c-e, 0.5; f,
0.25.
Diagnosis. Carapace covered with spines, small tubercles and
long club-like setae interspersed with pits. Integument of pereon
and pleon with pits and small spines. Antenna 1, maxilliped 3,
pereopods 1 and 2 and uropods with densely serrated margins.
Maxilliped 3 with merus second longest article, with long outer
spine, carpus with 3 outer teeth, dactylus with long simple setae
longer than article. Pereopod 1 with slender articles, with few
setae. Pereopod 2 with long dactylus with straight tip. Uropod
peduncle 2.25 times as long as pleonite 6 and 1.9 times as long
as exopod, exopod little longer than endopod.
Description. Body with integument densely pitted, with
bludgeon-like setae, small tubercles and spines on carapace
and with small spines on rest. (The only specimen, holotype, is
not in good shape, being a little dorsally compressed and
twisted). Length: 4.5 mm.
Carapace about half body length, 1.3 times as long as wide,
almost triangular in dorsal view, pair of small tubercles in
median region, bigger one on branchial part and larger ones
with spine on top at base of frontal lobe, integument densely
pitted, with long spines and club-like setae, ocular lobe with
spoon-like tip with crown of small spines, without eyes,
pseudorostral lobes straight. Lateral plates of pereon with long
spines, integument covered with pits and small spines. Pleon
with pits and spines (longer ones are broken).
Antenna 1 peduncle with strongly serrated articles, main
flagellum 3 -articulate, longer than distal article of peduncle,
accessory flagellum minute, uniarticulate.
Maxilliped 2 with strong plumose setae on inner margin of
basis and distally on merus, carpus with small tooth and 2
simple setae on inner margin, propodus 2nd longest article,
with robust seta on outer distal corner, longer than dactylar
teeth, dactylus with 3 teeth, median one minute. Maxilliped 3
with slender articles, basis less than half as long as appendage,
with plumose seta on inner margin and 2 longer ones on outer
distal corner, no outer process of basis, merus second longest
article, with strong tooth and plumose seta on outer distal
corner, carpus shorter than merus, with 3 teeth on outer
margin, propodus little longer than carpus, with 3 pappose
setae on inner margin, dactylus 0.75 of propodus length, with
long simple setae, longer than dactylus.
Pereopod 1 with slender articles, basis longer than rest of
articles combined, with 3 plumose setae on outer margin, ischium
with 3 teeth (one larger) on inner margin, merus little shorter
than carpus, with plumose seta on outer margin, carpus 2nd
longest article, with simple setae, propodus subequal to merus,
with simple setae, dactylus 0.57 of propodus length, with long
simple setae. Pereopod 2 basis more than 3rd of appendage
length, with plumose seta on distal inner comer, ischium and
merus with similar seta, carpus 1.7 times as long as merus, with
small tooth and long plumose seta on inner distal corner, dactylus
3.75 times as long as propodus, with numerous simple setae, and
1 terminal longer simple stout seta. Pereopods 3-5 with serrated
margins, progressively shorter, with plumose setae, progressively
shorter carpus (3 times as long as merus in 3rd pair), with
plumose seta on inner margin and stout simple one on distal
comer, dactylus with long stout simple terminal seta. Exopods
on maxilliped 3 and pereopods 1, 2; with serrated bases.
New nannastacid cumaceans from Museum Victoria
151
Uropod peduncle 2.25 times as long as pleonite 6 and 1.9
times as long as exopod, with highly serrated margins, exopod
little longer than endopod, with serrated margins and robust
terminal seta, endopod with serrated margins, with 2 long
microserrate setae on inner margin and a longer, more robust,
terminal one.
Etymology. The name of the species describes the serrated
appendages.
Distribution. Off Nowra, NSW; 770 m depth.
Remarks. Another species of Campylaspis with such highly
serrated appendages is C. echinata Hale, 1945 (described only
from the male). C. serrata has a differently textured carapace
integument, densely pitted, with long spines and bludgeon-like
setae, an ocular lobe with a spoon-like tip with a crown of small
spines, uropods with the exopod slightly longer than the
endopod, and the endopod with two microserrate long setae on
the inner margin instead of one.
Campylaspis setifera sp. nov.
Figures 33, 34
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°00.20'S, 148°37.70'E, 720 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin , 27 Jul 1986 (stn SLOPE 46),
NMV J52953.
Paratypes: 1 female, 1 dissected female (stn SLOPE 46), NMV
J52954; 1 female (stn SLOPE 46), MGAB CUM 1611.
Diagnosis. Carapace high and smooth. Antenna 1 with long
main flagellum. Dactylus of maxilliped 2 with short teeth.
Maxilliped 3 with enlarged articles, merus 2nd longest article.
Pereopod 1 with elongated articles with numerous plumose
setae, merus second longest article. Pereopod 2 with long
carpus, 2.5 times as long as merus, short dactylus with
digitiform extremity. Dactylus of pereopods 3-5 fused with
terminal stout seta. Long uropods, uropod peduncle 2.4 times
as long as pleonite 6 and twice as long as endopod, with 8
microserrate setae on inner margin, endopod little longer than
exopod, with 12 microserrate setae on inner margin.
Description. Body with highly calcified smooth integument.
Length: 4.7 mm.
Carapace bulky, 0.51 body length, 1.3 times as long as
high, oval in dorsal view, with short straight truncated
pseudorostrum, ocular lobe with large base and minute tip,
without lenses, antennal notch minute, lower margin smooth.
Pereon 0.07 of entire body length, last 2 segments more visible.
Pleon 0.41 body length, elongate.
Antenna 1 peduncle basal article the longest, rest
progressively shorter, main flagellum 3 -articulate, longer than
last 2 articles of peduncle combined, accessory flagellum tiny,
uniarticulate.
Maxilliped 2 basis and merus with strong plumose seta,
ischium visible, carpus with tooth and simple seta on inner
margin, propodus with inner tooth and simple seta, outer robust
seta on distal corner, dactylus with 3 equal teeth shorter than
propodal seta. Maxilliped 3 with enlarged articles, massive basis
less than half as long as appendage, with 2 pappose setae on
Figure 33. Campylaspis setifera sp. nov. female paratype: a, body,
lateral view; b, carapace and pereon, dorsal view; c, antenna 1; d,
maxilliped 2; e, maxilliped 3. Scale (in mm): a, b, 1; c, 0.2; d, e, 0.5.
Figure 34. Campylaspis setifera sp. nov. female paratype: a, pereopod
1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod 5; f,
pleonite 6 and left uropod. Scale (in mm): a-f, 0.5.
152
lorgu Petrescu
inner distal corner and an other 2, longer, on outer comer,
ischium with hyaline outer crest, merus 2nd longest article, with
2 plumose setae medially, serrated inner margin, tooth and
pappose seta on outer margin, carpus shorter than merus, with
stronger serration on inner margin, tooth and pappose seta on
outer margin, propodus shorter than carpus, with 4 teeth on inner
margin, dactylus shorter than propodus, with simple setae.
Pereopod 1 basis longer than half of appendage, with long
plumose seta on inner distal corner, ischium with tooth and long
plumose seta on inner margin, merus 2nd longest article, with
plumose setae on both margins, carpus as long as propodus,
with plumose setae on both margins, propodus with plumose
setae, dactylus almost half of propodus, with simple and
plumose setae. Pereopod 2 basis less than half as long as
appendage, with plumose seta on inner distal corner, merus
with a similar one, carpus long, 2.5 times as long as merus,
short dactylus (4 times as long as propodus), with digitiform
extremity with few simple short setae. Pereopods 3-5 basis
longer than half of appendage in 3rd pair, progressively shorter
in next 2, carpus progressively longer, twice as long as merus in
5th pair. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 2.4 times as long as pleonite 6 and twice
as long as endopod, with 8 microserrate long setae on inner
margin, exopod with a long microserrate stout seta, endopod 8
little longer than exopod, with 12 microserrate setae on inner
margin and longer terminal seta.
Etymology. The species is named “ setifera ” because of the
unusual feature for a female.
Campylaspis, numerous setae on the uropods typical of
males.
Distribution. Off Freycinet Peninsula, Tas.; 720 m depth.
Remarks. The new species is closer to C. thompsoni Hale, 1945
than to other species of the genus. It is strongly vaulted above
the carapace, has a massive maxilliped 3 with numerous teeth
on the margins of the merus and carpus, the pereopod 1 with
numerous plumose setae, the merus being the second longest
article, and the dactylus of pereopod 2 has a digitiform tip, and
long uropods. C. setifera differs in: the ocular lobe is without
lenses; antenna 1 has a longer main flagellum; maxilliped 2 has
three dactylar teeth instead of four; the propodus of maxilliped
3 also has inner teeth, longer carpus and shorter dactylus in
pereopod 2; and the uropod is totally different, almost male-
like, with numerous microserrate setae on its peduncle and
endopod (no setae on the peduncle and four setae on the
endopod in C. thompsoni ). C. setifera is the only species from
this material without a sulcus, tubercles or spines on the
carapace.
Campylaspis spinifera sp. nov.
Figures 35, 36
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°00.20'S, 148°37.70'E, 720 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin , 27 Jul 1986 (stn SLOPE 46),
NMV J52931.
Paratypes: 14 females, 10 immature males, 1 female, dissected,
NMV J52932; 2 females (stn SLOPE 46), MGAB CUM 1610.
Figure 35. Campylaspis spinifera sp. nov. female paratype: a, body,
lateral view; b, carapace, dorsal view; c, antenna 1; d, maxilliped 2;
e, maxilliped 3. Scale (in mm): a, b, 1; c, d, 0.2; e, 0.25.
Figure 36. Campylaspis spinifera sp. nov. female paratype: a, pereopod
1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod 5; f,
pleonite 6 and left uropod. Scale (in mm): a-f, 0.25.
New nannastacid cumaceans from Museum Victoria
153
Diagnosis. Carapace with a lateral sulcus, tubercles with spines
at the base of frontal lobe and near the anterolateral corner,
ocular lobe without lenses. Pereopod 1 basis less than half as
long as appendage, with slender articles. Pereopod 2 carpus 1.8
times as long as merus and with straight extremity of dactylus.
Uropod peduncle 1.4 times as long as pleonite 6 and its endopod,
with serrated margins, exopod little shorter than endopod, with
terminal robust long seta, endopod with 3 stout setae on inner
serrated margin and robust long terminal seta.
Description. Body elongated, with smooth integument. Length:
3.9 mm.
Carapace 0.47 of entire body length, longer than high, with
lateral sulcus, pair of tubercles provided with apical strong
spines at base of frontal lobe and near pointed anterolateral
corner, antennal notch small, ocular lobe small, without lenses,
pseudorostrum straight.
Antenna 1 peduncle with progressively shorter articles,
main flagellum 3 -articulate, longer than distal article of
peduncle, accessory flagellum minute, uniarticulate.
Maxilliped 2 plumose long seta on inner margin of basis
and merus, 2 simple setae and tooth on inner margin of carpus,
propodus with robust seta on outer margin, longer than dactylar
teeth, dactylus with 3 equal teeth. Maxilliped 3 basis half of
appendage, with 2 plumose setae on distal inner and outer
corners, merus second longest article, with tooth and plumose
seta on outer margin, shorter carpus, with 2 teeth and plumose
seta on outer margin, slender propodus as long as carpus, with
2 pappose setae on inner margin, dactylus half of propodus,
with simple terminal setae longer than dactylus.
Pereopod 1 basis less than half as long as appendage, with
slender articles, 6 strong serrations on inner margin of ischium,
merus and carpus, with fewer simple and plumose setae, merus
2nd longest article, progressively shorter articles from carpus
to dactylus, dactylus with short simple setae. Pereopod 2 basis
longer than one-third of rest of articles combined, merus with
plumose seta on inner and outer margin, carpus 1.8 times as
long as merus, with short robust seta and a simple one on inner
distal corner, dactylus 3.4 times as long as propodus, with
simple and plumose setae and straight extremity. Pereopods
3-5 with slender articles, progressively shorter basis and
longer carpus, twice as long as merus in 5th pair. Exopods on
maxilliped 3 and pereopod 1, 2.
Uropod peduncle 1.4 times as long as pleonite 6 and its
endopod, with serrated margins, exopod little shorter than
endopod, with terminal robust long seta, endopod with 2 stout
setae on inner serrated margin and robust long terminal seta.
Terminal setae with 1 subterminal setule in both rami.
Etymology. The name of the species reflects the presence of
characteristic spines on the carapace.
Distribution. Off Freycinet Peninsula, Tas.; 720 m depth.
Remarks. This species is more similar to C. lynseyae than
to others. It differs in: a pair of tubercles with a spine on top
near the anterolateral margin of the carapace; shorter dactylar
teeth of maxilliped 2; larger merus of maxilliped 3; a tapering
dactylus of pereopod 2; and shorter and stronger serrated
uropods.
Campylaspis tasmaniensis sp. nov.
Figures 37-39
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°02.20'S, 148°38.70'E, 800 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn SLOPE 45),
NMV J52947.
Allotype: male (stn SLOPE 45), NMV J52948.
Paratypes: 1 female, 2 mancas (stn SLOPE 45), NMV J52949.
Diagnosis. Carapace, pereon and pleon with numerous
tubercles. Basal article of main flagellum of antenna 1 enlarged.
Dactylar teeth of maxilliped 2 as long as robust seta of
propodus, median tooth much smaller. Maxilliped 3 basis
short, dactylus with long terminal simple setae. Pereopods with
slender articles. Pereopod 1 propodus as long as merus, longer
than carpus. Pereopod 2 with long dactylus with straight
extremity. Uropod peduncle 2.3 times as long as pleonite 6 and
endopod, with serrated margins, exopod little shorter than
endopod, with subterminal microserrate seta and longer robust
terminal seta, endopod with 3 stout setae on inner serrated
margin, a subterminal and a terminal microserrate robust seta.
Description of female. Body with tubercled reticulated
integument, without setae or spines. Length: 5.4 mm.
Carapace half body length, 2.1 times as long as high, with
lateral sulcus that rises from anterolateral part and fuses with
similar sulcus from other side, 2 parallel dorsal rows of acute
tubercles, 2 transverse parallel rows of tubercles at base of
frontal lobe, rows of tubercles delimiting the sulcus, 3 spines
on upper side of sulcus toward tip of pseudorostrum, small
antennal notch, anterolateral corner acute, short serrated
ventral margin, ocular lobe minute, without lenses,
pseudorostrum long and straight.
Antenna 1 proximal article of peduncle the longest, distal
article longer than median one, short main flagellum 3 -articulate,
basal one enlarged, accessory flagellum minute, 1-articulate.
Maxilliped 2 with plumose long seta on inner margin of
basis and merus, 2 simple setae and tooth on inner margin of
carpus, propodus with robust seta on outer margin, as long as
dactylar teeth, dactylus with 3 teeth, median one much shorter.
Maxilliped 3 basis less than half as long as appendage, with 1
plumose seta on distal inner and outer corners, merus 2nd
longest article, with long plumose seta on outer margin and
simple and plumose setae on inner margin, carpus half of
merus, with 3 teeth and long plumose seta on outer margin,
slender propodus 1.6 times as long as carpus, dactylus 0.6 of
propodus length, with simple long terminal setae (3.5 times as
long as dactylus).
Pereopod 1 basis less than half as long as appendage, with
slender articles, serration on inner margin of merus and carpus,
with fewer simple and plumose setae, propodus and dactylus
with simple setae, merus 2nd longest article, propodus as long
as merus and longer than carpus, dactylus with short simple
setae. Pereopod 2 basis more than one -third of rest of articles
combined, with plumose seta on distal inner and outer corner,
merus with plumose seta on inner and outer margin, carpus 1.8
times as long as merus, with short robust seta, a plumose long
seta on inner distal corner, dactylus 4 times as long as propodus,
154
lorgu Petrescu
Figure 37. Campylaspis tasmaniensis sp. nov. female holotype: a,
body, lateral view; b, body, dorsal view; c, antenna 1; d, maxilliped
2; e, maxilliped 3. Scale (in mm): a, b, 1; c, d, 0.2; e, 0.25.
Figure 38. Campylaspis tasmaniensis sp. nov. female holotype:
a, pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4;
e, pereopod 5; f, pleonite 6 and left uropod. Scale (in mm): a-e, 0.5;
f, 0.25.
Figure 39. Campylaspis tasmaniensis sp. nov. male allotype: a, body,
lateral view; b, body, dorsal view; c, maxilliped 3; d, pereopod 1; e,
pereopod 2; f, pereopod 3; g, pereopod 4; h, pereopod 5. Scale (in
mm): a, b, 1; c-h, 0.5.
with plumose setae and straight extremity. Pereopods 3-5 with
slender articles, progressively shorter basis and longer carpus,
twice as long as merus in 5th pair, 2 plumose setae on outer
margin of carpus in pair 3 and 4, dactylus with stout long
terminal seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 2.3 times as long as pleonite 6 and
endopod, with serrated margins, exopod little shorter than
endopod, with subterminal microserrate seta and longer simple
terminal seta, endopod with 3 stout setae on inner serrated
margin, subterminal and terminal microserrate robust setae.
Description of male. Body with integument as in female, last 3
pleonites missing. Carapace with more dorsal tubercles, lateral
sulcus shorter than in female, sulcus from each side separated,
carapace more compressed laterally than in female.
Maxilliped 3 basis shorter, a half of appendage, merus more
slender than in female, propodus with longer plumose seta on
outer margin, dactylus with long terminal setae as in female.
Pereopod 1 with more numerous plumose setae on all
articles including dactylus. Pereopod 2 dactylus shorter than
in female, 3 times as long as propodus (versus 4 times), with
more setae. Pereopods 3, 4 with large basis characteristic of
males, 3-5 with more plumose setae, dactylus fused with
terminal robust seta.
Etymology. The species bears the name of the type locality -
Tasmania.
New nannastacid cumaceans from Museum Victoria
155
Distribution. Off Freycinet Peninsula, Tas.; 800 m depth.
Remarks. The general body shape is reminiscent of C.
sagamiensis Gamo, 1967 but without pits and spines on the
integument, with a larger lateral sulcus, with a bulky basal
article of the main flagellum of antenna 1, an accessory
flagellum with one article, not with two. C. tasmaniensis also is
close to C. mosambica Ledoyer, 1988 but with more tubercles
on the carapace, and the dactylus of maxilliped 3 with a much
longer terminal seta.
Campylaspis thetidis Hale, 1945
Campylaspis thetidis Hale, 1945: 212, figs 47, 48.
Material examined. 5 females (stn SLOPE 1), NMV J52982; 1 female
(stn SLOPE 21), NMV J52983.
Distribution. Off NSW, 75 m depth.
Remarks. The species is recorded from a greater depth than
previously, now down to 209 m depth.
Campylaspis thompsoni Hale, 1945
Campylaspis thompsoni Hale, 1945: 183, figs 24, 25.
Material examined. 2 females, 6 males (stn SLOPE 1), NMV J54121;
1 female, NMV J52984; 3 females, 1 male (stn SLOPE 22), NMV
J52985; 6 females (stn SLOPE 27), NMV J52412; 2 females, 1 manca
(stn SLOPE 46), NMV J54126; 1 female (stn SLOPE 45), MGAB
CUM 1615; 1 female, 2 males (stn SLOPE 47), NMV J52987; 1 female,
1 male (stn SLOPE 48), NMV J52988.
Distribution. NSW, Vic., surface to 80 m depth.
Remarks. The new records extend the species’ range into
Victoria.
Campylaspis triplicata Hale, 1945
Campylaspis triplicata Hale, 1945: 200, figs 37, 38.
Material examined. 1 female, 1 male (stn SLOPE 6), NMV J52989; 3
females (stn SLOPE 19), NMV J52990; 1 female (stn SLOPE 22),
NMV J52991; 1 female, 2 males (stn SLOPE 45), NMV J52992; 2
females, 1 male (stn SLOPE 45), MGAB CUM 1616; 2 females (stn
SLOPE 46), NMV J53017.
Distribution. Qld, NSW, Tas.; 12-800 m depth.
Remarks. Previously known from Queensland in shallow water
the species’ range is extended to New South Wales and
Tasmania and to greater depths (800 m).
Campylaspis trisulcata sp. nov.
Figures 40, 41
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°02.20'S, 148°38.70'E, 800 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin , 27 Jul 1986 (stn SLOPE 45),
NMV J52950.
Diagnosis. Carapace large, with 3 lateral sulci. Propodus of
maxilliped 2 with robust outer seta longer than dactylar teeth.
Maxilliped 3 with 2 long plumose setae on outer distal corner,
\
Figure 40. Campylaspis trisulcata sp. nov. female holotype: a, body,
lateral view; b, carapace, dorsal view; c, antenna 1; d, maxilliped 2; e,
maxilliped 3. Scale (in mm): a, b, 1; c, 0.25; d, 0.3; e, 0.25.
Figure 41. Campylaspis trisulcata sp. nov. female holotype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and left uropod. Scale (in mm): a, b, 0.5; c-f, 0.25.
156
lorgu Petrescu
merus 2nd longest article, simple terminal setae twice as long
as dactylus. Pereopod 1 with short articles, with numerous
plumose setae. Pereopod 2 with carpus 1.3 times as long as
merus, dactylus 3 times as long as propodus. Uropod peduncle
2.3 times as long as pleonite 6 and 1.9 times as long as endopod,
with serrated inner margin, exopod shorter than endopod, with
robust terminal seta, endopod with 5 stout setae on serrated
inner margin, long robust terminal seta.
Description. Body with strongly calcified, smooth, integument.
Length: 5.1 mm.
Carapace 0.46 body length, twice as long as high, 1.1 times
as long as wide, with 3 lateral sulci with strongly elevated
margins, transverse lateral ridge at level of base of frontal
lobe, ocular lobe without eyes, antennal notch small,
anterolateral corner with short serration.
Antenna 1 short, peduncle with progressively shorter
articles, main flagellum 3 -articulate, little longer than last
article of peduncle, accessory flagellum minute, uniarticulate.
Maxilliped 2 strong, with long plumose setae on basis and
merus, tooth and 2 simple setae on carpus, propodus 2nd longest
article, with robust outer seta longer than dactylar teeth, dactylus
with 3 teeth, median one shorter. Maxilliped 3 basis little less
than half as long as appendage, with 2 long plumose seta on
outer distal comer, merus 2nd longest article, strongly serrated
on inner margin, with tooth and plumose seta on outer margin,
carpus 0.55 of merus length, long plumose seta on outer margin,
propodus 0.75 of merus length, with 3 pappose setae on inner
margin, simple terminal setae twice as long as dactylus.
Pereopod 1 with short articles, with numerous plumose
setae, basis little less than half as long as appendage, merus to
propodus with serrated margins, merus 2nd longest article,
carpus shorter than merus, propodus longer than carpus and
1.5 times as long as dactylus, dactylus with simple setae.
Pereopod 2 basis less than half as long as appendage, plumose
seta on inner margin of basis and merus, with carpus 1.3 times
as long as merus, with simple seta and plumose seta on distal
inner corner, dactylus 3 times as long as propodus, with simple
setae, terminal one subequal to dactylus. Pereopods 3-5 basis
longer than rest of articles combined, shorter in last pair,
carpus twice as long as merus, dactylus fused with terminal
seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 2.3 times as long as pleonite 6 and 1.9
times as long as endopod, with serrated inner margin, exopod
shorter than endopod, with robust terminal seta, endopod with
4 stout setae on serrated inner margin, single subterminal
setule and long robust terminal seta shorter than endopod.
Etymology. The name “ trisulcata ” reflects the presence of
three lateral sulci on the carapace.
Distribution. Off Freycinet Peninsula, Tas.; 800 m depth.
Remarks. C. trisulcata is similar to two other species with three
or four lateral ridges on the carapace delimiting depressions
(sulci): C. triplicata Hale, 1945 from Australia and C. sinuosa
Gamo, 1960 from Japan. C. triplicata has four lateral ridges
delimiting three depressions, and a massive merus on maxilliped
3. C. sinuosa is much closer, with four lateral ridges on the
carapace but bordering only two depressions, not three as in
C. trisulcata (more evident in dorsal view). It also differs in: the
dactylus of maxilliped 2 with four teeth (three in C. trisulcata)-,
a massive maxilliped 3 with teeth on the propodus and short
setae on the dactylus (versus smooth propodus and long terminal
setae of dactylus); shorter uropods; and an endopod with two
setae on its inner margin (versus four setae in C. trisulcata ).
Campylaspis uniplicata Hale, 1945
Campylaspis uniplicata Hale, 1945: 189, figs 29, 30.
Material examined. 2 females (stn SLOPE 1), NMV J52993; 2 females
(stn SLOPE 1), MGAB CUM 1617; 1 manca (stn SLOPE 2), NMV
J52995; 2 females (stn SLOPE 45), NMV J52994; 1 female (stn SLOPE
47), NMV J52996.
Distribution. East of Port Hacking, NSW and Tas.; 100-800 m
depth.
Remarks. Originally described from New South Wales, the
species is now known in Tasmanian waters and from greater
depths (800 m).
Campylaspis unisulcata Hale, 1945
Campylaspis unisulcata Hale, 1945: 187, figs 27, 28.
Material examined. 2 females (stn SLOPE 22), NMV J52997; 1 female
(stn SLOPE 45), NMV J54120; 7 females, 2 males (stn SLOPE 46),
NMV J52998; 2 females, 1 male (stn SLOPE 46), MGAB CUM 1618.
Distribution. East of Port Hacking, NSW, 100 m depth.
Remarks. The species is now recorded from Tasmania and at
greater depth, 800 m.
Procampylaspis Bonnier, 1896
Procampylaspis Bonnier, 1896: 541.— Hale, 1945 : 214.— Bacescu,
1992: 251.— Stoddart and Lowry, 2003: 373-418.
Type species. Procampylaspis echinata Bonnier, 1896.
Remarks. The genus is well-defined by the rake -like dactylus
of maxilliped 2 and by a long ischium on pereopod 1. One
species is known from Australia but three new species are
described in this study.
Key to species of Procampylaspis from Australian waters
1. Carapace without dorsal tubercles
P. australiensis sp. nov.
— Carapace with dorsal tubercles 2
2. Carapace with 1 dorsal tubercle
P. sordida Hale, 1945
— Carapace with dorsal tubercles in 2 parallel rows (5 in
female, 4 in male) P. spinifera sp. nov.
Procampylaspis australiensis sp. nov.
Figures 42-44
Material examined. Holotype female, NSW, off Nowra, 34°59.52'S,
151°05.94'E, 204 m, coarse shell, WHOI epibenthic sled, G.C.B. Poore
et al„ RV Franklin, 14 Jul 1986 (stn SLOPE 1), NMV J53000.
New nannastacid cumaceans from Museum Victoria
157
Figure 42. Procampylaspis australiensis sp. nov. female paratype: a,
body, lateral view; b, carapace, dorsal view; c, antenna 1; d,
maxilliped 2; e, maxilliped 3. Scale (in mm): a, 0.25; b, 0.5; c, 0.1; d,
0.2; e, 0.25.
Figure 44. Procampylaspis australiensis sp. nov. male allotype:
a, body, lateral view; b, pleonite 6 and right uropod. Scale (in mm):
a, 0.5; b, 0.25.
Figure 43. Procampylaspis australiensis sp. nov. female paratype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and right uropod. Scale (in mm): a-f, 0.25.
Paratypes: 1 immature male, 1 female (stn SLOPE 45),
NMV J53 001; 1 female, dissected (stn SLOPE 1), NMV J53 002.
Diagnosis. Carapace large, with smooth integument. Maxilliped
2 with 6 dactylar teeth. Merus of pereopod 1 as long as carpus,
shorter than propodus. Pereopod 2 with short and bulky articles,
dactylus twice as long as propodus. Uropod peduncle 1.4 times
as long as pleonite 6, 1.1 times as long as its endopod, endopod
with 3 inner setae, little longer than exopod.
Description of female. Body with smooth integument. Length:
2.6 mm.
Carapace 0.46 body length, 1.6 times as long as high, with
middorsal ridge, short ocular lobe devoid of lenses,
pseudorostral lobes 3 times as long as tip of ocular lobe,
antennal notch small, ventral margin smooth.
Antenna 1 peduncle with progressively shorter and robust
articles, main flagellum 3 -articulate, longer than distal article
of peduncle, accessory flagellum minute, uniarticulate.
Maxilliped 2 basis and merus with pappose seta on inner
margin, carpus and propodus with plumose seta on inner
margin, propodus second longest article, dactylus fused with
its 6 teeth, first 3 teeth progressively shorter toward distal end,
4th tooth longer, 5th tooth half of previous one, curved, 6th
tooth the longest, 1.7 times as long as 5th one, barely curved.
Maxilliped 3 basis half of appendage, with plumose seta on
inner distal corner, 2 longer setae on outer margin, merus with
tooth and plumose seta on outer margin, carpus with 3 teeth
and plumose seta on outer margin, propodus 2nd longest
158
lorgu Petrescu
article, with 2 pappose setae on inner margin, dactylus half of
propodus, with simple long setae.
Pereopod 1 merus as long as carpus, shorter than propodus,
dactylus half of propodus, with short simple terminal setae.
Pereopod 2 with short and bulky articles, basis 3rd of
appendage length, with plumose seta on inner and outer distal
corners, merus with tooth and plumose seta on inner margin,
carpus little longer than merus, with 2 short stout setae on
distal inner corner, dactylus twice as long as propodus with
few simple setae. Pereopods 3-5 with progressively shorter
basis (as long as rest of articles combined in 3rd pair), dactylus
with simple stout terminal seta. Exopods on maxilliped 3 and
pereopods 1, 2.
Uropod peduncle 1.4 times as long as pleonite 6, 1.1 times
as long as endopod, with serrated margins, exopod shorter
than endopod with terminal long stout seta, endopod with 3
inner setae, robust terminal seta.
Description of immature male. Body length: 3 mm.
Carapace, less vaulted than in female, 2.1 times as long as
high. Exopods, 5 pairs not fully developed.
Uropod longer than in female, peduncle 2.4 times as long
as pleonite 6, strongly serrated, 1.5 times as long as endopod,
exopod 0.7 of endopod length, with stout subterminal seta and
terminal seta twice as long, endopod with 3 microserrate setae
on strongly serrated inner margin, subterminal short
microserate seta and a terminal, more robust and longer seta.
Etymology. The name “ australiensis ” refers to Australia.
Distribution. Off Nowra, NSW, off Freycinet Peninsula, Tas.;
204-800 m.
Remarks. The new species differs from other species of
Pro campy laspis in the following combination of female
characteristics: the carapace is without spines or tubercles and
maxilliped 2 has six teeth on the dactylus. The only known
species of Procampylaspis from Australian waters is P. sordida
Hale, 1945 which has a dorsal conical tubercle on the carapace
and maxilliped 2 with five dactylar teeth; the first one, more
slender than in this species, is longer than the third tooth, and the
fifth tooth is shorter.
Procampylaspis poorei sp. nov.
Figures 45, 46
Material examined. Holotype female, Antarctica, Eastern Prydz Bay,
off the Larsemann Hills, 68°54.88'S, 76°37.03'E, 667-716 m,
epibenthic sled, P.M. O’Loughlin, RSV Aurora Australis, 18 Feb 1993
(stn A A 93 158), NMV J53009.
Diagnosis. Carapace with 3 dorsal tubercles, a pair medially
and 1 in posterior half, ocular lobe eyeless, close to anterior
extremity of pseudorostrum, with spine on top. Maxilliped 2
with 1st dactylar tooth longer than 2nd tooth and shorter than
3rd one, 5th tooth twice as long as 3rd tooth. An inner tooth on
ischium of maxilliped 3, 2 teeth on outer margin of merus and
carpus. Pereopod 1 with slender articles, merus as long as
carpus. Dactylus of 2nd pereopod 2 twice as long as propodus,
with plumose setae. Uropod peduncle 1.7 times as long as
Figure 45. Procampylaspis poorei sp. nov. female holotype: a, body,
lateral view; b, detail of integument of carapace; c, carapace, dorsal
view; d, antenna 1; e, maxilliped 2. Scale (in mm): a, 0.5; c, 1; d, e,
0.2.
Figure 46. Procampylaspis poorei sp. nov. female holotype: a,
maxilliped 3; b, pereopod 1; c, pereopod 2; d, pereopod 3; e,
pereopod 4; f, pereopod 5; g, pleonite 6 and right uropod. Scale (in
mm): a-f, 0.5; g, 0.25.
New nannastacid cumaceans from Museum Victoria
159
pleonite 6 and 1.4 times as long as endopod, endopod longer
than exopod, with 3 microserrate setae on inner margin.
Description. Body with highly calcified integument. Length:
3.8 mm.
Carapace 0.46 body length, twice as long as high, with 3
dorsal tubercles on midline, a close pair medialy and 1 in
posterior half, covered with pitted integument, ocular lobe
eyeless, close to anterior extremity of pseudorostrum, with
spine on top, antennal notch small, lower margin smooth.
Antenna 1 with progressively shorter articles of peduncle,
main flagellum 3 -articulate, longer than distal article of
peduncle, accessory flagellum minute, uniarticulate.
Maxilliped 2 with pappose seta on inner distal margin of
basis and carpus, propodus second longest article, with short
plumose seta on inner margin, 1st dactyl ar tooth longer than 2nd
tooth and shorter than 3rd one, 5th tooth twice as long as 3rd
tooth. Maxilliped 3 with 2 plumose setae on inner and outer
distal corner, inner tooth on ischium, 2 teeth and plumose seta
on outer margin of merus and carpus, merus 2nd longest article,
carpus as long as propodus, dactylus 0.68 times as long as
propodus, with long simple terminal setae.
Pereopod 1 with slender articles, basis less than half as
long as appendage, merus as long as carpus, propodus twice as
long as carpus, second longest article, dactylus half of
propodus. Pereopod 2 basis shorter than rest of articles
combined, merus with tooth and simple seta on inner margin,
carpus slightly longer than merus, with 2 simple setae on inner
distal corner, dactylus twice as long as propodus, with plumose
setae. Pereopods 3-5 with progressively shorter basis (as long
as half of appendage in 3rd and 4th pair), with plumose setae
on outer margin, dactylus with long stout terminal seta.
Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 1.7 times as long as pleonite 6 and 1.4
times as long as endopod, exopod shorter than endopod, with
subterminal microserrate seta and terminal robust longer seta,
endopod with 3 microserrate setae on inner margin and robust,
longer, terminal seta.
Etymology. The species is dedicated to Gary Poore, Principal
Curator, Museum Victoria, Melbourne, Australia, specialist in
Peracarida, as a sign of gratitude.
Distribution. Eastern Prydz Bay, off the Larsemann Hills,
Antarctica; 667-716 m depth.
Remarks. The only species of Procampylaspis with fewer than
five dorsal tubercles or spines on the carapace is P. compressa
Zimmer, 1907 which has two pairs of spines versus three dorsal
median tubercles as in P. poorei. The new species also differs
from P. compressa in: ocular lobe with one median spine on
the tip instead of two as in P. compressa ; antenna 1 with a
longer main flagellum; and the first and second dactylar teeth
of maxilliped 2 are separated up to the basis (versus not
separated) and the maxilliped 3 has fewer teeth on its articles.
Procampylaspis sordida Hale, 1945
Procampylaspis sordida Hale, 1945: 215, fig. 49.
Material examined. 1 female (stn SLOPE 27), NMV J53011; 3 females
(stn SLOPE 45), NMV J53012; 7 females, 1 male (stn SLOPE 46),
NMV J53013; 2 females (stn SLOPE 46), MGAB CUM 1620.
Distribution. Off Eden, NSW and Tas.; 60-800 m depth.
Remarks. Hale recorded the species from New South Wales.
New records are from Tasmania at 800 m depth.
Procampylaspis spinifera sp. nov.
Figures 47-50
Material examined. Holotype female, Tas., off Lreycinet Peninsula,
42°00.20'S, 148°37.70'E, 720 m, coarse shelly sand, WHOI epibenthic
sled, M.L. Gomon et al., RV Franklin, 27 Jul 1986 (stn SLOPE 46),
NMV J53 003.
Allotype male: (stn SLOPE 45), NMV J53004.
Paratypes: 2 immature males (stn SLOPE 46), NMV J53 006;
1 female, dissected (stn SLOPE 45), NMV J53007; 1 male, dissected
(stn SLOPE 46), NMV J53 008; 1 female (stn SLOPE 46),
MGAB CUM 1619.
Diagnosis. Carapace with 5 pairs of dorsal spines, pair of
spines on top of ocular lobe, anterolateral margin with strong
serration. Pair of dorsal spines on pereonites 3-5 and pleonite
1. Strongly serrated basal article of antenna 1 peduncle.
Maxilliped 2 with 5 dactylar teeth, 1st one as long as 3rd, 5th
tooth twice as long as 3rd tooth. Maxilliped 3 with strong tooth
on outer margin of merus. Pereopod 1 with merus longer than
carpus but shorter than propodus. Pereopod 2 with spine on
inner margin of merus, dactylus 2.8 times as long as propodus.
Dactylus of pereopods 3-5 fused with terminal seta. Uropods
with serrated peduncle, 2.27 times as long as pleonite 6, 1.9
times as long as endopod, exopod 0.7 endopod length, endopod
with 3 microserrate setae on inner margin.
Description of female. Body without setae on integument.
Length: 4.1 mm.
Carapace 0.43 body length, 2.1 times as long as high, with
5 pairs of dorsal spines, pair of spines on top of ocular lobe,
ocular lobe long, without eyes, anterolateral margin with
strong serration. A pair of dorsal spines on pereonites 3-5 and
pleonite 1.
Antenna 1 basal article of peduncle with 3 strong teeth on
inner margin, peduncle with progressively shorter articles,
main flagellum 3 -articulate, much longer than distal article of
peduncle, accessory flagellum minute, uniarticulate.
Maxilliped 2 with pappose seta on inner margin of basis and
merus, merus to propodus with serrated outer margin, propodus
2nd longest article, with plumose seta on outer margin, dactylus
with 5 teeth, 1st one as long as 3rd, 5th tooth twice as long as 3rd
tooth. Maxilliped 3 basis almost half of appendage, with
plumose seta on inner margin and 2 longer setae on outer
margin, ischium with tooth on inner margin, strong tooth on
outer margin of merus, 3 teeth on outer margin of carpus,
propodus shorter than carpus, dactylus shorter than propodus,
dactylus with simple long terminal setae.
Pereopod 1 basis less than half as long as appendage, with
merus longer than carpus, but shorter than propodus, dactylus
with simple terminal setae. Pereopod 2 basis shorter than rest
of articles combined, with spine on inner margin of merus,
carpus 1.4 times as long as merus, with 3 stout setae on distal
160
lorgu Petrescu
Figure 47. Procampylaspis spinifera sp. nov. female paratype: a,
body, lateral view; b, body, dorsal view; c, carapace, dorsal view; d,
antenna 1; d, maxilliped 2; e, maxilliped 3. Scale (in mm): a, b, 1; c,
0.2; d, 0.25; e, 0.25.
Figure 49. Procampylaspis spinifera sp. nov. male allotype: a, body,
lateral view; b, body, dorsal view; c, maxilliped 2. Scale (in mm): a,
b, 1; c, 0.2.
Figure 48. Procampylaspis spinifera sp. nov. female paratype:
a, pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4;
e, pereopod 5; f, pleonite 6 and right uropod. Scale (in mm): a-e,
0.5; f, 0.25.
Figure 50. Procampylaspis spinifera sp. nov. male allotype: a,
maxilliped 3; b, pereopod 1; c, pereopod 2; d, pereopod 3; e, pereopod
4; f, pereopod 5; g, pleonite 6 and right uropod. Scale (in mm): a-e,
0.5; f, g, 0.25.
New nannastacid cumaceans from Museum Victoria
161
inner corner, dactylus 2.8 times as long as propodus, with
simple setae. Pereopods 3-5 with progressively shorter basis,
with plumose seta on outer margin and longer carpus (carpus
twice as long as merus in 5th pair), dactylus fused with
terminal seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod with serrated peduncle, 2.27 times as long as
pleonite 6, 1.9 times as long as endopod, exopod 0.7 endopod
length, with short subterminal stout seta and longer terminal
robust seta, endopod with 3 microserrate setae on inner margin
and more robust and longer terminal seta.
Description of male. Body length: 5.4 mm. Carapace with
median dorsal small tubercle and pair of tubercles near posterior
extremity, pair of small tubercles on top of eyeless ocular lobe
not reaching extremity of pseudorostrum, lower margin shortly
serrated in anterior half.
Maxilliped 2 with 3rd tooth of dactylus more slender than
in female. Maxilliped 3 with more serrated basis and merus on
inner margin, carpus with 2 teeth on inner margin (3 in
female).
Pereopod 1 as in female, with larger basis. Pereopod 2 with
fewer stout distal setae on carpus, dactylus with terminal plumose
seta instead of simple one in female, larger basis. Pereopod 5,
with longer carpus than in female. Exopods, 5 pairs.
Uropod peduncle with serrated margins and numerous
plumose setae on inner margin, 1.7 times as long as pleonite 6,
1.8 times as long as endopod, exopod 0.8 of endopod length,
with subterminal and terminal stout simple seta, endopod with
6 stout setae on inner margin, seta subterminal and robust
longer terminal seta.
Etymology. The Latin name ii spinifera ,, describes the presence
of numerous spines on the body and appendages.
Distribution. Off Freycinet Peninsula, Tas.; 720-800 m depth.
Remarks. P. spinifera is close to P. spinosa Petrescu, 2001
from the south-eastern Pacific Ocean (American coast), both
having numerous dorsal spines on the carapace. These number
five pairs in females of P. spinifera and 12 in females of P.
spinosa. The setae on the dactylus of pereopod 2 are simple in
P. spinifera while plumose in P. spinosa.
Procampylaspis tasmaniensis sp. nov.
Figures 51, 52
Material examined. Holotype male, dissected, Tas., off Freycinet
Peninsula, 42°02.20'S, 148°38.70'E, 800 m, coarse shelly sand, WHOI
epibenthic sled, M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn
SLOPE 45), NMV J53010.
Diagnosis. Carapace with 2 dorsal parallel rows of 4 small
tubercles on posterior extremity. Lateral small spines on pereon
and pleon. Maxilliped 2 with 1st tooth of dactylus as long as the
3rd one, 5th tooth 1.3 times as long as 3rd tooth, 1st and 2nd
teeth enlarged. Maxilliped 3 with teeth on outer margin of
merus and carpus. Ischium of pereopod 1 2nd longest article.
Pereopod 2 with dactylus 3.7 times as long as propodus. Carpus
of 5th pereopod 3.7 times as long as merus. Dactylus of
pereopods 3-5 fused with its terminal seta. Uropod peduncle
2.7 times as long as pleonite 6, 2.4 times as long as exopod.
Figure 51. Procampylaspis tasmaniensis sp. nov. male holotype: a,
body, lateral view; b, body, dorsal view; c, antenna 1; d, antenna 2; e,
maxilliped 2. Scale (in mm): a, b, 1; c, 0.25; d, 0.5; e, 0.2.
Figure 52. Procampylaspis tasmaniensis sp. nov. male holotype: a,
maxilliped 3; b, pereopod 1; c, pereopod 2; d, pereopod 3;
e, pereopod 4; f, pereopod 5; g, pleonite 6 and left uropod. Scale (in
mm): a-g, 0.5.
162
lorgu Petrescu
Description. Body with calcified integument. Length 4.6 mm.
Carapace 0.39 entire body length, twice as long as high,
with parallel row of 4 dorsal small tubercles on posterior
extremity, a ridge starting behind the pseudorostrum obliquely
going backwards medially reaching the lower margin, antennal
notch marked, ocular lobe minute, eyeless, few fine setae on
integument, smooth lower margin.
Antenna 1 short and robust, articles of peduncle
progressively shorter, main flagellum 3 -articulate, longer than
distal article of peduncle, accessory flagellum minute,
uniarticulate. Antenna 2 of with several groups of setae on
outer margins of last 2 segments of peduncle.
Maxilliped 2 with strong long pappose seta on inner distal
corner of basis and carpus, propodus second longest article, 2
plumose setae on outer margin, 1st tooth of dactylus as long as
3rd one, 5th tooth 1.3 times as long as 3rd tooth, 1st and 2nd
teeth enlarged, separated up to basis. Maxilliped 3 basis longer
than rest of articles combined, with 2 teeth on inner distal
corner, 2 long plumose setae on outer distal corner, merus
second longest article, with 3 plumose setae on inner margin, 3
teeth and plumose seta on outer margin of merus and carpus,
propodus longer than carpus, dactylus 0.7 of propodus
Pereopod 1 basis less than half as long as appendage,
ischium 2nd longest article, merus as long as propodus, carpus
shorter than propodus, longer than dactylus, dactylus with
stout terminal long seta. Pereopod 2 basis shorter than rest of
articles combined, plumose seta on inner distal corner of basis
and merus, carpus 1.4 times as long as merus, 2 stout setae on
distal inner corner of carpus, dactylus 3.7 times as long as
propodus, with long terminal plumose seta. Pereopods 3 and 4
with progressively shorter basis, longer than rest of articles
combined, dactylus fused with terminal seta. Pereopod 5 with
shorter basis and longer carpus, 3.7 times as long as merus,
dactylus fused with terminal seta. Exopods on maxilliped 3
and pereopods 1-4.
Uropod peduncle 2.7 times as long as pleonite 6, with
plumose setae on inner margin, 2.4 times as long as exopod,
endopod broken off.
Etymology. The species bears the name of the collecting area
- Tasmania.
Distribution. Off Freycinet Peninsula, Tas.; 800 m depth.
Remarks. P. bonnieri Caiman, 1906 from the Mediterranean
and eastern Pacific (Petrescu, 2000) share with P. tasmaniensis
a carapace with spines on the posterior extremity and a minute
ocular lobe. The new species has fewer tubercles/spines on the
posterior extremity of the carapace, no row of spines near the
lower margin of the carapace, dactylar teeth of a different
shape, maxilliped 3 without strong teeth on the inner margin of
the merus, pereopod 1 with its ischium longer than the
propodus, and longer uropods.
Scherocumella Watling, 1991
Scherocumella Watling, 1991b: 754.— Stoddart and Lowry,
2003: 373-418.
Type species. Nannastacus longirostris Sars, 1879.
Remarks. The carapace is as in the genus Nannastacus and the
lenses grouped in two lateral pairs as always in Nannastacus.
The pseudorostral lobes meet each other in front of the eyelobe,
from base to top, similar to the genus Cumella and the uropodal
peduncle is as long or even longer than pleonite 6, as in Cumella.
Five species and one subspecies are known from Australia.
Key to species of Scherocumella from Australian waters
1. Peudorostral lobes long 2
— Peudorostral lobes short 3
2. Carapace with a dorsal hump behind frontal lobe
S. nasuta camelus Zimmer, 1914
— Carapace with no dorsal hump
S. nasuta nasuta Zimmer, 1914
3. Pereon and pleon with dorsal tubercles 4
— Pereon and pleon without tubercles 5
4. Carapace with serrated dorsal and lateral rows
S. vieta Hale, 1949
— Carapace without serrated rows
S. clavata Hale, 1945
5. Uropod peduncle longer than pleotelson
S. sheardi Hale, 1945
— Uropod peduncle shorter than pleotelson
S nichollsi Hale. 1949
Scherocumella nichollsi Hale, 1949
Nannastacus nichollsi Hale, 1949: 227, figs 1, 2.— Bacescu, 1992:
240.
Scherocumella nichollsi.— Watling, 1991b: 754.— Stoddart and
Lowry, 2003: 417.
Material examined. 1 female (stn MSL-EG 77), NMV J26637; 1 male
(stn MSL-EG 78), NMV J26638; 1 female (stn MSL-EG 104),
NMV J26639.
Distribution. Garden I., WA and Vic.; 6-25 m depth.
Remarks. Previously known from the type locality, the species
is now recorded from eastern Victoria, at 25 m depth.
Scherocumella sheardi Hale, 1945
Nannastacus sheardi Hale, 1945 : 156, figs 8, 9.— Bacescu,
1992: 243.
Scherocumella sheardi.— Watling, 1991b: 754.— Stoddart and
Lowry, 2003: 417.
Material examined. 3 females (stn MSL-EG 105), NMV J26640.
Distribution. Gulf St. Vincent, SA and eastern Vic.; surface to
4 m depth.
Remarks. The species was described by Hale from specimens
collected at Brighton, Gulf St Vincent, SA. The species has now
been discovered in eastern Bass Strait, Victoria from 27 m depth.
Schizotrema Caiman, 1911
Schizotrema Caiman, 1911: 341, 360-361.— Watling, 1991b:
755.— Bacescu, 1992: 259.— Stoddart and Lowry, 2003: 418.
Type species. Schizotrema depressum Caiman, 1911.
New nannastacid cumaceans from Museum Victoria
163
Remarks. In Schizotrema, the carapace is high as inNannastacus,
higher than in Cumella; the eyes are always as in Nannastacus,
but the pseudorostral lobes are completely separated from base
to top, not like in Nannastacus, Scherocumella or Cumella.
The uropodal peduncle is as long as or shorter than pleonite 6,
as in Nannastacus.
Key to species of Schizotrema from Australian waters
1. Exopod of uropod at least 0.3 times as long as endopodL^..
2
— Exopod of uropod at least 0.25 times as long as endopod
3
2. Last pereonite and 1st pleonite with a pair of strong dorsal
spines S. aculeatum Hale, 1936
— Last pereonite and 1st pleonite without spines
S. nudum Tafe and Greenwood, 1996
3. Carapace with dorsal and lateral spines
S. resimum Hale. 1949
— Carapace without spines S. leopardinum Hale, 1949
Schizotrema aculeatum Hale, 1936
Schizotrema bifrons aculeata Hale, 1936: 430, fig. 18.
Schizotrema aculeta.— Hale, 1945: 168, fig. 16.
Schizotrema aculeatum.— Lowry and Stoddard, 2003: 418.
Material examined. 1 female (stn SLOPE 2), NMV J45780.
Distribution. NSW, Gulf St Vincent, SA and WA: 2-503 m
depth.
Remarks. The species’ range is extended to New South Wales
and to greater depths.
Styloptocuma Bacescu and Muradian, 1974
Styloptocuma Bacescu and Muradian, 1974: 74.— Bacescu, 1992:
262.— Petrescu and Watling, 1999: 306.
Americuma Watling, 1991a: 580-581.
Cumella (Styloptocuma).— Waiting, 1991b: 752.
Type species. Styloptocuma antipai Bacescu and Muradian, 1974.
Remarks. Styloptocuma is characterised by pseudorostral lobes
reaching the extremity of the ocular lobe, the ocular lobe
without visual elements, and uropods with long peduncles,
twice or even longer than pleonite 6. The genus has not
previously been recorded from Australia.
Key to species of Styloptocuma from Australian waters
(females only)
1. Carapace dorsally and ventrally serrated 2
— Carapace without serration 4
2. Carapace only with ventral serration
S. granulosum sp. nov.
— Carapace with dorsal and ventral serration 3
3. Carapace with a lateral serrated row
S. spinosum sp. nov.
— Carapace without lateral serrated row
S. poorei sp. nov.
4. Dactylus of pereopod 2 twice as long as propodus
S. australiense sp. nov.
— Dactylus of pereopod 2 quarter length of propodus
S. nodosum sp. nov.
Styloptocuma australiense sp. nov.
Figures 53, 54
Material examined. Holotype female, NSW, off Nowra, 34°51.90'S,
151°12.60'E, 770 m, crinoid dominated, WHOI epibenthic sled, G.C.B.
Poore and C.C. Lu, RV Franklin, 15 Jul 1986 (stn SLOPE 6), NMV
J53 062.
Paratype: 1 female (stn SLOPE 6), NMV J53063.
Diagnosis. Carapace short, with long pseudorostrum, lower
margin smooth. Integument with few setae on carapace, smooth
on rest of body. Pereopod 1 with carpus as long as propodus.
Dactylus of pereopod 2 twice as long as propodus. Uropod
peduncle 2.8 times as long as pleonite 6, with serrated inner
margin 1.3 times as long as endopod, exopod shorter than
endopod, endopod with 4 setae on inner margin and robust curved
terminal seta.
Description. Body elongate, integument carapace covered with
sparse setae. Length: 3.2 mm.
Carapace shorter than one -third body length, 1.7 times as
long as high, long pseudorostrum (0.4 of entire carapace
length), antennal notch absent, lower margin smooth, ocular
lobe eyeless, reaching extremity of pseudorostral lobes, conical
in dorsal view. Pleon 0.55 body length.
Antenna 1 peduncle with progressively shorter articles,
median article with tubercle with 2 sensory setae, main
flagellum 3-articulate, longer than distal article of peduncle,
median and distal article with 1 aesthetasc, accessory flagellum
minute, uniarticulate.
Maxilliped 3 basis shorter than rest of articles combined,
with 2 plumose setae on inner margin and 4 (2 long, 2 short)
plumose setae on outer distal corner, short ischium, merus
shorter than carpus, merus and carpus with long plumose seta
on outer margin, propodus 2nd longest article, with 2 pappose
setae on inner margin, dactylus longer than merus, with 2 stout
curved terminal setae.
Pereopod 1 basis less than half as long as appendage, ischium
shorter than merus, carpus longer than merus, as long as
propodus, dactylus 0.4 propodus length, with long simple stout
terminal seta. Pereopod 2 less than half as long as appendage,
merus with seta on inner margin, carpus almost twice as long as
merus, with 1 seta on inner margin and 2 on outer margin, 2
stout setae on distal inner corner, dactylus twice as long as
propodus, with simple setae. Pereopods 3-5 with progressively
shorter basis and longer carpus (twice as long as merus in 5th
pair), propodus longer than merus, dactylus with a long stout
terminal seta. Exopods on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 2.8 times as long as pleonite 6, with
serrated inner margin, 1.3 times as long as endopod, exopod
shorter than endopod, with a stout terminal seta, endopod with
4 setae on inner margin and robust curved terminal seta.
Etymology. The species bears the name of type locality -
Australia.
164
lorgu Petrescu
Figure 53. Styloptocuma australiense sp. nov. female holotype: a,
body, lateral view; b, carapace , dorsal view; c, antenna 1; d,
maxilliped 3. Scale (in mm): a, b, 0.5; c, 0.1; d, 0.2.
Figure 54. Styloptocuma australiense sp. nov. female holotype:
a, pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4;
e, pereopod 5; f, pleonite 6 and left uropod. Scale (in mm): a-e,
0.25; f, 0.5.
Distribution. Off Nowra, NSW; 770 m depth.
Remarks. This is the first record of the genus Styloptocuma
from Australian waters. It was previously known from the deep
Atlantic (Bacescu, 1992) and Eastern Pacific (Petrescu, 1991;
Petrescu and Watling, 1999). Styloptocuma australiense is
morphologically similar to S. angustatum Jones, 1984 from the
Western Atlantic (Brazil): both have a carapace without spines
or tubercles, a similar anterolateral margin and almost similar
pereopods 1 and 2. The new species differs in having a longer
pseudorostrum, shorter carapace, and uropod endopod with
four setae on the inner margin instead of three.
Styloptocuma granulosum sp. nov.
Figures 55, 56
Material examined. Holotype female. Vic., S of Point Hicks, 38°25'S,
149°00'E, 1500 m, compacted clay, WHOI epibenthic sled, G.C.B.
Poore et al., RV Franklin, 22 Jul 1986 (stn SLOPE 27),
NMV J53067.
Diagnosis. Carapace with short pseudorostrum, antennal notch
large, lower margin serrated. Pleon with spines and setae.
Antenna 1 with basal article of peduncle longer than rest of
articles combined. Merus of maxilliped 3 with tooth on outer
margin. Carpus of pereopod 1 longer than propodus, propodus
twice as long as dactylus. Carpus of pereopod 2 twice as long
as merus, dactylus 3 times as long as propodus. Carpus of
pereopod 5 more than 3 times as long as merus. Uropod
peduncle 2.1 times as long as pleonite 6, 0.8 endopod length,
exopod shorter than endopod, endopod with 6 microeserrate
setae on inner margin.
Description. Body with hairy granulous integument. Length:
4.08 mm.
Carapace more than 3rd body length, more elevated on
posterior extremity, 1.9 times as long as high, ocular lobe
bulky, eyeless, reaching extremity of pseudorostrum, short
siphon, antennal notch large, anterolateral corner acute, lower
margin strongly serrated, numerous setae.
Antenna 1 proximal article of peduncle 1.65 times length
of rest of articles combined, median article without an evident
tubercle, main flagellum 3 -articulate, twice as long as distal
article of peduncle, accessory flagellum minute, uniarticulate.
Maxilliped 3 basis half of appendage, with 2 plumose
setae on inner distal corner and 2 much longer on distal outer
corner, merus and carpus with tooth and plumose seta on outer
margin, carpus longer than merus, propodus 2nd longest
article, dactylus half of propodus, with strong stout curved
terminal seta.
Pereopod 1 basis less than half as long as appendage,
carpus 2.1 times as long as merus, twice as long as propodus,
propodus twice as long as dactylus, dactylus with long robust
terminal seta, numerous simple setae especially on propodus
and dactylus. Pereopod 2 basis less than half as long as
appendage, carpus twice as long as merus, with stout seta on
inner distal corner, dactylus 3 times as long as propodus, with
long simple stout terminal seta. Pereopods 3-5 with
progressively shorter basis and longer carpus (carpus more
than 3 times as long as merus in last pair), dactylus fused with
New nannastacid cumaceans from Museum Victoria
165
Figure 55. Styloptocuma granulosum sp. nov. female holotype: a,
body, lateral view; b, antenna 1; c, maxilliped 3; d, pereopod 1.
Scale (in mm): a, 0.5; b, 0.2; c, d, 0.5.
Figure 56. Styloptocuma granulosum sp. nov. female holotype: a,
pereopod 2; b, pereopod 3; c, pereopod 4; d, pereopod 5; e, pleonite
6 and left uropod. Scale (in mm): a-d, 0.5; e, 0.25.
its stout terminal seta, with inner spine (see detail). Exopods
on maxilliped 3 and pereopods 1, 2.
Uropod peduncle 2.1 times as long as pleonite 6, with fine
setae on both sides, 0.8 endopod length, exopod slightly shorter
than endopod, with robust terminal seta (broken), endopod
with 6 microserrate setae on inner margin and terminal robust
short simple seta.
Etymology. In Latin “ granulosum ” means granular and
describes the integument.
Distribution. S of Point Hicks, Vic.; 1500 m depth.
Remarks. Styloptocuma granulosum differs mainly from other
species of Styloptocuma described in this paper in a longer
carapace with a smooth upper margin and serrated lower margin.
Styloptocuma cristatum Jones, 1984 from the Gulf of Biscay has
a longer carapace but with dorsal spines on the carapace, lateral
spines on the pleon and uropods with shorter rami.
Styloptocuma nodosum sp. nov.
Figures 57-60
Material examined. Holotype female, Vic., S of Point Hicks, 38°25'S,
149°00'E, 1500 m, compacted clay, WHOI epibenthic sled, G.C.B.
Poore et al„ RV Franklin , 22 Jul 1986 (stn SLOPE 27),
NMV J53064.
Allotype: male (stn SLOPE 27), NMV J53 065.
Paratypes: 3 females, 1 immature male, 7 mancas, 1 adult male,
dissected (stn SLOPE 27), NMV J53066.
Diagnosis. Carapace with short pseudorostrum, without antennal
notch, slightly serrated lower margin. Antenna 1 with tooth on
distal article of peduncle. Carpus and propodus of pereopod 1
subequal. Carpus of pereopod 2 3 times as long as merus,
dactylus with subterminal long seta. Uropod peduncle 2.9 times
as long as pleonite 6 and 1.3 times as long as its equal rami.
Description of female. Body with few hairs on carapace.
Length: 4.7 mm.
Carapace third of body length, 1.6 times as long as high,
with short upturned pseudorostrum (0.3 of carapace length),
with knotty surface and few long setae, antennal notch absent,
lower margin slightly serrated, ocular lobe eyeless. Pleon more
than half body length.
Antenna 1 slender, basal article of peduncle much longer
than rest of articles combined, median article with minute
tubercle, distal one with strong tooth, main flagellum shorter
than last article of peduncle, accessory flagellum tiny.
Maxilliped 2 with pappose long seta on inner distal corner
of basis and merus, carpus 2nd longest article, with pappose
setae on inner margin, dactylus half length of propodus, with
stout terminal seta. Maxilliped 3 basis longer than rest of
articles combined, with 4 plumose setae along distal margin (2
much longer), merus as long as carpus, with plumose seta on
outer margin of both, propodus 2nd longest article, dactylus
little less than half as long as propodus, with longer stout
terminal setae.
Pereopod 1 basis less than half as long as appendage, merus
more than twice length of ischium, carpus 1.8 times as long as
merus, as long as propodus, twice as long as dactylus, dactylus
166
lorgu Petrescu
Figure 57. Styloptocuma nodosum sp. nov. female holotype: a, body,
lateral view; b, antenna 1; c, maxilliped 2; d, maxilliped 3. Scale (in
mm): a, 0.5; b, d, 0.5; c, 0.2.
Figure 59. Styloptocuma nodosum sp. nov. male allotype: a, body,
lateral view; b, antenna 1; c, antenna 2; d, maxilliped 2. Scale (in
mm): a, 1; b, 0.25; c, 0.5; d, 0.2.
Figure 58. Styloptocuma nodosum sp. nov. female holotype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and left uropod. Scale (in mm): a-f, 0.5.
Figure 60. Styloptocuma nodosum sp. nov. male allotype: a,
maxilliped 3; b, pereopod 1; c, pereopod 2; d, pereopod 3; e, pereopod
4; f, pereopod 5; g, pleonite 6 and left uropod. Scale (in mm): a, c-g,
0.5; b, 0.5.
New nannastacid cumaceans from Museum Victoria
167
with terminal stout curved seta. Pereopod 2 basis 3rd of appendage
length, fine seta on inner margin of merus, carpus 3 times as long
as merus, with 2 stout setae on inner distal comer, dactylus one-
fourth as long as propodus, with subterminal long plumose seta
(less than half of dactylus). Pereopod 3 basis longer than half of
entire pereopod, carpus 3 times as long as merus, propodus little
longer than merus, with distal stout seta, dactylus with terminal
stout seta. Pereopod 4 with shorter basis and carpus longer than in
previous pair. Pereopod 5 much shorter than previous pairs, basis
half of appendage, terminal seta missing (limb is regenerated or
individual is first molt after manca 2). Exopods on maxilliped 3
and pereopods 1, 2; with 2nd article very long.
Uropod peduncle 2.9 times as long as pleonite 6 and 1.3
times as long as its equal rami, fine setae on both sides, exopod
with terminal stout seta (one-third of exopod length), endopod
with 2 pedunculate setae, 2 slender simple and 2 stout setae on
inner margin and longer terminal seta (as in exopod).
Description of male. Body with knotty surface only on pleonite
4. Length: 5.1 mm. Carapace, pseudorostrum with long setae,
large lateral swelling on branchial region, more pronounced
anterolateral corner, smooth lower margin.
Antenna 1 with similar peduncle, main flagellum longer
than in female. Antenna 2 (fig. 59c) with ranks of setae on
articles 2-5 of peduncle.
Maxilliped 2 with larger articles, more setulose. Maxilliped
3 with longer basis.
Pereopod 1 as in female, carpus and propodus equal, longer
than in female (3 times as long as merus), with shorter dactylus.
Pereopod 2 as in female, subterminal seta of dactylus simple, not
plumose. Pereopods 3 and 4 with large basis, longer than rest of
appendage in 3rd pair, shorter in 4th, carpus 3 times as long as
merus in 3rd pair, 4.5 times longer in 4th pair, long stout terminal
seta. Pereopod 5 with shorter and more slender basis than in
previous pairs, carpus 5 times as long as merus, propodus 0.4 of
carpus. Exopods, 5 pairs, stronger than in female.
Uropod peduncle with straight margins, 2.7 times as long
as pleonite 6, 1.4 times as long as endopod, exopod little
shorter than endopod, endopod with 5 setae on inner margin.
Etymology. The name “ nodosum ” means “knotty” in Latin and
describes the body surface.
Distribution. S of Point Hicks, Vic.; 1500 m depth.
Remarks. Styloptocuma nodosum differs from other species in
this paper in its knotty integument, a carapace without spines,
its slightly serrated lower margin of the carapace in females,
being without an antennal notch and with a more upturned
pseudorostrum. It resembles S. extans Jones, 1984 from the
deep Atlantic but has a shorter pseudorostrum, the integument
not knotty, and a spiny pleon.
Styloptocuma poorei sp. nov.
Figure 61
Material examined. Holotype female, Tas., off Freycinet Peninsula,
42°00.20'S, 148°37.70'E, 720 m, coarse shelly sand, WHOI epibenthic
sled, M.F. Gomon et al., RV Franklin , 27 Jul 1986 (stn SLOPE 46),
NMV J53068.
Figure 61. Styloptocuma poorei sp. nov. female holotype: a, body,
lateral view; b, antenna 1; c, maxilliped 3; d, pereopod 1; e, pereopod
2; f, pereopod 3; g, pereopod 4; h, pereopod 5; i, pleonite 6 and left
uropod. Scale (in mm): a, 0.5; b, 0.2; c, 0.1; d-i, 0.25.
Diagnosis. Carapace with short, little upturned pseudorostrum,
with several dorsal spines and strongly serrated lower margin.
Short serration on pleonite 5. Propodus of maxilliped 3 as long
as ischium, merus and carpus combined. Carpus of pereopod
1 longer than propodus. Dactylus of pereopod 2 3.8 times as
long as propodus. Uropod peduncle with serrated inner margin,
2.9 times as long as pleonite 6 and 1.3 times as long as endopod,
endopod twice as long as exopod, with 1 seta on inner serrated
margin.
Description. Body with weakly calcified smooth integument.
Length: 2.8 mm.
Carapace 0.31 body length, 1.4 times as long as high, short
pseudorostrum (0.18 carapace length), 7 dorsal median spines,
1 near posterior extremity, first 2 on frontal lobe, antennal
notch not marked, lower margin strongly serrated.
Antenna 1 basal article of peduncle little longer than other
2 articles combined, median article with small tubercle, main
flagellum longer than last article of peduncle, accessory
flagellum minute, uniarticulate.
Maxilliped 3 basis less than half as long as appendage, 2
long plumose setae on outer distal corner, a tooth and a plumose
seta on outer margin of merus and carpus, propodus 2nd longest
article, as long as ischium, merus and carpus combined, dactylus
with stout terminal curved seta, longer than dactylus.
Pereopod 1 basis less than half as long as appendage, carpus
longer than propodus. Pereopod 2 basis less than half as long as
appendage, carpus twice as long as merus, with 2 stout spines on
168
lorgu Petrescu
distal inner comer of carpus, dactylus 3 times as long as propodus,
with a stout terminal seta longer than dactylus. Pereopods 3-5
with progressively shorter basis and longer carpus (3.4 times
longer carpus than merus in 5th pair), dactylus fused with its
terminal seta. Exopods on maxilliped 3 and pereopods 1 , 2.
Uropod peduncle with a serrated inner margin, 2.9 times
as long as pleonite 6 and 1.3 times as long as endopod, endopod
twice as long as exopod, with 1 subterminal seta on inner
serrated margin.
Etymology. The species is dedicated to Gary Poore, Principal
Curator, Museum Victoria, Melbourne, Australia, well known
specialist in Peracarida, as a sign of gratitude.
Distribution. Off Freycinet Peninsula, Tas.; 720 m depth.
Remarks. The new species differs from other species of
Styloptocuma in this collection in the carapace with a few
dorsal spines, a strongly serrated entire lower margin, lacking
a lateral row of spines and a relatively smooth pereon and
pleon. Such a combination of characters may also be found in
S. subductum Jones, 1984 from the Atlantic but that species has
the lower margin of its carapace serrated only on the anterior
half and the uropod endopod has four setae on the inner margin
instead of one as in Styloptocuma poorei.
Styloptocuma spinosum sp. nov.
Figures 62, 63
Material examined. Holotype female, NSW, off Nowra, 34°57.90'S,
151°08'E, 503 m, bryozoa and shell, WHOI epibenthic sled, G.C.B.
Poore et al„ RV Franklin, 14 Jul 1986 (stn SLOPE 2), NMV J53018.
Paratype: 1 female (stn SLOPE 2), NMV J53019.
Diagnosis. Carapace with middorsal row of spines, another
lateral and lower margin densely serrated. Dorsal serrated crest
on pleonites 1-5. Pereopod 1 with propodus 2nd longest article.
Pereopod 2 with dactylus 3 times as long as propodus. Last 3
pairs of pereopods slender. Uropod peduncle serrated on inner
margin, twice as long as pleonite 6 and 1.5 times as long as its
equal rami.
Description. Body elongate, with weakly calcified integument.
Length: 2.36 mm.
Carapace 3rd of body length, 1.7 times as long as high,
pseudorostrum quarter of carapace length, slightly upturned,
siphon longer than carapace, a middorsal row of spines, 1
bigger spine on ocular lobe, ocular lobe eyeless, reaching
extremity of pseudorostral lobes, a median row of spines near
lower serrated margin, scattered setae. Pereon glabrous. Pleon
elongate, 0.58 of body length, with dorsal serrated crest on
pleonites 1-5, without setae.
Antenna 1 basal article of peduncle 2.5 times as long as
rest of articles combined, median article longer than distal
one, without visible tubercle, main flagellum longer than last
article of peduncle, accessory flagellum minute.
Maxilliped 3 basis half of appendage, with 2 plumose
setae on outer margin, merus and carpus with plumose seta on
outer distal corner, propodus 2nd longest article, twice as long
as dactylus.
Figure 62. Styloptocuma spinosum sp. nov. female holotype: a,
body, lateral view; b, carapace, lateral view, magnified; c, antenna 1;
d, maxilliped 3. Scale (in mm): a, 0.5; b, c, 0.5; d, 0.25.
Figure 63. Styloptocuma spinosum sp. nov. female holotype: a,
pereopod 1; b, pereopod 2; c, pereopod 3; d, pereopod 4; e, pereopod
5; f, pleonite 6 and right uropod. Scale (in mm): a-e, 0.25; f, 0.5.
New nannastacid cumaceans from Museum Victoria
169
Pereopod 1 basis shorter than rest of articles combined,
ischium with small tooth on inner margin, merus twice as long
as ischium, carpus 4 times merus and dactylus combined,
propodus 2nd longest article, twice as long as carpus, dactylus
with slender terminal long seta. Pereopod 2 basis one-third of
appendage length, simple seta on distal inner corner of basis
and merus, carpus twice as long as merus, with 2 unequal stout
setae on inner distal corner, dactylus 3 times as long as
propodus, with simple setae. Pereopods 3-5 slender, basis
longer than rest of articles combined in 3rd pair, progressively
shorter in pairs 4 and 5, carpus progressively longer in pairs
3-5, carpus 5 times as long as merus in 5th pair. Exopods on
maxilliped 3 and pereopods 1, 2.
Uropod peduncle serrated on inner margin, twice as long
as pleonite 6 and 1.5 times as long as its equal rami, exopod
with few fine setae on outer margin and a stout terminal seta,
endopod with 3 setae on inner margin and a terminal seta
longer and more robust than in exopod.
Etymology. The name of the species reflects the numerous
spines of carapace.
Distribution. Off Nowra, NSW; 503 m depth.
Remarks. Some features of the carapace, about 20 dorsal spines,
a lateral row of spines near the lower densely serrated margin
combined with dorsal serration of the pleon, make Styloptocuma
spinosum unique within Australian species of the genus. Such
carapace armature may also be found in S. antipai Bacescu and
Muradian, 1974 from the deep western Atlantic and S. echinatum
Jones, 1984 from the eastern Atlantic. The new species differ in
the glabrous pereon, absence of lateral rows of spinules on the
pleon and the shorter carpus of pereopod 2.
Styloptocumoides gen. nov.
Type species. Styloptocumoides australiensis sp. nov., here
designated.
Diagnosis. Carapace with short pseudorostrum, eyeless ocular
lobe reaches its extremity. 5 free pedigerous somites. No trace
of exopods in maxilliped 3 or pereopods in females.
Gender. Masculine.
Etymology. The new genus looks like Styloptocuma Bacescu
and Muradian, 1974.
Remarks. The closest genus to Styloptocumoides is
Styloptocuma Bacescu and Muradian, 1974, especially
regarding the carapace, with the ocular lobe reaching the tip of
the pseudorostrum. The main difference is the total absence of
exopods in the females of Styloptocumoides. A second
difference is five instead of four free thoracic segments.
Styloptocumoides australiensis sp. nov.
Figure 64
Material examined. Holotype female, NSW, off Nowra, 34°51.90'S,
151°12.60'E, 770 m, crinoid dominated, WHOI epibenthic sled, G.C.B.
Poore and C.C. Lu, RV Franklin, 15 Jul 1986 (stn SLOPE 6), NMV
J53060.
Figure 64. Styloptocumoides australiensis sp. nov. female holotype:
a, body, lateral view; b, antenna 1; c, maxilliped 3; d, pereopod 1; e,
pereopod 2; f, pereopod 3; g, pereopod 4; h, pereopod 5;
I, pleonite 6 and right uropod. Scale (in mm): a, 0.5; b-e, i, 0.2;
f-h, 0.25.
Diagnosis. Pseudorostrum 0.38 of carapace length, ocular lobe
eyeless, reaching its tip, few setae on carapace, no spines on
entire body, smooth lower margin of carapace. Basal article of
peduncle of antenna 1 shorter than rest of articles combined,
short main flagellum. Maxilliped 3 with a strong tooth on outer
margin of merus. Pereopod 1 with carpus 2nd longest article,
dactylus with long curved robust terminal seta. Pereopod 2
with carpus little longer than merus, with 2 robust short setae
on its distal inner corner, short dactylus. Pereopods 3-5
progressively shorter, with dactylus fused with its terminal
seta. No exopods. Short and robust uropods, peduncle twice as
long as pleonite 6 and its endopod, endopod slightly longer
than exopod, with 6 setae on inner margin.
Description. Body with weakly calcified integument. Length:
2.6 mm.
Carapace one-third body length, twice as long as high,
with pseudorostrum 0.38 of carapace length, ocular lobe
reaches its extremity, with some dorsal undulation viewed
from lateral side, with few setae, antennal notch marked, lower
margin smooth, siphon long.
Antenna 1 basal article of peduncle shorter than rest of
articles combined, main flagellum short, 3 -articulate, as long
as distal article of peduncle, with short aesthetascs, accessory
flagellum minute, uniarticulate.
Maxilliped 3 basis with 2 plumose setae on inner distal
corner, 3 on outer one (2 much longer), merus with strong
tooth and plumose seta on outer margin, carpus shorter than
170
lorgu Petrescu
merus, with plumose seta on outer margin, propodus slightly
longer than merus, with 2 pappose setae on inner margin, short
dactylus with stout long terminal seta.
Pereopod 1 basis less than half as long as appendage, carpus
2nd longest article, dactylus little less than half as long as
propodus, with terminal long curved robust seta. Pereopod 2
basis less than half as long as appendage, simple seta on inner
margin of merus, carpus little longer than merus, with 2 short
robust setae on inner distal corner, dactylus 1.6 times as long as
propodus, with long robust terminal seta. Pereopods 3-5 with
progressively shorter basis and longer carpus (3 times as long
as merus in last pair), dactylus fused with its stout terminal
seta. Exopods absent in maxilliped 3 and pereopods 1,2.
Uropod short and robust, peduncle twice as long as pleonite
6 and its endopod, exopod slightly shorter than endopod, with
robust terminal seta, endopod with 6 setae on inner margin
and terminal robust seta shorter than in exopod.
Etymology. The species bears the name of the collecting place
- Australia.
Distribution. Off Nowra, NSW; 770 m depth.
Remarks. The female of Styloptocumoides australiensis has
pereopods as in Styloptocuma poorei but maxilliped 3 has a
shorter propodus in S. australiensis-, maxilliped 3 and
pereopods 1 and 2 are without exopods, and obviously shorter
and larger uropods.
Vemacumella Petrescu, 2001
Vemacumella Petrescu, 2001: 1675-1677.
Figure 65. Vemacumella bacescui sp. nov. female holotype: a, body,
lateral view; b, body, dorsal view; c, antenna 1; d, maxilliped 3; e,
pereopod 1. Scale (in mm): a, 0.5; b, 0.5; c, d, 0.1; e, 0.2.
Type species. Vemacumella heardi Petrescu, 2001.
Remarks. The genus has the general aspect of Cumella but with
a higher carapace, eyeless ocular lobe like in Styloptocuma,
pseudorostral lobes meeting in front of ocular lobe like in
Cumella, not as in Styloptocuma-, median article of antennal
peduncle with a tubercle as in Styloptocuma but absent in
Cumella.
Vemacumella bacescui sp. nov.
Figures 65, 66
Material examined. Holotype female, Tas., off Freycinet Peninsula,
41°57.50'S, 148°37.90'E, 400 m, coarse shell, WHOI epibenthic sled,
M.F. Gomon et al., RV Franklin, 27 Jul 1986 (stn SLOPE 48),
NMV J53 069.
Paratype: 1 female (stn SLOPE 6), NMV J53070.
Diagnosis. Carapace with middorsal row of spines, anterolateral
corner strongly serrated, ocular lobe eyeless. Short and robust
antenna 1, median article of peduncle with small tubercle.
Basis and merus of maxilliped 3 with strong inner serration.
Carpus of pereopod 1 2nd longest article, as long as propodus
and dactylus combined. Carpus of pereopod 2 with long stout
seta on inner distal corner, short dactylus. Uropod peduncle
1.7 times as long as pleonite 6 and 1.3 times as long as endopod,
endopod longer than exopod, with 4 stout setae on inner
serrated margin.
Figure 66. Vemacumella bacescui sp. nov. female holotype: a,
pereopod 2 ; b, pereopod 3 ; c, pereopod 4 ; d, pereopod 5 ; e,
pleonite 6 and left uropod. Scale bar : a-e, 0.2
New nannastacid cumaceans from Museum Victoria
171
Description. Body strongly calcified, integument smooth.
Length: 2.4 mm.
Carapace 0.33 body length, 1.6 times as long as high, with
a median dorsal row of spines reaching frontal lobe, short
pseudorostrum meeting in front of ocular lobe, ocular lobe
eyeless, antennal notch marked, anterolateral corner strongly
serrated, lower margin smooth.
Antenna 1 proximal article of peduncle the longest, median
one the shortest, with tubercle provided with pedunculate seta,
main flagellum slightly shorter than distal article of peduncle,
3 -articulate, accessory flagellum 2-articulate, almost reaching
the extremity of basal article of main flagellum.
Maxilliped 3 basis less than half as long as appendage,
with 3 strong teeth on inner margin, 2 long plumose setae on
outer distal corner, without outer process, merus 2nd longest
article, with 2 strong teeth on inner margin, long plumose seta
on outer margin of merus and carpus, propodus longer than
carpus, with 2 pappose setae on inner margin, dactylus 0.5
propodus length, with 2 stout terminal setae.
Pereopod 1 basis shorter than rest of articles combined,
with strong serration on inner margin, tooth on inner margin
of ischium, merus longer than ischium, carpus 2nd longest
article, as long as propodus and dactylus combined, dactylus
with stout slender terminal seta. Pereopod 2 basis shorter than
rest of articles combined, merus with simple seta on inner
margin, carpus 1.7 times as long as merus, with long stout seta
on inner distal corner, dactylus 1.75 times as long as propodus,
with numerous short simple setae on both margins, long stout
simple terminal seta. Pereopods 3-5 with progressively shorter
basis (longer than half of appendage in 3rd pair) and longer
carpus (twice as long as merus in 5th pair), dactylus fused with
its terminal stout seta. Exopods on maxilliped 3 and pereopods
1 , 2 .
Uropod peduncle 1.7 times as long as pleonite 6 and 1.3
times as long as endopod, exopod 0.9 times length of endopod,
with long robust terminal seta, endopod with 4 stout setae on
inner serrated margin and robust terminal seta.
Etymology. The species is dedicated to the memory of the late
Mihai C. Bacescu (1908-1999), member of the Romanian
Academy, former director of “Grigore Antipa” National
Museum of Natural History from Bucharest, one of the highest
world authorities on Crustacea Peracarida, as a sign of homage
and gratitude for all he offered to his last student.
Distribution. Off Nowra, NSW, off Freycinet Peninsula, Tas.;
400-770 m depth.
Remarks. This is the first record of the genus in the western
Pacific (Australia) and second description of a species of
Vemacumella Petrescu. Vemacumella bacescui has in common
with Vemacumella heardi Petrescu, 2001 (described from the
south-eastern Pacific) a carapace longer than one -third of the
body length, five free pereonites, antenna 1 with a tubercle on
the median article of the peduncle, accessory flagellum 2-
articulate, and relatively short uropods with serrated margins
and unequal rami. Vemacumella bacescui differs in having
more numerous dorsal spines on the carapace and the basis of
maxilliped 3 without an outer process. Other morphological
characters are sexually dimorphic, the Australian species being
described from a female while the American one is from a
male.
Acknowledgements
I am more than grateful to Dr Gary Poore, Principal Curator
at Museum Victoria, Melbourne, Australia, for offering me the
possibility to work in his lab, the hospitality of his home, and
for the material lent for study; and to Lynsey Poore, his wife,
for the kindness and warmness she generously gave during my
staying in Melbourne. I extent my posthumous gratitude to my
mentor, the late Prof. Mihai Bacescu, for all he transmitted to
his last student in Crustacea.
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Cumiden, Vaunthompsoniiden, Nannastaciden und Lampropiden.
Zoologischer Anzeiger 31: 367-371.
Zimmer, C. 1914. Cumacea. Fauna Siidwest-Australiens 5: 184-187.
Zimmer, C. 1952. Indochinesische Cumaceen. Mitteilungen aus dem
Zoologischen Museum in Berlin 28: 5-36.
Appendix.
Station data from Museum Victoria collections. That which is
prefixed AA is from Prydz Bay, Antarctica. Those prefixed
MSL-EG were collected off East Gippsland, Victoria, by
tenth-metre-square Smith-Mclntyre grab from RV Sarda and
donated by the Marine Sciences Laboratories (see Coleman et
al., 1997). Those prefixed SLOPE were collected by WHOI
epibenthic sled during cruises on RV Franklin by Museum
Victoria on the south-eastern Australian continental slope (see
Poore et ah, 1994).
Antarctica
AA93 158, Eastern Prydz Bay, off the Larsemann Hills,
68°54.88'S, 76°37.03'E, 667-716 m, epibenthic sled, PM.
O’Loughlin, RSV Aurora Australis, 18 Feb 1993.
Victoria, eastern Bass Strait
MSL-EG 18, 14 km SW of Mario, 37°54'S, 148°25.7'E, 26 m,
sand-shell, 12 Aug 1989.
MSL-EG 20, 23 km SSW of Mario, 37°49'S, 148°25.7'E, 11 m,
sand-shell, 12 Aug 1989.
MSL-EG 26, 7.8 km ESE of eastern edge of Lake Tyers,
37°51.65'S, 148°10.60'E, 38 m, sand-shell, 25 Sep 1990.
MSL-EG 27, 11.2 km E of eastern edge of Lake Tyers,
37 0 51.41'S, 148°13.16'E, 32 m, sand-shell, 25 Sep 1990.
MSL-EG 28, 15.7 km E of eastern edge of Lake Tyers,
37°51.19'S, 148°16.28'E, 36 m, sand-shell, 25 Sep 1990.
MSL-EG 57, 5.4 km SW of Cape Conran, 37°51.28'S,
148°43.73'E, 50 m, sand-shell, 28 Sep 1990.
MSL-EG 62, 7.3 km SSW of Cape Conran, 37°52.67'S,
148°42.06'E, 48 m, sand-shell, 28 Sep 1990.
MSL-EG 67, 13.3 km E of eastern edge of Lake Tyers,
37°51.70'S, 148°14.60'E, 37 m, coarse sand, 4 Jun 1991.
MSL-EG 77, 11.7 km W of Pt Ricardo, 37°49.89'S, 148°30.13'E,
27 m, coarse sand, 4 Jun 1991.
New nannastacid cumaceans from Museum Victoria
173
MSL-EG 78, 11.7 km W of Pt Ricardo, 37°49.89'S, 148°30.13'E,
27 m, coarse sand, 4 Jun 1991.
MSL-EG 104, 11.7kmWof Pt Ricardo, 37°49.89'S, 148°30.13'E,
27 m, coarse sand, Feb 1991.
MSL-EG 105, 11.7km WofPt Ricardo, 37°49.89'S, 148°30.13'E,
27 m, coarse sand, Feb 1991.
MSL-EG 108, 15.5 km SW of Pt Ricardo, 37°53.14'S,
148°28.94'E, 45 m, medium sand, Feb 1991.
South-eastern Australian continental slope
SLOPE 1, NSW, off Nowra, 34°59.52'S, 151°05.94'E, 204 m,
coarse shell, G.C.B. Poore et al., RV Franklin, 14 Jul 1986.
SLOPE 2, NSW, off Nowra, 34°57.90'S, 151°08.00'E, 503 m,
bryozoa and shell, G.C.B. Poore et al., RV Franklin,
14 Jul 1986.
SLOPE 6, NSW, off Nowra, 34°51.90'S, 151°12.60'E, 770 m,
crinoid dominated, G.C.B. Poore and C.C. Lu, RV Franklin,
15 Jul 1986.
SLOPE 19, NSW, off Eden, 37°07.30'S, 150°20.20'E, 520 m,
grey coarse shell, G.C.B. Poore et al., RV Franklin, 20 Jul 1986.
SLOPE 21, NSW, off Eden, 36°57.40'S, 150°18.80'E, 220 m,
muddy shell, G.C.B. Poore et al., RV Franklin, 20 Jul 1986.
SLOPE 27, Vic., S of Point Hicks, 38°25.00'S, 149°00.00'E,
1500 m, compacted clay, G.C.B. Poore et al., RV Franklin,
22 Jul 1986.
SLOPE 45, Tas., off Freycinet Peninsula, 42°02.20'S,
148°38.70'E, 800 m, coarse shelly sand, M.F. Gomon et al., RV
Franklin, 27 Jul 1986.
SLOPE 46, Tas., off Freycinet Peninsula, 42°00.20'S,
148°37.70'E, 720 m, coarse shelly sand, M.F. Gomon et al., RV
Franklin, 27 Jul 1986.
SLOPE 47, Tas., off Freycinet Peninsula, 41°58.60'S,
148°38.80'E, trawl, 500-600 m, M.F. Gomon et al., RV
Franklin, 27 Jul 1986.
SLOPE 48, Tas., off Freycinet Peninsula, 41°57.50'S,
148°37.90'E, 400 m, coarse shell, M.F. Gomon et al., RV
Franklin, 27 Jul 1986.
Memoirs of Museum Victoria 63(2): 175-205 (2006)
ISSN 1447-2546 (Print) 1447-2554 (On-line)
http://www.museum.vic.gov.au/memoirs/index.asp
A new genus and new species of Desmosomatidae (Crustacea: Isopoda: Asellota)
from the deep sea of south-eastern Australia
Saskia Brix
Biocenter Grindel: Zoological Museum, University of Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany
(saskia.brix@uni-hamburg.de)
Abstract Brix, S. 2006. A new genus and new species of Desmosomatidae (Crustacea: Isopoda: Asellota) from the deep sea of
south-eastern Australia. Memoirs of Museum Victoria 63(2): 175-205.
A new genus and species, Chelantermedia composita sp. nov., and five new species of Desmosomatidae Sars, 1897
(Paradesmosoma australis sp. nov., Oecidiobranchus nowrae sp. nov., Disparella kensleyi sp. nov., Whoia victoriensis sp.
nov., Echinopleura cephalomagna sp. nov.) are described from the deep sea off south-eastern Australia. The species of
Desmosomatidae sampled in 1988 are the first record of this family in Australian waters.
Keywords Isopoda, Desmosomatidae, Australia, taxonomy, Chelantermedia, new genus, new species
Introduction
The new species described below were collected in the 1980s
as part of the SLOPE Program of Museum Victoria and the
then Victorian Institute of Marine Sciences. Samples were
taken off southern New South Wales and Tasmania and in
Bass Strait in 1988 from RV Franklin. Poore et ah, 1994
analysed depth and geographical diversity patterns of all
species of Isopoda. The present paper describes members of
the asellote family Desmosomatidae Sars, 1897, common
deep-sea benthic isopods with a slender body (Hessler, 1970;
Svavarsson, 1984, 1988; Wagele, 1989). In deep-sea areas that
are sampled for the first time, the fraction of isopod species
new to science ranges from 50-100% (Wilson, 1980; Poore et
ah, 1994; Park, 1999; Brandt et ah, 2004). The species of
Desmosomatidae described here are the first record of this
family in Australian waters. From a total of 21 desmosomatid
species new to science, six new species are described from the
material lodged in the Museum Victoria, Melbourne (Poore et
ah, 1994).
The specimens were compared with relevant type material
of similar species. For the drawings a Leitz MI85 compound
microscope (Biocenter Grindel and Zoological Museum,
Hamburg) and Olympus BX20 (Museum Victoria, Melbourne)
with a camera lucida were used. The dorsal and lateral habitus
drawings were made from the holotype in glycerine stained
with methylene green. All appendages were dissected from a
paratype (if not available, from the holotype) and deposited in
water-free glycerin jelly, stained and finally sealed. The total
body length was measured in dorsal and lateral views from the
anterior edge of the head to the posterior medial tip of the
pleotelson. Length-to-width ratios refer to the greatest length
and width of the limb articles or segments. The terminology
used in this study for the most important setal types follows
Hessler (1970). Following Wolff (1962) and Hessler (1970)
roman numerals refer to pereopods and Arabic numerals for
body segments and articles of appendages.
Abbreviations : A1 = antennula; A2 = antenna; MdL = left
mandible; MdR = right mandible; Mxl = maxillula; ip =
incisior process; lm = lacinia mobilis; mp = molar process;
Mx2 = maxilla; Mxp = maxilliped; Op = operculum; PI-VII =
pereopods I-VII; Pit = pleotelson; Pip 1-5 = pleopods 1-5;
Prn 1-7 = pereonites 1-7; Urp = uropods; ZMH = Zoological
Museum of Hamburg; AM = Australian Museum, Sydney,
NMV = Museum Victoria, Melbourne.
Chelantermedia gen. nov.
Type and only species. Chelantermedia composita sp. nov.
Diagnosis. Body elongated, length about 5 times longer than
width of pereonite 2. Lateral margins of pereonites straight,
Pereonites 6, 7 and pleotelson dorsally fused. Pereopod chelate,
enlarged, carpus ventrally with 1 slender seta midway and 1
slender seta proximal to claw-seta.
Etymology. The first part of the name refers to the chela of
pereopod I. Chela (Latin, a claw) and intermedia (Latin, in
between) reflect the intermediate set of characters between
Desmosomatidae and Nannoniscidae - as it occurs for example
in the chela of pereopod I.
176
Saskia Brix
Discussion. The new genus shows a mixture of characters of
Desmosomatidae Sars, 1897 and Nannoniscidae Hansen, 1916,
two families that are poorly differentiated. The new species, C.
composita, will be important in phylogenetic studies leading to
better understanding of the relationships within the family
Desmosomatidae. It possesses a combination of character states
not compatible with any existing genus. No desmosomatid
genus has pereonites 6 and 7 and the pleotelson fused as in
some nannoniscid genera, for example Rapaniscus Siebenhaller
and Hessler, 1981. All nannoniscid genera including species
with fused pereonites 6 and 7 ( Hebefustis Siebenhaller and
Hessler, 1977, Rapaniscus, Regabellator Siebenhaller and
Hessler, 1981, Nannoniscoides Hansen, 1916, Nannoniscus
Sars, 1870, Saetoniscus Brandt, 2002) have an antennula with
a bulbous distal article. The new genus does not possess such a
specialisation, the flagellar articles of the antennula tapering
towards the distal article. Pereopod I resembles most species of
Prochelator Hessler, 1970, Chelator Hessler, 1970 or
Oecidiobranchus Hessler, 1970. Prochelator is the only one of
these three that includes species with biramous uropods but
none has fused pereonites.
Chelantermedia composita sp. nov.
Figures 1-3
Material examined. Holotype. Female, preparatory, 2.1 mm; NMV
J18612. Australia, NSW, off Nowra (34°59.52'S, 151°5.94'E), 204.0 m,
WHOI epibenthic sled, G.C.B. Poore et al., RV Franklin, 14 Jul 1986
(stn SLOPE 1).
Paratypes (7 specimens: 3 females, 4 males). Same data as
holotype. NMVJ186121.
Description. (Holotype female). Habitus (fig. 1): body 2.1 mm
long, 5.17 times longer than width of pereonite 2. Pereonite 1
width 1.16 times cephalon width in dorsal view. Pereonite 1
length 1.11 pereonite 2 length, 0.96 pereonite 2 width. Pereonite
5 width 0.60 length, anterior margin straight. Pereonite 6 and
7 fused with pleotelson, lateral margins straight. Coxae 1-4
slightly produced, without setae. Pleotelson length as long as
wide, without posterolateral spines. Lateral margins straight,
posterior margin slightly rounded.
Antennula: 0.23 mm long, length 0.02 body length, 5
articles. Article 1 with 2 broom setae and 1 simple seta. Article
2 length 3.67 width, 1.94 article 1 length; with 2 long articulated
broom setae distally and 2 simple seta midway. Article 3 with 1
simple seta, article 4 with 2 long slender setae and article 5
distally with 1 aesthetasc and 2 long slender setae. Antenna:
length about one-third of body length, with 11 articles. Article
3 with 1 small seta. Article 4 with 2 small setae. Article 5
distally with 3 simple setae, 1 simple seta located midway.
Article 6 with 4 simple setae midway, distally with 2 small
broom setae and 6 long slender setae. Flagellar articles with
few setae.
Mandible: Without palp. Incisior process with 3 lobes.
Lacinia mobilis of left mandible with 4 teeth. Spine row
containing 4 spines. Molar process triangular with 7 setae.
Maxilliped: Epipodite lost during dissection. Endite with 2
coupling hooks and numerous fine setae on inner margin and
on distal tip. Palp of 4 articles, articles 1 and 2 clearly broader
than articles 3 and 4. Article 1 length 1.14 width. Edge of palp
article 2 with 4 long slender setae and inner margin with 2
long slender setae. Article 2 about 2 times width, article 3
length 5.00 width, article 4 length 3.50 width. Article 3 with 2
setae, article 4 with 3 setae.
Pereopod I: Chelate. Basis length 3.75 width, proximal to
ischium ventrally 1 large seta. Ischium length 1.77 width, with
1 ventral seta. Merus length 2.38 width, ventrally with 1 strong
unequally bifid seta, midway 1 slender seta, distodorsally 1
large simple seta. Carpus length 1.38 width, ventrally with
claw-seta, penultimate slender seta and 1 slender seta midway.
Claw-seta as long as propodus. Propodus length 2.69 width,
dorsally with 2 small setae, ventral margin with cuticular
membrane and 6 small slender setae. Dactylus length 3.6
width, without setae. Unguis (claw) of dactylus with 1 cuspidate
and 1 conate seta, 2 slender setae medially. Pereopods //-TV:
Similar. Pereopod III: Basis length 4.29 width, with 2 simple
setae and 1 broom seta. Ischium length 2.15 width, with 2
simple slender setae. Merus length 1.36 width, ventrally with
2 simple setae, distodorsally 1 simple seta. Carpus length 3.90
width, ventrally 3 distally setulate setae and combs between
insertion of setae, distodorsally 1 small simple seta. Propodus
length 3.90 width; ventral margin with 2 setae and combs
between them. Dactylus length 3.84 width, with 3 small slender
seta, unguis (claw) of 1 strong spittle -like formed conate seta
and 3 slender setae inserting ventrally. Pereopods V-VII:
Similar. Pereopod V: Basis length 3.00 width, with 7 broom
setae in irregular distances and ventrally 1 slender seta near
ischium. Ischium length 2 times width, with 1 dorsal and 1
ventral simple seta. Merus length 0.89 width, distodorsally
with 2 setae, ventrally 1 seta. Carpus length 2.80 width, with
ventral row of 4 long slender distally unequally bifid setae,
dorsally 3 slender setae and near propodus 1 small broom seta.
Propodus length 5.60 width, dorsally with 3 slender distally
unequally bifid setae increasing in length towards dactylus,
ventrally with row of 5 slender distally unequally bifid setae
increasing in length towards dactylus and 1 simple seta
between 4th and 5th seta of ventral row. Dactylus length 4.67
width, (slightly damaged) with unguis (claw) of 1 small robust
seta and 1 long conate seta broadened distally, 3 slender setae
inserting between them.
Pleopod 2 (operculum): Length 1.15 width. Form nearly
rectangular, lateral margins straight, distal margin straight.
Ventral surface without setae. Distal margin with 4 slender
setae. Pleopod 3: Endopod length 1.39 width, distally with 3
long plumose setae. Exopod length 1.46 of endopod length,
outer margin hirsute. Pleopod 4: Endopod oval and tapering to
tip, length 1.62 width. Exopod length 6.90 width, distally with
1 long plumose seta.
Uropods: Biramous. Exopod length 0.58 times endopod
length. Endopod 5.14 times longer than wide, 1 small broom
seta inserting medially, a bunch of 4 small broom setae and 1
slender seta inserting between the single small broom seta and
the tip of the endopod, distally with 3 long broom setae and 2
small simple setae. Exopod length 4.2 width, distally with 2
long slender setae. Protopod length 1.39 width, with 1 small
broom seta and 3 simple setae.
A new genus and new species of Desmosomatidae
177
Male. Habitus similar to female, 5.43 times longer than width
of pereonite 2. Antenna : 20 articles, flagellum basally swollen,
with 14 articles. Pleopod 7: 4.93 times longer than distal width,
tips triangular with 4 small slender setae on outer side. Pleopod
2: sympod length 2.03 times width, outer lateral margin slightly
convex, distally with 4 setae. Endopod inserting about 0.50 of
sympod length.
Etymology. From the latin word composita, composite. The
new species is composed of character states that occur in
Desmosomatidae and Nannoniscidae and are observed together
in the same species for the first time.
Distribution. Off southern NSW, Australia.
Figure 1. Chelantermedia composita sp. nov. Holotype female. NMV J18612. Dorsal and lateral views (A, B). Allotype male. NMV J186121.
Dorsal and lateral views (C, D).
178
Saskia Brix
Figure 2. Chelantermedia composita sp. nov. Paratype female. Allotype male. NMV J 186 121. Antennae, mouthparts, pereopod V, male pleopods.
A new genus and new species of Desmosomatidae
179
Figure 3. Chelantermedia composita sp. nov. Paratype female. NMV J 186121. Pereopods I-VII and pleopods (Op, Pip 3, 4, Urp).
180
Saskia Brix
Paradesmosoma australis sp. nov.
Figures 4-7
Material examined. Holotype. Female, preparatory, NMV J18608;
Type locality. - Australia, Vic., S of Point Hicks (38°17.70'S,
149°11.30'E), depth 400 m, WHOI epibenthic sled, RV Franklin, 24
Jul 1986 (stn. SLOPE 40).
Diagnosis. Body length about 4.4 times longer than width of
pereonite 2. Palp of left mandible with 2 articles, article 2
length 2.10 width, tapering to distal end, terminally 1 small
seta, margins hirsute. Pereonite 1 length 1.36 pereonite 2
length. Coxae 1-4 faintly produced, without setae. Pleotelson
anteriorly widest, about as wide as long, posterolateral spines
present, lateral margins slightly convex, posterior margin
slightly rounded. Uropods uniramous, endopod length 4.82
protopod length, 10.25 times longer than wide.
Description. Habitus: body 2.6 mm long, 4.36 times longer
than width of pereonite 2. Pereonite 1 width 1.44 times cephalon
width in dorsal view. Pereonite 1 length 1.36 pereonite 2 length,
0.97 pereonite 2 width. Pereonite 5 anterior margin straight,
lateral margins slightly concave. Coxae 1-4 faintly produced,
without setae. Pleotelson anteriorly widest, about as wide as
long, posterolateral spines present, lateral margins slightly
convex, posterior margin slightly rounded.
Antennula : About 0.26 mm long, length about 0.10 body
length, with 5 articles. Article 1 with 1 small slender seta and 4
small broom setae. Article 2 length 3.70 width, 1.48 article 1
length; marginally with 2 slender setae, distally with 2 long
articulated broom setae. Article 3 with 2 slender setae, article 4
distally with 1 broom seta, distal article with 1 aesthetasc, 1
broom seta and 2 long slender setae. Antenna (fig. 5): broken off.
Mandible : Palp of 2 articles. Article 1 without setae, article
2 length 2.1 width, tapering to distal end, terminally 1 small
seta, margins hirsute. Incisior process with 3 teeth. Lacinia
mobilis of left mandible with 3 teeth, right mandible not
dissected from specimen. Spine row containing 8 spines.
Molar process large, with 12 slender setae. Maxillula : Inner
lobe not dissected from specimen. Outer lobe broken off from
inner lobe, marginally with 6 ventral setae and 6 dorsal setae,
terminally with 12 strong spines. Maxilla : Medial lobe as
long, slightly broader than other lobes, terminally with 3
slender setulate setae, marginally with 14 setae, setae inserting
near base longest. Outer lobe terminally with 3 long slender
setulate setae, dorsal margin with 5 pairs of fine setae.
Maxilliped : Epipodite length 3.31 width, length 1.17 endite
length, outer margin hirsute. Endite with 2 coupling hooks,
with numerous fine setae. Edge of endite and palp articles 1-3
fringed with row of fine setae and 1 seta on distal corner. Palp
article 2 with 6 setae on inner margin, article 3 with 12 setae
on inner margin, article 4 with 3 setae, article 5 with 4 setae.
Article 1 length 0.37 width, article 2 length 1.03 width, article
3 length 0.85 width, article 4 length 1.5 width, article 5 length
1.50 width.
Pereopod I: Basis length 2.57 width, near coxa with 1
distally slender plumose seta, 5 simple slender setae and
proximal to ischium 1 slender seta. Ischium length 2.43 width,
ventrally with row of 6 simple setae, dorsolaterally with row of
7 simple setae. Merus length 0.72 width, distodorsally 1 simple
slender seta, ventrally with a row of 5 simple setae, 2 distally
setulate setae and 1 stout unequally bifid seta. Carpus length
1.44 width, with dorsolateral row of 5 simple slender setae,
distoventrally with claw-seta and a ventral row of setae of
irregular size and type: 3 robust unequally bifid setae and 5
slender setae (1 slender seta inserting proximal to propodus).
Propodus broadest at articulation to carpus, tapering towards
dactylus, length 3.05 width, ventrally fringed with fine setae
and 12 small setae inserted in cuticular membrane. Dactylus
length 5 times width, mediodistally with 3 small setae. Unguis
(claw) of 1 conate setae, 3 slender setae inserting ventrally.
Pereopod 11: Similar to pereopod III, different from pereopod
IV. Basis length 3.04 width, marginally with 7 slender setae
and 1 small broom seta, proximal to ischium ventrally with
bunch of 5 distally slender plumose setae. Ischium length 3
times width, ventrally with row of 23 distally slender plumose
setae, dorsally with 5 setae. Merus length 1.86 width, with
dorsolateral row of 9 long simple setae, ventrally with 6 simple
slender setae and 2 distally setulate setae. Carpus length 3.05
width, with ventral row of 8 robust unequally bifid setae
increasing in length towards propodus, distal seta of row as
long as propodus, dorsally with row of 12 simple setae.
Propodus length 4.18 width dorsally, ventrally 4 small slender
setae, dorsally 7 setae. Dactylus length 5.60 width,
mediodistally with 3 small seta. Unguis (claw) of 1 long conate
seta, 2 slender setae inserting ventrally. Pereopod IV: Basis
marginally with 8 distally slender plumose setae. Ischium
length 1.77 width, ventrally with row of 24 distally slender
plumose setae and dorsally 3 slender setae. Merus length 1.67
width, with 9 simple slender setae and ventral row of 13
distally slender plumose setae. Carpus length 2.41 width, with
dorsal row of 16 slender setae and a ventral row of 33 distally
slender plumose setae. Propodus length 1.66 width, dorsally
with 24 simple slender setae, ventrally with row of 25 distally
slender plumose setae, distally 2 small slender setae. Dactylus
width 0.13 propodus width, length 1.5 width, 3 small setae
mediodistally. Unguis (claw) with 1 conate seta, 2 slender
setae ventrally. Pereopod VI: Similar to pereopod V and
pereopod VII. Basis length 1.93 width, with few small slender
setae and 1 long slender seta ventrally proximal to ischium.
Ischium length 2.15 width, ventrally with 1 simple seta,
distodorsally 2 simple slender setae. Merus length 1.21 width,
with 2 ventral setae, distodorsally 1 simple slender seta.
Carpus length 3.53 width, ventrally with row of 6 long distally
setulate setae and 1 short unequally bifid seta, dorsally with
row of 7 simple slender setae. Propodus length 3.91 width,
ventrally with row of 11 long distally setulate setae, dorsally
with row of 4 simple slender seta, distally 1 small broom seta.
Dactylus length 6.75 width, proximal to unguis (claw) 4 simple
slender setae. Unguis (claw) of 1 long conate seta.
Pleopod 2 (operculum): Length 0.93 width, lateral margins
slightly convex, distal margin deeply concave, with 4 small
simple setae. Pleopod 3: Endopod length 1.62 width, distally
with 3 long plumose setae. Exopod length 0.75 of endopod
length, margins hirsute, distally with 1 small seta. Pleopod 4:
Endopod oval-shaped, length 2.31 width. Exopod length 5
width, distally 1 long plumose seta, outer margin hirsute.
A new genus and new species of Desmosomatidae
181
Uropods : Uniramous. Endopod length 4.82 protopod
length, 10.25 times longer than wide, marginally with 2 small
broom setae, distally with 4 slender setae and 5 broom setae.
Protopod length 1.7 width, with 3 simple slender setae.
Etymology. Australis (Latin), means “from the south”. The
name refers to the fact that this species is the first record of the
genus Paradesmosoma from the southern hemisphere.
Distribution. South-eastern Australia, off Vic.
Discussion. The new species is assigned to the genus
Paradesmosoma Kussakin, 1965 based on the characteristic
shape of pereopod IV, the kind of setation of this pereopod,
carpus and propodus “paddle-like” and surrounded by
numerous distally slender plumose setae (occurring in
Paradesmosoma only) and the characters of pereopod I:
carpus less enlarged than propodus, with ventral row of
irregular setae of varying types. P. australis sp. nov. is most
similar to P. orientale Kussakin, 1965 in the shape of palp
article 2 of the mandible. As in P. orientale, palp article 2 is
about 2.17 times longer than wide and tapers to the distal end
which is tipped by a small simple setae. In contrast to P.
orientale, the lateral margins of this article are hirsute in P.
australis. According to Kussakin’s (1965) drawings, neither P.
orientale nor P. conforme Kussakin, 1965 possesses
posterolateral spines on the pleotelson. P. australis has
posterolateral spines on the pleotelson. Unlike in the two
species from the northern hemisphere, the propodus of
pereopod I is posteriorly widest in P. australis and tapers
toward the dactylus.
Figure 4. Paradesmosoma australis sp. nov. Holotype female. NMV J18608. (A) Lateral and (B) dorsal views.
182
Saskia Brix
Figure 5. Paradesmosoma australis sp. nov. Holotype female. NMV J18608. Antennae, mouthparts and pereopod I.
A new genus and new species of Desmosomatidae
183
Figure 6. Paradesmosoma australis sp. nov. Holotype female. NMV J18608. Pereopods II-V.
184
Saskia Brix
Figure 7. Paradesmosoma australis sp. nov. Holotype female. NMV J18608. Pereopods VI and VII, pleopods (Op, Pip 3, Pip 4, Urp).
A new genus and new species of Desmosomatidae
185
Oecidiobranchus nowrae sp. nov.
Figures 8-10
Material examined. Holotype. Male, 1.4 mm, NMV J18606. Australia,
NSW, 65 km E of Nowra (34°55.52'S, 151°22.20'E), 2055 m depth,
G.C.B. Poore et al., RV Franklin , 23 Oct 1988 (stn. SLOPE 63).
Diagnosis. Body length 3.83 times longer than width of
pereonite 2. Pereonite 1 length 1.10 pereonite 2 length, similar
to pereonite 2 width. Pereonite 5 anterior margin slightly
convex, lateral margins straight. Lacinia mobilis of left
mandible with 5 teeth. Coxae 1-4 produced, tipped with stout
setae. PI carpus length 1.41 width, distodorsally with 1 long
robust simple seta, ventrally with the large claw-seta and a row
of distally setulate setae and setal combs inserted in a cuticular
membrane, propodus length 2.61 width, more enlarged than
carpus. Pleotelson with posterolateral spines.
Description. Habitus : body 1.4 mm long (measured without
appendages), 3.83 times longer than width of pereonite 2.
Pereonite 1 width 1.13 times cephalon width in dorsal view.
Pereonite 1 length 1.10 pereonite 2 length, similar to pereonite 2
width. Pereonite 5 width 1.36 length, anterior margin slightly
convex, lateral margins straight. Coxae 1^1 produced, tipped
with stout setae. Pleotelson length 0.71 width, posterolateral
spines present. Lateral margins convex, posterior margin
rounded.
Antennula : 0.31 mm long, length 0.22 body length, with 5
articles. Article 1 with 3 broom setae. Article 2 length 1.53
width, 2.36 article 1 length; distally with 3 broom setae and 1
small seta. Article 3 with 1 slender seta, article 4 with 2 small
broom setae, distal article with 1 aesthetasc, 1 broom seta and
4 long slender setae. Antenna : About 1 mm long, length about
0.71 body length, with 22 articles. Articles 3 with 3 small setae,
article 4 with 3 small setae. Article 5 with 4 slender setae and
distally 1 broom seta. Article 6 with 6 slender setae. Flagellum
basally swollen (sexual dimorphism), articles with 1 or 2
slender setae each, distal article with 5 long slender setae.
Mandible: Article 2 of palp ventrodistally with 4 small
setae, dorsally with rows of fine setae, apical article with 6
setae, distal one longest. Incisior process with 3 teeth. Lacinia
mobilis of left mandible with 5 teeth. Right mandible not
dissected from specimen. Spine row containing 5 spines.
Molar process with 6 fine slender setae. Maxillula: Inner lobe
slightly smaller than outer lobe, terminally with 5 setae. Outer
lobe dorsally with 4 fine setae, terminally with 11 strong
spines (4 spines with setules). Maxilla: Not dissected from
specimen. Maxilliped : Epipodite length 3.13 width, length
1.43 endite length. Endite with 2 coupling hooks, terminally
with 1 conate seta and numerous fine setae. Outer edge of
endite and palp articles 1-3 fringed with numerous fine setae,
distal corners tipped with 1 seta. Palp article 3 with 6 setae on
inner margin, article 4 with 2 setae and article 5 with 4 setae.
Article 1 length 0.5 width, article 2 length 1.16 width, article
3 length 1.07 width, article 4 length 1.1 width, article 5 length
1.6 width.
Pereopod I: Enlarged, chelate. Basis length 4.53 width,
marginally with 1 small broom seta and 5 simple setae,
proximal to ischium ventrally 1 long simple seta. Ischium
length 1.87 width, distodorsally 1 simple seta, ventrally 2
simple setae. Merus length 1.13 width, ventrally 2 setal
combs inserted in a cuticular membrane, distally 1 distally
setulate and 1 simple seta, distodorsally 2 setae. Carpus
length 1.41 width, distodorsally 1 long robust simple seta,
ventrally with the large claw-seta and a row of distally
setulate setae and setal combs inserted in a cuticular
membrane. Propodus length 2.61 width, more enlarged than
carpus, ventral and dorsal margin convex, distodorsally 2
small setae, ventrally fringed with fine setae and 9 small
setae inserted in a cuticular membrane. Dactylus length 7.50
width, folding to propodus, mediodistally with 3 small setae.
Unguis (claw) of dactylus with 1 cuspidate and 1 conate seta,
2 slender setae medially. Pereopod IIP. Pereopod II missing
from specimen, pereopod III similar to pereopod IV. Basis
length 5.70 width, marginally with 4 broom setae, 5 small
slender setae and proximal to ischium ventrally 1 large
simple seta. Ischium length 2.25 width, distodorsally 1 long
simple seta and distoventrally 1 long simple seta. Merus
length 2.50 width, dorsally with 2, ventrally with 1 simple
seta. Carpus length 4.30 width, with ventral row of 4 long
slender setae, dorsally with row of 4 setae. Propodus length
3.50 width, ventrally with row of 5 slender setae, dorsally
with 4 slender setae and 1 small broom seta. Dactylus length
4 times width, mediodistally with 3 small slender seta.
Unguis (claw) of 1 conate seta, 2 slender setae inserting
ventrally. Pereopod V: Similar to pereopod VI and VII. Basis
length 4 times width, marginally with 3 simple slender setae
and 3 broom setae. Ischium length 2.64 width, dorsally with
4 slender setae, ventrally with 2 small slender setae. Merus
length 1.89 width, 1 small slender seta distodorsally and 1
small slender seta distolaterally. Carpus length 3 times width,
ventrally with row of 5 long slender setae, and distodorsally
1 small broom seta. Propodus length 3.4 width, ventrally
with a row of 6 long slender setae, dorsally with 7 long
slender setae and 2 unequally bifid setae (1 midway, 1
distally). Dactylus length 4 times width, mediodistally with
3 small slender setae, unguis (claw) of 1 long conate seta, 2
slender setae inserting ventrally.
Pleopod 1: Length 2.04 width. Outer margins straight,
terminal margin rounded, with 3 small setae on each side.
Pleopod 2: Sympod oval-shaped, length 2.24 width. Outer
margin distally with 1 small seta. Endopod inserting 0.56 of
sympod length. Pleopod 3: Endopod length 1.4 width, distally
with 3 long plumose setae. Exopod length 0.52 of endopod
length, terminally tapering, with 1 simple seta, outer margin
hirsute. Pleopod 4 : Endopod oval-shaped, length 1.92 width.
Exopod missing.
Uropods: Uniramous. Endopod length 2.33 protopod
length, 3.5 times longer than wide, distally with 3 broom setae,
1 small seta and 4 slender setae. Protopod length 1.5 width,
with 3 simple slender setae.
Etymology. Nowrae (Lat.) means “from Nowra”. The name
refers to the sampling area on the southern Australian
continental slope 65 km E of Nowra.
Distribution. Off NSW, Australia.
186
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Discussion. Oecidiobranchus nowrae sp. nov. is assigned to
the genus Oecidiobranchus Hessler, 1970 on the basis of the
small and rounded breathing chamber. The male of the new
species differs from the other two species of the genus in the
characters of pereopod I. Unlike O. plebejum Hansen, 1916 and
O. nanseni Just, 1980, O. nowrae possesses a ventral row of
distally setulate seta on the carpus. Furthermore, the new
species possesses posterolateral spines at the pleotelson.
Neither O. plebejum, nor O. nanseni possess posterolateral
spines at the pleotelson.
Figure 8. Oecidiobranchus nowrae sp. nov. Holotype male. NMV J18606. (A) Lateral and (B) dorsal views, antennae.
A new genus and new species of Desmosomatidae
187
Figure 9. Oecidiobranchus nowrae sp. nov. Holotype male. NMV J18606. Mouthparts, pereopods I-V.
188
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Figure 10. Oecidiobranchus nowrae sp. nov. Holotype male. NMV J18606. Pereopods VI and PVII, pleopods (Pip 1, 2, 3, 4, Urp).
A new genus and new species of Desmosomatidae
189
Disparella kensleyi sp. nov.
Figures 11-14
Material examined. Holotype. Female, preparatory, 1.9 mm; NMV
J18605; Type locality. - Australia, NSW, 74 km E of Nowra (34°56.1 l'S,
151°28.06'E), 3150 m, box corer, G.C.B. Poore et al, RV Franklin ,
23 Oct 1988 (stn. SLOPE 64).
Diagnosis. Body length 3.98 times longer than width of
pereonite 2. Pereonite 1 length 1.05 pereonite 2 length, 0.93
pereonite 2 width. Pereonite 5 with spine-like ventral elongation.
Cephalon with 1 short anteriorly directed spine at insertion of
antennae. Palp of mandible of 2 articles, article 1 with 1 small
seta, article 2 terminally with 1 small seta. Incisior process with
4 teeth. Lacinia mobilis of left mandible with 4 teeth. Carpus of
pereopod I length 5 times width, with ventral row of 5 long
unequally bifid setae increasing in length towards the propodus
and 3 setal combs inserted in a cuticular membrane, dorsally
with 2 long simple setae. Pleotelson with posterolateral spines.
Distal margin of operculum slightly concave, with 6 slender
setae. Uropods biramous, exopod length 0.43 endopod length.
Description. Habitus : body 1.9 mm long (measured without
appendages), 3.98 times longer than width of pereonite 2.
Pereonite 5 with spine-like ventral elongation. Cephalon with 1
short anteriorly directed spine at insertion of antennae.
Pereonite 1 slightly wider than cephalon. Pereonite 1 length
1.05 pereonite 2 length, 0.93 pereonite 2 width. Pereonite 5
anterior margin straight, lateral margins slightly concave.
Coxae 1-4 produced, tipped with small stout setae. Pleotelson
length 0.71 width, posterolateral spines present, lateral margins
convex, posterior margin rounded.
Antennula: 0.3 mm long, length 0.16 body length, with 5
articles. Article 1 with 3 small broom setae. Article 2 length
3.83 width, 1.64 article 1 length; distally with 2 large articulated
broom setae. Article 3 with 1 small slender seta, article 4
distally with 2 small broom setae, distal article terminally with
1 aesthetasc, 1 broom seta and 2 long slender setae. Antenna :
About 1.1 mm long, length 0.58 body length, with about 12
articles (broken off after article 12). Article 3 with 2 small
slender setae. Article 4 with 2 slender setae. Article 5 with 3
small slender setae. Article 6 marginally with 2 and distally
with 3 slender setae. Flagellar articles 1-3 distally with 2
slender setae, following articles with long slender setae.
Mandible: Palp of 2 articles, article 1 with 1 small seta,
article 2 terminally with 1 small seta. Incisior process with 4
teeth. Lacinia mobilis of left mandible with 4 teeth, lacinia
mobilis of right mandible triangular and distally serrated (5
small teeth). Spine row containing 3 spines. Molar process
with 7 setae. Maxillula: Inner lobe smaller than outer lobe
(0.71 of outer lobe length), terminally with 5 setae. Outer lobe
5 times longer than wide, marginally with 3 pairs of fine setae,
terminally with 9 strong spines and 4 simple setae. Maxilla:
Not dissected from specimen. Maxilliped: Epipodite length
3.13 width, length similar endite length. Endite with 2 coupling
hooks, terminally with 2 conate setae and 5 small setae,
marginally few fine setae. Outer edge of endite and palp articles
1 and 2 with row of fine setae and 1 small seta on distal comers,
inner margin of article 3 with 5 setae, outer margin with 1 seta,
article 4 with 3 setae, article 5 with 2 setae. Article 1 length 0.6
width, article 2 length 0.93 width, article 3 length 1.07 width,
article 4 length 2.5 width, article 5 length 3 times width.
Pereopod I: Basis length 4.04 width, proximal to ischium
ventrally 1 long simple seta. Ischium length 0.63 width,
ventrally 1 small slender seta. Merus length 0.71 width,
dorsally 2 robust simple setae, ventrally 1 seta (broken off).
Carpus length 1.43 width, ventrodistally with large claw-seta
and slender proximal seta, ventrally 1 small seta midway.
Propodus length 3.47 width, ventrally fringed with fine setae
and row of 12 small setae inserted in a cuticular membrane.
Dactylus length 7.6 width, mediodistally 1 small seta. Unguis
(claw) of dactylus with 1 cuspidate and 1 conate seta, 2 slender
setae medially. Pereopod IP. Similar to pereopod III and
pereopod IV. Basis length 6.33 width, marginally with few
simple setae and 1 small broom seta, proximal to ischium
ventrally with 1 long simple seta. Ischium length 2.62 width,
distodorsally with 1 seta, ventrally 1 seta midway. Merus
length 1.45 width, distoventrally with 1 stout unequally bifid
seta and 2 small slender setae, distodorsally 1 composed seta.
Carpus length 5 times width, with ventral row of 5 long
unequally bifid setae increasing in length towards the propodus
and 3 setal combs inserted in a cuticular membrane, dorsally
with 2 long simple setae. Propodus length 5.38 width, ventrally
with 1 small slender seta midway, fringed with fine setae,
dorsally 2 long simple setae and distally 1 small broom seta.
Dactylus length 6.67 width, mediodistally 2 small setae.
Unguis (claw) of dactylus with 1 conate seta, 2 slender setae
ventrally. Pereopod V: Similar to pereopod VI and pereopod
VII. Basis length 6.30 width, marginally with 1 small and 2
broom setae. Ischium length 4.10 width. Merus length 1.60
width, with 3 slender setae. Carpus length 4.80 width, with
ventral row of 5 long slender setae and dorsally 2 long setae
(broken off), distally 1 small slender seta and 1 small broom
seta. Propodus length 4.13 width, ventrally with 1 small seta
and 3 long slender setae, dorsally with 2 slender setae. Dactylus
length 7.60 width, mediodistally 3 small setae, unguis (claw)
of 1 long conate seta, 2 slender setae inserting ventrally.
Pleopod 2 (operculum): Length 1.19 width. Lateral margins
slightly convex, distal margin slightly concave, distal margin
with 6 slender setae. Pleopod 3: Endopod length 1.44 width,
distally with 3 long plumose setae. Exopod length 0.74 of
endopod length, with 1 small terminal seta, outer margin
hirsute. Pleopod 4: Endopod length 2.66 width. Exopod length
8 times width, distally with 1 long plumose seta.
Uropods: Biramous. Endopod length 3.5 protopod length,
4.66 times longer than wide, distally with 5 broom setae, 2
small slender setae and 2 long slender setae. Exopod length
0.43 endopod length, 4 times width, terminally with 2 slender
setae. Protopod length similar width, with 1 slender seta.
Etymology. The name is in remembrance of Dr Brian Kensley.
Distribution. Off NSW, Australia.
Discussion. Disparella kensleyi sp. nov. shows affinity to
species of Chelator and Prochelator Hessler, 1970. The chela
of the new species is similar to the chela as found in Chelator
(ventral margin of carpus with small setae only), but Chelator
190
Saskia Brix
species possess uniramous uropods and a lacinia mobilis with
three teeth. A single midventral seta on the carpus together
with the claw-seta and the penultimate slender seta is also
known for the genus Prochelator but in Prochelator species
the midventral seta is always of composed setal type. Due to
the anteriorly directed spine at the antennular folds, which is
known for all species of Disparella, and the biramous uropods,
D. kensleyi fits best into Disparella. It is distinguished from the
other members of the genus by the spine-like ventral elongation
at pereonite 5 and the single small midventral seta on the carpus
of pereopod I together with the claw-seta and the penultimate
slender seta.
Figure 11. Disparella kensleyi sp. nov. Holotype female. NMV J18605. (A) Dorsal and (B) lateral views.
A new genus and new species of Desmosomatidae
191
Figure 12. Dispar ella kensleyi sp. nov. Holotype female. NMV J18605. Antennae, mouthparts, pereopod I.
192
Saskia Brix
Figure 13. Dispar ella kensleyi sp.nov. Holotype female. NMV J18605. Pereopods II-V.
A new genus and new species of Desmosomatidae
193
Figure 14. Disparella kensleyi sp. nov. Holotype female. NMV J18605. Pereopods VI and VII, pleopods (Op, Pip 3, 4) and pleotelson ventral view.
194
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Echinopleura cephalomagna sp. nov.
Figures 15-18
Material examined. Holotype. Female, preparatory, 1.8 mm; NMV
J18600 Australia, Vic., S of Point Hicks (38°17.70'S, 149°11.30'E), 400
m depth, WHOI epibenthic sled, G.C.B. Poore et al., RV Franklin, 24
Jul 1986 (stn. SLOPE 40).
Paratypes. 1 female, adult, 2.2 mm; NMV J18601; locality. -
Australia, NSW, 54 km ESE of Nowra (34°52.72’S, 151°15.04'E),
996-990 m, WHOI epibenthic sled, G.C.B. Poore et ah, RV Franklin,
22 Oct 1988 (stn. SLOPE 53). 1 female, preparatory; 2.1 mm; NMV
J53074. Australia, NSW, 54 km ESE of Nowra (34°52.72'S,
151°15.04’E), 996-990 m, WHOI epibenthic sled, G.C.B. Poore et ah,
RV Franklin, 22 Oct 1988 (stn. SLOPE 53).
Diagnosis. Body length 3.20 times longer than width of
pereonite 2. Cephalothorax highest part of body from lateral
view. Pereonite 1 slightly smaller than pereonite 2. Lateral
margins of pereonite 5, 6 and 7 as well as of pleotelson serrated.
Incisior process with 1 rounded tooth. Lacinia mobilis of left
mandible represented by 1 small bulge-like tooth.
Description. Habitus: body 2.2 mm long (measured without
appendages), 3.20 times longer than width of pereonite 2.
Cephalothorax highest part of body from lateral view. Pereonite
1 width 1.16 times cephalon width in dorsal view. Pereonite 1
length 0.94 pereonite 2 length, 0.91 pereonite 2 width. Pereonite
5 width 0.67 length, anterior margin straight, lateral margins
serrated. Coxae 1-4 produced, tipped with small stout setae.
Pleotelson length 1.06 width, lateral margins serrated, lightly
convex, posterior margin smooth, rounded.
Antennula: 0.27 mm long, length 0.12 body length, with 6
articles. Article 1 with 1 small slender seta and 3 broom setae.
Article 2 length 2.57 width, 0.69 article 1 length; distally with
4 articulated broom setae. Article 3 with 1 small seta, article 4
distally with 2 broom setae, distal article with 1 aesthetasc and
4 slender setae. Antenna (fig. 12): broken off.
Mandible: Article 1 of palp distally with 1 slender seta,
article 2 distoventrally with 2 small setae, fringed with rows of
fine setae, article 3 with 4 small and distally 1 setulate seta.
Incisor process with 1 rounded tooth. Lacinia mobilis of left
mandible represented by 1 small bulge -like tooth. Lacinia
mobilis absent at right mandible. Spine row containing 9
spines. Molar process with 10 setae. Maxillula: Inner lobe
smaller than outer lobe (0.63 of outer lobe length), distally
with 7 simple setae, dorsally with 12 fine setae and ventrally
with 5 pairs of fine setae. Outer lobe marginally with 21 pairs
of fine setae, terminally with 11 strong spines (7 spines with
setules). Maxilla (fig. 12): Medial lobe broader than other
lobes, terminally with 3 slender setae, ventrolaterally with 12
setae, setae near base longest. Outer lobe dorsally with 8 pairs
of fine setae, terminally with 3 long ventrally setulate setae.
Maxilliped (fig. 12): Epipodite length 3.08 width, length
similar endite length. Endite with 2 coupling hooks, terminally
with 1 fan seta and numerous fine setae, marginally with pairs
of fine setae. Edge of endite fringed with fine setae, palp article
1 with 2 setae on outer margin, article 2 with 3 setae on outer
margin and 3 setae on inner margin, article 3 with 7 setae on
inner margin, article 4 with 4 setae, article 5 with 3 setae.
Article 1 length 0.91 width, article 2 length 0.89 width, article
3 length 1.11 width, article 4 length 1.6 width, article 5 length
2.67 width.
Pereopod I: Basis length 5.94 width, marginally with 14
setae. Ischium length 3.17 width, ventrally with 7 small slender
setae, dorsally with 2 simple slender setae. Merus length 1.18
width, ventrally with a row of 4 simple setae, distodorsally
with 1 seta (broken off). Carpus length 3.20 width, ventrally
with dorsal row of 7 setae, distal and penultimate seta longest,
distally setulate, dorsally with 3 small slender setae. Propodus
length 8.00 times width, with few setae distally. Dactylus
length 4.20 width, mediodistally 3 small setae. Unguis (claw)
of dactylus with 1 cuspidate and 1 conate seta, 2 slender setae
medially. Pereopod II: Similar to pereopod III. Basis length
4.00 times width, marginally with 27 small setae, proximal to
ischium ventrally 1 long simple seta. Ischium length 3.13
width, ventrally 16 simple seta, dorsally 4 simple setae. Merus
length 1.38 width, ventrally with row of 6 setae, distodorsally
2 simple setae. Carpus length 3.57 width, with ventral row 11
long unequally bifid setae increasing in length towards the
propodus, dorsolaterally with row of 11 long simple setae,
dorsally with 6 small slender setae. Propodus length 2.70 times
width, ventrally with 1 small slender seta, 2 small stout
unequally bifid setae and 2 setal combs inserted in a cuticular
membrane, dorsally with row of 8 long simple setae. Dactylus
length 2.50 width, mediodistally 3 small setae. Unguis (claw)
of dactylus with 1 cuspidate and 1 conate seta, 2 slender setae
medially. Pereopod VI: Similar to pereopod VII, pereopod V
missing from specimen. Basis length 4.19 width, with 2 large
broom setae, marginally with 13 small setae. Ischium length
3.39 width, laterally with 9 small setae, dorsally with 6 slender
setae. Merus length 1.38 width, ventrally with 3 small slender
setae. Dorsally with 1 small seta and 1 long simple seta.
Carpus length 4.11 width, ventrally with row of 12 long slender
setae, dorsally with row of 9 slender setae and 4 small setae.
Propodus length 3.79 width, ventrally with row of 5 long
slender setae, dorsally with row of 6 setae. Dactylus length
8.50 width, distally with 1 slender seta, unguis (claw) of 1 long
conate seta, 2 slender setae inserting ventrally.
Pleopod 2 (operculum): Length 1.37 width. Lateral
margins straight, operculum tapering towards distal tip, setose
(ventral surface with about 34 setae), marginally with 58 setae.
Pleopod 3: Endopod length 1.8 width, distally with 3 long
plumose setae. Exopod length 0.44 of endopod length, margins
hirsute, distally with 1 simple seta. Pleopod 4: Endopod oval-
shaped, length 2 times width. Exopod length 7.80 width,
distally 1 long plumose seta.
Uropods: Uniramous. Endopod length 2.04 protopod
length, 8.17 times longer than wide, marginally with 6 slender
setae, distally with 5 long slender setae, 2 small setae and 5
broom setae. Protopod length 1.85 width, with 12 setae.
Etymology. The name refers to the extremely large
cephalothorax of the new species.
Distribution. South-eastern Australia, off Vic.
Discussion. Echinopleura cephalomagna sp. nov. is the second
species of the genus Echinopleura Sars, 1897. It belongs to this
genus because of the slender pereopod I and the features of the
A new genus and new species of Desmosomatidae
195
mandible and the serrated body margins from pereonite 5 to
the pleotelson. The simplified mandible is unique to
Ecliinopleura. The new species is easy to distinguish from the
only other species, E. aculeata Sars, 1897, by the presence of a
well developed mandibular palp. In E. aculeata the whole body
is serrated, even the cephalon and the coxae. E. aculeata
possesses a dorsal hook on the ischium of pereopod II, E.
cephalomagna lacks this hook. While the antennula of E.
aculeata consists of five articles, the antennula of E.
cephalomagna consists of six articles.
Figure 15. Echinopleura cephalomagna sp. nov. Holotype female. NMV J18600. (A) Dorsal and (B) lateral views.
196
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Figure 16. Echinopleura cephalomagna sp. nov. Paratype female. NMV J18601. Antennula, mouthparts.
A new genus and new species of Desmosomatidae
197
Figure 17. Echinopleura chephalomagna sp. nov. Paratype female. NMV J 18601. Anterior pereopods.
198
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Figure 18. Echinopleura cephalomagna sp. nov. Paratype female. NMV J18601. Posterior pereopods, pleopods (Op, Pip 3, Pip 4, Pip 5, Urp).
A new genus and new species of Desmosomatidae
199
Whoia victoriensis sp. nov.
Figures 19-23
Material examined. Holotype. Female, preparatory, 1.6 mm NMV
J18598; Type locality. - Australia, Vic., 76 km S of Point Hicks
(38°29.33'S, 149°19.98'E), 1840-1750 m depth, WHOI epibenthic sled,
G.C.B. Poore et al., RV Franklin , 26 Oct 1988 (stn SLOPE 69).
Paratype. Female, preparatory, 1.5 mm NMV J18599. Australia,
Vic., 67 km S of Point Hicks (38°23.95’S, 149°17.02'E), 1277-1119 m
depth, WHOI epibenthic sled, G.C.B. Poore et ah, RV Franklin, 25
Oct 1988 (stn. SLOPE 67).
Diagnosis. Body length 4 times longer than width of pereonite 2.
Pereonite 1 slightly smaller than pereonite 2. Pereonite 5 anterior
margin straight, lateral margins straight. Antennula with 6
articles. Incisor process with 2 teeth. Lacinia mobilis of left
mandible with 1 tooth. Coxae 1^1 angular, coxa 1 with small
stout seta, 2^1 without setae. Ischium, merus and carpus of
anterior pereopods laterally with numerous folds in which rows
of fine setae are inserted. Pereopod I and pereopod II similar in
setation, carpus with ventral row of 5 large robust unequally bifid
distally setulate setae increasing in length towards propodus,
distal seta of row reaching full length of propodus, dorsally with
a row of 5 slender distally setulate setae, propodus ventrally with
2 small stout unequally bifid setae and a row of 14 small setae
inserted between them, dorsally with a row of 5 slender distally
setulate setae and distally 1 small seta. Lateral margins of
pleotelson hirsute, form tapering to posterior margin. Urbiramous,
setose, exopod well developed (length 0.64 endopod length).
Description. Habitus : body 1.6 mm long (measured without
appendages), 4.02 times longer than width of pereonite 2.
Pereonite 1 width 1.13 times cephalon width in dorsal view.
Pereonite 1 length 0.60 pereonite 2 length, 0.91 pereonite 2
width. Pereonite 5 anterior margin straight, lateral margins
straight. Coxae 1-4 angular, coxa 1 with small stout seta, 2-4
without setae. Pleotelson tapering to posterior margin, length
1.19 width, without posterolateral spines, lateral margins
hirsute, convex, posterior margin triangularly convex.
Antennula: Length 0.23 body length, with 6 articles.
Article 1 with 1 small seta and 4 broom setae. Article 2 length
4.25 width, 1.90 article 1 length; distally with 4 articulated
broom setae and 1 small seta. Article 3 with 1 small seta,
article 4 with 2 slender setae and 1 small seta, article 5 with 1
slender seta, distal article terminally with 1 aesthetasc, 1
broom seta and 2 long slender setae. Antenna : broken off.
Mandible : 1st article of palp with 1 small seta, 2nd article
ventrodistally with 2 small setulate setae, dorsally with rows of
fine setae, apical article dorsally with 1 small seta and rows of
fine setae, ventrally with 5 setae, distal 1 longest. Incisor process
with 2 teeth. Lacinia mobilis of left mandible with 1 tooth,
lacinia mobilis of right mandible of the same shape as lacinia
mobilis of left mandible. Spine row containing 9 spines. Molar
process with 5 finely setulate setae. Maxillula: Inner lobe slightly
smaller than outer lobe, terminally with 7 setae, ventrally with 4
slender setae, dorsally with 5 pairs of fine setae. Outer lobe
marginally with 10 pairs of fine setae, terminally with 9 strong
spines (3 spines with setules). Maxilla : Medial lobe broader than
other lobes, distally with 7 simple setae, marginally with pairs of
fine setae, basally with 7 slender setae. Outer lobe terminally
with 3 setae, ventrally with 3 simple setae, dorsally with fine
setae. Maxilliped : Epipodite length 2.67 width, length 0.90
endite length. Endite with 2 coupling hooks, terminally with 1
fan seta and numerous small setae. Edge of endite and palp
articles 1.3 hirsute, distal comers with 1 small seta. Article 2
inner margin with 3 setae, article 3 inner margin with 7 setae,
article 4 with 4 setae, article 5 with 5 setae. Article 1 length 0.69
width, article 2 length similar to width, article 3 length 0.79
width, article 4 length 1.8 width, article 5 length 4 times width.
Pereopod I: Basis length 2.28 width, with few small setae
and proximal to ischium ventrally with 1 long simple seta.
Ischium length 2.14 width, ventrally with 3 slender distally
setulate setae and 2 robust unequally bifid distally setulate
setae, dorsally with 3 slender distally setulate setae. Ischium,
merus and carpus laterally with numerous folds in which rows
of fine setae are inserted. Merus length 0.42 width, ventrally
with 3 robust stout unequally bifid distally setulate setae,
distodorsally 1 simple slender seta and 1 robust unequally
bifid distally setulate seta. Carpus length 1.96 width, with
ventral row of 5 large robust unequally bifid distally setulate
setae increasing in length toward propodus, distal seta of row
reaching full length of propodus, dorsally with a row of 5
slender distally setulate setae. Propodus length 2.69 width,
ventrally with 2 small stout unequally bifid setae and a row of
14 small setae inserted between them, dorsally with a row of
5 slender distally setulate setae and distally 1 small seta.
Dactylus length 4.13 width, mediodistally with 3 small setae,
unguis (claw) of dactylus with 1 cuspidate and 1 conate seta, 2
slender setae medially. Pereopod IP. In setation similar to
pereopod I. Difference in length-to-width ratios: basis length
1.90 width, ischium length 2.10 width, merus length 0.35
width, carpus length 1.93 width, propodus length 2.94 width,
dactylus length 3.43 width. Unguis (claw) of dactylus with 1
cuspidate and 1 conate seta, 2 slender setae medially. Pereopod
VII: Similar to pereopod V and pereopod VI. Basis length
3.06 width, with few setae. Ischium length 2.19 width, ventrally
with 3 small slender setae. Merus length 0.73 width,
distodorsally 1 simple slender seta, ventrally 1 small and 1
simple slender seta. Carpus length 2.58 width, ventrally with
row of 9 long slender distally setulate setae, dorsally with a
row of 11 long slender distally setulate setae. Propodus length
3.79 width, ventrally with row of 4 long slender distally
setulate setae, dorsally with row of 10 long slender distally
setulate setae and 2 small unequally bifid setae, 1 midway, 1
distally. Dactylus length 4.2 width, unguis (claw) of 1 conate
seta, 1 slender seta inserted ventrally.
Pleopod 2 (operculum): Length 1.16 width. Surrounded by
64 setae, lateral margins slightly convex, distal margin straight.
Pleopod 3: Endopod length 1.62 width, inner margin hirsute,
distally with 3 long plumose setae. Exopod length 0.41 of
endopod length, outer margin hirsute, distally 1 slender seta.
Pleopod 4: Endopod oval-shaped, length 1.91 width. Exopod
length 7.5 width, outer margin basally hirsute, distally with 1
long plumose seta. Pleopod 5: Endopod only, length 4.18 width.
Uropods : Biramous. Endopod length 2.64 protopod length,
5.50 times longer than wide, marginally with 6 small broom
setae and 2 simple slender setae, distally with 2 broom setae, 1
200
Saskia Brix
small slender seta and 5 long slender setae. Exopod length
0.64 endopod length, 7 times width with marginally 2 simple
slender setae, distally with 5 long slender setae. Protopod
length 1.39 width, with 2 small slender setae and 4 long simple
slender setae.
Etymology. The name refers to the state Victoria in Australia.
Distribution. South-eastern Australia, off Vic.
Discussion. The new species is assigned to the genus Whoia
Hessler, 1970 because of its robust pereopod I with nearly
quadrangular articles and the similar size and shape of pereopods
I and II. Whoia victoriensis sp. nov. is in regard to the body
shape most similar to W. angusta Sars, 1899. Characters
distinguishing the new species from the other three species of
the genus are: lacinia mobilis only one tooth, pereopod I with
rows of extremely robust ventral setae on carpus and propodus,
propodus dorsally with row of setae, uropods biramous, exopod
well developed, reaching more than half of endopod length.
The new species shares an antennula consisting of six articles
with W. dumbshafensis Svavarsson, 1988, in W. variabilis
Hessler, 1970 and W. angusta the antennula has five articles. In
W. angusta the lacinia mobilis has four teeth and the lateral
margins of pereonite 5 are straight. In W. variabilis the uropodal
exopod is much smaller than in W. victoriensis sp. nov and the
uropods are less setose, pereopod I does not bear large robust
setae, the lacinia mobilis has four teeth. The lacinia mobilis of
W. dumbshafensis has only two teeth.
Figure 19. Whoia victoriensis sp. nov. Holoype female. NMV J18598. Dorsal and lateral views (A, B), paratype female, lateral view (C).
A new genus and new species of Desmosomatidae
201
Figure 20. Wlioia victoriensis sp. nov. Paratype female. NMV J18599. Antennae (A2 only articles 1-4), mouthparts.
202
Saskia Brix
Figure 21. Wlioia victoriensis sp. nov. Paratype female. NMV J18599. Anterior pereopods.
A new genus and new species of Desmosomatidae
203
Figure 22. Wlioia victoriensis sp. nov. Paratype female. NMV J18599. Posterior pereopods.
204
Saskia Brix
Figure 23. Wlioia victoriensis sp. nov. Paratype female. NMV J18599. Pleopods (Op, Pip 3, Pip 4, Urp).
A new genus and new species of Desmosomatidae
205
Acknowledgements
I am grateful to Dr Angelika Brandt for making this work
possible. I thank Dr Gary Poore for giving me the opportunity
to work with the collection of Museum Victoria, Melbourne.
Thanks to Dr G.D.F. (Buz) Wilson for helpful comments and
providing a working place to have a look at type material at the
Australian Museum, Sydney. I appreciate the motivating
support of Dr Joanne Taylor, Dr Robin Wilson and my fellow
PhD student Anna Syme during my stay in Melbourne. Lydia
Kramer and Bente Stransky are thanked for their help with the
final layout of the drawings. This work was supported by a
grant of the German Science Foundation (DFG) under contract
numbers Br 1121/ 22-1, 2 and 3.
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and Saetoniscus n. gen. from the deep sea of the Angola Basin.
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Brandt, A., Brokeland, W., Brix, S., and Malyutina, M. 2004. Diversity
of Southern Ocean deep-sea Isopoda (Crustacea, Malacostraca)- a
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J., Miihlenhardt-Siegel, U. and Wagele, J.-W. 2005. Diversity of
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of organisms at the deep-sea floor. Cambridge University Press:
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Park, J.-Y. 1999. A new isopod species from the abyssal South Pacific
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of the United Kingdom 79: 1061-1067.
Poore, G.C.B., Just, J., and Cohen, B.F. 1994. Composition and
diversity of Crustacea (Isopoda) of the southeastern Australien
continental slope. Deep-Sea Research 1 41 (4): 677-693.
Sars, G. O. 1870. Nye Dybvandscrustaceer fra Lofoten. Forhandlinger
i Videnskaps-selskabet I Christiania 1869 : 205-221.
Sars, G.O. 1897. Parts 7, 8. Desmosomidae, Munnopsidae (part). Pp.
117-144, pis. 149-164 in: An account of the Crustacea of Norway
with short descriptions and figures of all the species. Bergen
Museum: Bergen.
Sars, G.O. 1899. Parts 13, 14. Cryptoniscidae, Appendix. Pp. 233-270
in: An account of the Crustacea of Norway with short descriptions
and figures of all the species. Bergen Museum: Bergen.
Siebenhaller, J. F., and Hessler, R.R. 1977. The Nannoniscidae
(Isopoda, Asellota): Hebefustis n. gen. and Nannoniscoides
Hansen. Transaction of the San Diego Society of Natural History
19 (2): 17-44.
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Nannoniscidae (Isopoda, Asellota). Transaction of the San Diego
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brevicornh Hansen, 1916 (Isopoda, Asellota, Desmosomatidae)
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Svarvarsson, J. 1988. Desmosomatidae (Isopoda, Asellota) from
bathyal and abyssal depths in the Norwegian, Greenland and
North Polar Seas. Sarsia 73: 1-32.
Wagele, J.-W. 1989. Evolution und phylogenetisches System der
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sea. Systematics and Zoogeography of the Munnidae and the
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Isopoda Asellota. Galathea Reports 6: 320.
Memoirs of Museum Victoria 63(2): 207-213 (2006)
ISSN 1447-2546 (Print) 1447-2554 (On-line)
http://www.museum.vic.gov.au/memoirs/index.asp
A new species of Leongathus from the Tasman Sea collected during the 2003
NORFANZ Expedition (Crustacea: Amphipoda: Phoxocephalidae)
Joanne Taylor
Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia (jtaylor@museum.vic.gov.au)
Abstract Taylor, J. 2006. A new species of Leongathus from the Tasman Sea collected during the 2003 NORFANZ Expedition
(Crustacea: Amphipoda: Phoxocephalidae). Memoirs of Museum Victoria 63(2): 207-213.
Leongathus Barnard & Drummond, 1978 is rediagnosed and a new species Leongathus alannah sp. nov. described
from the Tasman Sea, collected by the 2003 NORFANZ Expedition. A discussion of the problematic placement of the new
species is included in light of the new species exhibiting a combination of characters from Leongathinae and
Harpiniinae.
Keywords Amphipoda, Phoxocephalidae, Harpiniinae, Leongathinae, Harpinia, Proharpinia, Leongathus, NORFANZ, Tasman Sea.
Introduction
A new species of phoxocephalid amphipod was discovered
from 700-800 m depth in the Tasman Sea by the 2003
NORFANZ Expedition. The subfamily and generic placement
of this species is unclear. It agrees with the diagnosis of the
Harpiniinae Barnard and Drummond, 1978, in possessing
article 2 of pereopod 5 of narrow form (3 times as long as
broad), a state unique to that subfamily. On the other hand, it
bears a partly trituritive molar as uniquely seen in the
monotypic Leongathinae, Leongathus nootoo Barnard and
Drummond, 1978. Using Barnard and Karaman’s (1991) key
to Harpiniinae, the new species keys closest to the genus
Proharpinia Schellenberg, 1931 based on the presence of
eyes, non-ensiform antenna 2, thin article 4 of pereopod 6,
ovate propodus of gnathopods and presence of spines on rami
of uropod 2. However, the new species differs from species
belonging to Proharpinia by the presence of setae on the
inner plate of maxilla 1, the unreduced flagellum of antenna
2, the presence of setae on epimeron 1-3, the presence of
apical nails on rami of uropods and partly trituritive molar.
The new species differs from the subfamily diagnosis of
Leongathinae only in the presence of the narrow article 2 of
pereopod 5, the harpiniin synapomorphy. It shares many
characters with the monotypic Leongathus including the
proximal confinement of the ventral setae of antenna 1 article
2, dissimilar gnathopods (gnathopod 2 enlarged) and the
cryptic carpus of gnathopod 2.
This species is an example of an increasing number of new
species of Phoxocephalidae that do not comply with described
genera (Taylor and Poore, 2001). The problem has been
overcome in the past by the publication of new genera that
bear little in the way of new or advanced characters but rather
a recombination of known traits. As a result, approximately
half of all phoxocephalid genera are monotypic. For example,
Linca Alonso de Pina, 1993 was erected based on a single
specimen from the Argentine continental shelf and although
showing some convergence with the Brolginae, its similarity
to Birubius Barnard and Drummond, 1978 best placed it in the
Birubiinae. A preliminary cladistic analysis of the
Phoxocephalidae does not support the monophyly of existing
subfamilies and highlights the incongruity between the
phylogeny and the current generic level classification (Taylor,
2003). Many species have been given generic or subfamily
status because of the few unusual traits they exhibit resulting
in numerous paraphyletic taxa remaining.
Results of the preliminary cladistic analysis by Taylor,
(2003) identified only one strict synapomorphy (pereopod 5
basis of narrow form) defining a clade that includes all genera
of the Harpiniinae. Although further phylogenetic work that
includes all species of this subfamily may indicate otherwise,
results indicate that there is no cladistic support for any of the
nine genera of Harpiniinae and only one, Harpinia can be
diagnosed.
The options for the new species, described here, are: to
erect a new genus of uncertain subfamily affinities on the basis
of a new combination of characters (ignoring potential
synapomorphies with existing genera); place it in Harpiniinae
{Proharpinia or Harpinia Boeck, 1876) on the basis of shared
pereopod 5; or, place it in the same genus as Leongathus
nootoo on the basis of shared partly trituritive molar and the
other characters given above. In the absence of convincing
evidence one way or the other, the last strategy is followed and
Leongathus is rediagnosed.
208
Joanne Taylor
Abbreviations are: A, antenna; H, head; MD, mandible;
MX, maxilla; MP, maxilliped; GN, gnathopod; P, pereopod;
EP, epimera; C, coxa; U, uropod; T, telson; r, right; tl., total
length; NMV, Museum Victoria, Melbourne; AM, Australian
Museum, Sydney, where material is lodged. All dissections
and illustrations follow the methods of Barnard and
Drummond, 1978 whereby the left side of the animal is
illustrated unless otherwise stated. Descriptions of the new
species closely follow that of other species described in
Barnard and Drummond (1978). Original illustrations were
scanned and inked using Adobe Illustrator following the
methods of Coleman, 2003.
Leongathus Barnard and Drummond
Leongathus Barnard and Drummond, 1978: 532. - Barnard and
Karaman, 1991: 617.
Type species. Leongathus nootoo Barnard and Drummond,
1978 (by original designation).
Diagnosis. Rostrum unconstricted. Eyes present, weak or
absent. Antenna 1 peduncular article 2 of medium length,
ventral setae confined proximally. Antenna 2 peduncular article
1 not or scarcely ensiform, article 3 with 2 facial setules, facial
robust setae on article 4 in 1 main row, all robust setae thick,
article 5 ordinary. Right mandibular incisor with 3 teeth, right
lacinia mobilis bifid, flabellate, denticulate, molar partly
triturative, with 7+ large teeth; palpar hump medium, apex of
palp article 3 oblique. Maxilla 1 inner plate with 4— 5 setae, palp
2-articulate. Maxillipeds ordinary, apex of palp article 3 not
strongly protuberant, dactyl elongate, apical nail not distinct.
Gnathopods dissimilar, gnathopod 2 strongly enlarged,
carpus of gnathopod 1 of ordinary length, free, of gnathopod
2 short, cryptic (posterior margin concealed by the abutment
of propodus and merus), palms oblique, gnathopods 1-2
propodus respectively thin and broadened, poorly setiferous
anteriorly. Pereopods 3-4 carpus without posteroproximal
robust setae, propodus with mostly thin armaments. Pereopod
5 basis of narrow form or broad form (basis equal to or greater
than twice width of ischium), but tapering distally, pereopods
5-6 merus-carpus medium to narrow; pereopod 7 unreduced,
article 3 not enlarged, dactyl well developed.
Epimera 1-2 with or without long facial brushes of setae,
with or without posterior setae, epimeron 3 bearing long
setae. Urosomite 3 without dorsal hook. Uropod 1 peduncle
without inter-ramal robust setae, without major displaced
robust seta (seta that is shifted onto the apical margin
disjunctly from the true inner margin), uropods 1-2 rami not
continuously setose to apex, without subapical robust setae or
nails, uropod 1 inner ramus with 1 row of marginal robust
setae. Uropod 2 inner ramus ordinary. Uropod 3 ordinary,
outer ramus longer than peduncle, bearing a 2nd article on
outer ramus (minute in L. alannah ), with 2 long apical setae.
Telson ordinary.
Remarks. To accommodate the new species the diagnosis of
Barnard and Karaman (1991) has been expanded to include the
traits eyes present; antenna 2 peduncular article 1 not ensiform;
right lacinia mobilis flabellate, denticulate; maxilla 1 inner
plate with 4-5 setae; pereopod 5 basis of narrow form or broad
form; pereopods 5-6 merus-carpus medium to narrow; epimera
1-2 with or without long facial brushes of setae, with or without
posterior setae.
Leongathus alannah sp. nov
Figures 1-4
Material examined. Holotype: Australia, Tasman Sea, West Norfolk
Ridge, Wanganella Bank (32°36.29'S, 167°43.59'E), 707 m, beam
trawl, P.B. Berents, 29 May 2003 (stn TAN0308/107), AM P68013
(female, tl. 20 mm).
Paratypes: Type locality, AM P73354 (1 female, tl. 16 mm).
Australia, Tasman Sea, North NorfolkRidge, (28°51.13’S, 167°42.32’E),
812 m, beam trawl, D.J. Bray, 15 May 2003 (stn TAN0308/029) NMV
J53346 (2 females, tl. 13-14 mm). Tasman Sea, North Norfolk Ridge,
(32°36.18'S, 167°47.26'E), 1029 m, beam trawl, P.B. Berents, 29 May
2003 (stn TAN0308/111), AM P66426 (female, tl. 18 mm).
Description. Female. Eyes present. Rostrum unconstricted,
exceeding apex of peduncular article 1 on antenna 1 . Antenna 1
peduncular article 1 about 1.5 times as long as wide, about 2.1
times as wide as peduncular article 2, ventral margin with 4
setae, dorsal apex with 3 setae; peduncular article 2 about 0.5
times as long as peduncular article 1, with 13 ventral and 3
apical setae; primary flagellum with 16 articles, about 0.8 times
as long as peduncle; accessory flagellum with 12 articles.
Antenna 2 not ensiform, peduncular article 4 with 11 robust
setae in 1 apical row, dorsal margin with 2 notches each bearing
1 robust seta and simple setae, ventral margin with 19 long
setae, 3 long ventrodistal robust seta; peduncular article 5 about
0.8 times as long as peduncular article 4, with 7 facial robust
seta, dorsal margin with 4 setae, ventral margin with 15 long
setae; flagellum 1.1 times as long as peduncular articles 4—5
combined, with 16 articles. Mandible, molar partly trituritive,
bearing 13 large teeth; right incisor with 3 teeth; left incisor
with 3 humps in 2 branches; right lacinia mobilis bifid, distal
branch shorter than proximal, flabellate, denticulate, proximal
branch simple; left lacinia mobilis with 5 teeth; right raker 10;
left rakers 11; molars composed of elongate lamina in form of
cone bearing 14 cusps and 2 short plusetae weakly disjunct,
molars covered with fine setae; palp article 1 short, article 2
with 10 medium inner apical setae, 6 outer setae and 4 facial
setae, article 3 about 1.1 times as long as article 2, apex oblique
with 19 robust-slender setae, with 6 basofacial setae. Maxilla 1
inner plate bearing 2 long apical pluseta, 3 apicolateral much
shorter seta; palp article 2 with 18 marginal robust setae.
Maxilla 2 plates extending equally, outer slightly broader than
inner, setation of inner and outer plate as illustrated. Maxilliped
inner plates with 2 large thick apical robust setae, 6 apicofacial
setae; outer plate large with 12 medial and apical robust setae,
13 apical and lateral setae; palp article 1 lacking apicolateral
setae, article 2 with 3 apicolateral setae, medial margin of article
2 strongly setose, article 3 protuberant, with 12 facial setae, 5
lateral setae, nail of article 4 obsolescent, with 2 accessory
setules. Coxa 1 slightly expanded distally; main ventral setae of
coxae 1^4=21-25-18^13, posteriormost seta of coxae 1-3
shortened; anterior and posterior margins of coxa 4 divergent,
posterodorsal corner rounded, posterodorsal margin short,
A new species of Amphipod of the genus Leongathus
209
Figure 1. Leongathus alannah sp. nov., female, holotype, tl. 20.0 mm.
210
Joanne Taylor
Figure 2. Leongathus alannah sp. nov., female, holotype, tl. 20.0 mm.
A new species of Amphipod of the genus Leongathus
211
Figure 3. Leongathus alannah sp. nov., female, holotype, tl. 20.0 mm.
212
Joanne Taylor
Figure 4. Leongathus alannah sp. nov., female, holotype, tl. 20.0 mm.
width-length ratio of coxa 4 almost = 5:6. Long posterior setae
on basis of gnathopods 1-2 and pereopods 3^1 =17-19-21-16,
short posteriors = 2-0-3-2, long anteriors = 12-20-12-6, short
anteriors = 1-0-2-0.
Gnathopod 1 carpus free, gnathopod 2 enlarged, carpus
cryptic. Width ratios of carpus-propodus on gnathopods 1-2 =
9:11 and 8:19, length ratios = 17:23 and 1:5; palmar humps
ordinary, palms oblique. Pereopods 3-4 similar, facial setae on
merus = 9 and 9, on carpus = 10 and 10; main spine of carpus
extending to M. 77 and M. 95, carpus lacking proximoposterior
robust setae; robust setae formula of propodus = 8+4 and 9+9;
acclivity on inner margin of dactyls of pereopods 3—4 weak,
obsolescent, emergent setule short, midfacial pluseta ordinary.
Coxae 5-7 posteroventral setae formula = 23-13-21; merus-
carpus of pereopod 5 medium of pereopod 6 narrow, facial
robust setae rows sparse, facial ridge formula on basis of
pereopods 5-7 = 0-1-1, anterior ridge of pereopod 7 long; width
ratios of basis, merus, carpus, propodus of pereopod 5 = 2:2:2: 1,
of pereopod 6 = 19:10:8:4, of pereopod 7 = 35:12:11:5, length
ratios of pereopod 5 = 15:6:6:8, of pereopod 6 = 18:22:19:22, of
pereopod 7 = 37:14:14:5; basis of pereopod 7 barely exceeding
apex of merus, with row of 9 facial setae along facial ridge, with
ventral setae. Pleopods 1-3 with 2 coupling hooks; articles on
outer rami = 24—22-24; articles on inner rami = 16-17-17.
A new species of Amphipod of the genus Leongathus
213
Epimeron 1 posteroventral corner rounded, anteroventral
margin with 26 medium setae, posterior margin with 12 setae;
epimeron 2 posteroventral corner rounded, with 3 rows of 5-7
anteroventral setae, posterior margin with 12 setae; epimeron
3 posteroventral corner rounded, ventral margin with 6 setae,
posterior margin with 18 setae. Urosomite 1 naked; urosomite
3 without dorsal hook. Uropods 1-2 rami with distinct apical
nails, uropod 1 outer ramus with 3 dorsal robust setae, inner
without robust setae, uropod 2 outer and inner rami with 1
dorsomedial robust seta; uropod 1 peduncle with 11 apicolateral
robust setae, with 3 sets of 1-3 basofacial slender setae, without
apical enlarged robust seta; uropod 2 peduncle with 9 dorsal
robust setae and 5 slender setae; apicolateral corners of
peduncles on uropods 1-2 without comb. Uropod 3 unreduced,
outer ramus longer than peduncle. Uropod 3 peduncle with 5
ventral robust setae, dorsally with 1 lateral robust seta; rami
feminine, inner extending to M. 80 on article 1 of outer ramus,
apex with 1 setae, medial margin with 4 slender setae, lateral
margin naked, article 2 of outer ramus very short, 0.05, bearing
2 long setae, apicomedial margin of article 1 with 4 slender
setae, lateral margin with 4 acclivities, robust setal formula =
1-2-2-2, si ender setae formula = l-0-0-0.Telson length-width
ratio = 17:19, almost fully cleft, each apex wide, rounded,
lateral acclivity broad, shallow, bearing 3 robust setae,
midlateral setules diverse.
Adult male. Unknown.
Distribution. Australia, Tasman Sea, Norfolk Ridge; 707-812 m
depth.
Etymology. Named for my daughter Alannah Taylor with
whom I was pregnant whilst preparing illustrations.
Remarks. The following variations from the holotype were
observed in the paratypes. The main ventral setae of coxae 1-4
= (20-21)-(21-25)-(18-22)-(42-46). Uropod 1 outer ramus
with 3-5 dorsal robust setae, inner ramus 0-3 dorsal robust
setae. Uropod 2 outer ramus with 1-6 dorsal robust setae, inner
ramus with 1 or without dorsal robust setae.
Acknowledgements
Thanks to Gary Poore for reading and suggesting improvements
to the manuscript and encouraging me to take time-out from
isopods to complete this paper. I am grateful to the Australian
Museum for the loan of material. Thanks to Anna McCallum for
assistance with illustrations and direction in Adobe Illustrator.
References
Alonso de Pina, G. M. 1993. Linca pinata , a new phoxocephalid genus
and species (Crustacea: Amphipoda) from the Argentine
continental shelf. Proceedings of the Biological Society of
Washington 106, 497-507.
Barnard, J.L., and Drummond, M.M. 1978. Gammaridean Amphipoda
of Australia, Part III: The Phoxocephalidae. Smithsonian
Contributions to Zoology 245: 1-551.
Barnard, J.L., and Karaman, G.S. 1991. The families and genera of
marine gammaridean Amphipoda (except marine gammaroids).
Parts 1 and 2. Records of the Australian Museum, Supplement 13:
1 - 866 .
Coleman, C.O. 2003. “Digital Inking”: How to make a perfect line
drawing on computers. Organisms, Diversity and Evolution. 3,
Electronic Supplement. 14: 1-14.
Taylor, J. 2003. The impact of phylogenetic analysis on the existing
classification of the Phoxocephalidae (Amphipoda). Unpublished
PhD Thesis, Department of Zoology, The University of Melbourne.
263 pp.
Taylor, J., and Poore, G. C. B. 2001. Descriptions of new species of
Birubius (Amphipoda: Phoxocephalidae) from Australia and
Papua New Guinea with comments on the Birubius -Kulgaphoxus-
Tickalerus-Yan complex. Memoirs of the Museum of Victoria
58(2): 255-295.
Memoirs of Museum Victoria 63(2): 215-255 (2006)
ISSN 1447-2546 (Print) 1447-2554 (On-line)
http://www.museum.vic.gov.au/memoirs/index.asp
Early Silurian phacopide trilobites from central Victoria, Australia
Andrew C. Sandford 1 and David J. Holloway 2
Abstract
Keywords
'PO Box 2682, Chf. ttf. nham , Victoria 3192, Australia (andrewsandiord@hotm ail com ).
2 Museum Victoria, PO Box 666, Melbourne, Victoria 3001, Ausiraha (dhouow@museum .vic.gov.au).
Sandford, A.C. and Holloway, D.J. 2006. Early Silurian phacopide trilobites from central Victoria, Australia. Memoirs of
Museum Victoria 63(2): 215-255.
A diverse component of trilobite faunas in lower Silurian marine strata of central Victoria are members of the suborder
Phacopina, here assigned to 19 species within the Phacopidae, Dalmanitidae and Acastidae. New genera are represented
by the blind acastid Berylacaste berylae gen. et sp. nov. and the phacopid Ivops wallanensis gen. et sp. nov. Other new
species are the phacopids Acernaspis georgei and Ananaspis kenleyi, and the dalmanitids Preodontochile springfieldensis
and Struverial plinthourgos. Two previously described species, Ananaspis typhlagogus Opik, 1953 and Dalmanites
athamas Opik, 1953 [= ‘ Dalmanitina ( EudolatitesY aborigenum Opik, 1953], are revised. Other species are assigned to
Bessazoon, Phacopidellal and indeterminate phacopid and dalmanitid genera. These trilobites are a significant
representation of early Silurian Phacopina on the world scale.
Trilobita, Silurian, Phacopidae, Dalmanitidae, Acastidae, Victoria, Australia, systematics, new taxa
Introduction
Trilobite assemblages occur in lower Silurian (Llandovery and
Wenlock) sedimentary sequences at various localities in
central Victoria, between Costerfield in the north and
Camberwell in metropolitan Melbourne in the south (fig. 1).
Despite the occurrence in these sequences of at least 45
trilobite taxa, 19 of which are members of the Phacopina, only
nine species have been described previously (Table 1). These
include ‘ Phacops ’ typhlagogus Opik, 1953, Dalmanites
athamas Opik, 1953, D. wandongensis Gill, 1948 and
‘ Dalmanitina ( Eudolatites)' aborigenum Opik, 1953. Fossil
collections in Museum Victoria permit the revision of Opik’s
poorly known species and the description of the 16 other
species of Phacopina recorded.
Strata and ages
Stratigraphic terminology employed here follows Rickards and
Sandford (1998), and differs significantly from thatof VandenBerg
et al. (2000) and VandenBerg (2003, figs 5.2, 5.7).
Trilobites of the suborder Phacopina occur at many
horizons in the Llandovery and Wenlock strata of central
Victoria. The stratigraphically lowest, and one of the earliest
recorded trilobites from Victoria (as Asaphus, see Selwyn,
1863), is Phacopidellal sp. from a siltstone underlying the
Lintons Creek Conglomerate Member of the Springfield
Formation at Keilor, at a similar horizon to beds yielding
graptolites of the late Llandovery (Telychian) Spirograptus
turriculatus-Monograptus crispus biozones (Rickards and
Sandford, 1998). Further north, on Deep Creek at Springfield,
the uppermost part of the Springfield Formation, which is
correlated with beds containing graptolites of the Monoclimacis
crenulata Biozone, has yielded a low diversity deep water
fauna of Phacopina dominated by Preodontochile
springfieldensis sp. nov. The overlying sandstones of the
Chintin Formation contain several more diverse trilobite
faunas including three representatives of the Phacopina,
Acernaspis georgei sp. nov., Ananaspis typhlagogusl (Opik,
1953) and Bessazoon sp., suggesting an age range for the unit
of latest Llandovery-early Wenlock.
Known from a single specimen from Camberwell , Struverial
plinthourgos sp. nov. is part of a poorly sampled trilobite fauna
occurring high in the Anderson Creek Formation in the eastern
suburbs of Melbourne, at similar horizons to those yielding
graptolites of the late Wenlock (early Homerian) Cyrtograptus
lundgreni-Testograptus testis Biozone (Rickards and Sandford,
1998). The only other trilobites known from the formation are
three specimens of the blind illaenid Thomastus jutsoni
(Chapman, 1912) and one of an undescribed homalonotid.
Opik (1953) documented a trilobite fauna from concretion-
bearing mudstones of the so-called ‘ Illaenus band’ in the
Costerfield area. Following Thomas (1937) this unit was
traditionally regarded as defining the base of the Wapentake
Formation, but Edwards et al. (1998) placed the ‘ Illaenus band’
at the top of the underlying Costerfield Siltstone. Although
216
Andrew C. Sandford and David J. Holloway
Table 1. List of trilobite taxa in the Llandovery and Wenlock beds of central Victoria. Taxa from the Yan Yean Formation marked with f occur
only in the upper horizons of the unit, which are earliest Ludlow in age.
Yan Yean Formation (late Wenlock-earliest Ludlow)
Proetidae indet.
Decoroproetus sp.f
Maurotarion euryceps (McCoy, 1876)
Encrinuridae indet.
Sthenarocalymene sp. f
Trimerus ( Trimerus ) vomer (Chapman, 1912) f
Ananaspis kenleyi sp. nov. f
Phacopidae gen. indet 1. f
Dalmanites wandongensis Gill, 1948
Kettneraspis hollowayi Sandford, 2000 f
Anderson Creek Formation (Wenlock)
Thomastus jutsoni (Chapman, 1912)
Trimerus? sp.
Struveria? plinthourgos sp. nov.
Bylands Siltstone (Wenlock)
Thomastus aops Sandford and Holloway, 1998
Proetidae indet.
Homalonotid indet.
Ananaspis sp. 1
Ivops wallanensis gen. et sp. nov.
Ananaspis typhlagogus ? (Opik, 1953)
Struveria sp. 2
Berylacaste berylae gen. et sp. nov.
Dicranurus sp.
Selenopeltinae sp.
Odontopleuridae indet.
Wapentake Formation (cont.)
Dalmanites athamas Opik, 1953
Struveria sp. 2
Odontopleuridae indet.
Chintin Formation (latest Llandovery-early Wenlock)
Proetidae sp.
Decoroproetus sp.
Radnor ia sp.
Encrinuridae indet.
Calymene sp.
Calymenella sp. 1
Trimerus ( Ramiotis ) rickardsi Sandford, 2005
Acernaspis georgei sp. nov.
Ananaspis typhlagogus (Opik, 1953)
Bessazoon sp.
Odontopleuridae indet.
Springfield Formation (mid-late Llandovery)
Encrinuridae indet.
Hadromeros? sp.
Calymenella sp. 2
Phacopidella ? sp.
Phacopidae gen. indet. 2
Phacopidae gen. indet. 3
Preodontochile springfieldensis sp. nov.
Costerfield Siltstone (late Llandovery?)
Dalmanitidae indet.
Wapentake Formation (Wenlock)
Thomastus thomastus Opik, 1953
Maurotarion sp.
Trimerus ( Ramiotis ) tomczykowae Sandford, 2005
Ananaspis typhlagogus (Opik, 1953)
Acernaspis ? sp..
Deep Creek Siltstone (early-mid Llandovery)
Calymenidae indet.
Uncertain stratigraphic assignment
Ananaspis sp. 2
Struveria sp. 1
Early Silurian phacopide trilobites from central Victoria, Australia
217
Figure 1. A, map of south-eastern Australia; approximate area of fig.
IB is indicated by a rectangle. B, map of central Victoria showing
localities cited in the text; the areas covered by maps in figs. 4 and 9
are indicated.
Thomas (1940, 1941, 1956) mapped the ‘ Illaenus band’ as a
distinct horizon extending around most of the Costerfield
Dome, Edwards et al. (1998: 50) stated that it is not an extensive
mappable unit. As noted by Rickards and Sandford (1998),
fossils are not confined to a narrow horizon in this part of the
sequence; indeed at localities on the western side of the
Heathcote-Nagambie road both Thomastus (attributed to
Illaenus by Thomas, 1937) and concretions occur 200 m
stratigraphically below and 100 m above the band as mapped
by Thomas (see fig. 9). Hence several concretion-rich beds
seem to be present over a fairly wide strati graphical interval,
and as there is otherwise no distinct lithological change in this
part of the sequence it seems an inappropriate level to define a
lithostratigraphical boundary. Rickards and Sandford (1998)
defined the base of the Wapentake Formation at the top of
sandstones lower in the sequence (the Costerfield Sandstone of
Reeder, 1990), and that is the boundary accepted here. We do
not recognise the ‘ Illaenus band’ as a distinct unit of the
Wapentake Formation, and we use the former term only when
referring to poorly localised material from old collections.
The fauna of the Wapentake Formation is dominated by
the illaenid Thomastus thomastus (see Sandford and Holloway,
1998) but Opik also described three species of Phacopina,
namely ‘ Phacops ’ typhlagogus, Dalmanites athamas and
‘ Dalmanitina ( EudolatitesY aborigenum. Opik considered
that shelly elements of the fauna indicated a late Llandovery
age but noted the post-Llandovery aspect of certain elements
including D. athamas. From a sandy concretion within the
beds Thomas in Opik (1953: 12) identified the graptolites
Monograptus cf. jaculum and Climacograptus hughesr,
however, these identifications cannot be confirmed as the
specimens are lost, so the postulated late Llandovery age
cannot be verified. Rickards and Sandford (1998) considered
that the presence of Ananaspis indicated a Wenlock age for
the fauna, and noted the similarity of the trilobite assemblage
to that of the Wenlock Anderson Creek Formation, with both
faunas having Thomastus as the dominant element and also
containing dalmanitid and homalonotid species.
At Wallan Rickards and Sandford (1998) distinguished a
massive siltstone from the underlying Chintin Formation and
named it the Bylands Siltstone. The unit contains several species
of Phacopina including the blind acastid Berylacaste berylae
gen. et sp. nov., the phacopids Ivops wallanensis gen. et sp. nov.
and Ananaspis typhlagogus ?, the dalmanitid Struveria sp. and,
at a slightly higher horizon, Ananaspis sp. 1. As in the case of
the Wapentake Formation assemblage, the presence of Ananaspis
and the similarity of the Wallan fauna to that of the Anderson
Creek Formation indicates a Wenlock age for the unit.
Dalmanites wandongensis dominates a low diversity
trilobite fauna from the middle horizons of the Yan Yean
Formation at Wandong and Kilmore. Graptolites from these
horizons belong to the latest Wenlock Colonograptus ludensis
Biozone. D. wandongensis ranges into the upper horizons of
the Yan Yean Formation, of earliest Ludlow age
( Neodiversograptus nilssoni Biozone), where it dominates a
more diverse fauna including the phacopid Ananaspis kenleyi
sp. nov. Farther south at Strathewen the upper beds of the Yan
Yean Formation have yielded a single specimen of a phacopid
218
Andrew C. Sandford and David J. Holloway
(Phacopidae gen. indet. 1 herein) with small eyes, a condition
that may be associated with life in a deep-water environment
(Fortey and Owens, 1997: 267; Whittington, 1997: 163).
Faunal comparisons
Trilobites are poorly represented in the Victorian Llandovery
(Table 1). None are known from Rhuddanian strata, and only
Hadromeros, Phacopidella?, an indeterminate flexicalymenine
and an indeterminate encrinurid from the Aeronian-lower
Telychian, making comparison difficult with faunas from this
interval elsewhere. Trilobites are more abundant in upper
Telychian strata. Calymenella occurs in the Calton Hill
Sandstone Member of the Springfield Formation. The low
diversity Preodontochile fauna from the uppermost beds of the
Springfield Formation on Deep Creek compares only to the
monospecific Preodontochile fauna from the Llandovery of
Spain. These Preodontochile faunas are interpreted as
atheloptic assemblages representing a deep water biofacies
whose relationship to other Llandovery deep water assemblages
is unclear. Edgecombe and Sherwin (2001) interpreted the
Aulacopleura-Raphiophorus Biofacies as inhabiting cold,
deep-water, offshore environments, and Chatterton and
Ludvigsen (2004) regarded the Maurotarion- dalmanitid
Biofacies as a deep water fauna representing depths not as great
as the Aulacopleura-Raphiophorus Biofacies. The latter is
represented in Australia by a low-diversity fauna from the early
Telychian Cotton Formation in central New South Wales
(Edgecombe and Sherwin, 2001), containing Aulacopleura,
Raphiophorus and Odontopleura ( Sinespinaspis ), and bears
no similarity to the Llandovery faunas of central Victoria. The
late Telychian to Sheinwoodian Chintin Formation yields the
oldest diverse trilobite faunas in the Silurian of central Victoria.
Eleven taxa are present including Acernaspis georgei sp. nov.,
Ananaspis typhlagogusl (Opik, 1953), Bessazoon sp., Trimerus
( Ramiotis ) rickardsi Sandford, 2005, Radnoria sp., Calymene
sp., Decoroproetus sp. and Calymenella sp. In composition the
Chintin fauna is closely comparable to that of the late Telychian
Richea Siltstone in south-west Tasmania (Holloway and
Sandford, 1993), which also contains species of Acernaspis,
Bessazoon, Trimerus {Ramiotis) and Decoroproetus. Taxa
represented in the Richea fauna but not known from the Chintin
fauna include Maurotarion, Dicranurus, Gravicalymene,
Anacaenaspis and Latiproetus?
The only other Llandovery trilobite faunas known from
Australia occur in the Broken River region of north Queensland.
The early Telychian fauna of the Poley Cow Formation contains
Gaotania, Gravicalymene?, Prostrix, Batocara, Coronocephalus,
Kosovopeltis, Proetus, Warburgella? , Otarion, Maurotarion,
Scharyia, Youngia, Sphaerexochus, Sphaerocoryphe and
Ceratocephala (Holloway, 1994). From the laterally equivalent
Quinton Formation, Lane and Thomas (1978) described
Rhaxeros, Sphaerexochus, Encrinurusl, a possible warburgelline
and an indeterminate calymenid, and Opik (in White, 1965: 43)
listed Encrinurus, Proetus? b Onycopyge or Sphaerexochus, and
undetermined scutelluids. These faunas thus show little similarity
in generic composition to the Llandovery faunas of Victoria and
Tasmania, and they further differ in lacking phacopids,
homalonotids and dalmanitids, and in the presence of scutelluids
and illaenids. These differences may reflect biogeographical
provincialism or environmental preferences. Rickards, Wright
and Sherwin {in Pickett et al., 2000: 131) noted an analogous
contrast between the Llandovery graptolite faunas of north
Queensland and those of central Victoria, the latter having much
stronger European affinities (98% of species) than the
Queensland faunas (75% of species). Rickards et al. considered
that the composition of the Victorian graptolite faunas indicate
stronger interaction with the Americas rather than with the
Uralian-Cordilleran Province.
Trilobites represented in the lower and middle Wenlock of
central Victoria include species of Thomastus, Maurotarion,
Decoroproetus, Trimerus {Ramiotis), Ivops gen. nov.,
Ananaspis, Dalmanites, Struveria, Berylacaste gen. nov.,
Dicranurus, and indeterminate Encrinuridae and
Odontopleuridae. Faunas of similar age are present in the
Rosyth Limestone and the laterally equivalent Boree Creek
Formation in the Orange district, central western New South
Wales, but of the 29 genera from those formations listed by
Holloway {in Pickett et ah, 2000: 163) only Ananaspis,
Decoroproetus and Dicranurus also occur in central Victoria,
and dalmanitids, homalonotids and acastids are conspicuously
absent from the New South Wales faunas.
Differences in faunal composition between central Victoria
and more northerly regions continued into the late Wenlock
and earliest Ludlow. The only trilobite genus in common
between the Yan Yean Formation and the Walker Volcanics of
the Canberra area (see Holloway in Pickett et al., 2000: 164) is
Sthenarocalymene {= Gravicalymene? of Holloway’s list), and
the latter fauna lacks phacopids and dalmanitids but includes
scutelluids, illaenids, harpetids, cheirurids and staurocephalids
that are absent from the Yan Yean. Diverse trilobite faunas of
mid-Wenlock to early Ludlow or possibly even late Ludlow
age occur in the Orange-Molong district in limestone bodies
within the Mirrabooka Formation, and in the laterally
equivalent Borenore and Molong limestones. Illaenids and
scutelluids dominate the fauna, which otherwise resembles
that from the underlying Boree Creek Formation except that
phacopids are absent (Holloway and Lane, 1998).
The general absence of phacopid and dalmanitid trilobites
in the Wenlock faunas of the Canberra and Orange districts
(apart from the occurrence of Ananaspis in the Boree Creek
Formation and Rosyth Limestone) is at least partly due to
environmental preferences. The Silurian sequences of those
districts were deposited on a series of offshore highs and
troughs whereas those of central Victoria were deposited on
the continental margin. The fauna of the Borenore Limestone
and its stratigraphical equivalents is similar to those occurring
in lithologically similar pure limestones of Silurian age
elsewhere in the world (see Lane, 1972; Thomas and Lane,
1998). Mikulic (1981) reported that in middle and late Silurian
reef limestones of North America the reef core is dominated
by illaenids, scutelluids, lichids and cheirurids, and the reef
fringes are characterised by assemblages of cheirurids,
calymenids and encrinurids. He interpreted rare phacopid and
dalmanitid trilobites in the reef fringes as temporary migrants
from non-reef areas.
Early Silurian phacopide trilobites from central Victoria, Australia
219
After the Wenlock members of the Phacopina were a
significant component of trilobite faunas in New South Wales
as in Victoria. The fauna of the late Ludlow Rosebank Shale in
the Yass district includes both phacopids and dalmanitids
[ Ananaspis crossleii (Etheridge and Mitchell, 1896), Dalmanites
meridianus (Etheridge and Mitchell, 1896)]. In the Early
Devonian, Echidnops, Paciphacops, Lochkovella, Odontochile,
several genera not belonging to the Phacopina, and at least one
species ( Sthenarocalymene sp. A, see Chatterton, Johnson and
Campbell, 1979; Holloway and Neil, 1982) were present in
both central Victoria and New South Wales; however, continuing
limitations on faunal exchange is evident in the absence of
homalonotids and acastids in New South Wales, and in the
absence of harpetids in central Victoria.
Systematic palaeontology
With the exception of the partly mineralized exoskeletons
preserved in concretions of the Wapentake Formation, all
trilobite specimens figured here are preserved in mudstones
and sandstones as internal and external moulds. For
photography, internal moulds were coated with colloidal
graphite, latex peels were made from external moulds, and all
were whitened with ammonium chloride. Specimens housed
in Museum Victoria are registered with the prefix NMV P, and
those housed in the collections of Geoscience Australia,
Canberra, are registered with the prefix CPC. Two missing
specimens of Ananaspis typhlagogus are documented with
registration numbers of the old Geological Survey of Victoria
collection, prefixed GSV. Trilobite localities with the prefix
PL are documented in the Museum Victoria invertebrate
palaeontology locality register.
Order Phacopida Salter, 1864
Suborder Phacopina Richter, Richter and Struve, 1959
Superfamily Phacopoidea Hawle and Corda, 1847
Family Phacopidae Hawle and Corda, 1847
Remarks. In his seminal review of the Phacopidae in which he
established the genera Acernaspis and Ananaspis, Campbell
(1967) proposed a classification of the family that he considered
to be ‘horizontal’ in the sense of Simpson (1961). His approach
was criticised by some workers (e.g. Eldredge, 1973: 292) as
creating paraphyletic and polyphyletic taxa, an outcome that
was acknowledged by Campbell (1977: 26). Nevertheless,
Acernaspis and Ananaspis have been universally recognised,
and the former has come to be accepted as monophyletic
(Ramskold and Werdelin, 1991: 61).
Ananaspis continues to present more problems than
Acernaspis because it embraces greater variability in
characters such as the width of the cheeks in relation to that of
the glabella, the relative lengths (exsag.) of L2 and L3, the size
of the eye and the position of its lower margin in respect to the
lateral border furrow, the depth of the vincular furrow medially
and the strength of its notching laterally, the glabellar sculpture
(but in relation to this character see remarks on Acernaspis
below), and the width and degree of taper of the pygidial axis.
These characters, many of which were regarded by Campbell
(1967) and later workers as diagnostic of Ananaspis, are
expressed to differing degrees and in a variety of combinations
in the species assigned. Ramskold and Werdelin (1991)
restricted Ananaspis to a small number of species of Ludlow
and Lochkovian ages [A. orientalis (Maksimova, 1968), to
which Ramskold and Werdelin ascribed a possible Ludlow
age, is from the Kokbaital Horizon of Central Kazakhastan,
now known to be early Lochkovian; Talent et al., 2001: 61].
Several other species previously assigned to Ananaspis, and of
late Llandovery-late Wenlock age, were referred to by
Ramskold and Werdelin as Ananaspis ’ or incertae sedis, and
said to represent ‘. . .a number of monospecific (or nearly so)
genera ... between Acernaspis and Ananaspis" (Ramskold and
Werdelin, 1991: 56). We are not in complete agreement
between ourselves on the relationships of these species which
include ‘ Phacops ’ typhlagogus Opik, 1953, redescribed below.
However, it is difficult to identify any consistently developed
characters that could be used to distinguish such species from
others assigned to Ananaspis s.s. by Ramskold and Werdelin,
or to be sure that the latter species are more closely related to
each other than to some of the species excluded from Ananaspis
by those authors. Consequently Ananaspis is more broadly
conceived herein than by Ramskold and Werdelin.
Acernaspis Campbell, 1967
=Eskaspis Clarkson et ah, 1977; -Murphycops Lesperance,
1968
Type species. Phacops orestes Billings, 1860 from the Jupiter
Formation (Llandovery), Anticosti I., Quebec, by original
designation.
Remarks. Chatterton and Ludvigsen (2004: 39) discussed the
possible synonymy of Acernaspis with Portlockia McCoy,
1846 (type species P. sublaevis) from the Wenlock of western
Ireland and gave notice of their intention to apply to the
International Commission of Zoological Nomenclature to
suppress the latter generic name as a nomen oblitum. We would
support such an application as being in the best interests of
nomenclatural stability.
Chatterton and Ludvigsen (2004) considered Acernaspis
mimica Lesperance and Letendre, 1982 to be a synonym of A.
orestes, and also noted that A. superciliexcelsis Howells, 1982
differs only in very minor respects from orestes and might be
considered synonymous. These conclusions are in accord with
the cladistic analysis of Acernaspis presented by Ramskold
and Werdelin (1991, fig. 5), in which these three taxa are
grouped together. Curtis and Lane (1998) considered a number
of other species to be synonyms of orestes including A.
quadrilineata (Angelin, 1851), A. konoverensis Mannil, 1970,
A. elliptifrons (Esmark, 1833) and A. sororia Ramskold, 1985;
however, Ramskold and Werdelin’s cladistic analysis shows
these species and orestes grouped quite separately. Chatterton
and Ludvigsen arrived at a similar conclusion, rejecting Curtis
and Lane’s synonymy and listing a number of characters
distinguishing orestes and quadrilineata, emphasising minor
differences in the lens formula and the proportions of the
glabella.
220
Andrew C. Sandford and David J. Holloway
Acernaspis rubicundula Ramskold, 1985, from the lower
Wenlock of Sweden, was interpreted by Ramskold as
intermediate between Acernaspis and Ananaspis with respect
to morphological trends that he recognised between his late
Llandovery species Acernaspis sororia and Ananaspis stokesii
(Milne Edwards, 1840) from the British Wenlock. He
emphasised the glabellar width, the short (exsag.) L2 and the
glabellar tuberculation as characters of rubicundula typical of
Ananaspis, and remarked that stokesii differs from rubicundula
in few features of which only the strongly forwardly expanding
glabella and deep lateral border furrow (on the fixigena) were
mentioned. Neotenic changes significant in the evolution of
Ananaspis from Acernaspis, including a decrease in the length
(exsag.) of L2 and the development of glabellar tuberculation,
were outlined by Ramskold (1988). In this context he
interpreted rubicundula as a sister-taxon to all other Acernaspis
species, exhibiting the first of these neotenic changes. In their
cladistic analysis of Acernaspis, Ramskold and Werdelin
(1991) discovered that their consensus tree was essentially
unresolved with the inclusion of rubicundula but on its removal
an almost fully resolved consensus tree was obtained.
Nevertheless they retained rubicundula in Acernaspis, and we
agree that the majority of its characters clearly ally the species
with that genus. However, in view of its unusual features, we
assign rubicundula to Acernaspis with question, together with
a somewhat similar species described below as A.l sp.
Acernaspis georgei sp. nov.
Figures 2, 3A-B, D-E, H-I
Acernasp is .— Rickards and Sandford, 1998: 750.
Type material. Holotype NMV P515 (cephalon) from PL1964,
Geological Survey locality B25, Springfield. Paratypes NMV P138259
(cephalon), NMV P138262 (pygidium), NMV P138278 (pygidium),
NMV P147765 (cephalon), NMV P147766 (cephalon), NMV P147796
(pygidium), NMV P312816 (cephalon) from PL256, Wallan. Paratype
NMV P138271 (cephalon) from PL598, type section of the Chintin
Formation, Springfield.
Other material. NMV P138260-P138261, NMV P147764, NMV
P147767-P147768, NMV P147788-P147795, NMV P147797-P147806
from PL256, Wallan. NMV P139447-P139453 from ‘Lancefield’
(unknown locality in the Parish of Goldie). For localities see Thomas
(1960), VandenBerg (1991), Sandford and Rickards (1999) (fig. 1),
Sandford (2005) (fig. 11) and (fig. 4).
Derivation of name. After ACS’s late father.
Diagnosis. Anterior cephalic border comprising 2.5% sagittal
cephalic length in dorsal view. Lateral glabellar furrows
moderately impressed, S2 extending close to axial furrow,
posterior branch of S3 weakly convex forward, anterior branch
placed posteriorly, with midlength opposite anterior margin of
palpebral lobe. Palpebral furrow moderately to deeply impressed,
distinct palpebral rim furrow, eye length (exsag.) 45% sagittal
cephalic length, postocular area with length (exsag.) 5% sagittal
cephalic length, visual surface with 15 files of lenses with up to
6 (mostly 5) lenses per file. Genal angle obtusely angular,
lacking prominent point or genal spine. Second and third
pygidial pleural furrows moderately impressed.
Description. Cephalon semicircular, in frontal view with anterior
margin weakly arched. Glabella weakly convex (sag., tr.), not
reaching anterior cephalic margin in dorsal view, maximum
width across frontal lobe approximately equal to cephalic length,
50% maximum cephalic width and 160% occipital width (tr.).
Axial furrow wide and very deep, weakly converging between
posterior margin and LI , diverging at 60° between LI and a point
opposite midlength of palpebral lobe, thereafter diverging
forward at 20°. Anterior margin gently rounded in front of
glabella, forming arc centred on posterior margin. Occipital ring
with length (sag.) 15% cephalic length and width (tr.) 33%
cephalic width, with exsagittal furrow impressed in anterior
margin defining obliquely directed lateral lobes. Occipital furrow
deeply impressed. LI high, lateral node isolated by deep exsagittal
furrow. SI deep laterally, weak across median part of glabella. S2
and S3 moderately impressed on external surface. S2 directed at
about 10° to transverse, straight, its inner end situated more or
less opposite 40% glabellar length from posterior. Posterior
branch of S3 with anteriormost point more or less opposite 55%
sagittal glabellar length from posterior. Anterior branch of S3
oriented at 40° to exsagittal line, weakly sigmoidal. L2 and L3
equal in length (exsag.). Anterior border furrow moderately
impressed. Anterior border wide. Eye placed with midlength
opposite 38% sagittal cephalic length from posterior. Palpebral
area high, convex. Palpebral lobe steeply inclined, flat (tr.), raised
above palpebral area. Visual surface large, with up to 71 lenses,
formula (NMV P138259, fig. 2D, H) 5 5 6 5 5 5 5 5 5 5 4 5 5 4 3.
Posterior border furrow deep adaxially, terminating abruptly
abaxially at a point in line (exsag.) with distal margin of eye.
Posterior border narrow adaxially, widening slightly abaxially.
Librigenal field concave below eye, not distinctly differentiated
from lateral border furrow. Posterior branch of facial suture
skirting posterior margin of eye abaxially, crossing genal area
directed anterolaterally at about 25° to transverse, reaching a
point opposite 33% sagittal cephalic length, deflected backwards
at same angle abaxially. Cephalic doublure with vincular furrow
deep throughout and strongly notched laterally. Cephalic
sculpture finely granulate.
Pygidium lenticular in dorsal outline. Axis comprising
85% sagittal pygidial length and 30% maximum pygidial
width anteriorly, with 6 rings that are well defined in anterior
part of axis and poorly defined in posterior part, terminal
piece merging with postaxial region. Pleurae with 5 pleural
furrows, anterior one narrow (exsag.), deep, reaching 75%
distance to margin, remaining pleural furrows successively
shallower, posteriormost one very weak. Interpleural furrows
very shallow. Border and furrow not defined.
Remarks. Acernaspis georgei occurs at several localities in the
Chintin Formation, although its relative abundance, faunal
associates, preservation and associated lithology vary markedly.
In Slab Hut Creek at PL256, Wallan, the species is common
(relative abundance 44%) and together with Calymene (relative
abundance 35%) dominates the fauna preserved almost entirely
as fragmentary cephala and pygidia. The trilobites at PL256
occur within bioclastic sandstone coquinas which contain
abundant disarticulated crinoid elements, gastropods, bryozoans
and clasts of siltstone incorporated into the coquina in the plastic
Early Silurian phacopide trilobites from central Victoria, Australia
221
Figure 2. Acernaspis georgei sp. nov. A-C, E-F, holotype NMV P515, cephalon (fragment), from PL1964, Springfield; A-C, x 5; E, x 6.5; F, x 6. D,
H, paratype NMV P138259, cephalon, from PL256, Wallan; D, x 5; H, x 4. G, paratype NMV P138261, pygidium, x 8, from PL256, Wallan. I,
paratype NMV P138271, cephalon (fragment), x 7, from PL598, Springfield. (C, D are internal moulds).
state, presumably as rip-up clasts. The coquinas are bedded but
form large irregular bodies within massive siltstones, with which
they have sharp contacts. At ‘Lancefield’ the species occurs
(relative abundance 14%) in a fauna dominated by Bessazoon
sp. and Calymene sp., preserved as isolated and generally
undamaged exoskeletal elements in medium- grained sandstone.
The species can be confidently assigned to Acernaspis.
Significant characters in this assignment include the low
glabellar profile, non-tuberculate cephalic ornament, long
(exsag.) L2 (equal in length to L3), wide anterior border visible
in dorsal view, poorly defined lateral cephalic border furrow,
short postocular area, strong notching of the vincular furrow
laterally and the weak expression of the pygidial interpleural
furrows and the fourth and fifth pleural furrows.
A poorly known, unnamed species of Acernaspis was
described by Holloway and Sandford (1993) from the late
Llandovery Richea Siltstone of Tasmania. The present species
differs from the Tasmanian one in having a longer postocular
area, a deeper palpebral furrow and a longer anterior cephalic
border. ‘ Phacops ’ macdonaldi Fletcher, 1950, from the upper
Llandovery to lower Wenlock of the Orange district, New
South Wales, was tentatively assigned to Acernaspis by
Sherwin (1971), but is now assigned to Ananaspis along with
its junior subjective synonym AcernaspisT oblatus [sic]
Sherwin (see discussion of Ananaspis typhlagogus).
Acernaspis georgei is most similar to the type species A.
orestes. The two species share a relatively long (sag.) anterior
cephalic border, a long postocular area and a non-spinose
genal angle. A. georgei has one fewer lens file in the eye but
otherwise shares a similar lens formula to small-eyed morphs
of orestes, which have 16 files of up to six (mostly five) lenses
per file. The species differ most notably in the arrangement of
the lateral glabellar furrows and in the deeper palpebral
furrows of georgei.
222
Andrew C. Sandford and David J. Holloway
Figure 3. A-B, D-E, H-I, Acernaspis georgei sp. nov. A, paratype NMV P147766, incomplete cephalon, x 4, from PL256, Wallan. B, NMV
P139448, damaged cephalon with cephalic doublure exposed, x 3.25, from ‘Lancefield’. D, I, paratype NMV P312816, incomplete cephalon from
PL256; D, x 5; I, x 4.5. E, NMV P139451, pygidium, x 4, from ‘Lancefield’. H, paratype NMV P138262, pygidium, x 5, from PL256, Wallan. C,
F-G, J-K, Ananaspis typhlagogus (Opik, 1953) from PL6361, Springfield. C, G, J-K, NMV P312813, cephalon; C, J, K x 3.75, G x 4.75. F, NMV
P312814, thorax, x 5. (A-B, H are internal moulds).
Species less closely related to A. georgei and A. orestes
include Mannil’s (1970) Estonian species A. semicircularis, A.
estonica, A. sulcata, A. rectifrons and A. incerta, and the
Scottish A. xynon Howells, 1982, all of which are easily
distinguished in having acutely angular genal angles or short
genal spines. In the depth and lateral extension of S2 (to a
point close to the axial furrow), A. georgei resembles A.
besciensis Lesperance and Letendre, 1982 from the Rhuddanian
of Anticosti Island, Canada {-A. salmoensis Lesperance, 1988
fide Chatterton and Ludvigsen, 2004), the oldest known
species of the genus. A. besciensis differs from A. georgei in
many other features, notably in having fewer lenses per file but
more files in the eye (up to four lenses in 16 files), a shorter (tr.)
and more forwardly convex S3, a narrow preoccipital ring and
a shallower preglabellar furrow.
Acernaspis ? sp.
Figures 5A-D
Material. NMV P139804 (incomplete cephalon) from PL385,
Costerfield. Wapentake Formation. For locality see fig. 9.
Description. Anterior and anterolateral cephalic margins
moderately arched upwards medially in anterior view. Glabella
weakly convex (sag., tr.), low anteriorly, sides of composite
lobe diverging at about 55° between level of S2 and outer end of
preglabellar furrow. SI curving forward adaxially, connected
medially by wide (sag.), shallow depression. S2 moderately
impressed, convex forwards, subparallel to posterior branch of
S3, its inner end connected to SI by shallow exsagittal furrow.
L2 about 75% length (exsag.) of L3 adaxially. S3 distinctly
shallower than S2; posterior branch of S3 not extending as far
Early Silurian phacopide trilobites from central Victoria, Australia
Figure 4. Fossil localities of the Springfield Formation, Chintin
Formation and Riddell Sandstone in the Springfield area, 65 km
NNE of Melbourne; the area covered by the map is indicated on fig.
1. The geology of this area given by Rickards and Sandford (1998)
(fig. 6) is incorrect. The presumed faunal similarities with the
Bylands Siltstone are not substantiated on description of the fauna
herein. On lithological and apparent faunal similarities Rickards
and Sandford erroneously correlated the siltstone at PL1369 with the
Bylands Siltstone and showed the Chintin Formation underlying it.
Additional field observations show that the Chintin Formation crops
out upstream from and hence overlies PL1369, which lies in the
uppermost beds of the Springfield Formation, as mapped by
VandenBerg (1991).
223
adaxially and abaxially as S2; anterior branch not quite meeting
posterior branch, oriented at about 60° to sagittal line and very
weakly curved, meeting axial furrow anteriorly. Preglabellar
furrow moderately impressed, continuous medially. Anterior
cephalic border of almost uniform length (sag., exsag.) except
abaxially. Preserved portion of eye (more than anterior half)
with 12 files of lenses, formula (from anterior) 456767666
6 7 6 ..., sclera depressed. Palpebral lobe raised high above
palpebral area, with shallow but distinct rim furrow; anterior
part of palpebral furrow deep and narrow. Librigenal field
concave below eye and merging with lateral border furrow.
Anterior branch of facial suture cutting across anterolateral
comer of glabella (fig. 5B-C). Vincular furrow moderately
impressed medially, deep laterally with strong notching. Medial
part of cephalic doublure not raised as high above vincular
furrow anteriorly as posteriorly, giving doublure a step-like
profile or ‘bevelled’ morphology. Composite glabellar lobe with
sculpture of low, perforate tubercles of small to moderate size.
Remarks. The species is represented by a single, incomplete
cephalon lacking the posterior portion more or less behind a
transverse line through the medial part of SI. The specimen
differs from Ananaspis typhlagogus from the same formation
in that the glabella is much more weakly convex, not as strongly
curved in anterior outline and does not overhang the anterior
border; the anterior border is wider (sag., exsag.); S2 is deeper,
more strongly curved (not transverse adaxially), runs subparallel
to the posterior branch of S3, and is connected to SI adaxially
by a shallow exsagittal furrow; the posterior branch of S3 is
narrower (tr.) and the anterior branch longer; the preglabellar
furrow and preserved portion of the palpebral furrow are
deeper; and the visual surface does not overhang the lateral
border as strongly in dorsal view and is higher, with more
lenses per file (up to seven instead of five as in typhlagogus).
Although it could be suggested that the low glabellar profile of
the specimen is due to tectonic flattening, we consider that this
is unlikely because of the other differences from typhlagogus
listed, especially those in the eyes.
Acernaspis ? sp. differs from most species of Acernaspis
except A.l rubicundula Ramskold, 1985, from the lower Wenlock
of Sweden, in having tuberculate sculpture on the composite
glabellar lobe (see discussion above under genus). A.l
rubicundula differs from A.? sp. in that S2 is not as curved, does
not extend as close to the sagittal line and is not connected to SI
adaxially by a weak longitudinal furrow; L2 is much shorter
(about half length of L3 adaxially); S3 is deeper, with a shorter
anterior branch not extending as close to the posterior branch;
and S2 and S3 have weakly raised rims. These differences
suggest that the Victorian species may not be most closely related
to rubicundula, despite the similarity in glabellar sculpture.
Ananaspis Campbell, 1967
Type species. Phacops fecundus Barrande, 1846 from the
Kopanina Formation (Ludlow), Kolednfk, Bohemia, by original
designation.
Diagnosis. Glabella moderately to strongly convex (sag., tr.),
vertical or slightly overhanging anteriorly in dorsal view.
224
Andrew C. Sandford and David J. Holloway
Figure 5. A-D, Acernaspisl sp., NMV P139804, cephalon, x 4, from PL385, Costerfield. E-G, Phacopidae gen. indet. 3, NMV P139353, enrolled
exoskeleton from PL1369, Springfield; E, x 4.0; F, x 4.5.
Composite lobe expanding strongly forward, maximum width
twice width at LI or a little more. L2 commonly significantly
shorter (exsag.) than L3. Eye of moderate to large size, lower
edge situated in or slightly above lateral border furrow anteriorly
and distant from lateral border furrow posteriorly, visual surface
lacking strongly raised sclera (may be slightly thickened dorsally).
Palpebral area as high as or higher than palpebral lobe. Fixigenal
portion of lateral border furrow deep and continuous with
posterior border furrow. Vincular furrow shallow to moderately
impressed medially, commonly rather weakly notched laterally.
Glabellar sculpture of bimodal tubercles lacking perforations and
with superimposed and interspersed granules, doublure finely
and densely granulate. Hypostome with short (sag.) posterior
border having 3 angular points on margin. Pygidial pleural
furrows deep and wide (exsag.), interpleural furrows distinct.
Remarks. This diagnosis distinguishes Ananaspis from a number
of closely related genera recognised since the work of Campbell
(1967). Echidnops Sandford, 2002 is the closest to Ananaspis
but differs not only in having characteristic occipital and thoracic
axial spines but also a much more deeply incised vincular furrow
medially. Paciphacops Maksimova, 1972 is distinguished
especially by the strongly raised sclera on the visual surface and
the perforate glabellar tubercles, but in addition LI is more
depressed medially, the glabella is more raised and subquadrate
in anterior profile, and the vincular furrow is deep medially.
Lochkovella Chlupac, 1972 (see also Sandford, 2004) differs
from Ananaspis in that the cephalic tuberculation is finer and of
more uniform size, the eye is situated very low on the cheek
with its lower edge indenting the lateral cephalic border, the
hypostome has a long (sag.) posterior border with five marginal
denticles, the pygidial interpleural furrows are not as deep, and
the pygidial granulation is coarser. Similar distinctions can be
made with Nephranomma Erben, 1952 (see Sandford, 2003)
which further differs from Ananaspis in lacking a vincular
furrow medially and in having a distinctive scaly sculpture on
the cephalic doublure. See the remarks on Ivops gen. nov. for
comparison of that genus with Ananaspis.
Early Silurian phacopide trilobites from central Victoria, Australia
225
Ananaspis kenleyi sp. nov.
Figures 6A-H, I?, J-K
Ananaspis .— Rickards and Sandford, 1998: 753.
Ananaspis ’ sp. nov.— Sandford, 2000: 199.
Type material. Holotype NMV P136821 (cephalon) from PL375,
Kilmore East. Paratype NMV P136815 (cephalon) from PL377,
Kilmore East. Paratypes NMV P136819 (cephalon), NMV P136820
(pygidium), NMV P140154 (enrolled thoracopygon), NMV P140155
(cephalon), NMV P140156 (cephalon) from PL375, Kilmore East. Yan
Yean Formation.
Other material. NMV P137165 from PL286, Williams locality
F22, Wandong. NMV P136801, NMV P137142 from “Wandong”
(exact locality unknown). NMV P136816 from PL377, Kilmore East.
NMV P136802, NMV P138648 from PL380, Geological Survey
locality Bbl8, Wandong. NMV P137146 from PL1692, Wandong.
NMV P140403 from PL1691, Kilmore East. NMV P139409 from
PL1342, Kilmore East. NMV P140598 from PL1699, Upper Plenty.
Yan Yean Formation. For localities see Taylor (1864), Williams (1964)
(fig. 2) and Sandford (2006) (figs 2-3).
Derivation of name. After Peter Kenley, formerly of the
Geological Survey of Victoria, who made valuable fossil
collections from the Silurian of central Victoria.
Diagnosis. Eye large, length (exsag.) 44% sagittal cephalic
length, placed with midlength opposite 45% sagittal cephalic
length from posterior, visual surface with about 20-21 files of
up to 7 lenses, postocular fixigenal field short, 7% sagittal
cephalic length. Posterior border furrow narrowing and
shallowing abaxially behind eye, terminating almost in line
(exsag.) with lateral extremity of visual surface. Lateral border
furrow indistinct, especially on librigena. Vincular furrow
wide and shallow medially. Glabella with low tubercles of
moderate size and density. Pygidium with 6-7 axial rings, 2nd
to 5th ones with successively diminishing pseudo-articulating
half rings. Five distinct pygidial pleural furrows, first 3 much
deeper than remainder, last one very shallow, interpleural
furrows very weak.
Description. Glabella narrow, maximum width approximately
80% sagittal length and 175% occipital width, placed level with
70% sagittal length from posterior. Axial furrow weakly
converging forward adjacent to occipital ring and LI, wide and
deep in front of occipital ring, diverging forward at about 65°
adjacent to composite lobe. Occipital ring comprising 12%
cephalic length sagittally, with small notch in anterior margin
defining obliquely oriented lateral lobe. Occipital furrow deep,
transverse medially. Medial part of LI as high as composite
lobe, lateral node small, depressed, isolated by deep exsagittal
furrow. Glabellar width at LI 85% occipital width. SI
shallowing rapidly adaxial to inner edge of LI node, expanding
forwards to become a wide (sag.), triangular depressed area. S2
and S3 variably impressed. S2 weakly arcuate, reaching axial
furrow, oriented at about 15° to transverse, anteriormost point
opposite 38% sagittal cephalic length from posterior. Posterior
branch of S3 strongly arcuate, anteriormost point opposite 55%
sagittal glabellar length. Anterior branch of S3 sinusoidal,
reaching axial furrow anteriorly. Length of L2 (exsag.) 70%
length of L3 and 11% sagittal cephalic length. Front of glabella
very high, strongly convex (tr.) in anterior view, in dorsal view
forming strongly rounded arc centred at glabellar midlength
(sag.) and overhanging anterior border. Preglabellar furrow
shallow and wide. Palpebral area high, of low convexity,
palpebral furrow moderately impressed, continuous with wide,
moderately impressed sutural furrow behind eye. Palpebral
lobe arcuate, of uniform width, raised above palpebral area.
Lens formula from anterior (NMV P140136) 3567677676
676???????, about 120 lenses, visual surface without
raised sclera. Librigenal field below eye concave, not distinctly
separated from border. Medial part of cephalic doublure 30%
sagittal cephalic length, anterior half very weakly convex (sag.)
and posterior half weakly concave, vincular furrow running
adjacent to anterior margin and separated from it by narrow
(sag.) rim. Hypostomal suture transverse medially.
Thoracic axis narrowing weakly backwards, comprising
about 30% segmental width (tr.). Axial rings with strong
lateral nodes defined by posteriorly divergent furrows that
notch front of segments. Pleurae horizontal and transverse
adaxial of fulcrum, steeply inclined lateral to fulcrum and
widening to well-rounded tips. Deep pleural furrow situated at
midlength (exsag.) of segment at fulcrum, on anterior segments
terminating distally at edge of articulating facet but extending
onto facet on posterior segments.
Pygidium lenticular in dorsal view. Axis comprising 80%
sagittal pygidial length and 25% maximum pygidial width
anteriorly, tapering uniformly backwards. Axial rings strongly
raised anteriorly, decreasing in height, length and definition
posteriorly. First 5 rings comprise 72% axial length. Posterior
end of axis broadly rounded and well defined. Pleural field
weakly convex with weakly defined border. Anteriormost
pleural furrow deep and wide, posterior furrows successively
narrower (exsag.) and shallower.
Remarks. Ananaspis kenleyi occurs in the uppermost beds of
the Yan Yean Formation at a number of localities between
Upper Plenty and Kilmore but is nowhere common. It occurs
together with the abundant Dalmanites wandongensis, and the
rare Trimerus ( Trimerus ) vomer Chapman, 1912 and
Sthenarocalymene sp.
Sandford (2000) considered Ananaspis kenleyi (as
‘ Ananaspis ’ sp. nov.) to be most closely related and possibly
ancestral to the distinctive A. woiwurrungi from the overlying
Melbourne Formation. Sandford listed differences between
the two species, kenleyi having stronger glabellar tuberculation
that extends further posteriorly, shorter eyes, a deeper medial
section of the vincular furrow and deeper pygidial pleural
furrows (in woiwurrungi only the first to third pleural furrows
are distinct on the external surface). Other differences in
kenleyi are the narrower glabella (cf. maximum glabellar
width 90% maximum cephalic width and 200% occipital
width in woiwurrungi ), shorter L2 (length 80% length of L3
and 15% sagittal cephalic length in woiwurrungi ), fewer lenses
in the eye (cf. 22 files with up to 8 lenses per file in woiwurrungi ),
slightly longer postocular fixigenal field (cf. length 5% sagittal
cephalic length in woiwurrungi ), indistinct lateral border
furrow (shallow in woiwurrungi ), shorter pygidial axis and
less distinct interpleural furrows.
226
Andrew C. Sandford and David J. Holloway
Figure 6. A-H, I?, J-K, Ananaspis kenleyi sp. nov. A, holotype NMV P136802, pygidium, x 6, from ‘Broadhursts Creek’, Wandong. B-C, paratype
NMV P136821, cephalon x 4, from PL375, Kilmore East. D, paratype NMV P140156, cephalon, x 4.5, from PL375, Kilmore East. E, paratype NMV
P140403, cephalon (fragment), x 2.5, from PL1691, Kilmore East. F, paratype NMV P137165, cephalon, x 3.5, from PL286, Wandong. G, paratype
NMV P140154, thoracopygon with pygidium displaced and inverted, view of pygidium, x 4.5. H, paratype NMV P136820, pygidium with circular
textural markings, x 4, from PL375, Kilmore East. I, paratype NMV P136815 cephalon, x 3, PL377, Kilmore East. J, paratype NMV P136819,
exoskeleton with displaced cephalon, view of doublure, x 4, PL375, Kilmore East. K, paratype NMV P138648, thoracopygon, x 2.5, from ‘Broadhursts
Creek’, Wandong. L-M, Phacopidae gen. indet. 1, NMV P136136 cephalon, x 5, from PL1368, Strathewen. (B-C, F-G, I-K are internal moulds).
Early Silurian phacopide trilobites from central Victoria, Australia
227
Many of the differences between kenleyi and woiwurrungi
can be interpreted in the context of ontogenetic and heterochronic
processes. Chlupac (1977) and Ramskold (1988) discussed a
number of features in the ontogeny of phacopids. Chlupac noted
that juvenile phacopids exhibit wider, more inflated glabellae
with deeper S2 and S3, continuous SI, fewer lenses in the eye,
deeper vincular furrows, stronger tuberculation and deeper
pygidial pleural and interpleural furrows. Ramskold’s more
detailed study showed that many of the characters of Ananaspis
could be recognised in the ontogeny of Acernaspis. He noted that,
compared with adults, juvenile Acernaspis exhibit a narrower,
longer occipital ring with a median node, a longer (sag., exsag.)
and less depressed LI, shorter L2, deeper and straighter S2 and
S3 with S2 extending closer to the axial furrow, a smaller and
more anteriorly placed palpebral lobe with deeper palpebral
furrows, stronger tuberculation on the glabella and cheeks, deeper
lateral border furrows, and the presence of genal spines.
Arguing that most of the juvenile characters are primitive for
phacopids (where this could be determined), Ramskold (1988)
concluded that Ananaspis was a paedomorphic genus descended
neotenically from Acernaspis. The opposite can be said for the
kenleyi-woiwurrungi lineage. Ananaspis kenleyi exhibits a more
paedomorphic or Ananaspis- like appearance compared to the
more Acernaspis- like woiwurrungi, the significant differences
being the stronger glabellar tuberculation, deeper vincular furrow,
smaller eye, shorter L2 and deeper pygidial pleural furrows of
kenleyi. The kenleyi-woiwurrungi lineage represents a reversal
from the Acernaspis-Ananaspis morphocline and indicates that
the radiation of phacopines in the Silurian was more complex
than supposed by Campbell (1967) or Ramskold (1988).
The relationships of the kenleyi-woiwurrungi lineage to
other Ananaspis species remains uncertain although, as noted by
Sandford (2000), there are similarities with the Wenlock species
Ananaspis stokesii from Britain, A. nuda (Salter, 1864) (=
Ananaspis sp. aff. A. stokesiV of Siveter, 1989, pi. 19, figs 1-27,
pi. 20, figs 8-10) from Ireland, and A. amelangi from Sweden
(note that although the name amelangi was originally
incorrectly derived, under the Code it cannot be corrected and
its proposed change to amelangorum by Ramskold and
Werdelin, 1991: 70 is an unjustified emendation; ICZN Articles
32.2, 32.3, 32.5.1). Of these species A. kenleyi is most like
nuda, exhibiting similar glabellar tuberculation and pygidial
pleural furrows. However, the indistinct lateral border furrow
on the fixigena and the shallower pygidial interpleural furrows
distinguish the kenleyi-woiwurrungi lineage from these northern
hemisphere species and others assigned to Ananaspis.
Ananaspis typhlagogus (Opik, 1953)
Figures 3C, F-G, J-K, 7-8, 710E-G, ?10J
Phacops typhlagogus.— Opik, 1953: 26, pi. 10, figs 81-84.—
Talent, 1964: 50.— Sherwin, 1971: 94.
Ananaspis typhlagogus.— Campbell, 1967: 32.— Ramskold, 1988:
312, 313.— Rickards and Sandford, 1998: 751.— Sandford and Holloway,
1998: 915.
‘ Phacops ’ typhlagonus [sic].— Chlupac, 1977: 77.
Ananaspis ’ typhlagoga [sic].— Ramskold and Werdelin, 1991:
72, 74.
Type material. Holotype CPC 686 (incomplete and partially enrolled
exoskeleton), figured Opik (1953: pi. 10, figs 81-84), figs 7C-E, from
the ‘ Illaenus band’ (exact locality unknown), Costerfield. Wapentake
Formation.
Other material. NMV P138228, NMV P138234-P138238,
GSV36558 (missing), GSV46658 (missing), unregistered Geoscience
Australia collection specimen from the 7 llaenus band’ (exact locality
unknown), Costerfield. NMV P138229 from PL385, Costerfield. NMV
P138279-P138289, NMV P140039, NMV P147835-P147837 from
PL1460 (Thomas locality F43A), Costerfield. NMV P140152 from
PL390, Costerfield. NMV P147049 from PL2263, Costerfield.
Wapentake Formation. NMV P312813-P312815 from PL6361,
Springfield. Chintin Formation. A single crushed cephalon NMV
P147055 (figs. 10E-G, J) from PL206, Wallan, in the Bylands Siltstone,
is tentatively assigned to the species.
Diagnosis. Glabella comprising a little more than 33%
maximum cephalic width posteriorly and about 58% across
frontal lobe. L2 not markedly shorter (exsag.) than L3 adaxially,
S2 transverse adaxially and curving backwards abaxially.
Composite lobe with densely and evenly distributed tubercles,
larger ones of moderate size. Eye large, posterior edge very
close to posterior border furrow, lower edge separated from
lateral border furrow anteriorly by narrow band of librigenal
field, visual surface with 16-18 rows of up to 5 lenses each.
Fixigenal lateral border furrow shallow and wide. Pygidium
with 6 axial rings and 4-5 pleural furrows.
Description. Exoskeleton of estimated maximum length 40 mm.
Cephalon semi-elliptical in outline, length (sag.) approximately
58% maximum width, in frontal view anterior margin broadly
arched upward adaxially with very weak downward sag medially.
Glabella of moderate to strong convexity (sag., tr.), slightly
overhanging anterior cephalic border in dorsal view, anterior
margin forming arc centred approximately level with inner ends
of S2. Width of glabella across frontal lobe almost twice width
across LI and approximately equal to sagittal cephalic length.
Axial furrow shallow opposite occipital ring, very deep in front
of occipital furrow and diverging forward at 55°-70° in front of
SI. Occipital ring of uniform length (sag. exsag.) except distally
where it is contracted, deflected slightly obliquely forward and
bears weak lateral lobes. Occipital furrow very deep behind
lateral node on LI, transverse and moderately impressed
medially. LI about 60% as long as occipital ring medially, with
small, subquadrate, depressed lateral nodes. SI deflected slightly
forward adaxial to lateral node on LI, expanding and shallowing
medially but continuous across glabella. L2 with maximum
length (exsag.) 70%-80% maximum length of L3. S2 placed
opposite midlength of eye, not reaching axial furrow. Posterior
branch of S3 oriented transversely but gently convex forwards,
inner end level with glabellar midlength (sag.). Anterior branch
of S3 oriented diagonally, weakly convex forward. Preglabellar
furrow moderately impressed. In anterior view anterior border
narrow medially, wider laterally. Eye 45% sagittal cephalic
length, distance of posterior edge from posterior border furrow
less than or equal to length (exsag.) of posterior border directly
behind. Visual surface on NMV P138236 (figs 7F-H, K) with
lens formula 3? 5? 5 4? 5 4? 4 4? 4 5 4 4 4 4 3? 4 3 2; NMV
P140152 (fig. 8J) with formula 2? 3? 4? 4 5 4 5 5? 5? 5? 4 5 5 5
43? Palpebral area weakly convex (tr., exsag.), palpebral furrow
228
Andrew C. Sandford and David J. Holloway
Figure 7. Ananaspis typhlagogus (Opik, 1953). A-B, J, NMV P138229, cephalon x 7, from PL385, Costerfield. C-E, holotype CPC 686, dorsal
exoskeleton, views of cephalon, x 3, from the ‘ Illaenus band’, Costerfield. F-H, K, NMV P138236, cephalon, x 4, from the ‘ Illaenus band’, Costerfield.
I (and fig. IOC), NMV P138237, dorsal exoskeleton, x 4.5, from the ‘ Illaenus band’, Costerfield. L-M, unregistered specimen in AGSO collection,
cephalon, view of hypostome, x 2.5, from the ‘ Illaenus band’, Costerfield. (I, L-M are latex casts).
Early Silurian phacopide trilobites from central Victoria, Australia
229
Figure 8. Ananaspis typhlagogus (Opik, 1953). A-B, D-E, NMV P147049, partially enrolled exoskeleton from PL2263, Costerfield; A, x 2.8; B, x
3; D-E, x 3.5. C, NMV P138237, view of pygidium, x 9. F, NMV P138287, pygidium, x 4.3, from PL1460, Costerfield. G, NMV P138233, thorax, x
2.5, from the ‘ Illaenus band’, Costerfield. H-I, NMV P138285, pygidium, x 4.5, from PL1460, Costerfield. J, NMV P140152, cephalon, x 3.9, from
PL390, Costerfield. (C, F, H-J are latex casts).
moderately impressed. Palpebral lobe crescentic in outline, very
weakly convex (tr.), slightly elevated above palpebral area, with
distinct rim furrow. Posterior border furrow deep, posterior
border expanding (exsag.) backward adaxially so that genal
angle is produced posteriorly. Librigenal field weakly concave
below eye, steeply inclined, indistinctly separated from very
weakly convex lateral border. Posterior branch of facial suture
directed slightly obliquely forward across genal field and
deflected backward across border to meet cephalic margin
approximately level with median part of occipital furrow. Medial
part of cephalic doublure steeply inclined backwards, weakly
concave (sag.) in posterior part and weakly convex anteriorly.
Hypostomal suture transverse medially. Vincular furrow
moderately impressed, separated from anterior margin by narrow
band, lateral notching moderately expressed.
Hypostome about 150% as wide anteriorly as long (sag.),
parabolic in outline behind large, equilaterally-triangular
anterior wings, with weakly defined shoulder opposite
midlength (sag.). Middle body ovate in outline, moderately and
evenly convex (sag., tr.), middle furrow and maculae indistinct.
Lateral border narrow, in lateral view sloping gently dorsally
from anterior wing to shoulder where it is gently deflected
ventrally. Posterior border expanding slightly medially,
posterior border furrow shallower than lateral border furrow.
Thorax of 11 segments. Axis wider (tr.) than pleurae,
strongly convex (tr.), rings with lateral lobes weakly defined
by slight expansion, by shallow notch in anterior margin, and
by slightly oblique orientation of articulating furrow. Pleurae
steeply inclined beyond fulcrum, well rounded distally, with
deep pleural furrow terminating distally at edge of articulating
facet, approximately 25% distance from fulcrum to tip.
Pygidia all incomplete or poorly preserved, with large,
strongly oblique articulating facet and broadly rounded posterior
margin. Axis gently tapering backwards and decreasing in
height, bluntly rounded posteriorly, 1st ring standing higher
than remainder and with shallow medial embayment in posterior
230
Andrew C. Sandford and David J. Holloway
Figure 9. Fossil localities about 1 km south of Costerfield township; the
area covered by the map is indicated on fig. 1.
edge to accommodate pseudo -articulating half ring on 2nd
segment, 2nd and 3rd inter-ring furrows slightly expanded
sagittally but without pseudo -articulating half-rings, 6th and
7th inter-ring furrows very poorly defined close together. Axial
furrow deeper alongside axis than behind it. Pleural furrows
short (exsag.) and rather sharply incised, 1st interpleural furrow
shallow but distinct, remaining interpleural furrows very weak.
Border very poorly defined.
Remarks. Ananaspis typhlagogus is not very common in the
Wapentake Formation. Opik (1953) reported that he had
numerous fragments of the species but that only the holotype
was well enough preserved to be used for description. Specimens
such as the holotype that are preserved in the siliceous nodules
of the Wapentake Formation are undeformed and the exoskeleton
may be partly preserved as a mineralised crust (see Sandford
and Holloway, 1998: 921). In contrast, moulds from mudstones
of the Wapentake are crushed and fractured (fig. 8J), and those
from a sandstone clast are fragmentary (figs 8F, H-I).
Opik (1953) placed typhlagogus with the Phacops orestes
group (i.e. Acernaspis) but noted that the latter species differs
in ‘many important points’ including the lower glabellar
profile and stronger vincular notching. Campbell (1967)
assigned typhlagogus to Ananaspis on the basis of its glabellar
tuberculation and the bevelled profile of the cephalic anterior
margin and doublure.
Ananaspis typhlagogus is very closely related to A.
macdonaldi Fletcher, 1950 [=Acernaspisl oblatus [vie]
Sherwin, 1971; see Holloway, 1980: 64; Ramskold and
Werdelin, 1991:73], from the upper Llandovery to lower
Wenlock of central-western New South Wales, and to
Ananaspis’ sp. of Waisfeld and Sanchez, 1993, from strata of
presumed similar age in Argentina. Sherwin (1971) stated that
typhlagogus has more definite tuberculation and a more strongly
inflated glabella than macdonaldi. However, the significance of
the latter difference is difficult to assess in view of the wide
variation in glabellar profile in typhlagogus, and we cannot see
any clear difference between the species in sculpture. Ramskold
and Werdelin (1991) regarded macdonaldi and typhlagogus as
possible synonyms, but we consider that typhlagogus can be
distinguished by a higher visual surface of the eye with a greater
number of lenses (five in the longest file instead of four in
macdonaldi ), a lower eye socle, a larger LI, and a more strongly
curved posteromedial outline of the hypostome. Ananaspis’ sp.
from Argentina, known only from an internal mould of a single
cephalon (Waisfeld and Sanchez, 1993, pi. 1, figs 7-10), is difficult
to distinguish from typhlagogus but it has an eye with up to six
lenses per file and a vincular furrow that appears to be slightly
deeper medially and has a distinct flexure anterolaterally.
Also similar morphologically to A. typhlagogus is A.
aspera (Hawle and Corda, 1847) from the Ludlow of the Czech
Republic (see Chlupac, 1977, pi. 5, figs 9-25). The similarities
include: relatively narrow (tr.) cheeks; L2 not markedly shorter
(exsag.) than L3; adaxial half of S2 transverse; lower edge of
eye situated above lateral border furrow anteriorly; visual
surface very steeply inclined; and first order glabellar tubercles
of moderate size. A. aspera differs from A. typhlagogus in that
the cheeks are approximately as wide as the glabella posteriorly
instead of slightly narrower than it; the eyes are much shorter
(exsag.); the palpebral furrow is deeper; the lateral border
furrow is deeper on the fixigena; and the pygidium has a
greater number of more strongly defined segments and a
narrower, subparallel-sided axis.
Ananaspis sp. 1
Figures 10A, 10D, 101
Material. NMV P139354 (partly disarticulated cephalothorax), NMV
P139355 (cephalon), NMV P139356 (right cheek) from PL1338, Wallan.
Bylands Siltstone. For locality see Sandford and Rickards (1999: fig. 1).
Description. Cephalon subsemicircular, length 50% width.
Glabella strongly convex (tr.), with maximum width situated at
about 66% sagittal glabellar length and equal to 56% maximum
cephalic width, 110% sagittal cephalic length and 170%
occipital width. Outline of front of glabella forming arc centred
at 33% sagittal cephalic length from posterior. Axial furrow
wide and very deep in front of occipital ring, diverging strongly
at about 75° between SI and anterior end of palpebral furrow,
thereafter diverging forward at about 45° to widest part of
Early Silurian phacopide trilobites from central Victoria, Australia
231
Figure 10. A, D, I, Ananaspis sp. 1 from PL1338, Wallan. A, D, NMV P139354, cranidium, x 6.5. I, NMV P139356 cephalon (fragment),
enlargement of eye, x 9. B-C, Ananaspis sp. 2, NMV P136142, damaged cephalothorax, from PL1371, Coburg; B, x 1.4; C, x 1.2. E-G, J,
Ananaspis typhlagogusl (Opik, 1953), NMV P147055, cephalon (crushed) from PL206, Wallan; E-G, x 3; J, x 2.5. H, Ivops wallanensis gen. et
sp. nov., paratype NMV P139326, thoracopygon, x 3, from PL206, Wallan. K, Phacopidae gen. indet. 3, NMV P312077, partly disarticulated
thoracopygon, x 4.5, from PL1369, Springfield. (A, C, E-H, J-K are internal moulds).
frontal lobe. Occipital ring 30% maximum cephalic width,
short, 10% glabellar length sagittally and slightly shorter
laterally, lacking lateral lobes defined by notches in anterior
margin. Occipital furrow deep, transverse. LI as high as
composite lobe medially, lateral node of moderate size and
height, isolated by deep exsagittal furrow from preoccipital
ring, width of glabella across LI 75% occipital width. SI deep,
directed anteromedially from inner end of lateral node on LI,
at about 22° to transverse, connected medially by a shallower,
wide (sag.), transverse furrow. L2 65% length (exsag.) of L3
232
Andrew C. Sandford and David J. Holloway
and 11% sagittal cephalic length. S2 and S3 moderately
impressed. S2 directed at 45° distally, curving abruptly to
transverse direction proximally, anteriormost point opposite
40% sagittal glabellar length from posterior. Posterior branch
of S3 weakly convex forward, oriented transversely opposite
glabellar midlength (sag.). Anterior branch of S3 straight, not
connected to posterior branch, placed anteriorly opposite
anterior margin of eye, directed anterolaterally at 60° to
exsagittal line. Frontal lobe high. Preglabellar furrow very
shallow. Anterior border very short (sag.). Eye large, 50%
sagittal cephalic length, placed with midlength of eye opposite
33% sagittal glabellar length, occupying entire length of genal
field. Palpebral area low, weakly convex, palpebral furrow
moderately impressed. Palpebral lobe steeply inclined, raised
above palpebral area, with shallow rim furrow. Visual surface
large, higher anteriorly than posteriorly, with more than 18 files
of up to 7 lenses each, without raised sclera. Posterior border
short (exsag.) adaxially, approximately 300% as long distally.
Posterior border furrow deep and wide, continuous with deep
and wide fixigenal lateral border furrow. Genal angle obtuse,
with small point at angle. Lateral border moderately convex,
wide (tr.) posteriorly, narrowing strongly anteriorly. Librigenal
lateral border furrow moderately impressed. Posterior branch
of facial suture skirting back of eye to lateral border furrow,
strongly convex forward across lateral border, anteriormost
point opposite 25% cephalic length. Cephalic doublure with
extremely shallow vincular furrow anteriorly, very shallow and
very weakly notched laterally. Cephalic tubercles variable in
size from small to moderate, with superimposed granules,
densely distributed on composite lobe of glabella, LI and
palpebral area, remainder of exoskeleton with granulose
sculpture.
Remarks. Ananaspis sp. 1 is the only trilobite known from
PL1338. Stratigraphically, the locality is about 50 m above
PL206, which lies 750 m further to the south and yields a more
diverse fauna.
Although the above description is based only on one
complete cephalon and two fragments, the material is sufficient
to distinguish the species from other Victorian Silurian
phacopids. The species is most easily distinguished by its
fine, dense cephalic tuberculation, by the large size of the eye
which extends the entire length of the genal field, and by the
very shallow vincular furrow medially. Assignment to
Ananaspis is indicated by features including short L2 (relative
to the length of L3), a narrow glabella, the variably sized
glabellar tubercles, a deep lateral border furrow continuous
with the posterior border furrow and the obtuse genal angle
with a small point. The eye is larger than those of other
Ananaspis, although eye length is a character that varies
significantly between species here assigned to the genus
(eye length up to about 47% sagittal cephalic length in A.
amelangi, 45% in A. fecunda and A. crossleii, 40% in A.
decora, 37% in A. calvescens, 30% in A. aspera). Ananaspis
sp. 1 most closely resembles amelangi, sharing large eye size,
similar cephalic ornament and cephalic proportions. The
species differ in that amelangi has a glabella that does not
always extend to the anterior margin of the cephalon, the
occipital ring has more strongly defined lateral lobes, the
visual surface has only six lenses per file and 15-16 files, the
preglabellar and vincular furrows are deeper and the vincular
notching is deeper. Ananaspis sp. 1 differs from the type
species in having finer cephalic tuberculation, a larger eye and
fewer lens files in the visual surface (19-21 in fecunda). In
lens formula A. sp. 1 is most similar to A. guttulus (16 files of
up to seven lenses per file) and decora (16-17 files of seven,
rarely eight, lenses per file).
Ananaspis sp. 2
Figures 10B-C
Material. NMV P136142 (cephalothorax) from PL1371, about 50 m
SW of corner of Murray Road and Elizabeth Street, Coburg. Dargile
Group, precise stratigraphic horizon uncertain.
Description. Glabella wide, maximum width 110% sagittal
length and 65% maximum cephalic width, overhanging anterior
border. Axial furrow very shallow and slightly convergent
forwards adjacent to occipital ring, very deep adjacent to
composite lobe, straight and diverging forwards at about 65°.
Occipital ring 50% maximum glabellar width, medial section
raised, separated from short (exsag.) lateral section by deep
notch in anterior margin. Occipital furrow transverse medially,
deep. LI short (exsag.) 13% sagittal cephalic length, lateral
node low, globular, isolated from remainder of LI. SI deep
laterally, shallower medially where it forms posterior edge of a
slightly depressed triangular area between LI and composite
lobe. Length of L2 16% sagittal cephalic length. S2 and
posterior branch of S3 of moderate depth, wide, uniformly
arcuate, almost reaching axial furrow. S2 directed obliquely
inwards-forwards at about 15° to transverse. Posterior branch
of S3 oriented transversely opposite glabellar midlength (sag.),
anterior branch directed parallel to axial furrow. Posterior
border furrow deep and wide (exsag.), widening abaxially,
continuous with wide and deep lateral border furrow on
fixigena. Postocular area 14% sagittal cephalic length. Glabellar
tubercles large to very large, low and flat-topped, distributed
irregularly with moderate density. Tuberculation subdued on
genal field.
Remarks. The formation to which the strata at PL1371 belong
is difficult to determine as the site is no longer exposed. Fossils
are uninformative in this respect as the trilobite is the only
fossil known and is not known from elsewhere.
The glabellar tuberculation, depth and placement of the
lateral glabellar furrows, depth of the posterior and lateral
cephalic border furrows and length of the postocular area are
comparable to Eastern European and Central Asian species of
Ananaspis. In the greater width of the glabella the Coburg
species is closest to A. crossleii from the Yass district of New
South Wales (maximum glabellar width 60% maximum
cephalic width; see Sherwin, 1971, pi. 2, figs 6-9, pi. 3, figs
1-7). The Coburg specimen differs from crossleii in having a
wider glabella with larger, flat and less densely distributed
tubercles, and a longer (exsag.) postocular area (postocular
length 14% sagittal cephalic length versus 10% in crossleii ).
Early Silurian phacopide trilobites from central Victoria, Australia
233
Ivops gen. nov.
Type species. Ivops wallanensis sp. nov. from the Bylands
Siltstone (Wenlock), central Victoria.
Derivation of name. After the late Professor Ivo Chlupac,
Charles University, Prague. Gender masculine.
Diagnosis. LI with lateral node very small and depressed, L2
short (exsag.). S2 containing deep pit adjacent to axial furrow,
forming a notch in side of glabella. Anterior branch of S3 long,
subparallel to axial furrow. Front of glabella overhanging anterior
border. Eye relatively small, length (exsag.) approximately 30%
sagittal cephalic length, its lower edge lying just above lateral
border furrow anteriorly, visual surface without raised sclera.
Lateral border furrow on fixigena continuous with posterior
border furrow. Vincular furrow with weak notching laterally.
Pygidium with weakly tapering axis constituting about 30%
maximum pygidial width anteriorly, and with deep pleural and
shallow but distinct interpleural furrows. Dense sculpture of
bimodal tubercles on glabella.
Remarks. Ivops is known only from the type species. The genus
closely resembles Ananaspis in glabellar and pygidial proportions,
the short L2 (exsagittal length 80% that of L3), the continuity of
the posterior border furrow with the lateral border furrow on the
fixigena, the weak notching in the lateral part of the vincular
furrow, and the depth of the pygidial pleural and interpleural
furrows. The relatively small eye with its lower edge placed a
short distance above the lateral border furrow and with a small
number of lenses in the dorsoventral files are features comparable
with species such as A. aspera and A. calvescens. Despite these
shared features Ivops is unlike Ananaspis and other known
Silurian phacopines in that S2 is deep laterally and contains an
apodemal pit where it meets the axial furrow. This feature
contrasts with the more typical phacopine morphology of a
shallow S2 that is isolated from the axial furrow. A laterally deep
S2 similar to that of Ivops is present in ‘ Paciphacops' microps
Chatterton, Johnson and Campbell, 1979 (type species of Kainops
Ramskold and Werdelin, 1991), from the upper Lochkovian to
lower Pragian of New South Wales, but in that species S2 does
not contain an apodemal pit and does not meet the axial furrow in
all specimens. The relatively small eye of Ivops (the smallest of
known Wenlock phacopids) is interpreted as an adaptation to a
deep-water environment; see discussion in remarks on Berylacaste
berylae gen. et sp. nov.
Ivops wallanensis sp. nov.
Figure 11
Ananaspis .— Rickards and Sandford, 1998: 752.
Ananaspis .— Sandford and Holloway, 1998: 915.
Type material. Holotype NMV P139323 (enrolled cephalothorax).
Paratypes NMV P139324 (dorsal exoskeleton, pygidium displaced),
NMV P139325 (enrolled cephalothorax), NMV P139326 (thoracopygon),
NMV P139328 (partly enrolled dorsal exoskeleton), NMV P138230
(dorsal exoskeleton), NMV P138231 (incomplete cephalothorax). All
from PL206, Wallan. Bylands Siltstone.
Other material. NMV P138232, NMV P139327, P147056 from
PL206, Wallan. For locality see Sandford and Holloway (1998: text-
fig. 1).
Derivation of name. In reference to the type locality.
Diagnosis. As for genus.
Description. Exoskeleton of known maximum length 30 mm.
Cephalon about 60% as long (sag.) as wide, anterior outline
with greatest curvature medially and in front of anterior
extremity of eye. In frontal view anterior margin moderately
arched upward medially, glabella strongly convex and evenly
rounded. Glabella comprising about 30% maximum cephalic
width posteriorly, width across frontal lobe about twice width at
occipital ring and equal to sagittal length. Axial furrow diverging
forward at about 70° from SI to anterior end of palpebral
furrow, thereafter diverging slightly more weakly to widest part
of frontal lobe. Occipital ring high medially and with small,
well-developed, obliquely oriented lobe laterally. Occipital
furrow deep, transverse medially. LI slightly narrower (tr.) than
occipital ring, as high medially as composite lobe, lateral node
smaller than lateral lobe of occipital ring. SI expanding (exsag.)
adaxially or bifid, posterior section transverse and weakly
continuous medially, anterior section curving forward and
rapidly dying out. L2 almost as short (exsag.) laterally as LI
and about 60% length of L3. S2 meeting axial furrow more or
less opposite posterior edge of palpebral lobe, directed
anteromedially and shallowing rapidly, weakly arcuate. S3 with
anterior and posterior branches confluent opposite about 40%
glabellar length (sag.) from posterior, anterior branch extending
forward to opposite about 60% glabellar length (sag.) from
posterior. Preglabellar furrow distinct across entire width (tr.)
of glabella on internal moulds, decreasing in length (exsag.)
adaxially in front of lateral part of frontal lobe. Posterior border
and border furrow deflected rather strongly backward beyond
fulcrum to genal angle. Palpebral lobe narrow (tr.), lenticular in
outline, raised above palpebral area, palpebral furrow weakly
curved. Visual surface with lenses arranged regularly in files
anteriorly but less regularly posteriorly, lens formula (NMV
P139328, figs 11H-I, K) from front 456652565473?. Librigenal
field subvertical below eye, lateral border steeply inclined.
Posterior branch of facial suture transverse across genal field
and deflected backward at about 45° across border. Cephalic
doublure flat sagittally, vincular furrow forming distinct
bevelled edge anteriorly, deep posterolaterally. Hypostomal
suture transverse medially.
Thorax of 11 segments. Axis comprising 30% segmental
width (tr.), strongly convex, rings transverse medially, with
short exsagittal incisions in anterior margin defining obliquely
forwardly directed lateral lobes. Pleurae with well-rounded
tips and deep pleural furrows that are situated at midlength
(exsag.) of segment at fulcrum and terminate distally at edges
of articulating facets.
Pygidium lenticular, length (sag.) about 60% maximum
width. Axis with 5 well defined rings and 1 or 2 very short
(sag.) and weak ones posteriorly, axial terminus well rounded.
Pleurae weakly convex (tr.), with 4 distinct pleural furrows
and 2 very weak ones posteriorly. Border not present.
Remarks. The above description is based mainly on the least
deformed specimens (e.g. figs 11B-C, E, J). S2 and S3 appear
to vary markedly in depth, being very weak on most specimens
234
Andrew C. Sandford and David J. Holloway
Figure 11. Ivops wallanensis gen. et sp. nov., from PL206, Wallan. A, G, paratype NMV P138230, dorsal exoskeleton; A, x 4.3; G, x 3.2. B, F, J, L,
paratype NMV P139324, exoskeleton with displaced pygidium; B, view of pygidium, x 6.5; F, view of cephalon, x 3.8; J, view of pygidium, x 5.4; L,
view of cephalic doublure, x 3.8. C, E, holotype NMV P139323, enrolled exoskeleton, view of cephalon, x 3.8. D, paratype NMV P139325, cephalon,
x 3.8. H-I, K, paratype NMV P139328, partly enrolled exoskeleton, view of cephalon, x 4.3. (A, C-F, J-K are internal moulds).
Early Silurian phacopide trilobites from central Victoria, Australia
235
but moderately impressed on both internal and external moulds
of NMV P138230 (figs 11 A, G). However, the apodemal pit in
the lateral part of S2 is always present.
Phacopidella Reed, 1905
Type species. Phacops glockeri Barrande, 1846 from the upper
part of the Motol Formation (upper Wenlock), Czech Republic,
by original designation.
Phacopidella ? sp.
Figure 12A
Asaphus.— Selwyn, 1863: map note.
Phacops crosslei.— Chapman, 1913: 210, 229.— Chapman, 1915:
169.
Phacopidella!.— Rickards and Sandford, 1998: 749.
Material. NMV P1218 (dorsal exoskeleton with displaced cephalon),
probably from PL1393, Geological Survey locality Ba5, Keilor.
Springfield Formation, in siltstone underlying the Lintons Creek
Conglomerate Member; late Llandovery, Telychian ( Spirograptus
turriculatus-Monograptus crispus biozones). For locality see Selwyn
(1863), Rickards and Sandford (1998) (fig. 5).
Description. Exoskeleton, of estimated original length about 35
mm, has been flattened tectonically and slightly sheared. The
preserved portion of the cephalon consists of the glabella behind
about the middle (sag.) of the frontal lobe together with the
posteromedian part of cheeks. The glabella is slightly narrower
across LI than across the occipital ring and expands gently
forwards in front of LI, the axial furrow here diverging at about
40°. The occipital ring is almost twice as long sagittally as
exsagittally, with lateral lobes defined by very short exsagittal
incisions in the anterior edge. The median portion of the occipital
furrow is arched forward. LI has large, quadrate nodes laterally
and is slightly shorter (sag., exsag.) than the occipital ring
medially. SI is deep laterally, is deflected forwards adaxial to the
short exsagittal furrow defining the lateral node of LI and
shallowing rapidly, becoming indistinct medially. S2 is short
(exsag.), very faint and weakly convex forward, directed slightly
obliquely backward laterally. L2 is about as long (exsag.) as LI.
The posterior branch of S3 is indistinctly preserved on the left
side of the glabella, running subparallel to S2. Axial furrow
deep. Left palpebral lobe possibly partly preserved, seemingly
relatively long (exsag.) and obliquely oriented. Thorax with 11
segments. Axial rings with short, deep exsagittal incisions in
anterior margin defining large, quadrate lateral lobes. Pleurae
with deep pleural furrow situated at middle (exsag.) of segment
at fulcrum; distal parts of pleurae not preserved but overall width
of pleural lobe appears not to exceed width of axis. Pygidium
large, approximately 160% as wide as long (sag.). Axis about
30% maximum pygidial width anteriorly and about 66% sagittal
pygidial length, with 5 well defined rings and weak 6th ring.
Pleurae with 5 deep pleural furrows and 2 weak ones posteriorly;
interpleural furrows moderately impressed. Dorsal surface of
pleurae not intact abaxially on internal mould, so appearance of
wide, smooth border is largely an artefact.
Remarks. The lithology of this specimen is identical to that of
a specimen of Hadromeros from Keilor, one of two trilobites
collected by C.D. Aplin in the 1850s that were noted on the
geological quarter sheet by Selwyn (1863) as Cheirurus and
Asaphus. NMV PI 218 is apparently the second trilobite
collected by Aplin, and that recorded by Chapman (1913) as
Phacops crossleii Etheridge and Mitchell, 1896 from Keilor.
However, the museum label with the specimen records the
locality as ‘Saltwater River, one mile west of Gisborne’, as
published by Chapman (1915). Chapman (1915) noted the
lithology of the trilobite specimen as ‘bearing a strong
resemblance to the Keilor.. .mudstones’. Chapman, the museum
palaeontologist at the time, appears to have confused Gisborne
with Keilor. Only Early to Middle Ordovician graptolitic beds
outcrop in the Gisborne area, with a lithology quite unlike that
of the trilobite specimen.
The low pygidial convexity (largely a result of tectonic
flattening) and well defined pygidial segmentation explain
Selwyn’s (1863) preliminary assignment of the specimen to
Asaphus. The glabellar segmentation, especially SI that is
continuous though very weak medially and the shallowness
and orientation of S2 and the posterior branch of S3, as well as
the overall form of the pygidium, indicate that the specimen
belongs to the Phacopidae. The gentle forward expansion of
the glabella, the relatively long (sag., exsag.) LI with large
lateral nodes, the well-defined nodes on the occipital ring and
thoracic axial rings, and the relatively large and well-segmented
pygidium are all consistent with assignment to Phacopidella.
A characteristic feature of Phacopidella (and of Eophacops )
is the abrupt shallowing and adaxial flexure of the axial furrow
at the anterolateral extremity of L3. This part of the glabella is
not preserved in the specimen; however, so that the presence
of this feature and assignment to Phacopidella cannot be
confirmed. This uncertainty aside, the specimen differs from
the type species of Phacopidella, P. glockeri, in having a
wider axis on the thorax and pygidium, the pygidial axis
extends closer to the posterior margin, and the pygidial pleural
and interpleural furrows are deeper.
Phacopidae gen. indet. 1
Figures 6L-M
Lochkovella! .— Rickards and Sandford, 1998: 753.
Material. NMV P136136 (cephalon) from PL1638, Williams locality
W25, Strathewen. Yan Yean Formation. The locality occurs in strata
mapped just below the boundary of the Anderson Creek Formation
and the Dargile Formation (Garratt, 1972). Strata mapped at this
horizon in some areas around Melbourne were reassigned by Rickards
and Sandford (1998) to the Yan Yean Formation. The presence of a
distinctive encrinurid at PL1638 that otherwise occurs in the upper
beds of the Yan Yean Formation at Heathcote (at PL2259, Thomas
locality F41, Parish of Heathcote) and at Upper Plenty (PL1697,
PL1699) supports the assignment of strata at PL1638 to the Yan Yean
Formation. For localities see Thomas (1940), Williams (1964) (fig. 2)
and Sandford (2006) (figs 2, 4).
Description. Cephalon semi-elliptical in outline, sagittal length
about 60% maximum width. Glabella gently to moderately
convex (sag., tr.), frontal lobe low, width across frontal lobe
approximately twice width across LI and a little more than
sagittal length. Axial furrow wide and very deep, diverging at
236
Andrew C. Sandford and David J. Holloway
Figure 12. A, Phacopidellal sp., NMV P1218, dorsal exoskeleton, x 2.7, from PL1393, Keilor. B-C, Struveria sp. 1, NMV P139357, partly
disarticulated exoskeleton with hypostome and cephalic doublure exposed, from PL1386, Broadmeadows; B, x 3.4; C, x 4. D-F, Phacopidae gen.
indet. 2, NMV P312076, cephalon, x 8, from PL1369, Springfield. (A, E-F are internal moulds).
about 60° in front of SI. Occipital ring comprising 40%
maximum cephalic width, raised high medially, shorter (exsag.)
and obliquely directed laterally but without well defined lateral
lobes. Occipital furrow deep. LI of almost uniform length
(sag., exsag.) across glabella, high medially, inner edge of
lateral node obscured. SI very deep and wide laterally, directed
slightly forward of transverse, shallowing abruptly close to
sagittal line. S2 deep, weakly convex forward, its outer end
opposite posterior edge of eye, running parallel to SI and very
close to it. S3 not clear but posterior branch apparently directed
posteromedially at about 30° to transverse and with inner end
more or less level with glabellar midlength (sag.) and close to
Early Silurian phacopide trilobites from central Victoria, Australia
237
S2. Eye small and obliquely oriented, situated almost its own
length (exsag.) from posterior border furrow and with midlength
opposite glabellar midlength (sag.). Palpebral furrow
moderately impressed, weakly arcuate, continuous with
postocular furrow that is of similar depth and reaches lateral
border furrow. Palpebral lobe not raised above palpebral area.
Visual surface not well preserved but appears greatly reduced,
ovate in outline with lower edge distant from lateral border
furrow, and with about 10 lenses. Posterior border furrow deep
adaxially, continuous with wide, deep lateral border furrow.
Posterior border uniformly short (exsag.) except distally. Genal
angle rounded. Lateral border weakly convex, wider behind
intersection of postocular furrow than in front. Sculpture
obscured by preservational pitting covering most of cephalic
surface.
Remarks. Tentative assignment of this specimen to Lochkovella
by Rickards and Sandford (1998) was based on the small and
forwardly placed eye, the deep and continuous posterior and
lateral cephalic border furrows, the deep postocular furrow,
and the apparent absence of coarse tuberculation on the
glabella. These characters as well as the narrow cheeks invite
comparison with the Czech Early Devonian (Pragian) species
‘ Phacops ( Phacops ?)’ hanusi Chlupac, 1977 and ‘P. (P?)’
veles Chlupac, 1972, which were assigned to Lochkovella by
Sandford (2004). However, other characters of the specimen
are incompatible with assignment to Lochkovella. Such
characters include the glabella of low convexity that does not
overhang anteriorly, and the very small, elliptical visual
surface of the eye with its lower edge distant from the lateral
border furrow. It is possible that the specimen belongs to an
undescribed genus.
The monotypic Orygmatos Sandford, 2000, from a slightly
higher stratigraphic horizon low in the Melbourne Formation
at Yan Yean, 10 km to the north-east, also has very small eyes
with an elliptical visual surface. Compared with the present
specimen from Strathewen, Orygmatos has an occipital ring
that is more expanded (sag., exsag.) and prominent medially,
LI is very short (sag., exsag.) and markedly depressed, the
composite glabellar lobe expands more strongly forward and
is coarsely tuberculate, the palpebral furrow meets the
postocular furrow in an broad curve rather than an abrupt
angle, and the posterior border furrow and fixigenal portion of
the lateral border furrow are very wide. Denckmannites
Wedekind, 1914 has even more greatly reduced eyes than the
Strathewen specimen, much weaker palpebral and postocular
furrows, a more elongated glabella, wider cheeks, and a longer
anterior border. The small-eyed Denckmannites rutherfordi
Sherwin, 1968 from the Ludlow-Pndolf of central western
New South Wales was assigned to Lochkovella by Chlupac
(1977) but Sandford (2004) noted that the presence of strong
notching in the lateral part of the vincular furrow excluded the
species from that genus, a conclusion supported by the apparent
absence of granular sculpture on the exoskeleton. The very
low cephalic profile (sag., tr.), the more pentagonal outline of
the glabella, the subparallel alignment of S2 and S3, and the
broadly arcuate union of the palpebral and postocular furrows
distinguish rutherfordi from the Strathewen specimen.
Phacopidae gen. indet. 2
Figures 12D-F
Material. NMV P312076 (cephalon) from PL1369, Deep Creek,
Springfield. Springfield Formation. For locality see fig. 4.
Remarks. This very small cephalon (3 mm in length), and two
other small specimens documented below as Phacopidae gen.
indet. 3, are the only phacopids known from PL1369. The
cephalon undoubtedly belongs to a juvenile individual, and its
size is in the range of late meraspides and early holaspides of
Acernaspis (cephalic lengths 1.3 mm to 3.3 mm) documented
by Ramskold (1988). It is characterised by a strongly convex
glabella that overhangs anteriorly, a short (exsag.) L2, a very
large eye occupying almost the entire length of the genal field,
a well defined fixigenal lateral border furrow joining the
posterior border furrow, and a granulose sculpture. The
glabellar convexity, length of L2 and distinct lateral border
furrow suggest assignment to Ananaspis, but the last two
characters could also be attributed to the small size of the
specimen as Ramskold (1988) observed that in juveniles of
Acernaspis L2 is shorter and the lateral border furrow deeper
than in adults. The glabella lacks the tuberculation characteristic
of Ananaspis, whereas the sculpture is normally relatively
coarser in juveniles than adults.
Phacopidae gen. indet. 3
Figures 5E-G, 10K
Material. NMV P139353 (partially enrolled exoskeleton), NMV
P312077 (partly disarticulated thoracopygon) from PL1369,
Springfield. Springfield Formation.
Remarks. These two small specimens are considered to belong
to the same species because of similarities in the pygidia, which
have six axial rings, two pseudo-articulating half rings
(although the third ring is damaged in the thoracopygon), four
distinct pleural furrows, and well-defined interpleural furrows
extending almost to the margin. The partially enrolled
exoskeleton has a cephalon with a moderately convex (sag.)
glabella that does not overhang the preglabellar furrow and
anterior border in dorsal view, very weak S2 and S3, a shallow
fixigenal border furrow, an eye of moderate size situated well
in front of the posterior border furrow, a weakly curved
palpebral lobe, and a finely granulose sculpture. Compared to
the cephalon described above as Phacopidae gen. indet. 2,
which is from the same locality, the present specimen represents
a larger individual (cephalic length 7.5 mm as opposed to 3
mm) and has a more weakly convex glabella and much smaller
eye. The difference in the eyes cannot be attributed to
ontogenetic changes as eye size increases during ontogeny
rather than decreases, as documented by Ramskold (1988) in
Acernaspis. We therefore conclude that Phacopidae gen. indet.
2 and Phacopidae gen. indet. 3 belong to different taxa.
Superfamily Dalmanitoidea Vodges, 1890
Family Dalmanitidae Vodges, 1890
Dalmanites Barrande, 1852
238
Andrew C. Sandford and David J. Holloway
Type species. Trilobus caudatus Briinnich, 1781 from the
Coalbrookdale Formation ( Wenlock) of England.
Dalmanites athamas Opik, 1953
Figures 13A-K
Dalmanites sp.— Thomas, 1937: 66.
Dalmanites athamas Opik, 1953 : 28, pi. 10, figs 88-91; text-fig. 9
(IV).— Holloway and Sandford, 1993: 97.— Sandford and Holloway,
1998: 916.— Rickards and Sandford, 1998: 751.
Dalmanitina ( Eudolatites ) aborigenum Opik, 1953: 26 ( partim .),
pi. 10, fig. 85, text-fig. 9 (III) (non pi. 10, figs 86-87 = Struverial sp.).
Dalmanites ? athamas.— Talent, 1964: 50.
‘D almanitin a' aborigenum.— Talent, 1964: 50.
Type material: Holotype NMV P52484 (pygidium, figured Opik,
1953: pi. 10, fig. 88) and paratype NMV P52485 (fragment of cephalon,
figured Opik, 1953: pi. 10, fig. 89) from PL2262, Thomas locality F43,
Costerfield. Paratype NMV P52486 (fragment of cephalon, figured
Opik, 1953: pi. 10, figs 90-91) from PL2269, Thomas locality F51,
Costerfield. Wapentake Formation.
Other material. NMV P138215-P138216 from the ‘ Illaenus band’
(exact locality unknown), Costerfield. NMV P138217 from PL389,
Costerfield. NMV P52482 (holotype of ‘ Dalmanitina (EudolatitesY
aborigenum ), P138218-P138219, NMV P138223 from PL2269,
Costerfield. NMV P138220 from PL2263, Thomas locality F44,
Costerfield. NMV P139805 from PL386, Costerfield. Wapentake
Formation. For localities see Thomas (1940) and fig. 9.
Diagnosis. Dalmanites with very short (sag., exsag.) anterior
cephalic border lacking median expansion or process. Glabella
about 77% as wide across frontal lobe as long (sag.), frontal
lobe comprising a little more than half sagittal length of
glabella. Pygidium with 14-15 axial rings and 10 pleural
furrows. Axis narrow, comprising 23% maximum pygidial
width anteriorly, with apodemes on 1st 10-12 segments. Pleural
furrows flat-bottomed, as long (exsag.) as or longer than
preceding anterior pleural band, posterior bands much shorter
except adaxially and dying out distally on more posterior
segments, pleural nodes absent.
Description. Glabella with widths across LI, L3 and frontal
lobe approximately in ratio 1:1. 6:1. 8. Occipital ring, LI and L2
of equal length, together comprising 35% sagittal glabellar
length. SI and S2 transverse, SI with apodemal pit extending
almost to axial furrow, S2 with slightly narrower (tr.) apodemal
pit and shallow extension to axial furrow. L3 as long (exsag.)
adaxially as LI and L2, twice as long at axial furrow. S3 deepest
proximally, expanding abaxially and subsequently contracting
again toward axial furrow. Frontal lobe transversely elliptical,
width 150% length, with short longitudinal depression in
posterior half and forwardly expanding median field of muscle
scars occupying slightly more than anterior half. Anterior
border slightly shorter sagittally than laterally, weakly concave.
Anterior branch of facial suture running slightly closer to
preglabellar furrow than to anterior cephalic margin. Posterior
border furrow lanceolate in outline, gently deflected backward
distally, not meeting lateral border furrow.
Thoracic axis weakly convex (tr.), gently tapering
backward. Axial rings very weakly convex (sag., exsag.),
slightly bowed forward medially. Axial furrow deep. Pleural
furrows expanding (exsag.) abaxially to fulcrum, deep, anterior
slope much steeper than posterior slope. Anterior pleural
bands strongly convex (exsag.), posterior bands becoming
flattened toward fulcrum and inclined, poorly differentiated
from posterior slope of pleural furrow. Pleural bands with
sculpture of small, sparse granules.
Pygidium triangular in outline, shape of posterior termination
unknown. Axis with length about 70% maximum pygidial width,
tapering uniformly backward, 1st ring with subrectangular
medial embayment in posterior edge accommodating pseudo-
articulating half ring on 2nd segment, inter-ring furrows
shallowing medially and expanded (sag.) except towards back
of axis where they become increasingly more poorly defined.
Axial furrow narrow but distinct on external surface. 1st pleural
furrow straight except distally where it is deflected strongly
backwards, subsequent furrows successively more evenly
curved and more posteriorly directed, 10th furrow directed
exsagittally. 7th pleural rib with distinct forward deflection
proximally. Interpleural furrows sharply impressed externally.
Doublure narrow, moderately convex abaxially and with a low
upturned lip adaxially, with densely distributed small granules.
Remarks. The synonymy of ‘ Dalmanitina ( Eudolatites)'
aborigenum with Dalmanites athamas was discussed by
Sandford and Holloway (1998: 916). The paratype of
aborigenum, a pygidial fragment, is not congeneric with the
holotype and we assign it to Struveria sp. 2.
New material of athamas has become available since
Opik’s (1953) original description but no cephala that are more
complete. Opik considered the species to be close to and
possibly a subspecies of Dalmanites wandongensis from the
overlying Dargile Formation, but comparison of the available
material indicates that the two species are distinct. D.
wandongensis was revised by Sandford (2006) and differs
from D. athamas in that LI is shorter than L2 instead of equal
in length; the pygidial pleural furrows are shorter (exsag.) and
not as flat-bottomed in cross section but form a continous slope
with the succeeding posterior pleural band; and the interpleural
furrows are weaker.
Ramskold (1985) recognised a closely related group of
European and North American Wenlock-early Ludlow species
of Dalmanites including the type species, D. caudatus
(Briinnich, 1781), as well as D. corrugatus (Reed, 1901), D.
limulurus (Green, 1832), D. myops (Konig, 1825), D. nexilis
Salter, 1864, D. obtusus (Lindstrom, 1885) and D.
platycaudatus Weller, 1907. These species share prominent
lateral nodes on the first, sixth and seventh thoracic axial rings,
and nodes on the pygidial pleural ribs. D. athamas lacks the
thoracic and pygidial nodes characterising this group, which also
differs from athamas in having a medially expanded anterior
cephalic border, and pygidia with shorter (exsag.) pleural furrows
and longer (exag.) posterior pleural bands that are subequal in
length to the anterior bands.
Bessazoon Curtis and Lane, 1998
Type species. Dalmanites weaveri var. tenuimucronata
Whittard, 1938 from the Hughley Shales (upper Llandovery) of
Shropshire, England, by original designation.
Early Silurian phacopide trilobites from central Victoria, Australia
239
Figure 13. A-K, Dalmanites athamas Opik, 1953. A, NMV P138215, thoracopygon, x 1.5, from the ‘ Illaenus band’, Costerfield. B, NMV P138219,
pygidium, x 3.5, from PL2269, Costerfield. C, paratype NMV P52485, cranidium, x 2, from PL2262, Costerfield. D, G, NMV P138217, pygidium, x
2, from PL389, Costerfield. E, paratype NMV P52486, cranidium, x 2, from PL2269, Costerfield. F, NMV P138218 pygidium, x 2, from PL2269,
Costerfield. H, holotype NMV P52484, pygidium, x 1.5, from PL2262, Costerfield. I, NMV P138223, thorax, enlargement showing granulation, from
PL2269, Costerfield. J, NMV P139805, teratological pygidium, x 2, from PL386, Costerfield. K, NMV P52482, incomplete fixigena, holotype of
‘ Dalmanitina (Eudolatites) aborigenunC Opik, 1953, x 2, from PL2269, Costerfield. L, Struveria sp. 2, NMV P52483, pygidium, paratype of
‘ Dalmanitina ( Eudolatites ) aborigenurri’ Opik, 1953, x 3, from PL2269, Costerfield. (B, D are latex casts)
240
Andrew C. Sandford and David J. Holloway
Remarks. Of the characters regarded by Curtis and Lane (1998:
62) as diagnostic of Bessazoon, Chatterton and Ludvigsen
(2004: 47) questioned the taxonomic value of the size of the
eyes and the loss of the pygidial mucro in large specimens. In
the type species and B. tigerense (Holloway and Sandford,
1993) the eyes are very large and occupy almost the entire
length of the genal field, but in Curtis and Lane’s species B.
buttingtonense and the cephalon they illustrated as B. cf. B.
tenuimucronatum (their pi. 9, fig. 1) the eyes are smaller and do
not extend very close to the border furrows anteriorly and/or
posteriorly. In regard to the loss of the mucro, Curtis and Lane
illustrated three pygidia assigned to B. tenuimucronatum with
the posterior termination preserved. Two of these (Curtis and
Lane’s pi. 8, figs 6, 8) have a mucro, that on the larger specimen
being considerably shorter than the one on the smaller specimen.
The third pygidium, which is very much larger that the others
(Curtis and Lane’s ‘type 2’ pygidium, pi. 8, fig. 2), does not
have ‘a shorter mucral spine’ as stated by Chatterton and
Ludvigsen (2004) but an embayment in the margin
posteromedially. Because of this difference, as well as the
differences in size and the much greater number of axial rings
and pleural furrows, it is not possible to be confident that this
pygidium is correctly assigned to B. tenuimucronatum.
However, we note that it has a distinctive pleural structure in
which the posterior bands expand distally and the anterior
bands are pinched out, and that this structure appears to be
shared by the smaller pygidium in pi. 8, fig. 6 (the structure is
not clear in the third pygidium as the dorsal surface is broken
away distally to reveal the doublure). Although we believe that
these pygidia are correctly assigned to the Dalmanitidae, their
pleural structure is unusual for Silurian and Devonian
representatives of the family, in most of which it is the anterior
rather than the posterior pleural band that is dominant distally
(Holloway, 1981: 710), as is also the case in B. buttingtonense
(Curtis and Lane, 1998, pi. 9, figs 2, 4, 7), B. tigerense (Holloway
and Sandford, 1993, fig. 6) and in the unnamed species from
Victoria described below. The only pygidium of B.
buttingtonense with the posterior termination preserved (Curtis
and Lane, pi. 9, fig. 2a, b) is of about the same size as the ‘type
2’ pygidium of B. tenuimucronatum but has neither a mucro
nor a posteromedian embayment; instead the margin is rather
truncated in dorsal view and arched upwards in posterior view.
There is no evidence that the mucro is lost in B. tigerense but
all known pygidia are smaller than that of buttingtonense and
the ‘type 2’ pygidium of tenuimucronatum.
Some of the other characters listed by Curtis and Lane as
diagnostic of Bessazoon cannot in our view be used to
distinguish Bessazoon either. The size of the palpebral area is
determined by the size of the eye, and thus in comparison with
other Silurian and Devonian dalmanitids is not particularly
large in B. buttingtonense and B. cf. B. tenuimucronatum. We
can see no difference from Dalmanites and other closely
related genera in the form of the palpebral lobe, which rises
steeply from the palpebral furrow and becomes flat towards
the outer margin. Finally, a posterior cephalic border furrow
that fails to meet the lateral border furrow distally, an epiborder
furrow on the lateral cephalic borders and genal spine, and a
straight-sided pygidial axis are not unusual features for
dalmanitids but are present in most Silurian and Devonian
representatives. The discrimination and composition of
Bessazoon are in need of review, but in the meantime we apply
the name here to dalmanitids differing from species commonly
assigned to Dalmanites (e.g. see Ramskold, 1985) in lacking a
well-developed anterior cephalic process (although in B.
tenuimucronatum the cephalic margin is deflected slightly
forwards medially) and tubercles on the glabella, and in having
a pygidium with a curved posterolateral outline, a slender,
narrow-based mucro merging anteriorly with a strong postaxial
ridge, and a very wide doublure extending adaxially beyond
the distal ends of the pleural and interpleural furrows.
Bessazoon sp.
Figures 14, 17D-F
Dalmanites— Rickards and Sandford, 1998: 750.
Material. NMV P139427-P139439, NMV P138276 from PL1452,
Goldie. NMV P139470-P139487 from ‘Lancefield’ (exact locality
unknown). NMV P147769-P147770 from PL256, Wallan. NMV
P312817 from PL6361, Springfield. Chintin Formation. For locality
PL1452 see Thomas (1960), marked as ‘shelly fossils, Dalmanites’.
Description. Anterior cephalic margin parabolic in outline (fig.
141). LI about 75% length of L2. SI deflected forwards slightly
at adaxial end, S2 more or less transverse; S3 shallow, oriented
at about 65° to sagittal line, expanding (exsag.) abaxially. Inner
ends of S1-S3 in line (exsag.). Palpebral lobe large, length
(exsag.) about 40% sagittal glabellar length, almost semicircular
in dorsal outline, anterior margin opposite anterior margin of
L3 and posterior margin reaching almost as far back as occipital
furrow. Palpebral furrow weak (fig. 17E), palpebral area wide
(tr.) and weakly concave.
Hypostome approximately as wide across anterior wings
as long (sag.), subparallel sided from back of anterior wing to
outer end of posterior border furrow, thereafter narrowing
backward and parabolic in outline. Anterior wings small,
triangular, length (exsag.) about 15% sagittal length of
hypostome. Middle body comprising 80% sagittal length of
hypostome, weakly convex transversely and flat sagittally.
Maculae indistinct swellings behind weak depressions placed
opposite 40% hypostomal length from posterior and halfway
between sagittal line and lateral border furrow. Lateral border
narrow, approximately 6% width of hypostome at midlength
(sag.), lateral border furrow weak in anterior half and
moderately impressed in posterior half. Posterior border long,
15% hypostomal length sagittally, posterior border furrow
semicircular in outline.
Pygidium triangular in outline, relatively undeformed
specimens with length (excluding mucro) approximately 75%
estimated maximum width. Axis about 25% maximum
pygidial width anteriorly, tapering uniformly backward, with
14 axial rings of which last 6 are poorly defined. Inter-ring
furrows 1-9 with deep apodemes extending approximately.
33% width, all inter-ring furrows very shallow medially. Axis
continuous posteriorly with postaxial ridge and mucro. Axial
furrow deep. Mucro slender, at least 75% length (sag.) of
remainder of pygidium. 10 pleural furrows that are directed
Early Silurian phacopide trilobites from central Victoria, Australia
241
Figure 14. Bessazoon sp. A, NMV P139479, pygidium, x 2, from ‘Lancefield’. B, NMV P139481, pygidium, x 3.5, from ‘Lancefield’. C, NMV
P139475, pygidium, from ‘Lancefield’. D, NMV P139474, pygidium, x 2.5, from ‘Lancefield’. E, NMV P139477, pygidium with doublure exposed,
x 6, from ‘Lancefield’. F, NMV P139471, crushed cephalon, x 2, from ‘Lancefield’. G, NMV P139427, hypostome, x 3, from PL1452, Lancefield.
H, J, NMV P139438, pygidium, from PL1452, Lancefield; H, enlargement of anterolateral region showing granular sculpture, x 8; J, x 2. 1, NMV
P139439, crushed cephalic doublure, x 1.9, from PL1452, Lancefield. K, thoracic segment, enlargement showing granulose ornament on pleural
tip, x 8, from ‘Lancefield’. L, NMV P139470, fragment of cephalon, x 3, from ‘Lancefield’. M, NMV P139473, pygidium, x 2, from ‘Lancefield’.
(E-F are internal moulds).
242
Andrew C. Sandford and David J. Holloway
successively more strongly backward, last one parallel to
sagittal axis. Interpleural furrows moderately incised,
widening distally. Anterior pleural bands expand slightly and
very gradually abaxially, at fulcrum approximately as long
(exsag.) as succeeding pleural furrow, slightly elevated above
posterior bands distally. Posterior pleural bands subparallel
sided except distally where they taper, at fulcrum comprising
about 75% length (exsag.) of anterior bands.
Densely distributed small granules present on preserved
parts of external surface including occipital ring, thoracic
pleural tips and doublure, dorsal surface of pygidium and
pygidial doublure.
Remarks. This species can be distinguished from other Silurian
dalmanitids from central Victoria by its pygidial pleural
morphology, with relatively narrow, trench-like pleural furrows,
and anterior bands that are slightly longer (exsag.) than the
posterior bands at the fulcrum and only slightly elevated above
the posterior bands distally. Dalmanites athamas differs in
having longer (exsag.) pleural furrows and anterior pleural
bands that are much longer than the posterior bands, whereas
D. wandongensis has anterior bands that are slightly shorter
(exsag.) than the posterior bands at the fulcrum and strongly
elevated above the posterior bands distally. D. wandongensis
also differs from the present species in having a pygidium with
a well defined axial terminus and a shorter mucro that is very
broad at the base, and a hypostome with deeper middle, lateral
border and posterior border furrows, and a posterior margin
that is transverse in outline medially instead of parabolic.
The specimens are poorly preserved and mostly fragmentary,
especially the cephala. Assignment to Bessazoon is based on
the finely granulose cephalic ornament, the entire anterior
cephalic margin, the curved posterolateral pygidial margin and
the long, slender mucro merging with the axial terminus. The
type species B. tenuimucronatum differs from the present one in
having slightly inflated lateral glabellar lobes, a more strongly
curved (exsag.) palpebral lobe, a deeper palpebral furrow, and
the posterior pleural bands on the pygidium more expanded and
prominent distally than the anterior bands rather than the
reverse. B. tigerense from the upper Llandovery of Tasmania
has a more strongly curved palpebral lobe, a deeper palpebral
furrow, and less robust postaxial ridge and mucro.
Preodontochile Degardin and Pillet, 1984
Type species. Dalmanites ( Preodontochile ) camprodonensis
Degardin and Pillet, 1984 from the central Pyrenees, Spain, by
original designation. The precise age of D. ( P .) camprodonensis
is uncertain, as Degardin and Pillet stated (p. 87) that the species
occurs with graptolites of the early Wenlock Monograptus
riccartonensis Biozone, but elsewhere (fig. 4) they showed its
stratigraphical range as lying in the upper Llandovery.
Remarks. Degardin and Pillet (1984) erected the monotypic
Dalmanites ( Preodontochile ) for a poorly known species
represented by few and poorly preserved specimens including a
single crushed cranidium, an isolated thoracic segment and a
number of pygidia that are mostly incomplete posteriorly. The
material permits only a limited assessment of Preodontochile,
but as diagnostic of their subgenus Degardin and Pillet listed
the small eye, the course of the anterior branch of the facial
suture, and the multisegmented pygidium with a short, blunt
mucro. They likened the facial suture both to that of Dalmanites
in being situated in close proximity to the glabella and to that of
Odontochile in being separated from the glabella by the
preglabellar furrow and a narrow band of the anterior cephalic
border. However, as noted by Whittington and Campbell (1967),
the distinction between Dalmanites and Odontochile on the
basis of the anterior cephalic morphology is not as clear-cut as
stated by Richter, Richter and Struve (1959). Several Wenlock-
Ludlow species of Dalmanites, including the Swedish D.
imbricatulus (Angelin, 1851) (see Ramskold, 1985), the North
American D. puticulifrons Whittington and Campbell, 1967
and D. rutellum Campbell, 1967, and the Australian D.
wandongensis, have a narrow band of the anterior border
enclosed by the facial suture on the cranidium. This condition
is also present in the Victorian Llandovery dalmanitid described
below as Preodontochile springfieldensis (see Fig. 16A), which
further resembles P. camprodonensis in the greatly reduced eye
situated far forwards opposite L3, the robust genal spine, the
short, bluntly pointed mucro and the finely granulose sculpture.
We consider this combination of characters to be of generic
significance. With better understanding of P. camprodonensis
other cephalic features of P. springfieldensis may also prove to
be diagnostic of the genus, such as the uniformly narrow (sag.,
exsag.) anterior border that is less than half the width of the
lateral border and lacks a median projection. In its pygidium
with a large number of segments and narrow axis, P.
camprodonensis resembles late Silurian-Devonian species
assigned to Odontochile and closely related genera. Degardin
and Pillet (1984) considered a large number of pygidial
segments as diagnostic of Preodontochile, but in view of the
otherwise close similarity of the type species with the more
poorly segmented P. springfieldensis the number of pygidial
segments is here regarded as only of specific significance.
Other dalmanitid genera known from the Llandovery
include Bessazoon (see above), Daytonia Holloway, 1981 and
Prodontochile Kobayashi and Hamada, 1971, all of which are
easily distinguished from Preodontochile by their much larger
eyes. In addition, Bessazoon differs from Preodontochile in
having the facial suture tightly enclosing the glabella, and a
long, slender mucro connected to the pygidial axis by a
postaxial ridge; Daytonia has the occipital ring markedly
reduced in length abaxially, SI bifurcate adaxially and
converging slightly with S2 abaxially, the anterior cephalic
border as wide as the lateral border, short and slender genal
spines, and a pygidium with parabolic outline and a tiny mucro
joined to the axis by a postaxial ridge; and Prodontochile has
a narrower lateral cephalic border and slender genal spines.
Wenlock-early Ludlow species of Dalmanites assigned by
Ramskold (1985) to his group around the type species D.
caudatus differ from Preodontochile in having a large eye
extending from opposite L3 to opposite LI, an anterior cephalic
border with a well-developed medial process, tuberculate
sculpture on the glabella, prominent lateral nodes on some
thoracic axial rings, pleural nodes on some pygidial segments,
and a long pygidial mucro.
Early Silurian phacopide trilobites from central Victoria, Australia
243
Preodontochile springfteldensis sp. nov.
Figures 15, 16
Type material. Holotype NMV P312070 (cephalon with broken and
displaced genal field). Paratypes NMV P312071 (incomplete
cephalothorax showing damage to genal field), NMV P139350
(incomplete and partly disarticulated cephalothorax showing cephalic
doublure and hypostome), NMV P312074 (incomplete pygidium),
NMV P312075 (incomplete and partly disarticulated thoracopygon).
All from PL1369, Springfield.
Other material. NMV P139350-P139352, NMV P312072-
P3 12073, from PL1369, Springfield.
Diagnosis. Preodontochile with eye oriented slightly obliquely
to exsagittal line and visual surface having approximately 40
lenses arranged in about 15 files of up to 4 lenses each. Posterior
cephalic border markedly expanding (exsag.) abaxially, genal
spine very broad proximally and tapering strongly distally.
Pygidial axis about 25% maximum pygidial width anteriorly,
with 10 axial rings, first 5 well defined by inter-ring furrows
that contain apodemal pits laterally and are shallow medially,
posterior rings poorly defined by shallow inter-ring furrows
that are not continuous medially. Pleurae with about 7 shallow
pleural furrows and very weakly impressed interpleural furrows.
Mucro with length (measured from terminus of axis) comprising
about 25% sagittal pygidial length. Pygidial doublure wide.
Description. Cephalon (excluding genal spines) semi-circular
in outline, anterior margin with very weak convex-downward
medial flexure in anterior profile. Glabella weakly convex,
slightly narrower across LI than across occipital ring, thereafter
expanding moderately forward, a little more strongly in front of
S3 than behind, width across frontal lobe 170-180% occipital
width and about 80% sagittal length of glabella. Occipital ring
raised slightly higher than remainder of glabella, width (tr.)
about 30% cephalic width across same transverse line, length
about 12% cephalic length sagittally, shorter (exsag.) laterally.
Median section of occipital furrow gently convex forward and
very shallow, lateral section (approximately 30% total width)
with arcuate, slit-like apodemal pit not quite reaching axial
furrow distally. SI and S2 defined only by slit-like apodemal
pits that are equal in width (tr.) to occipital apodemal pit and
placed directly in front of it; SI gently arcuate, equidistant from
occipital furrow and S2; S2 transverse, situated at 33% cephalic
length from posterior. L3 as long (exsag.) as LI and L2
adaxially, 160% as long abaxially. Adaxial portion of S3
oriented at about 20° to transverse, abaxial portion very wide,
lenticular in outline, oriented at about 35° to transverse. Frontal
lobe comprising half sagittal length of glabella, elliptical in
outline, with short, shallow sagittal depression in posterior
half. Axial furrow moderately impressed, preglabellar furrow
shallow. Anterior cephalic border of uniform length (sag.,
exsag.), comprising about 8% sagittal cephalic length,
subhorizontal, weakly concave. Lateral border furrow defined
mainly by change in slope, lateral border very wide (tr.), twice
width of anterior border, gently sloping, weakly concave.
Posterior border furrow deep, adaxial portion transverse,
abaxial portion directed posterolaterally, terminating before
reaching lateral border furrow. Genal spine long, with
longitudinal furrow dividing horizontal lateral portion from
steeply sloping adaxial portion. Eye with length about 20%
sagittal cephalic length, anterior edge placed opposite front of
L3 and posterior edge just in front of level of S2. Palpebral lobe
not greatly raised above palpebral area, crescentic in outline,
narrow (tr.). Palpebral furrow weak. Visual surface with lower
margin subtending an angle of about 80° in dorsal view.
Anterior branch of facial suture diverging gently forward
towards widest part of frontal lobe, posterior branch weakly
sinusoidal, meeting lateral cephalic margin more or less
opposite posterior edge of eye. Librigenal field weakly convex,
about 66% width of lateral border.
Cephalic doublure flat anteromedially, approximately 33%
sagittal cephalic length, crossed by weak arcuate furrow
defining crescentic area adjacent to hypostomal suture. Lateral
to hypostomal suture, doublure with upturned inner flange that
is continuous posteriorly with adaxial surface of genal spine.
Only available hypostome is incomplete posteriorly. Anterior
wing small, middle body with parabolic posterior outline,
middle furrow indistinct, posterior portion of lateral border
furrow and posterior border furrow shallow.
Anterior part of thoracic axis subparallel-sided and
comprising about 33% segmental width (tr.) (fig. 15A-B, D,
F), posterior part of axis relatively narrower (fig. 16G). Axial
rings weakly convex (sag.), with indistinct lateral lobes that
lack nodes. Axial furrow moderately impressed. Pleural
furrows sigmoidal, short (exsag.) and sharply impressed, at
fulcrum dividing segment into anterior and posterior bands of
equal length (exsag.) and height. Distal ends of pleurae rounded
anteriorly and angular posteriorly.
Pygidium moderately convex transversely, subtriangular
in outline with weakly convex sides converging posteriorly at
about 60°. Axis tapering uniformly backward, 2nd segment
with broad, subrectangular pseudo-articulating half ring,
much smaller and weaker pseudo-articulating half rings
possibly present on next two segments. Mucro triangular,
postaxial ridge absent.
Exoskeleton finely granulate, lacking tubercles.
Remarks. The poor preservation of the only available cranidium
of P. camprodonensis does not permit detailed comparison
with P. springfieldensis, but the latter differs in the less
obliquely oriented eye and the greater abaxial expansion of the
posterior border. Differences in the form of SI and S2, which
in springfieldensis are isolated and very narrow (exsag.) but in
camprodonensis appear wider (exsag.) and deeper, with SI said
to be connected medially (Degardin and Pillet, 1984, p. 87, fig.
5), can be attributed to crushing in the latter. The most striking
difference between P. springfieldensis and P. camprodonensis
is in pygidial segmentation. Pygidia of the type species have
about 20 axial rings, at least 15 deep and medially continuous
inter-ring furrows, at least 11 wide and deep pleural furrows,
and very strongly incised interpleural furrows. In contrast,
pygidia of springfieldensis have only ten axial rings, five
continuous inter-ring furrows that are shallow medially, seven
narrow (exsag.), shallow to moderately impressed pleural
furrows, and very weak interpleural furrows.
The occurrence of P. springfieldensis at PL1369 together
with another relatively small-eyed form (Phacopidae gen.
indet. 3) is significant. Eye reduction in phacopids and other
244
Andrew C. Sandford and David J. Holloway
Figure 15. Preodontochile springfieldensis sp. nov., from PL1369, Springfield. A-B, G, paratype NMV P312071, incomplete cephalothorax with
damaged genal field; A-B, x 2.2; G, x 2.7. C, E (and figs 16A, E-F), holotype NMV P312070, cephalon with left cheek broken and displaced; C,
x 2; E, x 1.8. D, F (and fig. 16H), paratype NMV P139350, cephalothorax; D, x 1.8, with glabella showing tool marks from preparation; F, x 2.0,
showing cephalic doublure and hypostome. (A, C, E, G are internal moulds).
Early Silurian phacopide trilobites from central Victoria, Australia
245
Figure 16. Preodontochile springfieldensis sp. nov., from PL1369, Springfield. A, E-F, holotype NMV P312070, x 2. B, paratype NMV P312074,
pygidium, x 2. C, G, paratype NMV P312075, thoracopygon; C, x 3; G, x 2.7. D, paratype NMV P312072, cranidium and displaced cheek, x 5.
H, paratype NMV P139350, enlargement of eye, x 8. 1, NMV P139351, three thoracic segments, x 1.5. (B-C, E are internal moulds).
trilobites has been widely interpreted as an adaptation to deep
water, subphotic environments, although Alvaro and Vizcaino
(2003) noted that high turbidity may create subphotic
environments in shallower settings. The taphonomy of the
Deep Creek trilobite population is indicative of a deep water
facies rather than turbid conditions. The degree of articulation
is high (isolated tergites 40%), and although there are no fully
articulated exoskeletons, specimens have been found with the
hypostome only slightly displaced along the hypostomal suture
(fig. 15F), and with incomplete thoraces attached to the
cephalon or only slightly detached from it or from the pygidium
(figs 15A-B, D, G, 16G). This evidence suggests deposition at
depths below normal wave base where bottom current activity
is negligible, preventing displacement, winnowing and
concentration or reworking of the exoskeletal elements. These
deep water beds correlate in age with the late Telychian (mid
Monoclimacis crenulata Biozone) eustatic highstand (event 4)
documented from six palaeocontinents by Johnson (1996).
The abrupt transition from these beds to the shallower water
facies of the overlying Chintin Formation may reflect the
influence of an extensive latest Telychian Gondwanan
glaciation event documented from Brazil (Grahn and Caputo,
1992). Otherwise, the Chintin Formation appears to correspond
closely in age to the subsequent regressive phase that
culminated in an earliest Wenlock lowstand.
Struveria Rickards, 1965
Type species. Struveria howgillensis Rickards, 1965 from the
upper Wenlock-lower Ludlow of northern England and North
Wales, by original designation.
Remarks. Apart from the type species and the Victorian
specimens described below, Struveria includes S. orba
(Barrande, 1852) from the upper Wenlock of the Czech
Republic, and S. simrica (Hede, 1915) from the upper
246
Andrew C. Sandford and David J. Holloway
Llandovery?-Wenlock of southern Sweden (see Laufeld et al.,
1975) and the upper Wenlock of Poland (Tomczykowa, 1991).
S. howgillensis is the most completely known species (Rickards,
1965: pi. 85, figs l-6)butneitherthisnorthe other representatives
have been extensively illustrated, and most are rather poorly
preserved. Consequently the genus is not very well known
morphologically, but distinctive characters include: a very short
(sag., exsag.) anterior cephalic border; almost transverse SI
and S2, both containing apodemal pits; a moderately large eye
extending very close to the lateral border furrow anteriorly and
laterally; a long, flattened genal spine; a pygidium that is curved
in outline posterolaterally and lacks a convex border or mucro;
pygidial axial rings bearing oblique muscle impressions
laterally, except for the last few; a narrow, rather sharp postaxial
ridge; short (sag., exsag.), sharply impressed pygidial pleural
and interpleural furrows, the pleural ones terminating distally
a short distance from the margin whereas the interpleural ones
reach the margin; and posterior pleural bands that are slightly
more raised distally than the anterior bands towards the back of
the pygidium.
Struveria orba was tentatively assigned to Delops by
Snajdr (1982), but the lectotype cranidium and other specimens
figured by Snajdr (1982: pi. 2, figs 5-10) resemble the type
species of Struveria in the strongly transverse frontal lobe and
distinctly concave lateral outline of the glabella behind a
transverse line through the inner end of S3, and they lack the
tuberculate sculpture and expanded (exsag.) pygidial pleural
furrows characteristic of Delops. Two pygidia assigned to orba
by Barrande, one of them a paralectotype (Barrande 1852: pi.
26, fig. 38; Snajdr 1982: pi. 2, fig. 1), are not conspecific with
the lectotype but belong to Delops dermolac (see also Budil,
1996). ‘ Calymenel daviesii Salter, 1865, known only from an
internal mould of a pygidium from the upper Wenlock of
Wales, was tentatively assigned to Struveria by Morris (1988:
223) but we cannot assess this assignment from the woodcut
illustration of Salter (1865) (fig. 23).
As pointed out by Rickards (1965: 549) and Bergstrom (in
Laufeld et al., 1975: 219), Struveria shows similarities to
Eudolatites, a genus largely restricted to the Caradoc apart
from the late Ashgill subgenus E. (. Deloites ) Destombes, 1972,
which was regarded as an independent genus by Tomczykowa
(1991) and Vanek and Vokac (1997). Eudolatites differs from
Struveria in that SI is more obliquely oriented; S2, though
commonly rather deep, seems to lack an apodemal pit (Destombes,
1972: pi. 3, fig. la, pi. 4, fig. la); the librigenal field is much
broader anterior and lateral to the eye; there is either no genal
spine present, or it is short, strongly tapered, and rounded in cross
section distally rather than flattened (see Rabano in Gutierrez
Marco and Rabano, 1987: pi. 1, fig. lc); and the pygidium lacks a
sharply defined postaxial ridge.
Struveria sp. 1
Figures 12B-C
Material. NMV P139357 (partly disarticulated dorsal exoskeleton with
displaced hypostome, thorax and pygidium) from PL1386, Geological
Survey locality B3 (exact locality unknown), Moonee Ponds Creek,
Broadmeadows. Precise stratigraphic horizon uncertain.
Remarks. The cephalon of the specimen has the glabella except
for the anteriormost part obscured by the displaced hypostome
and medial part of the doublure. Much of the left fixigena is
preserved, including the large palpebral lobe and the posterior
border, the latter curving backwards abaxially to the base of
the genal spine. The pygidium is more completely preserved
than the cephalon, is almost twice as wide as long (sag.), with
six or seven axial rings and pleural furrows, and the axis
comprising a little more than 25% maximum pygidial width
anteriorly. The specimen can be assigned to Struveria with
confidence on the basis of the pygidial morphology.
The hypostome of Struveria has not previously been
illustrated. That of the present specimen is similar to
hypostomes of other dalmanitids (e.g. see Ramskold, 1985, pi.
10, fig. 12).
Struveria sp. 2
Figures 13L, 17A, 17C
Dalmanitina {Eudolatites) aborigenum Opik, 1953: 26 ( partim .),
pi. 10, figs 86-87 {non pi. 10, fig. 85 = Dalmanites athamas Opik,
1953).
Material. NMV P52483 (fragmentary pygidium), from PL2269,
Thomas locality F51, Costerfield. Wapentake Formation. For locality
see Thomas (1960). NMV P139337 (cranidium), NMV P138209
(pygidium), from PL206, Wallan. Bylands Siltstone.
Remarks. Compared to the pygidium of Struveria sp. 1 from
Broadmeadows, the pygidium from Wallan (fig. 17A) is longer
(maximum width about 150% sagittal length), with a wider
axis (almost 33% maximum pygidial width anteriorly) and a
greater number of axial rings (nine or ten) and pleural furrows
(ten or 11). It is unlikely that these differences can be explained
by the larger size of the Wallan pygidium, and we therefore
consider it to belong to a separate species. The cranidium from
Wallan (fig. 17C) is small and indifferently preserved but its
morphology is consistent with assignment to Struveria.
The paratype of ‘ Dalmanitina {Eudolatites)' aborigenum
is a fragmentary pygidium from Costerfield, consisting of the
anterior part of the axis and the adjacent part of the right
pleural lobe (fig. 13L). It differs from pygidia of Dalmanites
athamas, to which the holotype of aborigenum belongs, in
that the inter-ring furrows do not shallow markedly medially
but are deeply impressed across the entire width of the axis;
the pleural furrows are not expanded but are short (exsag.),
sharply impressed and similar in appearance to the interpleural
furrows; and the anterior and posterior pleural bands are flat-
topped. These characters suggest that the specimen belongs to
Struveria, and it may be conspecific with the Wallan specimens
which come from a similar stratigraphical level.
Struveria 1 ! plinthourgos sp. nov.
Figure 18
Eudolatites sp.— Rickards and Sandford, 1998: 751.
Type material. Holotype NMV P79125 (internal mould of dorsal
exoskeleton) from PL1374, old Camberwell brick pit. Rose Street,
Camberwell. Anderson Creek Formation.
Early Silurian phacopide trilobites from central Victoria, Australia
247
Figure 17. A, C, Struveria sp. 2, from PL206, Wallan. A, NMV P138209, pygidium, x 2. C, NMV P139337, cranidium, x 5.5. B, Dalmanitidae indet.,
NMV P127957, partial view of incomplete thorax, x 0.9, from old Costerfield Antimony Mine, Costerfield. D-F, Bessazoon sp. D, NMV P147769,
pygidium (fragment), x 1.8, from PL256, Wallan. E, NMV P312817, cranidium (fragment), x 2.2, from PL6361, Springfield. F, NMV P147770,
pygidium (fragment), x 2.4, from PL256, Wallan.
Derivation of name. Greek, ‘brickmaker’, in reference to the
type locality.
Diagnosis. Glabella subpentagonal in outline, expanding
strongly and rather uniformly forward, width across frontal lobe
about twice width across occipital ring and approximately equal
to sagittal length. Eye short (exsag.), with posterior edge opposite
S3. Pygidium with strongly convex (tr.) axis of 10 or 11 rings, 9
pleural furrows, and relatively broad, concave border on which
pleural and interpleural furrows are very weakly expressed.
Description. Cephalon with length (sag.) about 60% maximum
width. Glabella comprising a little less than 30% cephalic
width posteriorly. Occipital furrow shallowing and deflected
forward medially. SI and S2 transverse, meeting axial furrow
distinctly, adaxial portions with slit-like apodemal pits situated
in line exsagittally, S2 comprising 33% glabellar width at this
level and with apodemal pit occupying half width (tr.) of furrow.
LI approximately 80% length (exsag.) of L2 laterally. S3
diverging forward at about 125°, shallow adaxially and
deepening slightly abaxially but not expanding appreciably. L3
a little more than twice as long (exsag.) abaxially as adaxially.
Frontal lobe comprising approximately half sagittal length of
glabella, rhombic in outline, apparently with short (exsag.)
longitudinal depression at about midlength. Eyes and palpebral
lobes not preserved, palpebral furrow apparently shallow,
weakly curved and oriented slightly oblique to exsagittal line.
Posterior branch of facial following gently sigmoidal course,
directed slightly forward across most of genal field and
deflected backward laterally. Posterior border furrow deeply
impressed, apparently deflected forward slightly abaxially and
dying out distally before reaching lateral border furrow.
Backward deflection of posterior cephalic margin distally
indicates that genal spine was present (fig. 18D) but its length is
unknown. Fixigenal field densely pitted.
Thorax of 11 segments. Axis strongly convex (tr.),
increasing slightly in width from 1st to 4th or 5th segment and
thereafter narrowing at similar rate, width on 1st segment
equal to that on 8th segment and almost 120% that on last
segment. Axial rings gently convex (sag., exsag.), without
lateral lobes. Axial furrow deeply impressed. Pleural lobe
slightly wider (tr.) than axis in anterior half of thorax and
becoming increasingly wider towards back. Pleurae strongly
downturned beyond fulcrum, with large articulating facets
and pointed tips becoming backwardly deflected towards back
of thorax. Pleural furrows very deep, extending weakly onto
articulating facets distally where they curve slightly forward.
Posterior termination of pygidium unknown; if it is
rounded rather than mucronate or angular, pygidium is about
150% as wide as long (sag.) with parabolic posterior outline;
anterolateral corner well-rounded. Axis strongly convex (tr.),
width 25% maximum pygidial width anteriorly, possibly not
tapering uniformly backward but a little more strongly across
first 3 segments than next 2 or 3, thereafter at similar rate as
initially. 1st ring with broad, shallow median excavation in
posterior margin for pseudo-articulating half ring on 2nd
segment, much weaker excavations present on 2nd and 3rd
rings. Inter-ring furrows 1-6 with apodemal pits laterally,
248
Andrew C. Sandford and David J. Holloway
Figure 18. StruvericP. plinthourgos sp. nov., holotype NMV P79125, internal mould of dorsal exoskeleton, from PL1374, Camberwell; A, x 1;
B-E, x 1.25.
remaining inter-ring furrows very weak. Pleural field rather
strongly convex (tr.), 1st segment with wide articulating facet
extending 66% width of pleura and with deep pleural furrow
not extending onto facet distally, subsequent segments with
successively shallower pleural furrows more closely
approaching interpleural furrows in depth and length (exsag.).
Concave border comprising almost 33% pleural width
anteriorly, not bounded adaxially by distinct border furrow,
outer margin not rolled in transverse section.
Remarks. The only known specimen is an internal mould of a
very large and substantially complete dorsal exoskeleton almost
130 mm long (assuming the pygidium lacks a mucro). The
cephalon is crushed, especially the frontal lobe of the glabella
which bears radiating fractures, so that its flattened profile is
not indicative of the original convexity. The anterior outline of
the glabella in dorsal view is formed by an arcuate fracture,
below which the cephalon slopes steeply downward to a lower
margin that is a smoothly curved line (figs 18C-D); it is not
clear whether this line is the facial suture or the hypostomal
suture, but the former is more likely. The lateral part of the
right cheek has been pushed downward and inward along a
posterolaterally-directed fracture that has destroyed the
palpebral lobe, and of the right librigena only the border and
possibly a small portion of the adjacent field are preserved. On
the left cheek the palpebral lobe has been broken off along the
Early Silurian phacopide trilobites from central Victoria, Australia
249
palpebral furrow, the flattened appearance of the genal spine
base may not be indicative of its original shape in cross-section,
and all that remains of the librigena is the lateral outline (fig.
18D). The distal parts of the last few thoracic segments are
missing on the left side, together with most of the left pleural
lobe and posterior termination of the pygidium.
Generic assignment of the species is problematic, partly
because certain important characters cannot be determined,
including the form of the cephalic borders and genal spine, and
the nature of the pygidial termination. The specimen was
assigned to Eudolatites by Rickards and Sandford (1998),
although that genus is not known to range above the Ordovician,
but features such as the transverse orientation of SI and the
presence of an apodemal pit in S2 are incompatible with such an
assignment. On the basis of these characters and the form of the
pygidial pleural and interpleural furrows we tentatively assign
plintliourgos to Struveria. The species differs from other
members of the genus in that the glabella is subpentagonal in
outline rather than club-shaped (i.e. it is not concave in lateral
outline behind a transverse line through the inner end of S3), the
eye is shorter (exsag.) and does not extend as far backward, the
palpebral furrow is more weakly curved, the genal spine appears
to be smaller at its base, and the pygidium has a concave border;
however ,we can not be certain that the last feature is not caused
by compression of the dorsal exoskeleton onto the doublure.
Dalmanitidae indet.
Figure 17B
Dalmanitina sp. cf. Dalmanitina ( Eudolatites ) aborigenum.—
Opik, 1953 : 28, pi. 10, fig. 92.
Trilobita indet.— Talent, 1964: 51 (pars).
Material. NMV PI 27957 (incomplete thorax) from old Costerfield
Antimony Mine, Costerfield. Costerfield Siltstone. Locality marked
Thomas (1941).
Remarks. This poorly preserved thorax with seven or possibly
eight segments distinguishable is the only fossil recorded from
the Costerfield Siltstone. Based on its large size (about 60 mm
wide) and the form of the pleural furrows, Opik (1953, p. 29)
suggested that it probably belongs to Dalmanitina, a genus
then interpreted more broadly than now, and that it may be
related to his species ‘ Dalmanitina ( Eudolatites )’ aborigenum
from beds higher in the sequence. However, Talent (1964),
regarded the specimen as too poorly preserved for identification
even to family level.
We agree with Opik that the specimen probably belongs to
the Dalmanitidae, and that the short (exsag.), sharply incised
pleural furrows preclude assignment to Dalmanites or its close
allies. However, the holotype of ‘ Dalmanitina ( Eudolatites )’
aborigenum, the species to which he thought it may be related,
belongs to Dalmanites athamas (see above), and the fragmentary
pygidium also included in aborigenum by Opik is here assigned
to Struveria sp. 2. In size, the abaxially subquadrate outline of
the axial rings and the form of the pleural furrows, the poorly
preserved thorax from Costerfield is not unlike Struveria ?
plithourgos described above, and it is possible that the two
forms are related.
Superfamily Acastoidea Delo, 1935
Family Acastidae Delo, 1935
Berylacaste gen. nov.
Type species. Berylacaste berylae gen. et sp. nov.
Derivation of name. After ACS’s mother. Gender feminine.
Diagnosis. Glabella expanding strongly forward, about twice
as wide across frontal lobe as across occipital ring, strongly
inflated, overhanging anterior cephalic border. LI almost as
long (exsag.) as L2, S2 and S3 very shallow. Visual surface
absent, palpebral lobe very small, situated towards front of
genal field, merging with eye ridge that reaches axial furrow,
palpebral suture straight and obliquely oriented. Genal spine
very small, thorn-like. Thoracic pleural tips with small
posterolaterally directed spines. Pygidial axis with 3 rings and
poorly segmented posterior part, pleurae with 4 pleural furrows
and poorly defined border furrow, posteromedian margin
rounded, lacking spine or point.
Remarks. Berylacaste gen. nov. is known only from the type
species. The genus is distinguished from others in the
Acastoidea most notably by the strong forward expansion of
the glabella, the frontal lobe that overhangs the anterior
cephalic border, the vestigial palpebral lobe situated far
forward, and the absence of a visual field. Other distinctive
characters are the very weak impression of S2 and S3, the
presence of genal and thoracic terminal pleural spines, and the
very weak pygidial segmentation.
Edgecombe (1993) recognised the monophyletic
superfamily Acastoidea s.s. comprising the Calmoniidae and
the Acastidae, but excluding more primitive genera such as
Phacopidina, Baniaspis and Kloucekia which he referred to as
Acastoidea s.l. without familial assignment. Berylacaste gen.
nov. can be assigned to his Acastoidea s.s. as it exhibits a
‘shouldered’ anterior cephalic margin (i.e. the median part of
the margin in front of the glabella projects in front of the
outline of the cheeks) and an abrupt change in depth between
the anterior and posterior inter-ring furrows in the pygidial
axis. The Acastidae was considered by Edgecombe (1993) to
consist of a monophyletic Acastidae s.s. together with some
other loosely related genera (Llandovacaste and Australoacaste)
that were assigned to Acastidae s.l. He gave a brief diagnosis
for the Acastidae s.s. that included the following
synapomorphies: SI transverse or anteromedially directed,
lacking proximal bifurcation; LI much shorter (exsag.) than
L2; S2 transverse. Berylacaste conforms to this diagnosis in
the orientation of SI and S2, but SI appears to be slightly
expanded or incipiently bifurcate at its proximal end in some
specimens (e.g. figs 19F-G), and LI is not markedly shorter
than L2. We note, however, that there is some variation in the
length of LI within the Acastidae s.s., and that it may be almost
as long as L2 (e.g. see Richter and Richter, 1954, pi. 3, fig. 37;
Tomczykowa, 1991, pi. 7, fig. 23, pi. 9, figs 7-12, 14, 16-17).
Ramskold and Edgecombe (1993: 265) identified a
‘Wenlock group’ of acastid genera including Acaste, Acastoides
and Acastocephala, some species of which persist into the late
Silurian. Berylacaste is contemporaneous with most members
250
Andrew C. Sandford and David J. Holloway
of that group but shares few of their stated characteristics,
such as a distinctly convex forward S2, a deep sagittal
impression at the back of the glabellar frontal lobe, a deep
preglabellar furrow, and a genal angle that is rounded or bears
a tiny node. The wide glabella considered characteristic of the
‘Wenlock group’ is not as strongly expanded as that of
Berylacaste. In the form of the genal spine and the apparent
lack of a sagittal impression on the frontal glabellar lobe
Berylacaste is more similar to Ludlow-Pndolf species of
Acastella, but in the other cephalic characters listed above
Berylacaste is easily distinguished from those species, which
also differ in having a strongly segmented pygidium with a
spine or blunt point posteriorly. Scotiella (see Shergold, 1967,
pi. 2, figs 1-8, pi. 3, figs 1-8) and Ewacaste Ramskold and
Edgecombe, 1993 (see Schrank, 1972, pi. 21, figs 6-9;
Tomczykowa, 1991, pi. 7, figs 21-24) are like Berylacaste in
the very weak S2 and S3 but have less inflated glabellae that
expand only weakly forward and large eyes, and Scotiella also
has a mucronate pygidium. Berylacaste cannot be considered
primitive; rather, it is a highly derived form evolved
independently of and along different lines to other Acastidae.
Reduction of the visual surface and palpebral lobe also
occurs in the Devonian calmoniid genera Typhloniscus (see
Cooper, 1982), Punillaspis Baldis and Longobucco, 1977 and
Tormesiscus Waisfeld et al., 1994. The eye of Berylacaste
most closely resembles that of Tormesiscus, in which it is also
reduced to a laterally directed palpebral ridge placed anteriorly
on the genae. The glabella of Tormesiscus is similarly
expanded strongly forward and is inflated anteriorly,
overhanging the anterior border. The genera differ in thoracic
and pygidial features, the depth of the glabellar axial furrows
and the course of the facial suture, the latter apparently being
submarginal in Tormesiscus.
Berylacaste berylae gen. et sp. nov.
Figures 19-20
Acastidae n. gen.— Rickards and Sandford, 1998: 752.
Type material. Holotype NMV P138224 (dorsal exoskeleton) and
paratypes NMV P138225 (dorsal exoskeleton), NMV P138226
(cephalon), NMV P139330 (dorsal exoskeleton), NMV P139331
(thoracopygon with displaced pygidium), NMV P139332 (cephalon
with displaced librigena), NMV P139333 (cephalon), NMV P139334.
All from PL206, Bylands Siltstone.
Other material. NMV P138227 from PL206, Wallan. Bylands
Siltstone.
Derivation of name and diagnosis. As for genus.
Description. Exoskeleton elliptical in outline, estimated
maximum length 20 mm. Cephalon strongly convex (sag., tr.),
about twice as wide as long (sag.), anterior margin with
maximum curvature medially in dorsal view and with outline
interrupted anteromedially by slightly protruding glabella.
Glabella rounded-pentagonal in dorsal view, about as wide
across frontal lobe as long (sag.), maximum width situated at
approximately 66% sagittal glabellar length from posterior.
Axial furrows deep, diverging forward at about 55°. Occipital
ring comprising about 30% maximum glabellar width,
approximately twice as long sagittally as distally, with median
node situated close to posterior edge. Occipital furrow
moderately impressed medially and forming convex forward
arc, increasing in depth abaxially and curving slightly forward
distally. LI about as long (exsag.) distally as occipital ring,
apparently with weakly defined lateral node. SI very deeply
impressed laterally and directed slightly obliquely inward and
backward, shallowing abruptly adaxially and curving forward
slightly subparallel to occipital furrow, apparently very weakly
impressed medially. S2 and S3 more distinct on internal moulds
than on exterior of exoskeleton, S2 transverse, reaching axial
furrow, S3 directed obliquely inwards and backwards from
axial furrow in concave-forward curve, inner end situated
opposite glabellar midlength (sag.). Frontal lobe transversely
elliptical or rhombic, lacking sagittal furrow. Anterior border
subvertical. Genal field moderately convex (tr.), steeply sloping
laterally and in front of palpebral lobe. Palpebral lobe slightly
raised, placed opposite anterior half of L3, palpebral furrow
shallow, oblique and weakly curved, continuous adaxially with
weak furrow behind eye ridge. Anterior branch of facial suture
diverging forward in gentle curve subparallel with axial furrow,
not cutting across anterolateral comer of glabella, posterior
branch directed posterolaterally across genal field and inner
part of lateral border, deflected sharply backward across outer
part of border to meet cephalic margin opposite LI. Librigena
lenticular in outline with long posterior projection. Posterior
border narrow proximally, gently widening abaxially to about
midpoint (tr.), thereafter uniform in width and gently curving
forward. Posterior border furrow deep proximally, moderately
impressed distally and continuous with shallower lateral border
furrow. Lateral border weakly convex, steep and poorly defined
anteriorly. Genal spine diverging slightly from line of lateral
margin. Cephalic doublure strongly convex laterally, widening
and flattening medially, no vincular furrow.
Hypostome unknown.
Thorax of 11 segments. Axis weakly tapering backward,
comprising about 30% segmental width at front of axis and
about 25% toward the back. Axial rings strongly convex (sag.),
flexed forward distally, lateral lobes not defined. In transverse
profile pleurae convex adaxially, concave abaxially. In dorsal
view pleurae convex forward adaxially, concave forward
abaxially, posterior edge of segment curving slightly forward
distally to spinose tip. Pleural furrows short (exsag.) and deep,
extending onto articulating facet distally. Anterior pleural band
approximately as long (exsag.) as posterior band at fulcrum.
Pygidium lenticular, about twice as wide as long (sag.).
Axis as wide as pleura anteriorly, comprising about 75%
sagittal pygidial length, weakly tapering backwards and
broadly rounded posteriorly. 1st axial ring high, 2nd ring well
defined, 3rd ring poorly defined. Pleurae moderately convex
(tr., exsag.), comprising at least 5 segments, 1 st 4 four with
posterior pleural bands slightly more raised than anterior bands
on internal moulds and extending onto border distally. Only 2
pleural furrows distinct, 1st sharply incised, not extending
onto articulating facet distally, 2nd reaching halfway to margin.
1st 2 interpleural furrows weaker than pleural ones. Pygidial
margin smooth on exterior of exoskeleton, weakly scalloped
on internal mould. Pygidial doublure narrow, convex.
Early Silurian phacopide trilobites from central Victoria, Australia
251
Figure 19. Berylacaste berylae gen. et sp. nov., from PL206, Wallan. A, paratype NMV P138224, dorsal exoskeleton, x 4. B-C, paratype NMV
P139332, cephalon with displaced librigena, x 5. D-E (and fig. 20D), paratype NMV P139331, thoracopygon; D, x 8; E, x 4. F-H, paratype NMV
P139333, cephalon, x 4. I-J (and fig. 20C), paratype NMV P138225, dorsal exoskeleton with down-flexed pygidium, x 5. K, holotype NMV
P138224, cephalothorax, x 5. (A, C, F, H, I, K are internal moulds).
Remarks. The specimens are rather poorly preserved and affected
by compression, shearing and fracturing. As a result there is
apparent variation in the depth of some furrows, such as the
medial part of SI (compare fig. 19A with 19B-C, G) and the
lateral cephalic border furrow (compare figs 19B-C with 19H).
Some other morphological features are unclear, such as the extent
to which lateral nodes are defined on LI by adaxial expansion
and/or bifurcation of SI (compare figs 19F-G with 19K).
252
Andrew C. Sandford and David J. Holloway
Figure 20. Berylacaste berylae gen. et sp. nov., from PL206, Wallan. A-B, E, paratype NMV P138226, cephalon; A-B, x 4; E, enlargement of
genal angle showing thorn-like genal spine. C, paratype NMV P138225, enlargement of pygidium, x 10. D, paratype NMV P139331, enlargement
of pygidium, x 8.
The blind condition of Berylacaste reinforces other
evidence for a deep-water, possibly sub-photic environmental
setting for the trilobite fauna at PL206, Wallan. Berylacaste
berylae is well-represented, but the fauna is dominated by the
blind, effaced styginid Thomastus aops Sandford and Holloway,
1998. Together these blind trilobites represent 63% of the
trilobite population and 22% of the faunal diversity. Furthermore
Ivops wallanensis, representing 19% of the trilobite population,
bears an eye markedly smaller than other Wenlock phacopids.
The taphonomy of the fauna at PL206 contains a high
proportion of partly articulated exoskeletons (60%) and
indicates assignment to trilobite taphofaciesTIV (see Sandford,
2002), considered to represent depths below maximum storm
wave base. Exoskeletons of T. aops showing partly displaced
librigenae may be moult assemblages characteristic of
taphofacies TIV, although as the amount of displacement is
small it is more likely due to post-depositional compaction.
Four other acastids are known from central Victoria, all in
the Lower Devonian. Acaste lokii Edgecombe, 1993 and
Acastella frontosa Shergold, 1968 are from the Humevale
Siltstone at Lilydale, and Acastella sp. was recorded by
Holloway and Neil (1982) from the Mt Ida Formation at
Heathcote. These species are easily distinguished from
Berylacaste berylae by their large eyes and deeply impressed
S2 and S3. A distinctive, as yet undescribed acastid with long
occipital and thoracic axial spines occurs in the Humevale
Siltstone at Yarra Junction and at Kinglake West.
Acknowledgements
We thank Dr Greg Edgecombe (Australian Museum, Sydney),
Dr Catherine Cronier (Universite des Sciences et Technologies
de Lille 1), and Prof. Tony Wright (University of Wollongong)
for their helpful reviews of the manuscript.
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Memoirs of Museum Victoria 63(2): 257-287 (2006)
ISSN 1447-2546 (Print) 1447-2554 (On-line)
http://www.museum.vic.gov.au/memoirs/index.asp
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004
(Echinodermata: Asteroidea)
P. Mark O’Loughlin 1 and Francis W.E. Rowe 2
honorary Associate, Marine Biology Section, Museum Victoria, GPO Box 666, Melbourne, Vic. 3001, Australia
(pmo@bigpond.net.au)
2 Research Associate, Australian Museum, Sydney, NSW, Australia (fwer.goldbrookenn@btinternet.com). Private address:
Beechcroft, Norwich Road, Scole, Diss, Norfolk, IP21 4DY, U.K.
Abstract O’Loughlin, P. Mark and Rowe, Francis W.E. A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004
(Echinodermata: Asteroidea). Memoirs of Museum Victoria 63(2): 257-287.
The Indo-west Pacific Aquilonastra O’Loughlin is reviewed. Eleven species are retained in Aquilonastra : A. anomala
(H.L. Clark); A. batheri (Goto); A. burtonii (Gray); A. cepheus (Muller and Troschel); A. corallicola (Marsh); A. coronata
(Martens); A. iranica (Mortensen); A. limboonkengi (Smith); A. minor (Hayashi); A. rosea (H.L. Clark); A. scobinata
(Livingstone). Asterina lorioli Koehler is reassigned to Aquilonastra. Thirteen new species are described: A. byrneae; A.
colemani; A. conandae; A. doranae; A. halseyae; A. marshae; A. moosleitneri; A. oharai; A. richmondi; A. rowleyi; A.
samyni; A. watersi; A. yairi. The four subspecies of Asterina coronata Martens are junior synonyms: Asterina coronata
cristata Fisher; Asterina coronata euerces Fisher; Asterina coronata fascicularis Fisher; Asterina coronata forma
japonica Hayashi. The 13 fissiparous Red Sea specimens described by Perrier as Asteriscus wega are the syntypes.
Asteriscus wega Perrier is a junior synonym of Asterina burtonii Gray. The provisional referral to Aquilonastra of three
species is discussed: A. rosea; A. rowleyi; A. scobinata. No fissiparous species develops into a non-fissiparous pentaradiate
form. A key to the species of Aquilonastra and map of type localities are provided.
Introduction
A revision of Aquilonastra O’Loughlin (in O’Loughlin and
Waters, 2004) was anticipated by O’Loughlin and Rowe
(2005), and its scope has grown through loans and donations
of an abundant array of asterinid specimens and photos from
the Indo-Pacific, Red Sea and Mediterranean regions.
This revision and description of 13 new species are based
on morphological observations. O’Hara et al. (in preparation)
are currently working on a molecular phylogeny analysis of
species of Aquilonastra, and will examine congruity with the
morphological observations. Byrne (2006) reported that for
Aquilonastra a phylogeny potentially provides evidence of a
life history transformation series, from planktonic feeding to
planktonic non-feeding to the benthic non-feeding mode of
development, all three modes clustered in Aquilonastra with
A. minor a terminal taxon. Three species are referred to
Aquilonastra provisionally: A. rosea (H.L. Clark, 1938), A.
rowleyi sp. nov. and A. scobinata (Livingstone, 1933).
We recognize limitations in this revision. Images for
fissiparous specimens on the Seychelles suggest another new
species, but no material was available (see figs 3k, 31). Sloan et
al. (1979) reported Asterina burtoni for Aldabra Atoll as a
species with fissiparous and non-fissiparous forms. They
described the non-fissiparous form as having five rays and single
madreporite. We recognize throughout this work two such
forms as distinct species. No material was examined to establish
the status of these species. Indeed, we are unable to support the
view that pluriradiate, fissiparous asterinids represent juveniles
of larger, non-fissiparous pentaradiate adults (see Clark, 1967b
for review). James (1975, 1982, 1985, 1989) reported Asterina
burtoni from Lakshadweep (Laccadives) and Indian and Sri
Lankan seas. Again, no material was examined to confirm
systematic status. Sastry (1991) also reported Asterina burtoni
from Lakshadweep, that appears to have represented non-
fissiparous and fissiparous forms. No material was examined to
confirm systematic status. The evidence in this paper indicates
that Aquilonastra species have local geographical ranges. Thus
some species reported here with extensive distributions (such as
Oman to South Africa, and northern Australia to China) may
prove with further analysis to be more than one species.
258
P. Mark O’Loughlin and Francis W.E. Rowe
Body form, numbers of spinelets and spines per plate, and
size of spinelets and spines are all related to specimen size for the
25 Aquilonastra species. And all of these characters show some
variation for specimens of the same species at the same size, and
on the same specimen. Preservation history is another factor
affecting size and form of the various morphological characters.
Species diagnostic characters are always given here for nominated
specimen sizes, and for dried specimens. All of the Aquilonastra
species show mottled live colour, and within a species the mottled
colour can vary greatly. Colour is sometimes uniform on a
specimen. However, some colours predominate in some species.
Colour appears to vary with substrate colour. A full generic
diagnosis for Aquilonastra is given, and these morphological
characters are not repeated in each species diagnosis. Terminology
follows O’Loughlin and Waters (2004).
Abbreviations for institutions are: AM— The Australian
Museum, Sydney; HUJ— The Hebrew University of Jerusalem;
MNHN— Museum National d’Histoire Naturelle, Paris;
MRAC— Museum of the Republic of Central Africa, Brussels;
NHM— The Natural History Museum, London; NMV—
Museum Victoria, Australia; NSMT— National Science
Museum, Tokyo; TAU— Tel-Aviv University; TM— Tasmanian
Museum, Australia; UF— The University of Florida; WAM—
The Western Australian Museum. Specimen registration
number prefixes are: AM J; MNHN EcA; NMV F; TM H;
WAM Z. MAU is MNHN collection code for Mauritius.
Photography for many figures was performed using a Leica
MZ16 stereomicroscope, DC300 Leica digital camera, and
“Auto -Montage” software for composition of images.
Table 1. Species of Aquilonastra O’Loughlin, 2004, with type localities.
Species Type localities
A. anomala (H.L. Clark, 1921)
A. batheri (Goto, 1914)
A. burtonii (Gray, 1840)
A. byrneae sp. nov.
A. cepheus (Muller and Troschel, 1842)
A. colemani sp. nov.
A. conandae sp. nov.
A. corallicola (Marsh, 1977)
A. coronata (Martens, 1866)
A. doranae sp. nov.
A. halseyae sp. nov.
A. iranica (Mortensen, 1940)
A. limboonkengi (Smith, 1927)
A. lorioli (Koehler, 1910)
A. marshae sp. nov.
A. minor (Hayashi, 1974)
A. moosleitneri sp. nov.
A. oharai sp. nov.
A. richmondi sp. nov.
A. rosea (H.L. Clark, 1938)
A. rowleyi sp. nov.
A. samyni sp. nov.
A. scobinata (Livingstone, 1933)
A. watersi sp. nov.
A. yairi sp. nov.
Australia, Torres Strait, Murray I.
Japan
Red Sea
Australia, Great Barrier Reef, One Tree I.
Indonesia, Jakarta (as Batavia)
SE Papua New Guinea, China Straits
W Indian Ocean, La Reunion I.
W Pacific Ocean, Caroline Is., Palau
Indonesia, Molucca and Flores Is.
S Japan, Ryukyu Is., Okinawa, Henza Island
N Indian Ocean, Maidive Is.
Iranian Gulf
China, Amoy
Pakistan, Karachi
Red Sea
Japan, Honshu, Kushimoto
N Indian Ocean, Maidive Is.
S Japan, Ryukyu Is., Okinawa, Seragaki
Tanzania, Ras Kimbiji (central coast)
SW Australia, off Perth, Rottnest I.
SE Africa, Sodwana Bay
Arabian Sea, Oman, Masirah I.
Tasmania
Arabian Sea, Oman, Masirah I.
E Mediterranean Sea, Israel, Michmoret
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
259
Key to Aquilonastra O’Loughlin species
1. Typically 5 equal or subequal rays, sometimes 6; form
symmetrical; single conspicuous madreporite, rarely 2,
very rarely 3 2
— Typically more than 5 rays, up to 9, in unequal size groups;
form asymmetrical; always more than 1 inconspicuous
madreporite 18
2. Gonopores actinal 3
— Gonopores abactinal 5
3. Abactinal plates paxilliform; spinelets in dense, frequently
crescentiform, clusters; spinelets pencil-like
A. scobinata (SE Australia)
— Abactinal plates not paxilliform; spinelets not in dense
clusters; spinelets not pencil-like 4
4. Abactinal plates with low rounded elevations; spinelets
subpaxilliform; maximum R = 9 mm A. minor (Japan)
— Abactinal plates lacking rounded elevations; spinelets not
subpaxilliform; maximum R = 15 mm
A. byrneae (NE Australia, Mariana Is.)
5. Abactinal spinelets in dense round paxilliform clusters
^ A. rosea (SW Australia)
— Abactinal spinelets not in dense paxilliform clusters 6
6. Projecting abactinal pedicellariae with conspicuous
toothed valves, longer than spinelets; oral plate with up to
10 spines A. rowleyi (SE Africa)
— If abactinal pedicellariae present, valves not longer than
spinelets; oral plate with up to 8 spines 7
7. Abactinal spinelets on rays differentiated on plates into
apically thick and marginally thin; some irregularly
distributed paxilliform plates 8
— Abactinal spinelets may be of variable form, but not
differentiated into two distinct forms; lacking paxilliform
abactinal plates 9
8. Abactinal radial plates with central subglobose spinelets,
peripheral short conical to subgranuliform spinelets;
lacking pedicellariae; actinal central interradial plates
each with about 3 spines A. lorioli (N Indian Ocean)
— Abactinal radial plates with central digitiform spinelets,
peripheral short conical spinelets; pedicellariae present;
actinal central interradial plates each with about 5
spines A. coronata (Japan to N Australia)
9. Pedicellariae with differentiated valves in abactinal
proximal interradii 10
— Lacking pedicellariae 13
10. Abactinal spinelets thick, up to about 12 on each proximal
carinal plate (at R = 21 mm) A. iranica (Persian Gulf)
— Abactinal spinelets thin or small, up to more than 24 on
each proximal carinal plate (at R = 17 mm) 11
11. Abactinal proximal spinelets up to more than 40 per plate;
superomarginal plates each with up to about 20 spinelets
(at R - 19 mm) A. bather i (Japan)
— Abactinal proximal spinelets fewer than 25 per plate;
superomarginal plates each with fewer than 10 spinelets
(at R - 19 mm) 12
12. Proximal abactinal spinelets small, thick, frequently of two
forms, subgranuliform apically on plates; abactinal distal
interradial plate spinelets splayed and overlapping adjacent
plate spinelets (at R = 20 mm); actinal interradial plates each
with up to 5 spines (at R = 20 mm); size up to R = 25 mm ....
A. richmondi (E Africa coast, Madagascar, Mauritius)
— Proximal abactinal spinelets thin, similar form; abactinal
distal interradial spinelets not overlapping adjacent plate
spinelets if splayed; actinal interradial plates each with up
to 10 spines (at R = 19 mm); size up to R = 19 mm
A. watersi (Arabian Sea, Mauritius)
13. Abactinal spinelets sacciform, short, wide globose basally,
tapered to sharply pointed apically; up to about 12
spinelets on each proximal abactinal plate (at R = 19 mm);
predominantly 2 actinal interradial spines on each plate ....
A. halseyae (Maldives)
— Abactinal spinelets not widely globose basally, not tapered
to a sharp point apically; more numerous than 14 on each
proximal abactinal plate (at R> 12 mm); predominantly >
3 actinal interradial spines on each plate 14
14. Proximal abactinal spinelets short, thick, columnar or
conical 15
— Proximal abactinal spinelets long, thin, subsacciform 16
15. Rays long, subdigitiform; spinelets mostly spread over
exposed plate surface; predominantly 6 spines per actinal
interradial plate (at R = 16 mm); actinal interradial spines
short, thick, bluntly conical, sacciform. A. samyni
(Arabian Sea to SE Africa, Madagascar, La Reunion)
— Rays short, strongly tapered; spinelets mostly concentrated
over projecting proximal plate edge; predominantly 3
spines per actinal interradial plate (at R = 16 mm); actinal
interradial spines conical to digitiform .., 5r6 ^
A. marshae (Red Sea, Gulfs of Aqaba and Suez)
16. Rays short, merging with disc; lacking doubly-papulate
carinal plates; spinelets not clustered into groups on
plates; spinelets frequently splay -pointed
A. oharai (Okinawa)
— Rays long, discrete; some doubly-papulate carinal plates
present; spinelets frequently clustered into groups on
plates; spinelets not splay-pointed 17
17. Rays tapered; abactinal plates angled over papulae;
spinelets long, thin, sub-sacciform to sacciform, tapering
to fine point, rugose, subacicular; spinelets frequently
projecting proximally over papulae
A. cepheus (Indonesia to N Australia)
— Rays digitiform; abactinal plates not angled over papulae;
spinelets long, thick, conical to subsacciform, with
numerous (5-6) points on distal sides and end of spinelets;
spinelets not projecting proximally over papulae
I A. limboonkengi (China)
18. Actinal interradial spines predominantly 1 per plate
A. conandae (Mascarene Is.)
— Actinal interradial spines predominantly > 1 per plate
19
19. Spinelets of 2 distinct forms, long thick digitiform apically
on upper ray and marginal plates
* A. corallicola (NE Indian to central W Pacific Oceans)
— Spinelets of one form 20
20. Spinelets elongate, not subgranuliform (at R = 5 mm) ....... 21
— Spinelets truncate, subgranuliform (at R = 5 mm) 23
260
P. Mark O’Loughlin and Francis W.E. Rowe
21. Spinelets long, frequently distinctly splay-pointed;
pedicellariae present; size up to R = 12.5 mm
A. anomala (central W Pacific)
— Spinelets not long, not distinctly splay-pointed; lacking
pedicellariae; size up to R = 5 mm 22
22. Rays narrow at base, long, elevated; spinelets thick columnar
or conical; actinal interradial spines up to 2 per plate
.. L .. A. colemani (Papua New Guinea, Indonesia)
— Rays wide at base, short, not elevated; spinelets thin
digitiform or conical with distally long spines; actinal
interradial spines up to 5 per plate
A. doranae (Okinawa)
23. Size up to R = 18 mm; some central abactinal plates
atypically large and irregular
A. burtonii (Red and Arabian Seas)
— Size up to R = 9 mm; central abactinal plates not unusually
large and irregular 24
24. Abactinal spinelets up to 16 per plate; spinelets splay-
pointed; suboral spines up to 4 per plate
, 1; _, A. yairi (Red and Mediterranean Seas)
— Abactinal spinelets up to 10 per plate; spinelets not splay-
pointed; suboral spines up to 2 per plate
A. moosleitneri (Maidive Is.)
Asterinidae Gray, 1840
Remarks. For recent revision of Asterinidae see O’Loughlin
and Waters (2004). For addition of new genus Ailsastra see
O’Loughlin and Rowe (2005).
Aquilonastra O’Loughlin, 2004
Aquilonastra O’Loughlin, in O’Loughlin and Waters, 2004: 5
(key), 13-15, tables 1 and 2.— O’Loughlin and Rowe, 2005: 181. —
Saba and Fujita, 2006: 270,-Byrne, 2006: 244, 245, 248, 250, 251.
Diagnosis ( emended from O’Loughlin and Waters, 2004). Rays
discrete, broad or narrow at base, tapering, rounded distally,
interradial margin deeply incurved; fissiparous or non-fissiparous;
fissiparous species with more than 1 inconspicuous madreporite,
up to 9 rays, rays frequently unequal in length, form frequently
asymmetrical; non-fissiparous species with 1 conspicuous
madreporite, rarely 2, predominantly 5 rays, sometimes 6, form
symmetrical, typically stellate; body flat actinally, high convex
abactinally; disc variably distinct in non-fissiparous species,
sometimes delineated by 5 transversely elongate radial and 5
short interradial plates; abactinal plates predominantly irregular
on upper rays, in longitudinal series on sides of rays, series not
perpendicular to margin; papulate areas extensive, plates
predominantly with single notch for papula in papulate areas,
papulae predominantly single per space, large, in longitudinal
series along sides of rays; abactinal plates with glassy convexities;
abactinal spinelets small, glassy, subgranuliform to digitiform,
columnar or conical or sacciform or splay -pointed sacciform, in
bands or tufts, numerous (1CM-0 per proximal abactinal plate);
superomarginal and inferomarginal plates in regular series;
actinal plates in longitudinal, not oblique, series; suboral spines
present; adradial actinal spines in complete series; superambul acral
plates present; superactinal plates present.
Type species. Asteriscus cepheus Muller and Troschel, 1842
(by original designation in O’Loughlin and Waters, 2004).
Other species. For all 25 species see Table 1 and Fig 1.
Remarks. O’Loughlin and Waters (2004) referred 12 species to
the new genus Aquilonastra. They noted that A. heteractis had
some characters that were exceptional to those shared by the
other species of Aquilonastra. More recently O’Loughlin and
Rowe (2005) reassigned this species to their new genus in the
recombination Ailsastra heteractis (H.L. Clark, 1938). The
remaining 11 original species are retained here in Aquilonastra.
Asterina lorioli Koehler, 1910 was incertae sedis in O’Loughlin
and Waters (2004). It is added here in the new combination
Aquilonastra lorioli (Koehler, 1910). Thirteen new
Aquilonastra species are described. The emended generic
diagnosis refines the description but does not change any
diagnostic characters.
Aquilonastra anomala (H.L. Clark, 1921)
Figures 1, 2a, 7a
Asterina anomala H.L. Clark, 1921: 95-96, pi. 7 fig. 8, pi. 23 fig.
5, pi. 26 figs 2, 3.— H.L. Clark, 1938: 143-144.-H.L. Clark, 1946:
133-134.— A. M. Clark and Rowe, 1971: 68, fig. 17g, tbl. 1.— Marsh,
1974: 92.— Marsh, 1977: 270-271, fig. 7, tbl. 2.-Oguro, 1983: 222-
224, figs 5, 6, 12, 13.— A. M. Clark, 1993: 207,-Rowe and Gates,
1995: 33-34. -Waters et al„ 2004: 874, 876, 877, tbl. 1, figs 1, 2.
Aquilonastra anomala.— O’Loughlin and Waters, 2004: 11, 13-
15, fig. 1.
Material examined. N Australia, Torres Strait, Darnley I., 30 Apr
1977, WAM Z6849 (2); Kimberley, 17 Jul 1988, Z6843 (1); Papua New
Guinea, Bismark Archipelago, 1-34 m, 29 Jun 2003, UF 2283 (1);
15-22 m, 26 Jun 2003, UF 2217 (1); West New Britain, 3-37 m, 17 Jun
2003, UF 2270 (1); Christmas I., 13 Feb 1987, Z6851 (1); Lord Howe
I., AM J6169 (21); 15 Feb 1979, J16574 (2); H.L. Clark, Apr 1932,
NMV F95593 (6); 20 May 2003, F97690 (6); 15 Mar 2002, F96699
(1); Solomon Is. 4-6 m, 20 Jun 2004, F94607 (1); 3-5 m, 26 Jun 2004,
F94616 (3); Caroline Is. Z6845 (2); Palau, 26 Jul 1999, UF 1740 (3);
Samoa, F96698 (2).
Diagnosis. Fissiparous Aquilonastra species; up to 8 rays,
predominantly 7, broad basally, rounded distally; up to R =
12.5 mm (H.L. Clark, 1938); pedicellariae with differentiated
conical valves larger than spinelets sometimes present in
proximal interradial angle; up to 4 inconspicuous madreporites
seen; abactinal gonopores present.
At R = 8 mm, r = 4 mm, lacking carinal series of plates,
upper rays with 2 irregularly arranged longitudinal series of
singly papulate plates; plates domed, angled over papulae
more than notched; single papula per plate, rarely 2; some
secondary plates intergrade with primary plates; spinelets
long, thin conical pointed to prominently splay-pointed
sacciform, up to about 20 spinelets over projecting surface of
each proximal abactinal plate, predominantly in transverse
double band, rare clustering of spinelets; distal interradial
plates with up to about 6 sacciform splay-pointed spinelets;
superomarginal and inferomarginal plates subequal;
superomarginal plates each with up to about 8 spinelets,
inferomarginal plates each with up to about 16 slightly larger
spinelets, marginal spinelets long, splay-pointed sacciform.
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
261
Spines per actinal plate up to: oral 7, suboral 5, furrow 6,
subambulacral 5, actinal interradial 7 (variable, predominantly
3-4); interradial spines glassy, thin, conical, pointed, spinous,
in transverse series or tufts on plates.
Colour (live). “Green, prettily variegated with white and rusky,
with traces of red and yellow along the margins” (H.L. Clark,
1921); “green is common as a tinge, if not as a ground colour,
and orange and brown are very generally evident; white
blotches or markings may occur” (H.L. Clark, 1946); brown
with reddish tinge apically, green tinge radially, white
interradially, some orange marginally (photo by G. Paulay).
Distribution. N and NE Australia; Papua New Guinea; Lord
Howe I.; Solomon Is.; Caroline Is., Palau; Marshall Is.; Fiji;
Tonga; Samoa; Cook Is.; 0-37 m (Marsh, 1974; A.M. Clark,
1993; this work).
Remarks. A distinguishing character for the fissiparous A.
anomala is the relatively long, frequently splay-pointed,
abactinal spinelets. Green is commonly present in the mottled
live colours.
Aquilonastra batheri (Goto, 1914)
Figures 1, 2b, 7b
Asterina penicillaris (part).— Sladen, 1889: 393 (two Challenger
specimens from Kobe, Japan, non Asterina penicillaris (Lamarck,
1816), according to Goto (1914); Asterina penicillaris of uncertain
identity and validity, according to A.M. Clark (1993)).
Asterina batheri Goto, 1914: 651-656, pi. 19 figs 275-278.—
Hayashi, 1940: 119, pi. 13 figs 5, 6.-Hayashi, 1973: 71, pi. 12 fig.
2.— A.M. Clark, 1993: 207,-Fujita and Saba, 2000: 169, pi. 1C, pi.
3D, F.-Waters et al„ 2004: 874, 876, 877, tbl. 1, figs 1, 2.
Aquilonastra batheri.— O’Loughlin and Waters, 2004: 11, 13-15,
figs 1, 9e.— Saba and Fujita, 2006: 286.— Byrne, 2006: 245, tbl. 2.
Material examined. Japan, Toyama Bay, NMV F97441 (1); AM J1 1564
(2).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, broad
basally, narrowly rounded distally; up to R = 34 mm, r = 14 mm
(Goto, 1914); abactinal proximal interradial pedicellariae, pairs
of tooth-like differentiated valves; gonopores abactinal; direct
development into brachiolaria stage (Hayashi, 1973).
Figure 1. Indo-west Pacific distribution of type localities for the species of Aquilonastra O’Loughlin.
262
P. Mark O’Loughlin and Francis W.E. Rowe
Figure 2. Photos of live colour and form of Aquilonastra species, a, A. anomala (H.L. Clark), Lord Howe I. (photo N. Coleman); b, A. batheri
(Goto), Japan, Honshu, Hiroshima, Takehara (NSMT E-4136; photo T. Fujita, see Fujita and Saba, 2000); c, A. burtonii (Gray), Red Sea, Egypt
(photo J. Hinterkircher); d, A. burtonii (Gray), Oman (photo G. Paulay); e, A. byrneae sp. nov., Qld, One Tree I. (photo M. Byrne); f, A. cepheus
(Muller and Troschel), Papua New Guinea, Loloata I. (photo N. Coleman); g, A. cepheus (Muller and Troschel), Qld, Lady Elliot I. (photo N.
Coleman); h, A. cepheus (Muller and Troschel), WA, Exmouth (photo N. Coleman); i, A. colemani sp. nov., Indonesia, Mayo I. (NMV F109374;
photo S. Uthicke); j, A. colemani sp. nov., Papua New Guinea, Milne Bay (photo N. Coleman); k, A. conandae sp. nov., Indian Ocean, La Reunion
I. (photo A. Barrere); 1, A. coronata (Martens), Japan, Kyushu, Goto Is., Fukue I. (NSMT E-3683; photo T. Fujita).
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
263
Figure 3. Photos of live colour and form of Aquilonastra species, a, A. doranae sp. nov., Japan, Okinawa (UF 3913, holotype; photo G. Paulay);
b, A. halseyae sp. nov., Indian Ocean, Maidive Is. (photo N. Coleman); c, A. marshae sp. nov.. Red Sea, Egypt (photo J. Hinterkircher); d, A.
minor (Hayashi), Japan, Honshu, Hiroshima, Takehara (NSMT E-4102; photo T. Fujita, see Fujita and Saba, 2000); e, A. oharai sp. nov., Japan,
Okinawa (photo G. Paulay); f, A. rosea (H.L. Clark), WA, Ludlow (photo N. Coleman); g, A. samyni sp. nov., SE Africa, Sodwana Bay (photo Y.
Samyn); h, A. samyni sp. nov., Oman (photo G. Paulay); i, A. scobinata (Livingstone), Victoria (photo L. Altoff); j, A. watersi sp. nov., Oman
(photo G. Paulay); k, unidentified Aquilonastra fissiparous species, Indian Ocean, Seychelles Is. (photo N. Coleman); 1, unidentified Aquilonastra
fissiparous species, Seychelles Is., La Digue I. (photo M. Richmond).
264
P. Mark O’Loughlin and Francis W.E. Rowe
Figure 4. Photos of live colour and form of Aquilonastra species, a, A. conandae sp. nov., Indian Ocean, La Reunion I., on under-surface of coral
slab (photo A. Barrere); b, A. corallicola (Marsh), Indian Ocean, Cocos (Keeling) I. (photo N. Coleman); c, A. lorioli (Koehler), Pakistan (photo
Qaseem Tahera); d, A. marshae sp. nov.. Red Sea, Egypt (photo J. Hinterkircher); e, A. richmondi sp. nov., Tanzania, Mnazi Bay (photo M.
Richmond); f, A. yairi sp. nov., Israel, Achziv (photo M. Tsurnamal, 5 July 1964).
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
265
Figure 5. Photos of Aquilonastra species, a, A. burtonii (Gray), Zanzibar, spinelets (NHM 2004.2831); b, A. byrneae sp. nov., Qld, One Tree I.,
spinelets and papulae (photo M. Byrne); c, A. cepheus (Muller and Troschel), Jakarta (R = 20 mm; holotype MNHN EcAsl471); d, A. cepheus
(Muller and Troschel), Papua New Guinea, spinelets (R = 17 mm; UF 2332); e, A. colemani sp. nov., Papua New Guinea, spinelets (UF 3284,
holotype); f, A. conandae sp. nov.. La Reunion I., spinelets (NMV F107414); g, A. corallicola (Marsh), Cocos (Keeling) I., pedicellaria (arrow)
and spinelets (UF 745); h, A. corallicola (Marsh), Cocos (Keeling) I., marginal spinelets (UF 745); i, A.halseyae sp. nov., Maidive Is., spinelets
(NHM 1965.6.1.84, holotype); j, A. iranica (Mortensen), Iranian Gulf (R = 13 mm; WAM Z6868); k, A. iranica (Mortensen), Iranian Gulf,
spinelets and pedicellariae (arrows) (R = 14 mm; WAM Z6868); 1, A. limboonkengi (Smith), China, Amoy, spinelets (NHM 1926.12.22.35-36,
syntype).
266
P. Mark O’Loughlin and Francis W.E. Rowe
Figure 6. Photos of Aquilonastra species, a, A. limboonkengi (Smith), China, Amoy, spines (NHM 1926.12.22.35-36, syntype); b, A. lorioli
(Koehler), Bombay, atypically two conspicuous madreporites, not fissiparous (R = 22 mm; NHM 1960.10.4.11-16); c, A.lorioli (Koehler),
Pakistan, Karachi, spinelets (MNHN EcAs2662, syntype); d, A. marshae sp. nov.. Red Sea, Egypt, spinelets (R = 13 mm; NMV F109382); e, A.
marshae sp. nov.. Red Sea, marginal spinelets (TM H1814); f, A. richmondi sp. nov., SE Africa, Sodwana Bay, spinelets (MRAC 1737); g, A.
richmondi sp. nov., SE Africa, Sodwana Bay, distal interradial overlapping spinelets (R = 23 mm; MRAC 1737); h, A. rosea (H.L. Clark), WA,
Jurien Bay, circular paxilliform spinelet clusters (R = 7 mm; WAM Z31162); i, A. rowleyi sp. nov., abactinal view (MRAC 1736, holotype, partly
dissected, R = 23 mm); j, A. samyni sp. nov., SE Africa, Sodwana Bay, marginal spinelets (MRAC 1741); k, A. scobinata (Livingstone), Tasmania,
Tamar R. mouth, crescentiform paxilliform spinelet clusters (R = 14 mm; NMV FI 12176); 1, A. watersi sp. nov., Arabian Sea, Oman, spinelets
and pedicellaria (arrow) (R = 15 mm; partly cleared paratype UF 3283).v
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
267
Figure 7. Photos of Aquilonastra species, a, A. anomala (H.L. Clark), Lord Howe I., marginal spinelets (NMV F97690); b, A. batheri (Goto),
Japan, doubly papulate carinal plates with groups of spinelets (R = 19 mm; NMV F97441); c, A. burtonii (Gray), Red Sea, disc with five
inconspicuous madreporites (arrows), five rays, large irregular proximal plates (R = 11 mm; lectotype NHM [18]40.3.23.54); d, A. burtonii
(Gray), Oman (UF 1126); e, A. byrneae sp. nov., Qld, Tryon I., actinal interradius with two distal gonopores (R = 14 mm; NMV F109373); f, A.
colemani sp. nov., Papua New Guinea (UF 3284, holotype).
268
P. Mark O’Loughlin and Francis W.E. Rowe
At R = 19 mm, r = 10 mm, up to 4 proximal doubly-papulate
carinal plates, each with up to 4 tufts each with 6-12 splayed
spinelets, up to about 40 spinelets on proximal abactinal plates;
plates with low domes for spinelets; few secondary plates;
spinelets conical to digitiform, thick, subsacciform, splay-
pointed; superomarginal plates smaller than inferomarginals,
each series with up to about 20 spinelets per plate, slightly
more stout on inferomarginals.
Spines per actinal plate up to: oral 7, suboral 5, furrow 7,
subambulacral 9, actinal interradial 12 proximally 6 distally;
interradial spines thin conical, to digitiform to subsacciform.
Colour (live). Disc and proximal apical area reddish-orange,
madreporite white, proximal rays and interradii mottled with
dark and light reddish-orange and white, distally pale reddish-
orange (photo from M. Komatsu); disc and upper rays red, with
sides of rays mottled mauve and brown, madreporite off-white
(photo from T. Fujita).
Distribution. “Common in the middle and southern regions of
Japan” (Hayashi, 1973); sublittoral (Hayashi, 1940) to 92 m
(Sladen, 1889).
Remarks. O’Loughlin and Waters (2004) determined material
from Oman (UF70) as A. batheri. This material is redetermined
in this work as A. watersi sp. nov. (below). The potentially
large size, presence of numerous interradial pedicellariae with
differentiated valves, and grouping of spinelets into tufts on
proximal abactinal plates are distinguishing characters. Red is
commonly present in the mottled live colours.
Aquilonastra burtonii (Gray, 1840)
Figures 1, 2c-d, 5a, 7c-d
Asterina burtonii Gray, 1840: 289.— Gray, 1866: 16.— Gray, 1872:
118.— H.L. Clark, 1923: 283 (status of species; possibly part).— Smith,
1927a: 641-645 (part).— A. M. Clark, 1952: 207 (possibly part;
fissiparous; possibly A.yairi sp. nov. below).— Tortonese, 1960: 20-21
(probably part).
Asterina burtoni.— Tortonese, 1966: 3, fig. 1.— A.M. Clark, 1967a:
146, tbl. 1 (part; Red Sea material).— James and Pearse, 1969: 84-85
(part).— A.M. Clark and Rowe, 1971: 38 (part), 68, fig. 17a.— Tortonese,
1977: 281-282.— Price, 1983: 47-48, fig. 14 (part).— Archituv and
Sher, 1991: 670.— Mladenov and Achituv, 1999: 152 (part).— Karako
et al., 2002: 139-144 (part, El Fauz Red Sea population).
Asteriscus wega Perrier, 1869: 102.
Asterina wega— Perrier, 1875: 318.— Achituv, 1969: 329-341
(part, “pluriradiate” form).— Achituv, 1973a: 333-336 (part, Akhziz
lagoon and Haifa populations).
Asterinides burtoni.— We rrill, 1913: 482.
Asterina gibbosa.— Tortonese, 1957: 190 (non Asterina gibbosa
Pennant, 1777; see Tortonese 1966).
Aquilonastra burtoni.— O’Loughlin and Waters, 2004: 11, 13
(part), 14.
Material examined. Lectotype (judged to be Gray’s type by G.A.
Smith, 1927a). Red Sea, NHM [18]40.3.23.54 (dry).
Asteriscus wega Val., Red Sea, M. Botta, 1837, MNHN EcAs2713
(1) (labelled “type”; see Remarks; not A. burtonii ).
Red Sea, coll. Michelin, 1868, EcAsl566 (6); Ras Muhammad, 8
Aug 1968, HUJ SLR1917 (5); Gulf of Suez, Et Tur, 20 Sep 1967, HUJ
SLR845 (2); 20 Sep 1967, TAU NS2090 (1, fissiparous form); 24 Dec
1965, TM H1815 (2); lie Abulat, Calypso , EcAsll842 (1); La Sicherie,
EcAsll841 (1); S Sinai, El Fauz, Nov 2003, NMV F109383 (4); Gulf
of Aqaba, near Dahab, 4 Nov 1981, TAU NS24408 (2, fissiparous
forms); Sep 1976, TAU NS24501 (1); 18 Dec 1967, HUJ SLR1127 (1);
19 Feb 1968, SLR1341 (2).
Oman, Masirah I., Bar al Hikman peninsula, coral rubble, 3-4 m, 7
Nov 1999, UF 1126 (2); 0-1 m, 7 Nov 1999, UF3280 (1); 3^1 m, 23 Jan
2005, UF 4240 (1); UF 4239 (2); 1-8 m, 18-24 Jan 2005, UF 4237 (1).
Zanzibar, Prison I., seagrass bed, 6 Aug 1995, NHM 2004.2831 (1).
Diagnosis. Fissiparous Aquilonastra species; rays up to 8,
frequently 7, form frequently asymmetrical post-fissiparity;
form of larger specimens sometimes symmetrical with 5 equal
rays; most interradii with inconspicuous madreporite; rays
narrow basally, tapering, narrow rounded distally, digitiform; up
to R = 18 mm; at R = 12 mm, r = 5 mm; gonopores not seen.
At R = 12 mm, lacking carinal plates; numerous secondary
plates, intergrade with primary plates, frequently large
irregular proximal abactinal plates; abactinal plates arched
over papulae, rarely notched; spinelets small, variable in form,
conical to columnar to digitiform, distally spinous, some
splay-pointed, spread sparsely over plate surface, rarely up to
about 15 spinelets on proximal plates, up to about 7 on mid-
interradial plates; superomarginal plates each with up to about
6 spinelets, inferomarginal plates each with up to about 12
larger spinelets.
Spines per actinal plate up to: oral 6, suboral 3 frequently 2,
furrow 6, subambulacral 4, actinalinterradial 1-4 (predominantly
3); interradial spines conical, sacciform.
Colour (live). “Greenish gray colour on the aboral side; a large
and irregular, purplish brown blotch is on the centre, and is
surrounded by red spots on the basal parts of the arms; the
latter are usually darker (greenish) near their extremity, where
a pale median line is to be observed” (Tortonese, 1966; Haifa
specimens; the red spots are presumed here to be the
madreporites); “variegated yellowish, brown and red”
(Tortonese, 1977; Aqaba specimen); majority of 14 colour
morphotypes from Elat were “mottled brown and orange”
(pers. comm. Y. Achituv); Akhziz specimens were
predominantly “mottled brown and orange” (pers. comm. Y.
Achituv); “pale grey, mottled red and brown” (label with
Zanzibar specimen); mottled browns, off-white (photos by G.
Paulay).
Distribution. Eastern Mediterranean, Akhziz lagoon and Haifa
populations; Red Sea, Gulf of Suez, Gulf of Aqaba; Arabian
Sea, Oman; Arabian Gulf; NW Indian Ocean, Zanzibar; 0-8 m.
Remarks. Smith (1927a) discussed in detail the historical
confusion surrounding Gray’s Asterina burtonii, and gave a
full synonymy. Gray’s type was reported lost, but Smith
asserted that he had found two of Gray’s type specimens of
Asterina burtonii. We follow Smith (1927a), and accept that the
larger of these two specimens (NHM [18]40.3.23.54) is the
type for Asterina burtonii. This lectotype for Asterina burtonii
has five equal rays, but five small madreporites (R = 11 mm).
We judge that the smaller of these two types (NHM
[18]40.3.23.55) is conspecific with a second Red Sea fissiparous
species Aquilonastra yairi sp. nov. (below).
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
269
A Paris Museum specimen of Asteriscus wega Val.
(MNHN EcAs2713) labelled “type” was collected from the
Red Sea by M. Botta in 1837. The Valenciennes manuscript
name was not published, and this specimen has no type status.
The name Asteriscus wega was published by Perrier who
described (1869, 1875) 13 fissiparous specimens (syntype
series) collected from the Red Sea by M. Botta in 1858. These
fissiparous specimens, up to R = 15 mm (Perrier reported a
diameter of 2-3 centimetres), are not conspecific with the
Valenciennes non-fissiparous “type” specimen that has been
examined here and has six equal rays and one conspicuous
madreporite. It is referred below to Aquilonastra marshae sp.
nov. Based on Perrier’s 1869 and 1875 descriptions, Asteriscus
wega Perrier, 1869 is judged to be conspecific with Asterina
burtonii Gray, 1840, of which it becomes a junior synonym.
This decision supports the opinion of a synonymy by A.M.
Clark (1952, 1967b) and A.M. Clark and Rowe (1971).
Achituv (1973a) studied large numbers of four eastern
Mediterranen populations of small fissiparous asterinids from
Acre, Akhziz pool, Akhziz lagoon and Haifa. For the first two
populations maximum R = 8 mm; for the latter two populations
maximum R = 17 mm. These results are closely consistent
with data for the two fissiparous species from the Red Sea, A.
yairi sp. nov. with R up to 7 mm (below) and A. burtonii with
R up to 18 mm. The invasion of the Mediterranean by both
Red Sea fissiparous species is judged to be the best explanation
for this data.
Mladenov and Achituv (1999, abstract) reported genetic
studies of four populations of Asterina burtoni :
1. non-fissiparous Red Sea population from Elat
2. sympatric fissiparous Red Sea population from Elat
3. two allopatric fissiparous Mediterranean populations
They observed a high genetic difference between the
fissiparous and non-fissiparous Elat populations, but not as high
as between some fissiparous populations. On the assumption
that the fissiparous populations were the same species, they
concluded that there were not separate species in the four
populations. We judge that there are two species represented by
the fissiparous populations, and that both species occur in the
eastern Mediterranean and one of them at Elat. This would
explain the high genetic differences observed by Maldenov and
Achituv (1999). To us, the Mladenov and Archituv (1999)
evidence supports a conclusion of three species. This conclusion
is congruent with morphological differences.
Karako et al. (2002) reported similar genetic studies and
results for four fissiparous populations fromAkhziv, Shikmona,
and Mikhmoret on the coast of Israel, and El Fauz at the mouth
of the Gulf of Aqaba, and a non-fissiparous population at Elat at
the northern end of the Gulf of Aqaba. “Pentaradiate”, rather
than having a large conspicuous madreporite, was used to
determine non-fissiparous specimens and can be misleading as
the larger fissiparous specimens frequently have five equal rays
but continue to have more than one inconspicuous madreporite.
We judge, as above, that the Karako et al. results support an
hypothesis of two fissiparous species and one non-fissiparous
species. The three Mediterranean populations appear to us to be
Aquilonastra yairi sp. nov. (below), and the El Fauz population
A. burtonii. We refer the non-fissiparous species at Elat to
Aquilonastra marshae sp. nov. (below). Specimens from all
locations in the Karako et al. study, except Shikmona, were
examined by us, and provide morphological support for the
systematic decision that there are three separate species.
Tortonese (1936, 1966) first recorded A. burtonii in the
Mediterrnean from Massawa (as A. wega) in 1936. Mortensen
(1926) reported a fissiparous specimen from the Gulf of Suez
as Asterina burtonii , but gave insufficient detail to judge here
whether the specimen was A. burtonii or A. yairi sp. nov. In
O’Loughlin and Waters (2004), material TM H1815 was
referred to A. burtoni and that determination is confirmed
here. A specimen TM H1814 was also assigned to A. burtoni,
but is redetermined in this work as Aquilonastra marshae sp.
nov. (below). There is no evidence in this study of a second
smaller fissiparous species ( Aquilonastra yairi sp. nov. below)
in the Gulf of Aqaba.
Soliman (1999) reported studies of two asteroid populations
in the Arabian Gulf, understanding them to be both Asterina
burtoni. There is no indication in the report that either
population had fissiparous individuals, and we assume that
Soliman studied non-fissiparous populations that were thus not
A. burtonii. In this review three other asterinid species occur in
the region: A. samyni sp. nov. (below, Oman) reaches R = 27
mm; A. water si sp. nov. (below, Oman) reaches R = 19 mm; A.
iranica (Mortensen, 1940, Arabian Gulf) reaches R = 35 mm.
One Soliman (1999) population was up to R = 26 mm in size,
and we hypothesize that it was A. samyni. The other was up to
R = 16 mm, and we hypothesize that it was A. water si. Soliman
(1995) also reported an asterinid population study in the
Arabian Gulf at Qatar. The largest individual was R = 26 mm.
Again, we hypothesize that the population was A. samyni.
A.M. Clark (1974) and A.M. Clark and Courtman-Stock
(1976) reported Asterina burtoni for SE Africa, and referred
to both single madreporite pentamerous and fissiparous
specimens from Mozambique. A. burtonii is reported here
from Zanzibar, and possibly occurs off Mozambique. H.L.
Clark (1923) reported Asterina burtonii in the fauna of South
Africa, but referred only to a specimen from Mozambique.
Non-fissiparous specimens are not A. burtonii. Jangoux (1984),
and Jangoux and Aziz (1984, 1988) reported Asterina burtoni
for New Caledonia, La Reunion, Seychelles, Mineures and
Maldives. We found no evidence to confirm A. burtonii in any
of these localities. Following many authors, Walenkamp
(1990) listed A. cepheus Muller and Troschel, 1842, A. wega
Perrier, 1869, A. cephea var iranica Mortensen, 1940 and ? A.
anomala H.L. Clark, 1921 as junior synonyms of A. burtonii
Gray, 1840. We consider only A. wega to be a junior synonym.
Walenkamp’s (1990) material is referred to A. richmondi sp.
nov. (see below).
Perrier (1875), H.L. Clark (1923), Mortensen (1926), Smith
(1927), A.M. Clark (1952) and Tortonese (1960) retained the
original spelling of Gray {burtonii). Recent authors have used
A. burtoni. Walenkamp (1990) argued for a restoration of the
orignial spelling. We agree.
The characters distinguishing A. burtonii from A. yairi sp.
nov. are detailed in the Remarks for A. yairi below. The
mottled live colours for A. burtonii are red, orange, yellow,
brown, grey, off-white.
270
P. Mark O’Loughlin and Francis W.E. Rowe
Aquilonastra byrneae sp. nov.
Figures 1, 2e, 5b, 7e
Aquilonastra new. sp.— Byrne, 2006: 245, 248, 251, tbl. 2.
Material examined. Holotype. NE Australia, Great Barrier Reef, near
Heron I., One Tree I., rocky shallows, Maria Byrne, Jan 2002, NMV
F98748 (alcohol).
Paratypes. Type locality, Dec 2004, F98747 (1, alcohol); interridal,
12 Apr 2006, FI 11326 (2, alcohol); 27 May 2004, F109358 (1, dry,
dissected).
Other material. Tryon I., 15 Sep 1970, FI 09373 (2, dry); Mariana Is.,
Guam I., Asan Point, reef flat, under rock, 1 1 Apr 1996, UF 894 (dry).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5,
narrow basally, broadly to narrowly rounded distally; up to R =
15 mm, r = 8 mm; gonopores actinal, interradial pairs frequently
close, near margin; protandric hermaphrodite (Maria Byrne,
pers. comm.).
At R = 15 mm, 0-3 proximal doubly -papulate carinal plates,
each with up to 4 tufts with 3-5 spinelets per tuft; numerous
proximal secondary plates, 0-3 per space; single large papula per
space, rarely 2; disc variably distinctly bordered; spinelets small,
short, form variable from conical to digitiform, distally pointed
or blunt, some sacciform, not splay-pointed; up to about 16
spinelets across projecting edge of proximal abactinal plates,
frequently 1-2 on bare distal mid-plate; up to 12 on proximal
surface of distal interradial plates; superomarginal plates smaller
than inferomarginals (subequal on Guam specimen),
superomarginals with up to about 12 short conical pointed
spinelets per plate, inferomarginals with up to about 20 spinelets,
more stout and longer on inferomarginals.
Spines per actinal plate up to: oral 7, suboral 6, furrow 7,
subambulacral 6, actinal interradial 8, predominantly 5;
interradial spines thick, bluntly conical, subsacciform.
Colour (live). Disc sometimes cream centrally surrounded by
dark brown ring, abactinally mottled with proximal upper rays
predominantly olive-green to greenish-brown, interradii and
distal rays predominantly cream; actinal surface cream with
green patches (photo and pers. comm. M. Byrne); disc red or
brown, thin boundary of white plates, proximal upper rays
crimson red, remaining abactinal surface mottled with
predominantly dark and pale brown, red and white (Guam
photos by G. Paulay as Asterina cepheus).
Distribution. NE Australia, Great Barrier Reef, near Heron I.,
Tryon I. and One Tree I.; (possibly) W Pacific Ocean, Guam;
rocky shallows.
Etymology. Named for Maria Byrne, Professor of
Developmental and Marine Biology in the University of
Sydney, and Director of One Tree Island Research Station on
the Great Barrier Reef, in appreciation of her contribution of
specimens for this work and her research on life history
diversity and evolution in Asterinidae.
Remarks. A significant diagnostic character of A. byrneae is
the presence of actinal gonopores. The dominant mottled live
colours are green, brown and red, with cream or white. At R =
12 mm, the material from the Great Barrier Reef has
predominantly 5 actinal interradial spines per plate, the Guam
specimen predominantly 3. This variation (on its own) is judged
to be an inadequate basis for separating the material into two
species, but this and colour differences suggest the possibility
of two species. We identify the Guam specimen as A. byrneae
with some hesitation.
Aquilonastra cepheus (Miiller and Troschel, 1842)
Figures 1, 2f-h, 5c-d
Asteriscus cepheus Muller and Troschel, 1842: 41-42.— Dujardin
et Hupe, 1862: 375-376.-Perrier, 1869: 99.
Asterina cephea.— Perrier, 1875: 315-317 (part; type).— Mobius,
1880: 50.— Doderlein, 1888: 825,-Sluiter, 1889: 307-308.
Asterina cepheus.— Martens, 1866: 85.— H.F. Clark, 1915: 95.—
Fisher, 1919: 411, pi. 115 fig. 4.— Fisher, 1925: 79-80.— Fiao and A.M.
Clark, 1995: 130, pi. 15, figs 10, 11.— Rowe and Gates, 1995: 34.—
A.M. Clark and Mah, 2001: 335.
Asterina burtoni.—H.L. Clark, 1921: 96-97, pi. 6 fig. 2.— Smith,
1927b: 276.— H.F. Clark, 1938: 144-145.-H.F. Clark, 1946: 133.-
Marsh, 1974: 91-92 (non Asterina burtonii Gray, 1840).
Asterina burtoni cepheus.— A.M. Clark and Rowe, 1971: 68-69,
fig. 17h, table 1, pi. 9 figs 4-5. -Fiao, 1980: 171, figs 2:1, 2:2, 4, pl.4
figs 6-7.— A.M. Clark, 1993: 208 (non Asterina burtonii Gray, 1840)
Aquilonastra cepheus.— O’Foughlin and Waters, 2004: 11, 13-15.
Material examined. Holotype. Indonesia, Jakarta (as Batavia), M.
Reynaud, 1829, MNHN EcAsl471 (dry).
Other material. Indonesia, Sulawesi, 2 Jan 2000, UF 2623; New
Guinea, Trobriand Group, AM J22934 (1); 1 Jun 1998, UF 2332 (1); 1
Jul 1998, UF 2415(1); Philippines, Bohol I., 25 Mar 2004, NMV
F106973 (2); Australia, Queensland, Heron I., 14 Jul 1973, F95594 (2);
One Tree I., J23331 (1); Tryon I., 15 Sep 1970, F95789 (4); Northern
Territory, Darwin, 11 Oct 1976, F95799 (1); Western Australia,
Abrolhos Is., 2 Sep 1972, WAM Z6778 (7); J8321 (5); 11 Mar 1972,
F95793 (21); 12 May 1972, F95795 (6); Exmouth Gulf, F95794 (4);
F95790 (2); F95792 (1); F95787 (1); F95788 (1); Aug 1972, F95790
(2); Quobba, 20 Jun 1972, F95791 (3).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, rarely
4 or 6, long, broad basally, strongly tapered, narrowly rounded
distally, predominantly subequal, form sometimes
asymmetrical; up to R = 25 mm; gonopores abactinal, some
close pairs under same plate.
At R = 20 mm, r = 10 mm, abactinal plates angled up over
papulae, crescentiform appearance; 0-3 proximal doubly
papulate carinal plates; some large secondary plates, frequently
1 per space, up to 2; predominantly 1 papula per space, up to
3; disc frequently distinct, bordered by 5 large radial plates 5
small interradials; spinelets glassy, rugose, long, thin, finely
pointed distally, not splay-pointed, subacicular, subsacciform
to sacciform, up to about 32 spinelets in double series on large
radial plates, up to about 24 in double series on proximal
abactinal plates (frequently fewer), spinelets on anterior edge
of plates, frequently angled over papula, sometimes in tufts;
up to about 15 on mid-interradial plates; superomarginal plates
with variably up to 8-10 spinelets per plate, inferomarginals
with variably up to 16-24 thicker, longer spinelets per plate.
Spines per actinal plate variably up to: oral 6-9, suboral
2-8, furrow 6-8, subambulacral 4-8, actinal interradial 4-8;
interradial spines thick, conical, blunt, subsacciform.
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
271
Colour (live). Wide range of mostly mottled colours with pink,
mauve-red, grey, brown, grey-brown, reddish-brown, dark
brown, black flecking, green, cream, white; disc frequently
with broken white border (photos from N. Coleman).
Distribution. Indo-Malayan Region, Ceylon, Southern China,
Philippines, Indonesia, W Pacific Ocean, northern Australia;
0-70 m (Liao and A.M. Clark, 1995; Marsh, 1974 as Asterina
burtonr, Rowe and Gates, 1995).
Remarks. As noted in the diagnosis, the numbers of spinelets
and spines per plate is more variable than in other Aquilonastra
species. The distinctive diagnostic character of A. cepheus is
the thin long finely-pointed subsacciform spinelets projecting
over papulae. Smith (1927b), Liao (1980) and Liao and A.M.
Clark (1995) all noted this as a distinguishing character.
O’Loughlin and Waters (2004) redetermined as Aquilonastra
cepheus a specimen from Hong Kong (NHM 1981.2.6.25). It is
redetermined in this work as A. limboonkengi (below). In his
comprehensive synonymy for A. burtonii, Walenkamp (1990)
showed the position of many authors who considered A. cepheus
to be a junior synonym of A. burtonii. Liao and A.M. Clark
(1995) restored the species status of A. cepheus. Asterina
cepheus was reported for Thursday I. (N Australia) by Bell
(1884, Alert collections); for Zanzibar by Bell (1903); and for
Saya de Malha and Seychelles Is. by Bell (1909). None of this
material was examined. We found no evidence of A. cepheus in
the western Indian Ocean. The live colour of A. cepheus is
sometimes uniform, frequently mottled, and varies greatly.
Aquilonastra colemani sp. nov.
Figures 1, 2i-j, 5e, 7f
Material examined. Holotype. SE Papua New Guinea, China Straits,
Samarai I., on rubble, 150°48'E, 9°40'S, 10 m, J. Starmer, 6 Jun 1998,
UF 3284 (dry).
Paratypes. Type locality and date, UF 2419 (8, dry).
Other material. Indonesia, Flores Sea, West Sumbawa Regency,
Mayo I., underside of coral rubble, shallow sublittoral, 14 Nov 2005,
NMV FI 12173 (1); F109374 (4).
Diagnosis. Fissiparous Aquilonastra species; up to 7 rays,
predominantly 6, narrow base, rounded distally, subdigitiform;
up to R = 5 mm, r = 3 mm; high elevation apically, sides of rays
steep; up to 2 inconspicuous interradial madreporites seen, up
to 3 anal pores; gonopores not evident.
At R = 5 mm, upper ray plates irregular in size and form,
longitudinal series of large papulae along sides of rays, single
papula per plate; secondary plates present; spinelets thick
short conical to columnar, spinous surface, not splay-pointed,
up to about 12 spinelets on free surface of proximal abactinal
plates; superomarginal and inferomarginal plates subequal;
superomarginal plates each with up to about 8 spinelets,
inferomarginal plates each with up to about 12 spinelets.
Spines per actinal plate up to: oral 6, sub oral 3, furrow 5,
subambulacral 3, actinal interradial 2; interradial spines
glassy, rugose, bluntly pointed conical to digitiform.
Colour (live). Abactinally very dark brown on disc and upper
rays, white margin (photos from N. Coleman, S. Uthicke).
Distribution. SE Papua New Guinea, China Straits; Indonesia,
Flores Sea, Mayo I.; 0-10 m.
Etymology. Named for Neville Coleman, with gratitude for his
generous assistance in making available to us his many live
colour slides of Indo -Pacific asterinids.
Remarks. The distingusihing features of A. colemani are the
small size (up to R = 5 mm), fissiparous habit, and dark brown
with white margin colouration
Aquilonastra conandae sp. nov.
Figures 1, 2k, 4a, 5f, 8a-b
Asterina burtoni— Kojadinovic et al., 2004: 225-229 (part,
fissiparous specimens; non Asterina burtonii Gray, 1840).
Asterina’ sp. 2.— Rowe and Richmond, 2004: 3287-3288 (part,
not Riviere Banane or Zanzibar specimens), fig. 5 (colour).
Material examined. Holotype (in alcohol). Indian Ocean, Mascarene
Is., La Reunion I., Trou d’Eau, rocky shallows, C. Conand, 17 Sep
2004, MNHN EcAhll904.
Paratypes. Type locality and date, AM J24288 (23, in alcohol);
EcAhll905 (26, in alcohol); NMV F107411 (1, dry, dissected);
FI 0741 2 (29, in alcohol); F107413 (4, dry, dissected).
Other material. La Reunion I., type locality, Sep 2003, F107414
(5); 12 Jun 2002, NMV F109368 (2); 22 Mar 2003, F109366 (1); 16
Feb 2006, F109379 (1); Rodrigues I., lie aux Fous, coralline substrate.
Shoals of Capricorn Programme, 21 Sep 2001, NHM 2004.2815-2824
(10); Antonio’s Finger, 9 m, 21 Sep 2001, J24289 (1); Antonio’s Finger
Reef, off Grande Baie, NHM 2004.2826-2829 (4); Passe Coco, under
rubble, 6 m, NHM 2004.2830 (1); Trou Malabar, coral rubble, 10 m,
NHM 2004.2825 (1); Agalega Is., 25 Feb 1979, WAM Z6871 (1);
Mauritius (lie Maurice), MAU-74, 27 lots, Peyrot-Clausade, 1974,
EcAsl 1877-1 1903 (381); EcAs2578 (6); (lie Bourbon), Maillard, 1862,
EcAsl563 (1).
Diagnosis. Fissiparous Aquilonastra species; rays 5-7,
predominantly 6, largest specimens with 5; rays discrete, variably
narrow to wide basally, tapering, rounded distally, sometimes
digitiform; up to R = 10 mm, r = 5 mm (holotype, in alcohol);
contiguous spinelets over papulae possibly act as pedicellariae,
spinelets not differentiated as valves; abactinal gonopores.
At R = 8 mm, plates with proximal notch or indentation for
papula, rarely doubly notched; secondary plates present; single
large papula per papular space; 2 longitudinal series of single
papulae along each side of rays; abactinal spinelets glassy,
rugose, on upper rays short thick columnar to conical,
subgranuliform, up to rarely 12 per plate, readily lost, on sides
of rays short conical, in distal interradius thin, some splay-
pointed; marginal plates in regular series, subequal;
superomarginal spinelets up to about 5 per plate, short conical;
inferomarginal spinelets up to about 10 per plate, distal larger,
some splay-pointed.
Spines per actinal plate up to: oral 6 (rare), suboral 3 (rare),
furrow 4, subambulacral 2, actinal interradial 3 (predominantly
1); interradial spines short, thick, conical to subsacciform,
pointed distally.
Colour (live). Variably mottled with red, cream, brown and
green (photos from C. Conand).
272
P. Mark O’Loughlin and Francis W.E. Rowe
Figure 8. Photos of Aquilonastra species, a, A. conandae sp. nov.. La Reunion, actinal surface (NMV F107414); b, A. conandae sp. nov.. La
Reunion, interior with sup erambul acral (left arrow) and superactinal (right arrow) plates (NMV F107414); c, A. corallicola (Marsh), Cocos
(Keeling) I., fissiparous (up to R = 13 mm, UF 745); d, A. coronata (Martens), Australia, Darwin, side of ray, some paxilliform plates with
differentiated spinelets (R = 22 mm; NMV F95796); e, A. doranae sp. nov., Okinawa (R = 5 mm; holotype UF3913); f, A. halseyae sp. nov.,
Maidive Is., actinal spines (NHM 1965.6.1.84, holotype).
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
273
Etymology. Named for Chantal Conand, Professor Emeritus,
University of La Reunion, in appreciation of her considerable
contribution to echinoderm research and this paper.
Distribution. Indian Ocean, Mascarene Is., Agalega Is.,
Mauritius, Rodrigues Is., La Reunion I., rocky and coralline
substrate, 0-10 m.
Remarks. Kojadinovic et al. (2004) determined La Reunion
material to be Asterina burtoni, and discussed both small
asymmetrical fissiparous forms with more than one
inconspicuous madreporite and slightly larger symmetrical
pentaradiate sexual forms with a single madreporite. Maximum
size was R = 9 mm, consistent with A. conandae material
observed in this study. But none of the many specimens from
La Reunion seen here had only a single madreporite. Some of
the largest specimens were symmetrical and pentaradiate, but
all had more than one inconspicuous madreporite. Only a few
pentaradiate symmetrical specimens from La Reunion with a
single conspicious madreporite have been seen in this study.
Two are referred to Aquilonastra samyni sp. nov. (below), and
a few to an undescribed species of Tegulaster. A specimen
from Zanzibar (NHM 2004.2831), referred by Rowe and
Richmond (2004: 3288) to Asterina ’ sp. 2, is determined here
as Aquilonastra burtonii (above).
The distinguishing diagnostic characters for A. conandae
amongst western Indo-Pacific fissiparous species of
Aquilonastra are: predominantly single actinal interradial
spine per plate; high proportion of specimens with fewer rays
(80% with 5-6 rays); size difference. With R up to 10 mm, A.
conandae is larger than A. moosleitneri sp. nov. below (R up
to 9 mm) and A. yairi sp. nov. below (R up to 7 mm), and
smaller than A. burtonii (R up to 18 mm) and A. corallicola
below (R up to 16 mm).
Aquilonastra corallicola (Marsh, 1977)
Figures 1, 4b, 5g-h, 8c
Asterina corallicola Marsh, 1977: 271-275, figs 8, 9, this 1, 2.—
Oguro, 1983: 224-225, figs 7-9, 14.-A.M. Clark, 1993: 208.
Asterina anomala.— Fujita et al. 2001: 319, pi. 2G (non Asterina
anomala H.L. Clark, 1921).
Aquilonastra corallicola.— O’Loughlin and Waters, 2004: 11,
13-15.
Material examined. Paratypes. Caroline Is, Palau, 10 m, M.
Yamaguchi, 9 Feb 1971, WAM Z1704 (3); AM J10257 (2).
Other material. Caroline Is, Palau, 6 m, 7 Sep 1995, UF 3209 (1);
0-4 m, 6 Mar 2003, UF 1715 (1); 6 m, 7 Sep 1995, UF 3223 (1); 2 m,
6 Jun 1995, UF 324 (1); 23 m, 8 Jun 1995, UF 2440 (2); Jun 1975,
Z6845 (2); Marianas Is, Guam, 2-4 m, 11 Sep 1997, UF 895 (7); 2-3
m, 27 Nov 1998, UF 862 (12); 10-15 m, 14 Jan 2000, UF 678 (1); 18
m, 30 May 1997, UF 1119 (1); 1-2 m, 16 Apr 1999, UF 864 (6); 2 m, 5
Nov 1998, UF 827 (4); 10 m, 20 Jan 2000, UF 675 (1); Saipan I., 3-12
m, 18 Aug 2003, UF 3528 (1); Singapore, 1-3 m, 3 Nov 2000, UF 2714
(1); Indonesia, Sulawesi, 4-32 m, 26 Sep 1999, UF 1827 (1); Indian
Ocean, Cocos (Keeling) I., under dead coral rubble, 15 m, 8 Dec 1999,
UF 745 (7); 10-20 m, 6 Dec 1999, UF 750 (3); 1-2 m, 8 Dec 1999, UF
768 (2); UF 1648 (1); NE Australia, Great Barrier Reef, One Tree I.,
NMV F98746 (6); Fiji, 5-8 m, 11 Oct 2001, UF 1007 (1).
Diagnosis. Fissiparous Aquilonastra species; up to 8 rays,
predominantly 6 or 7, rays elongate, wide basally, tapered,
narrowly rounded distally; up to R = 16 mm (Sulawesi), 12 mm
(Palau), 13 mm (Cocos); some abactinal pedicellariae, with 2-
3 differentiated thick valves; abactinal gonopores.
At R = 12 mm, r = 5 mm; upper ray plates irregular in
arrangement, rarely a few doubly papulate carinal plates
proximally; 1-3 papulae per papular space, 3 rare; papulae in
2 longitudinal series along each side of ray; 0-2 secondary
plates per papular space proximally; upper ray plates
paxilliform with 2 forms of spinelets; 1-3 thick, digitiform,
blunt apical spinelets on each plate, up to about 12 thin conical
pointed spinelets peripherally on each plate, spinelets not
splay-pointed; superomarginal plates each with up to about 6
thin pointed spinelets; inferomarginal plates each with 1-4
thick digitiform apical spinelets, up to about 8 smaller thin
conical pointed spinelets.
Spines per actinal plate up to: oral 6, suboral 6, furrow 5,
subambulacral 5, actinal interradial 5; interradial spines
rugose, some digitiform, most thin conical pointed; actinal
spines generally of 2 forms, tall digitiform, short thin.
Live colour. Mottled red and yellow (photos from G. Paulay).
Distribution. W Pacific Ocean, Marianas Is., Caroline Is., Fiji,
NE Australia; E Indian Ocean, Cocos (Keeling) I.; Sulawesi;
Singapore; 0-32 m.
Remarks. The distinugishing characters for this large fissiparous
species are the long tapering rays, two forms of spinelets on
paxilliform upper ray plates, and golden red mottled colour.
Three small specimens from Madagascar (MNHN EcAsll876;
R = 4 mm), collected by Cherbonnier on 6 April 1960, are
damaged but appear to have the diagnostic characters of A.
corallicola. The colour and form of the material from Malaysia
referred to Asterina anomala by Fujita et al. (2001) indicate
that it is A. corallicola.
Aquilonastra coronata (Martens, 1866)
Figures 1, 21, 8d
Asterina coronata Martens, 1866: 73-74.— Fisher, 1918: 108-
110.— A.M. Clark and Rowe, 1971: 68, pi. 9 fig. 6.— VandenSpiegel et
al., 1998: 452-453, fig. 37A-D, pi. 3 fig. 8.-A.M. Clark, 1993: 208.-
Rowe and Gates, 1995: 34.— Chao, 1999: 407-408.— Fujita et al.,
2001: 319-320, pi. 2H. -Waters et al., 2004: 876-877, tbl. 1, figs 2, 3.
Asterina spinigera Koehler, 1911: 20-21, pi. 4 figs 11, 12 (junior
synonym by VandenSpiegel et al., 1998).
Asterina novae-zealandiae.— Goto, 1914: 643, pi. 19 figs 279-281
(non Asterina novaezelandiae Perrier, 1875 = A. coronata Martens,
1866, according to Fisher, 1919: 413).
Asterina cristata Fisher, 1916: 27-28.
Asterina cristata euerces Fisher, 1917: 91.
Asterina coronata cristata Fisher, 1918: 111, pi. 13.— Fisher, 1919:
411-414, pi. 115 fig. 3, pi. 131 figs 4, 4a.-Fisher, 1925: 80.-A.M.
Clark, 1993: 208-209.— O’Loughlin and Waters, 2004: 15 (junior
synonym, this work).
Asterina coronata euerces Fisher, 1918: 110.— Fisher, 1919: 414-
416, pi. 115 figs 1-2, pi. 116 figs 1-2, pi. 131 figs 5, 5a. -A.M. Clark,
1993: 209.— O’Loughlin and Waters, 2004: 15 (junior synonym, this
work).
274
P. Mark O’Loughlin and Francis W.E. Rowe
Asterina cor onata fascicular is Fisher, 1918: 110.— Fisher, 1919:
414.-H.L. Clark, 1928: 390.-H.L. Clark, 1938: 145-148, pi. 12 fig.
1.— A.M. Clark, 1993: 209 (junior synonym by Rowe and Gates,
1995).
Asterina coronata coronata.— Fisher, 1918: 110.— Fisher, 1919:
414.— A.M. Clark, 1993: 208.
Asterina coronata formajaponica Hayashi, 1940: 119-120, pi. 11
figs 5-7, pi. 13, fig. 7.— Hayashi, 1973: 72, pi. 12 fig. 3 (junior synonym,
this work).
Aquilonastra coronata.— O’Loughlin and Waters, 2004: 11, 13-
15.— Byrne, 2006: 245, tbls 1, 2, fig. 1.
Material examined. Australia, NT, Darwin, H.L. Clark, Jul 1929,
NMV F95797 (2); 11 Jun 1976, F95798 (1); 13 Jun 1976, F95796 (4);
AM J6188 (10); J6613 (18); J8206 (2); Caroline Is., J13660 (1); Taiwan,
Sanshi, 28 Feb 2003, UF 1425 (2); J19956 (1); Japan, F96700 (1);
Kushimoto, J 1 1 563 (2).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, broad
basally, tapering, narrowly rounded distally; up to R = 32 mm;
gonopores abactinal; numerous abactinal pedicellariae
frequently present, 2-4 curved pointed differentiated valves.
At R = 30 mm, r = 14 mm; abactinal surface very uneven
with irregularly distributed high paxilliform plates; abactinal
spinelets of 2 forms on paxilliform plates, up to about 6 thick
long digitiform or pointed spinelets on apex of each plate, up to
about 10 thin short pointed spinelets around margin of plate;
thin spinelets subacicular, not splay-pointed; disc clearly
delineated by 5 radial 5 interradial plates; lacking doubly-
papulate carinal plates; upper ray with irregular zig zag series of
small primary and secondary plates, lacking papulae; 0^1
secondary plates per papular space on upper rays, intergrade
with primary plates; 1^1 papulae per papular space;
superomarginal plates smaller than inferomarginals,
superomarginals with up to about 10 thin, rugose, pointed, not
splay-pointed spinelets per plate; inferomarginals with up to
about 4 thick long digitiform or pointed spinelets on apex of
each plate, up to about 12 thin spinelets around margin of plate.
Spines per actinal plate variably up to: oral 8, suboral 8,
furrow 7, subambulacral 7, actinal interradial 7, predominantly
5; interradial spines conical, pointed, frequently two forms on
each plate, thick centrally thin peripherally.
Colour (live). “The normal coloration is mottled olive-greens,
light and dark, with more or less dark dull red, usually in
irregular blotches. One specimen was very largely bright rust-
red, over most of the dorsal surface. Some specimens occur
with no trace of green dorsally; these are more or less fawn-
colour mottled with brown and have a distinct red tinge. Most
secimens have red markings but the shade may be very deep; in
a few cases it was replaced by black. A common feature is a
blotch of carmine at the base of each arm; in one specimen this
was nearer vermilion.” (H.L. Clark, 1938); mottled grey-brown,
with orange and white markings (photo from M. Komatsu);
mottled grey-brown, with dark brown, orange and white
markings (photo from T. Fujita).
Distribution. Northern Australia to Japan, Singapore to
Caroline Is.
Remarks. Fisher (1916, 1917, 1918, 1919, 1925) established and
maintained three subspecies of Asterina coronata, but stated
his own uncertainty. H.L. Clark (1928) recorded his doubt
about the subspecies. Hayashi (1940) added a fourth subspecies
from Japan. Rowe and Gates (1995) made A. coronata
fascicularis a junior synonym of A. coronata. Material from
Taiwan and northern Australian are morphologically
conspecific, including the presence of abundant pedicellariae.
In his key to four subspecies Fisher (1919) used the presence of
pedicellariae to key A. coronata euerces from the other three.
This is erroneous, since abundant pedicellariae are present in
the northern Australian and northern Pacific material. These
pedicellariae closely resemble those illustrated by Fisher (1919,
pi. 131 fig 5a) for A. coronata euerces. Variation in spine
number per plate is not an adequate basis for upholding the
subspecies. All are judged here to be junior synonyms.
O’Loughlin and Waters (2004) redetermined two
specimens from Bombay (NHM 1960.10.4.11-16), previously
identified as Asterina lorioli, as Aquilonastra coronata. That
decision is reversed here.
The distinguishing character of A. coronata is the
irregularly distributed high paxilliform abactinal plates with
two forms of spinelets.
Aquilonastra doranae sp. nov.
Figures 1, 3a, 8e
Material examined. Holotype (in alcohol). Japan, Okinawa, Yonashiro
Marine Road causeway to Henza I., 26°20'N, 127°56'E, intertidal
seagrass, 25 Jul 2004, G. Paulay, UF 3913.
Diagnosis. Fissiparous Aquilonastra species; rays 6, subequal,
short, broad basally, narrowly rounded distally; R = 5 mm, r =
3.5 mm; 3 inconspicuous interradial madreporites; gonopores
not evident.
Abactinal plates strongly imbricate, notched for papula;
upper ray plates with 1 papula, 2 irregular longitudinal series;
plates on sides of rays with 1 papula, 1 longitudinal series,
beginnings of second longitudinal series (0-3 plates); spinelets
conical or digitiform, distally spinous, up to about 10 spinelets
across raised proximal edge of proximal abactinal plates, plate
surfaces bare distally; superomarginal plates each with up to
about 6 spinelets, inferomarginal plates each with up to about
12 longer splay-pointed spinelets.
Spines per actinal plate up to: oral 5, suboral 3, furrow 4,
subambulacral 3, actinal interradial 5 (predominantly 3);
interradial spines conical, long, thin, finely tapered.
Colour (live). Mottled red, green, grey, white abactinally;
proximal disc area crimson red; proximal rays and interradii
dark grey-green to yellow-green; distal abactinal surface
mottled pale green, mauve grey, white (photo by G. Paulay).
Distribution. Japan, Okinawa, Henza I., seagrass shallows.
Etymology. Named for Ruth Doran, with gratitude for her
assistance in providing asterinid distribution maps.
Remarks. Although only one specimen is available, the
fissiparous habit, geographical isolation, green colouration,
short rays, thin digitiform or conical spinelets with long spines
distally, and up to five actinal interradial spines per plate
together support the erection of a new species.
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
275
Aquilonastra halseyae sp. nov.
Figures 1, 3b, 5i, 8f
Asterina burtoni.— A.M. Clark and Davies, 1966: 599, 603.—
Jangoux and Aziz, 1984: 861, 872, 873 (part; non Asterina burtonii
Gray, 1840).
Asterina cepheus.— Moosleitner, 1997: 12-13, fig. 23b (only) (fig.
23a is a photo of a Red Sea Aquilonastra sp., pers. comm. H.
Moosleitner (= A. marsliae sp. nov., see below); non Asteriscus
cepheus Muller and Troschel, 1842).
Asterina burtoni cepheus.— A.M. Clark and Rowe, 1971: 68-69,
fig. 17, tbl. 1.— A.M. Clark, 1993: 208 (non Aster iscus cepheus Muller
and Troschel, 1842).
Material examined. Holotype (in alcohol). Indian Ocean, Maldives,
Addu Atoll, Gan I., lagoon reef, 10 m, R Spencer Davies, 11 Feb 1964,
NHM 1965.6.1.84.
Paratypes. Type locality and date, NHM 1965.6.1.85a (1, dry);
NHM 1965.6.1.85b (1, part dissection, in alcohol).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, broad
basally, narrowly rounded distally; up to R = 19 mm, r = 9 mm;
gonopores not seen.
At R = 19 mm, rare proximal doubly papulate carinal
plates; rarely 1 secondary plate per papular space; spinelets
distinctly sacciform, sharp point distally; up to about 15
spinelets per proximal plate, sometimes in discrete groups;
primary superomarginal plates frequently separated by smaller
plate; superomarginals with 2-4 thin, sacciform pointed
spinelets; inferomarginal plates with up to about 12 spinelets,
proximally similar to superomarginal spinelets, more stout
and longer distally.
Spines per actinal plate up to: oral 6, suboral 5, furrow 6,
subambulacral 4, actinal interradial 4 (predominantly 2);
interradial spines stout, short, subsacciform, pointed distally,
on raised proximal edge of plates.
Colour (live). “Various shades of brown, sometimes with darker
or lighter spots” (Moosleitner, 1997; fig. 23b); disc variably
outlined in white; abactinally mottled pink, pale red-brown,
white, with dark red-brown flecks; or mottled dark and light red-
brown, white, bright red star apically (photos by N. Coleman).
Distribution. Indian Ocean, Maldives, 1-30 m (Moosleitner,
1997).
Etymology. Named for Sheila Halsey of the Natural History
Museum in London, who has graciously assisted the authors
with literature searches and facilitated the loan of materials for
echinoderm research.
Remarks. The distinguishing character of A. halseyae is the
distinctive sacciform, conical, sharply-pointed form of the
abactinal spinelets. A.M. Clark and Rowe (1971) judged that
the fissiparous and non-fissiparous asterinds occurring in the
Maldives were Asterina burtoni Gray, 1840, and referred the
fissiparous form to the subspecies A. burtoni burtoni and the
non-fissiparous form to the subspecies A. burtoni cepheus. We
refer the non-fissiparous Maldives asterinid to A. halseyae sp.
nov. (here), and the fissiparous Maldives asterinid to A.
moosleitneri sp. nov. (below).
Aquilonastra iranica (Mortensen, 1940)
Figures 1, 5j-k
Asterina cephea var. iranica Mortensen, 1940: 65-66, pi. 1 figs 1-4.
Asterina burtoni.— A.M. Clark and Rowe, 1971: 68, 69, tbl. I —
Price, 1983: 47-48, fig. 14 (part, non-fissiparous).
Asterina burtoni burtoni var. iranica.— AM. Clark, 1993: 207, 208.
Aquilonastra iranica. — O’Loughlin and Waters, 2004: 11, 13-15.
Material examined. Syntype, Iranian Gulf, S of Bushire, coral reef, 18
Feb 1937, AM J17891 (1, dry). Other material. Bahrain, Jufair, mudflat,
WAM Z6868 (6); W Pakistan, Balochistan coast, Gwader, 25°N62°E,
2 Dec 1977, NMV F112182 (1); F112183 (2).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, long,
broad basally, tapering, rounded distally; up to R = 35 mm, r =
15 mm (Mortensen, 1940); abactinal proximal interradial
pedicellariae, pairs of tooth-like differentiated valves; abactinal
gonopores.
At R = 20 mm, r = 11 mm, from 0 to 5 proximal doubly
papulate carinal plates; lacking secondary plates; spinelets on
abactinal plates in small groups, up to 4 per group, or in
transverse single or double series; up to about 12 spinelets on
proximal abactinal plates; spinelets short, thick, conical to
subsacciform; distal interradial plates with 4-6 splayed
spinelets; superomarginal plates smaller than inferomarginals,
up to about 6 spinelets per superomarginal plate, up to about
12 thicker spinelets per inferomarginal plate.
Spines per actinal plate up to: oral 8, suboral 5, furrow 7,
subambulacral 9 (sometimes in 2 series), actinal interradial 5
(predominantly 3); interradial spines long, conical.
Colour (live). “Grayish, with reddish or bluish-gray spots; oral
side lighter, uniformly coloured” (Mortensen, 1940).
Distribution. Iranian Gulf; Arabian Sea, W Pakistan, soft and
hard substrate.
Remarks. O’Loughlin and Waters (2004) raised A. cephea var.
iranica to species status. A. iranica is distinguished by: large
size (up to R = 35 mm); pedicellariae with differentiated valves;
few short thick conical to subsacciform spinelets on proximal
abactinal plates.
Aquilonastra limboonkengi (Smith, 1927)
Figures 1, 51, 6a, 9a
Asterina limboonkengi Smith, 1927b: 273-276, figs 1-3.— Liao,
1980: 171, figs 3:1, 3:2, 5.-A.M. Clark, 1982: 490-491.-A.M. Clark,
1993: 211.— Liao and A.M. Clark, 1995: 66, 130-131, pi. 18 figs 6-7.
Aquilonastra limboonkengi.— O’Loughlin and Waters, 2004: 11,
14, 15.
Material examined. Syntypes. China, Amoy, C. Ping, NHM
1926.12.22.35-36 (2, alcohol; very damaged).
Other material. SE Hong Kong, North Rocks, near Ninepins I., 15
m, NHM 1981.2.6.23-25 (1).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, long,
narrow to broad basally, tapering, rounded distally (rays
digitiform on syntypes), up to R > 25 mm (Liao and A.M.
Clark, 1995); gonopores abactinal.
276
P. Mark O’Loughlin and Francis W.E. Rowe
Figure 9. Photos of Aquilonastra species, a, A. limboonkengi (Smith), China, Amoy, ray (NHM 1926.12.22.35-36, syntype); b, A.lorioli
(Koehler), Pakistan, Karachi, disc (MNHN EcAs2662, syntype); c, A. lorioli (Koehler), Pakistan, New Pachha, atypically eight rays, two
conspicuous madreporites (arrows), asymmetrical (non-fissiparous; photo Qaseem Tahera); d, A. marshae sp. nov.. Red Sea, actinal (TM H1814);
e, A. moosleitneri sp. nov., Maidive Is. (R = 7 mm; holotype NHM 1902.3.13.27); f, A. oharai sp. nov., Okinawa (R = 12 mm; holotype UF
3285).
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
277
At R = 18 mm, r = 7 mm (larger syntype), from 0 to 4
proximal doubly-papulate carinal plates; 0-1 secondary plates
per papular space; disc bordered with 5 radial 5 smaller
interradial plates; spinel ets on proximal abactinal plates in up
to 4 groups, up to 10 spinelets per group, up to more than 30
spinelets on proximal abactinal plates; spinelets long, thick,
conical to subsacciform, with numerous (5-6) points on distal
sides and end of spinelets, not splay-pointed; mid-interradial
plates with up to about 20 long, thin, pointed spinelets,
sometimes larger group over anterior edge of plate, smaller
group distally, groups splayed and overlapping spinelets on
adjacent plates; superomarginal plates with up to about 12 thin
spinelets per plate, up to about 24 thick spinelets per
inferomarginal plate.
Spines per actinal plate up to: oral 7, suboral 6, furrow 7,
subambulacral 7, actinal interradial 10 (frequently about 6);
interradial spines long, conical.
Colour (live). “Dark brown with irregular red, purple or light
brown spots” (Liao and A.M. Clark, 1995).
Distribution. SE coastal China, Guangdong and Fujian
Provinces (Liao and A.M. Clark, 1995).
Remarks. Liao (1980) and Liao and A.M. Clark (1995)
distinguished Asterina limboonkengi from Asterina burtoni
cepheus (= A. cepheus here) by: abactinal spinelets squat,
opaque with 5-6 terminal points (not slender, transparent with
1-3 terminal points); actinal spines stout, rugose with many
points distally (not slender, smooth with few points distally);
thin abactinal plates with large papular spaces with numerous
secondary plates in large specimens (not thick with small
papular spaces lacking secondary plates). We do not agree
with all of these distinctions. We considered the actinal spines
on the type specimen of A. cepheus to be short, thick, conical,
blunt, and those on the syntypes of A. limboonkengi to be long,
conical, finely tapered. We observed secondary plates on the
types of both species. We thus have some uncertainty about
the status of A. limboonkengi in China waters. O’Loughlin
and Waters (2004) determined material from Oman (UF 68,
UF 246, UF 1645) to be Aquilonstra limboonkengi. This
material is referred here to the new species Aquilonastra
samyni (below).
Aquilonastra lorioli (Koehler, 1910) comb. nov.
Figures 1, 4c, 6b-c, 9b-c
Asterina lorioli Koehler, 1910: 129-131, pi. 19 figs 5-8.— H.L
Clark, 1915: 95. -A.M. Clark and Rowe, 1971: 38, 67. -A.M. Clark,
1993: 211.— O’Loughlin and Waters, 2004: 11, 37.
Palmipes sarasini de Loriol, 1897: 12.— Koehler, 1910: 129 (part;
non Palmipes sarasini de Loriol, 1897).
Material examined. Syntypes. Pakistan, Karachi, MNHN EcAs2662
(1, dry, ray broken off); Mergui Archipelago, Cheduba I., WAM Z6848
(1, dry).
Other material. Bombay, BMNH 1960.10.4.11-16 (2); Native
Jetty, Karachi, BMNH 1967.11.1.4 (2); Pakistan, Buleji, 24°N, 66°E, 9
Sep 2006, NMV F112184 (9).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, rarely
6 (Koehler, 1910, pi. 19 fig. 7), broad basally, narrowly rounded
distally, up to R = 22 mm, r = 13 mm; gonopores abactinal.
At R = 22 mm, variable series of doubly-papulate carinal
plates from none to half ray length, each with one cluster or up
to 4 tufts of spinelets, up to about 20 spinelets on proximal
carinal plates; disc defined by 5 transversely elongate series of
subpaxilliform spinelets on radial plates, 5 small interradial
plates; some high paxilliform clusters of spinelets on rays;
spinelets vary significantly in form from large globose to thick
digitiform apically on plates, to small pointed conical on
margin of plates and on secondary plates; papular spaces large,
predominantly single large papula per space, frequently 2 per
space, up to 5 small secondary plates per space; superomarginal
and inferomarginal plates subequal, superomarginal plates
with up to about 10 short conical spinelets per plate,
inferomarginal plates with up to about 20 spinelets.
Spines per actinal plate up to: oral 7, suboral 4, furrow 6,
subambulacral 5, actinal interradial 6, predominantly 3;
interradial spines thick, digitiform to bluntly conical.
Colour (live). Abactinal colour variably mottled with grey-
brown, green-brown, yellow-brown, red-brown, red, brown
(photos from Qaseem Tahera).
Distribution. Arabian Sea, Karachi, Bombay, Ceylon.
Remarks. Specimen EcAs2662 (R = 18 mm) is from the type
locality (Karachi), and the label is written in Koehler’s
handwriting (Tim O’Hara pers. comm.). It is judged here to be a
syntype. Specimen WAM Z6848 (R = 7 mm) is from the type
locality (Cheduba I.), and the label records “Exchange Zoological
Survey India”. It is judged to be one of the small specimens (7-
11 mm) referred to by Koehler (1910), and also a syntype.
O’Loughlin and Waters (2004) listed Asterina lorioli as
incertae sedis, because type material and information about
internal skeletal structure were not available. Type material is
available for this study, and the species is reassigned as the
new combination Aquilonastra lorioli (Koehler, 1910). The
variety of spinelet form, and paxilliform clusters of spinelets
along rays, are similar to the spinelet arrrangement in
Aquilonastra coronata (Martens, 1886). A. lorioli is
distinguished by the globose to subgranuliform spinelets. A
specimen (photo from Qaseem Tahera, figure 9c) has eight
rays and two conspicuous madreporites, but the spinelets are
distinctively those of A. lorioli.
Aquilonastra marshae sp. nov.
Figures 1, 3c, 6d-e, 9d
Asterina cephea.— Perrier, 1875: 315-317 (part. Red Sea; non
Asteriscus cepheus Muller and Troschel, 1842).
Asterina burtonii.— Mortensen, 1926: 121 (part).— A.M. Clark,
1952: 207 (part; non-fissiparous).— Tortonese, 1960: 20-21 (part; non
Asterina burtonii Gray, 1840).
Asterina burtoni.— Achituv, 1969: 329-341 (part, “pentaradiate”
form).— James and Pearse, 1969: 84-85 (part).— A.M. Clark and
Rowe, 1971: 68, tbl. 1 (part. Red Sea non-fissiparous).— Achituv,
1973b: 547-553.— Tortonese, 1977: 281-282 (part).-Price, 1982: 7
(part. Red Sea non-fissiparous).— Mladenov and Achituv, 1999: 152
278
P. Mark O’Loughlin and Francis W.E. Rowe
(part).— Karako et al., 2002: 139-144 (part, Elat Gulf of Aqaba non-
fissiparous population; non Asterina burtonii Gray, 1840).
Asterina cepheus.— Moosleitner, 1997: 12-13, fig. 3a (only) (non
Asteriscus cepheus Muller and Troschel, 1842).
Material examined. Holotype. Red Sea, Jousseaume, 1892, MNHN
EcAsll907 (dry).
Paratypes. Type locality and date, EcAsl0316 (7, dry).
Other material. Red Sea, M. Botta, 1837, EcAs2713 (1; labelled as
“type” of Asteriscus wega Val.; discussed under A. burtonii above);
NMV FI 12169 (12); S Sinai, El Fauz, Nov 2003, F109382 (5); Ras el
Misalla, 22 Sep 1970, HUJ SLR3030 (3); Ras Matarma, 31 Jan 1969,
SLR2199 (1); 22 Sep 1981, TAU NS24413 (1); Egypt, under rocks,
shallows, 15 Jul 2005, F107430 (1); Feb 2003, F109362 (1); 15 Jul
1966, TM H1814 (1); Gulf of Suez, Mission Dollfus, 1928, EcAsll839
(1); 27 Oct 1971, NS8560 (1); et Tur, 11 Sep 1968, HUJ IEC.57/141-
198 (11); Gulf of Aqaba, 8 Oct 1968, TAU NS4130 (5); Jez Tiran, 25
Sep 1981, WAMZ6877(1).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5,
sometimes 6, short, broad basally, tapered, narrowly rounded
distally; up to R = 16 mm, r = 7 mm; predominantly 1
conspicuous madreporite, sometimes 2; gonopores abactinal;
some paired thick spinelets probably act as pedicellariae, not
differentiated as valves.
At R = 16 mm, 0-3 doubly-papulate carinal plates; plates
angled over papulae, proximally crescentiform, 1 papula per
space, frequently 1 secondary plate per papular space; disc
bordered or not by 5 radial 5 interradial plates; spinelets thick,
short, conical to subdigitiform, rugose, distally with multiple
long fine points and frequently splay-pointed; up to about 14
spinelets across raised proximal edge of abactinal plates and
angled over papula, frequently 1-2 on central bare distal
surface of plate; up to about 8 spinelets per plate in mid-distal
interradius; up to about 7 conical pointed spinelets per
superomarginal plate, up to about 16 digitiform spinelets per
inferomarginal plate, thick distally on plates.
Spines per actinal plate up to: oral 7, suboral 3, furrow 6,
subambulacral 5, actinal interradial 5, predominantly 3; actinal
interradial spines thick, sacciform, conical to digitiform.
Colour (live). Variably mottled with combinations of dark and
pale mauve, dark brown to black, pale brown, red, green, white;
sometimes predominantly pink-mauve or white (photos from
H. Moosleitner and J. Hinterkircher; Moosleitner (1997)).
Distribution. Red Sea, Gulfs of Suez and Aqaba.
Etymology. Named for Loisette Marsh, Western Australian
Museum, with appreciation of her generous assistance with
loan material and her significant research into echinoderm
systematics in the Indo-Pacific region.
Remarks. Two fissiparous species occur in the Red Sea, and
are recognized in this work as Aquilonastra burtonii (above)
and Aquilonastra yairi sp. nov. (below). Aquilonastra marshae
has frequently been reported in the literature as a larger non-
fissiparous pentaradiate growth stage of Asterina burtonii (see
synonymy). Achituv (1973b) reported on the genital cycle of
Asterina burtoni from the Gulf of Elat. Fissiparous and non-
fissiparous forms were not distinguished. Maximum R was 14
mm. We judge that the material was probably principally A.
marshae. Aquilonastra samyni sp. nov. (below) occurs on the
coast of the Arabian Sea and is morphologically similar to A.
marshae. The diagnostic characters that distinguish A.
marshae from A. samyni are discussed in the Remarks for A.
samyni below.
Aquilonastra minor (Hayashi, 1974)
Figures 1, 3d
Asterina minor Hayashi, 1974: 41-44, fig. 1.— A.M. Clark, 1993:
211.— Fujita and Saba, 2000: 169-170, pi. 3E.— Waters et al., 2004:
873, 876, 877, tbl. 1, figs 1, 2.
Aquilonastra minor.— O’Loughlin and Waters, 2004: 11, 13-15,
fig. 1.— Saba and Fujita, 2006: 270-272, 286, fig. 15.— Byrne, 2006:
245, 251, tbls 1, 2, fig. 1.
Material examined. Japan, NMV F96697 (2).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, short,
broad basally, narrowly rounded distally; up to R = 9 mm;
gonopores actinal; direct development into brachiolaria stage
(Hayashi, 1974).
At R = 7 mm, r = 5 mm, rare proximal doubly papulate
carinal plates, rare secondary plates, abactinal plates with low
domes, spinelets in tufts or bands, subpaxilliform, described
as “paxilliform” by Hayashi (1974); up to about 20 spinelets
on proximal abactinal plates; spinelets thin, columnar, distally
spinous; superomarginal plates smaller than inferomarginals,
up to about 8 spinelets per superomarginal plate, up to about
12 spinelets per inferomarginal plate.
Spines per actinal plate up to: oral 6, suboral 4, furrow 5,
subambulacral 3, actinal interradial 3; interradial spines
digitiform.
Colour (live). “Pale greenish white, pale green or brownish
grey with small greenish white patterns and small dark brown
scattering spots, and reddish colour intermixed in the marginal
zone; madreporite yellowish orange” (Hayashi, 1974); disc
cream, rays and interradii pale mottled pink, brown, cream
white (photo from M. Komatsu).
Distribution. Japan, southern Honshu, Kushimoto, littoral
(Hayashi, 1974).
Remarks. The largest of the many specimens observed by
Hayashi (1974) were up to R = 6 mm. The two specimens
donated by Mieko Komatsu to Jon Waters, and subsequently to
Museum Victoria, are up to R = 7 mm (preserved). Fujita and
Saba (2000) reported R up to 9.4 mm for Takehara (assumed
not dried). Saba and Fujita (2006) reported R up to 7.7 mm for
Sagami Bay. We wonder whether there is more than one species
being referred to A. minor. Distinctive characters for A. minor
are: rounded low domes on some abactinal plates; actinal
gonopores.
Aquilonastra moosleitneri sp. nov.
Figures 1, 9e
Asterina burtoni.— A.M. Clark and Davies, 1966: 599.— A.M.
Clark, 1967b: 146, fig. lB.-Jangoux and Aziz, 1984: 861, 872, 873
(part; non Asterina burtonii Gray, 1840).
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
279
Asterina burtoni burtoni.— A.M. Clark and Rowe, 1971: 68, fig.
17e.— A.M. Clark, 1993: 207-208 (non Asterina burtonii Gray, 1840).
Asterina anomala.— Moosleitner, 1997: 12, fig. 22 (non Asterina
anomala H.L. Clark, 1921).
Material examined. Holotype. Maidive Is., Male, Hulule, J.S Gardiner,
1899-1900, NHM 1902.3.13.27 (alcohol). Paratypes. Type locality
and date, NHM 1902.3.13.28-33 (30).
Other material. Maidive Is., Eryadoo I., WAM Z6854 (1; photo in
Moosleitner, 1997).
Diagnosis. Fissiparous Aquilonastra species; rays up to 7,
predominantly 6, form frequently asymmetrical post-fissiparity,
form of larger specimens sometimes symmetrical with 5 equal
rays, most interradii with inconspicuous madreporite; rays
narrow basally, tapering, narrow rounded distally; up to R = 9
mm, r = 4 mm; abactinal gonopores on largest pentaradiate
specimens.
At R = 9 mm, lacking carinal plates; some secondary
plates; 3 longitudinal series of papulae on sides of mid-ray;
spinelets granuliform, short conical to columnar, rugose,
blunt; up to about 10 spinelets over each plate, readily detached;
superomarginal plates each with up to about 5 spinelets,
inferomarginal plates each with up to about 10 larger
spinelets.
Spines per actinal plate up to: oral 5, suboral 2, furrow 5,
subambulacral 3, actinal interradial 3 (predominantly 3);
interradial spines short, conical, pointed.
Colour (live). “Reddish, speckled with darker and lighter spots”
(Moosleitner, 1997).
Distribution. Maidive Is.
Etymology. Named for Horst Moosleitner, with gratitude for his
assistance in providing photos and specimens for this work, and
with appreciation of his work on the asteroids of the Maldives.
Remarks. Moosleitner (1997) noted that in the absence of a
connecting growth series this small fissiparous species was
probably a separate species from the non-fissiparous asterinid on
the Maldives. We agree that there are two species. A.M. Clark
(1967b) determined the small fissiparous asterinid from the
Maidive Is as Asterina burtoni. The 31 specimens (NHM
1902.3.13.27-33) are similar to Aquilonastra conandae in size,
number of rays, and spinelet and spine form, but are similar to
Aquilonastra burtonii in frequently having more than one
interradial actinal spine per plate. The species size (up to R = 9
mm) is significantly smaller than A. burtonii (up to R = 18 mm).
Aquilonastra oharai sp. nov.
Figures 1, 3e, 9f
Material examined. Holotype. Japan, Okinawa, Seragaki, under rock
on reef, 1-2 m, G. Paulay, 26 Jul 2004, UF 3285 (alcohol).
Paratypes. Type locality and date, UF 3916 (1, alcohol); Kunigami,
reef flat, under rock, G. Paulay, J. Geller, M. Malay, Y. Hiratsuka, 4 Jul
2004, UF 3914 (1, alcohol; BOKI-14, d GP 595, 596).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, wide
basally, tapered, rounded distally; up to R = 12 mm, r = 7 mm;
gonopores abactinal.
At R = 12 mm, lacking proximal doubly papulate carinal
plates; some secondary plates, frequently 1 per proximal
papular space; single large papula per space; spinelets conical,
thin, fine point to splay-pointed, subsacciform, in double
splayed series across proximal edge of plates, rarely in tufts,
up to about 14 spinelets on proximal abactinal plates, up to
about 8 on distal interradial plates; superomarginal plates
smaller than inferomarginals, superomarginals with up to
about 7 spinelets per plate, inferomarginals with up to about
14 spinelets, more stout on inferomarginals.
Spines per actinal plate up to: oral 6, suboral 2, furrow 4,
subambulacral 4, actinal interradial 6, predominantly 4-6
proximally; interradial spines conical, thick basally, pointed
distally.
Colour (live). Variable, disc dark brown, mottled abactinally
with mauve-violet, dark and pale brown, greenish brown, off-
white; or disc dark brown, mottled abactinally with pale and
dark green, and off-white (photos by G. Paulay).
Distribution. Japan, Okinawa, 0-2 m.
Etymology. Named for Tim O’Hara, Senior Curator of Marine
Invertebrates, Museum Victoria, in appreciation of his
contribution to this work and to echinoderm systematics and
biogeography.
Remarks. A. oharai is distinguished from A. cepheus and A.
limboonkengi by the shorter rays, absence of any proximal
doubly papulate carinal plates, absence of clustering of the
abactinal spinelets on the plates, and the splay-pointed form of
some abactinal spinelets. A. oharai is described from only three
specimens, with R up to 12 mm. The morphological characters
used to distinguish the new species may be variable within the
species, and size related. In the absence of adequate comparative
material we have some uncertainty about the status of A. oharai
in relation to A. cepheus and A. limboonkengi.
Aquilonastra richmondi sp. nov.
Figures 1, 4e, 6f,g, 10a
Asterina burtoni.— Jangoux, 1973: 35-38, fig. 13 (probably part,
Mozambique material; non Asterina burtonii Gray, 1840).
Asterina burtonii.— Walenkamp, 1990: 67-72, figs 30, 31 (part,
Mozambique material; non Asterina burtonii Gray, 1840).
Asterina coronata.— Jangoux, 1973: 38-39 (part, 3 Mozambique
specimens; non Asterina coronata Martens, 1866 according to
Walenkamp, 1990; non Asterina burtonii Gray, 1840).
Asterina’ sp. 1.— Rowe and Richmond, 2004: 3287 (part. Grand
Pate specimen), fig. 4 (colour).
Material examined. Holotype. Tanzania, Ras Kimbiji, lower littoral,
under boulders, M. Richmond, 4 Sep 2004, NHM 2005.37 (alcohol).
Paratypes. Type locality and date, NHM 2005.35 (1); NHM
2005.36 (1).
Other material. Cotes dArabie, St. XLVII, 1901-1904, MNHN
EcAsll043 (1); Kenya, Kiunga Marine Reserve, 8 m, Apr 1999,
MRAC 1739 (1); Zanzibar, Rousseau, 1841, EcAh3884 (1); Tanzania,
Mnazi Bay, Ruvula Peninsula, rocky, lower littoral, 3 Feb 2004, NHM
2004.2832 (1); Mombassa, NHM 1972.8.22.3-17 (2 of 14 seen);
Comoros, Mayotte, Mission Cherbonnier 25, littoral reef, 1959,
MNHN EcAsll862 (1); Mission Cherbonnier 27, 1959, EcAsll863 (1);
280
P. Mark O’Loughlin and Francis W.E. Rowe
Figure 10. Photos of Aquilonastra species, a, A. richmondi sp. nov., SE Africa, Sodwana Bay, disc (MRAC 1737); b, A. rowleyi sp. nov., SE
Africa, Sodwana Bay, disc (MRAC 1736, holotype); c, A. rowleyi sp. nov., SE Africa, Sodwana Bay, pedicellariae (MRAC 1736, holotype); d, A.
samyni sp. nov., SE Africa, Sodwana Bay, disc (MRAC 1741); e, A. watersi sp. nov., Rodrigues I. (NHM 2004.2813-2814); f, A. yairi sp. nov.,
Mediterranean Sea, Israel, Michmoret (R = 7 mm; holotype NMV FI 12174).
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
281
NW Madagascar, Nossi-Bi I. (Nosy Be), Plante Collection, 1965-
1970, EcAs 11865 (2); littoral reef, 2 Dec 1959, EcAsll858 (5);
Cherbonnier, 6 Oct 1959, EcAsll859 (7); SW Madagascar, Mission
Cherbonnier 201, Station Platier, 1962,EcAsll861 (2); NE Madagascar,
lie Sainte Marie, lie aux Nattes, 26 Jun 1960, EcAsll860 (1); SE
Madagascar, Fort Dauphin, Mission Decary, 1932, EcAsll864 (1);
Madagascar, Gruvel Collection, 1923, EcAsl0379 (1); Mauritius,
Rodrigues I., Grand Pate, on coral, 20 m, 23 Sep 2001, AM J24287 (1);
E South Africa, Sodwana Bay, 11 m, 10 Feb 2001, MRAC 1737 (1); 14
m, Aug 1999, MRAC 1738 (1).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, rarely
6, broad to narrow basally, tapered, narrowly rounded distally ; up
to R = 25 mm, r= 12 mm; form frequently asymmetrical, one ray
shorter than other 4; gonopores abactinal; pedicellariae sometimes
evident on upper rays and in interradii, 2-3 short, thick valves not
significantly differentiated from adjacent spinelets.
At R = 20, r = 10 mm, abactinal plates closely imbricate,
proximal edge projecting, spinelets subpaxilliform; 0-5
proximal doubly-papulate carinal plates; few proximal
secondary plates, 0-2 per space; predominantly single large
papula per space, sometimes 2; spinelets short, conical, bluntly
pointed, subsacciform, not splay-pointed, frequently of two
forms, thin on outer margin of plates, thick short on apex of
plates (columnar to subgranuliform); up to about 20 spinelets
on each proximal abactinal plate, predominantly over surface
of plate, some in clusters of 3-4; distal interradial plates with
up to about 8 long, thin, pointed spinelets, frequently splayed
in distinctive widely radiating cluster per plate, overlapping
spinelets of adjacent plates on larger specimens ; superomarginal
plates smaller than inferomarginals, superomarginal plates
with up to about 7 spinelets per plate, inferomarginals with up
to about 14 spinelets per plate, thin proximally thick distally.
Spines per actinal plate up to: oral 8, suboral 4, furrow 7,
subambulacral 5, actinal interradial 5, predominantly 2;
interradial spines thick, rugose, bluntly conical, subsacciform.
Colour (live). Sharply mottled with green, maroon, pale brown,
pink, white, black (photos by M. Richmond); variable and
changeable, mottled with red, pink, orange, white, grey, blue
(Walencamp, 1990; with colour photos).
Distribution. Cotes dArabie, E Africa, Kenya, Tanzania,
Comoros, Madagascar, Mauritius, E South Africa; under
boulders and in crevices in rocks and live coral; 0-20 m.
Etymology. Named for Matt Richmond, with appreciation of
his contribution of material from Tanzania and Rodrigues I.
Remarks. Two distinctive features of A. richmondi are:
frequently two forms of spinelets on the abactinal plates, thicker
apically; splayed overlapping spinelet clusters on the distal
abactinal plates. The detailed description and colour photos of
A. burtonii by Walenkamp (1990) indicate that the Mozambique
Inhaca material was not A. burtonii and is conspecific with A.
richmondi. Ludwig (1899) reported Asterina cepheus for
Zanzibar. There is no evidence in this study of A. cepheus
occurring in the western Indian Ocean, but the morphologically
similar A. richmondi is reported here for Zanzibar.
Aquilonastra rosea (H.L. Clark, 1938)
Figures 1, 3f, 6h
Paranepanthia rosea H.L. Clark, 1938: 161-162, pi. 22 fig. 8.—
H.L. Clark, 1946: 137. -A.M. Clark, 1993: 223,-Marsh and Pawson,
1993: 285.— Rowe and Gates, 1995: 39.
Asterinopsis rosea.— Cotton and Godfrey, 1942: 203.
Aquilonastra rosea . — O’Loughlin and Waters, 2004: 11, 13-15.
Material examined. Paratypes. SW Western Australia, Rottnest I.,
AM J6171 (3).
Other material. Rottnest I., 18 Aug 2004, WAM Z31171 (1); Perth,
4 Aug 2004, Z31174 (1); Jurien Bay, J7437 (4); 13 m, 27 Apr 2005,
Z31162 (1); Abrolhos Is., 110 m, 6 Dec 1970, Z21265 (1).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, long
(at R = 17 mm), wide basally, tapering to narrowly rounded
distally, up to R = 17 mm, r = 9 mm.
At R = 17 mm, regular doubly papulate carinal plates
proximally; abactinal plates not in 2 “fields”; abactinal plates
with high raised rounded column or ridge, concave papular
notch; longitudinal series of plates and papulae along sides of
rays; predominantly single large papula per space; very few
secondary plates; disc delineated by 5 wide radial plates, 5
short interradial plates; spinelets sacciform, long, splay-
pointed; spinelets in paxilliform dense splayed round tufts of
more than 30 per plate; superomarginal and inferomarginal
plates with subequal tufts of spinelets.
Spines per actinal plate up to: oral 9; suboral 12; furrow 7;
subambulacral 12; actinal interradial 20; spines on
subambulacral and actinal plates in tufts; interradial spines
sacciform, splay-pointed.
Colour (live). “Rose-red, more or less variegated with cream-
colour” (H.L. Clark, 1938).
Distribution. SW Australia, Abrolhos I. to Rottnest I.; 0-110 m.
Remarks. O’Loughlin and Waters (2004) reassigned
Paranepanthia rosea to Aquilonastra. This reassignment is
upheld here provisionally on the basis of the arrangement of
abactinal plates and papulae, the presence of internal
superambulacral plates, and the arrangement of actinal plates.
However, the paxilliform tufts of long thin abactinal spinelets
and actinal spines, and high numbers of spinelets and spines
per plate, are atypical of Aquilonastra. A. scobinata (below) is
morphologically similar to P. rosea in having paxilliform tufts
of numerous long thin abactinal spinelets, superambulacral
plates, and actinal plates in series parallel to the furrow. The
paxilliform tufts of spinelets in A. rosea are round, those in A.
scobinata are frequently crescentiform.
Aquilonastra rowleyi sp. nov.
Figures 1, 6i, 10b, c
Material examined. Holotype (dry). South Africa, KwaZulu Natal,
Sodwana Bay, 11 m, Y. Samyn and I. Tallon, 10 Feb 2001, MRAC
1736.
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, not
discrete or elongate, wide basally, narrowly to broadly rounded
distally, R = 23 mm, r = 14 mm; form shallow concave
282
P. Mark O’Loughlin and Francis W.E. Rowe
interradially; pedicellariae present in proximal interradii, 2
valves differentiated with inner teeth, taller than adjacent
spinelets; abactinal gonopores.
At R = 23 mm, lacking abactinal carinal series of plates,
broad band of upper ray plates irregularly arranged,
longitudinal and transverse series along sides of rays creating
lattice-like appearance, plates with concave papular notch;
single small papula per space; 6 longitudinal series of papulae
along each side of mid-ray; secondary plates numerous on
irregular upper ray; disc delineated by continuous curved
dense band of spinelets on 5 wide radial plates, each with up to
about 100 spinelets, 5 small interradial plates; spinelets glassy,
elongate, thin, pencil-like; up to about 30 spinelets in short
crescentiform band across projecting edge of each plate on
proximal ray; irregular lumpiness more than glassy convexities
on plates; superomarginal and inferomarginal plates subequal,
in regular series; superomarginal plates with up to about 8
pencil-like spinelets; inferomarginal plates with up to about 16
spinelets, proximally subequal with superomarginal spinelets,
distally longer and thicker.
Integument evident actinally; non-plated area in one
proximal actinal interradius.
Spines per actinal plate up to: oral 10 long, thin; suboral 5
long, 5 short; furrow 7; subambulacral 7; actinal interradial 4,
predominantly 3; interradial spines glassy, long, sacciform, in
webbed combs.
Colour (live). No record.
Distribution. East African coast, KwaZulu Natal, Sodwana
Bay, 11m.
Etymology. Named for Chris Rowley (Marine Biology Section,
Museum Victoria), in appreciation of the photography and
curatorial assistance that have been graciously provided in
support of asterinid systematic research.
Remarks. The new species has many of the diagnostic characters
of Aquilonastra, but is assigned with reservations because the
numerous spinelets are pencil -like and not typical of
Aquilonastra. They resemble those of Patiriella oliveri (Benham,
1911), Callopatiria granifera (Gray, 1847), and Aquilonastra
scobinata (below). The rays are distinct, but the form is closer to
subpentagonal than the discrete-rayed form of Aquilonastra. In
an absence of adequate material we are unwilling to erect another
asterinid genus. The tall pedicellariae with distinctive inner-
toothed valves are unique amongst asterinids.
Aquilonastra sarnyni sp. nov.
Figures 1, 3g, h, 6j, lOd
Material examined. Holotype. Oman, Masirah I., 1-7 m, under rocks,
G. Paulay, 6 Nov 1999, UF 246 (alcohol).
Paratypes. Type locality, 15-18 m, G. Paulay, 5 Nov 1999, UF 68
(8, alcohol); Bar al Hikman, under rocks, 0-1 m, M. Bouchard, 7 Nov
1999, UF 1645 (2).
Other material. Oman, Bar al Hikman peninsula, 1-3 m, under
rocks, 18-20 Jan 2005, UF 4210 (2); 2-3 m, UF 4201 (3); 2-4 m, under
rocks, Jan 2005, UF 4143 (3); Muscat, Qurm, intertidal, 26 Jan 2005,
UF 4147 (6); UF 4251 (1); UF 4252 (1); Bandar Khayran, under rocks,
0-5 m, 29 Oct 1999, UF 1378 (1); Madagascar, MNHN EcAsll853 (1);
NW Madagascar, Nossi Be (Nosi Be), littoral reef, 24 Sep 1964,
EcAsll848 (1); 2 Dec 1959, EcAsll849 (5); 1962, EcAsll850 (1); 6
Oct 1959, EcAsll851 (3); 3 Dec 1959, EcAsll852 (2); SW Madagascar,
Tulear, EcAsll847 (1); lie Sainte-Marie, 11 Feb 1979, WAM Z6870
(2); La Reunion I., Tobogan, 10 Sep 2001, NMV F109365 (1); South
Africa, Sodwana Bay, 10 m, Jul 2000, MRAC 1741 (1); 13 m, Aug
1999, MRAC 1740 (1); MRAC 1743 (1); Bangha Nek, 16 m, Aug 1999,
MRAC 1742 (1).
Diagnosis. Non-fissiparous Aquilonastra species; rays
predominantly 5, rarely 6, narrow basally, slight taper, rounded
distally, subdigitiform, up to R = 27 mm, r = 11 mm; single
conspicuous madreporite, 2 rare, 3 very rare; gonopores
abactinal.
At R = 23 mm, r = 9 mm, 0-3 proximal doubly papulate
carinal plates; disc frequently well-delineated by 5 radial 5
interradial plates; up to about 30 spinelets in irregular double
transverse series across proximal abactinal plates, some
spinelets in poorly defined clusters (spinelets granuliform on
small specimens); 0-2 secondary plates per papular space
proximally, rarely 2 papulae per space; spinelets small, short,
thick, conical to columnar, sometimes splay-pointed,
sometimes with long thin distal point; up to about 14 on distal
interradial plates, not overlapping adjacent plate spinelets if
splayed; superomarginal plates smaller than inferomarginals,
superomarginals with up to about 12 spinelets per plate,
inferomarginals with up to about 24 larger spinelets per plate.
Spines per actinal plate up to: oral 8, suboral 6, furrow 7,
subambulacral 8, actinal interradial 14, predominantly 8-12;
interradial spines short, thick, rugose, sacciform, bluntly conical
(lower spine numbers per plate on South Africa material).
Colour (live). Variable; disc white, apically around disc bright
red, rays mottled mauve, brownish-red, off-white, brown
(photos from G. Paulay and Y. Sarny n).
Distribution. Oman (Masirah I.); Madagascar; La Reunion I.;
E South Africa (Sodwana Bay); 0-18 m.
Etymology. Named for Yves Samyn, of the Royal Belgian
Institute of Natural Sciences, for his contribution of specimens
used in this work and in appreciation of his research on
echinoderm systematics.
Remarks. Pentaradiate, single madreporite, non-fissiparous
specimens as small as R = 6 mm have been examined, evidence
that this species does not have a small pluriradiate fissiparous
growth stage. Large specimens have mostly come from the
deeper sublittoral (3-18 m), while the numerous small
specimens have mostly come from the intertidal and shallow
sublittoral (0-2 m). If this species occurs in deeper sublittoral
habitats this might account for the absence of the species in
collections from most of the eastern African coast. It might
also account for the absence of large specimens from
Madagascar, as the collecting appears to have been littoral.
A. sarnyni is morphologically similar to A. marshae
(above), but is distinguished from A. marshae (characters in
brackets) by: rays long, mostly subdigitiform, only slightly
tapered (not short, mostly strongly tapered); abactinal surface
is predominantly flat (plates not raised proximally and angled
over papulae); spinelets are predominantly spread over plate
surface (not predominantly concentrated over raised proximal
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
283
plate edge); distal plate surfaces with spread spinel ets (not
bare with 1-2 central spinelets); at R = 16 mm, predominantly
6 spines per actinal plate (not 3); actinal spines short, blunt,
sacciform (not conical to digitiform).
For population studies by Soliman (1995, 1999) see
Remarks under A. burtonii above.
Aquilonastra scobinata (Livingstone, 1933)
Figures 1, 3i, 6k
Asterina scobinata Livingstone, 1933: 1-2, pi. 5 figs 9-12, 15.—
H. L. Clark, 1938: 149-1 50. -Cotton and Godfrey, 1942: 201.-H.L.
Clark, 1946: 132,-Dartnall, 1969: 55,-Dartnall, 1970a: 73 , 76.-
Dartnall, 1970b: 19-20, figs 1, 2.— Dartnall, 1980: 8, 34, 66.— Zeidler
and Shepherd, 1982: 412, fig. 10.6e.-O’Loughlin, 1984: 134. -A.M.
Clark, 1993: 213.— Rowe and Gates, 1995: 35.— Waters et al., 2004:
873, 876, figs 1, 2.
Aquilonastra scobinata.— O’Loughlin and Waters, 2004: 11, 13-
14, fig. 1.— Byrne, 2006: 245, 248, tbls 1, 2, fig. 1.
Material examined. Holotype. Tasmania, AM J1241.
Other material. Tas., Port Arthur, 25 Nov 1968, NMV FI 12178
(2) ; Eaglehawk Neck, J9060 (3); Tamar R. mouth. North Head, 28 Oct
1978, FI 12176 (1); King I., 10 Mar 1980, FI 12177 (1); Vic., Inverloch,
28 Mar 1981, F72975 (1); Point Leo, 20 Apr 1935, F58683 (1); Phillip
I. , NMV F72998 (1); Flinders ocean platforms, 7 Sep 1994, FI 12180
(3) ; Cape Otway, 3 Apr 1983, F58682 (2); Killarney, F72997 (10); Port
Fairy, F72985 (2); SA, Port MacDonnell, 8 Jan 1988 FI 12179 (1).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, wide
basally, tapered, narrowly rounded distally, up to R = 18 mm, r
= 8 mm; form frequently asymmetrical with one ray shorter;
actinal gonopores, hermaphrodite.
At R = 14 mm, r = 7 mm, proximal ray with zig-zag carinal
series of singly papulate plates, bordered laterally by subequal
non-papulate plates; singly papulate plates in longitudinal
series along sides of rays; papulate plates not notched, slightly
crescentiform, non-papulate plates round to oval to irregular;
rarely 2 papulae per space; secondary plates rare; disc small,
variably delineated by 5 long radial 5 short interradial plates;
spinelets elongate, thin, pencil-like, splay-pointed sacciform;
up to more than 30 per plate, in splayed tuft or crescentiform
or straight band across plate; superomarginal and
inferomarginal plates subequal, in regular series;
superomarginal plates with up to about 10 pencil-like splay-
pointed spinelets; inferomarginal plates with up to about 20
paxilliform similar spinelets.
Spines per actinal plate up to: oral 7; suboral 6; furrow 6;
subambulacral 6; actinal interradial 7; interradial spines long,
rugose, digitiform, some pointed distally.
Colour (live). Dark brown to greyish-brown to cream, with
some dark flecking (O’Loughlin, 1984).
Distribution. Tas., Bass Strait, Cape Conran (eastern Vic.,
NMV collections) to Port MacDonnell (SA); rocky lower
littoral and shallow sublittoral.
Remarks. A. scobinata is provisionally maintained in
Aquilonastra because of the arrangement of abactinal plates
and papulae, the presence of internal superambulacral plates,
and the arrangement of actinal plates. The paxilliform tufts of
long thin pencil-like abactinal spinelets and actinal spines, and
high numbers of spinelets and spines per plate, are atypical of
Aquilonastra. Morphological similarities of A. scobinata to A.
rosea and A. rowleyi have been noted above.
Dartnall (1970b) reported actinal gonopores and
hermpahrodite reproduction. Mieko Komatsu (pers. comm.)
confirmed that a specimen (FI 12180) was hermaphroditic.
Aquilonastra watersi sp. nov.
Figures 1, 3j, 61, lOe
‘Asterina’ sp. 1.— Rowe and Richmond, 2004: 3287 (part, Trou
Malabar specimens).
‘ Asterina ’ sp. 2. —Rowe and Richmond, 2004: 3287 (part. Riviere
Banane specimen).
Asterina cephea.— Loriol, 1885: 69-71, pi. 21 figs 1-5 (non
Asteriscus cepheus Muller and Troschel, 1842).
Material examined. Holotype. Oman, Masirah I., shore, under rocks,
G. Paulay, 5 Nov 1999, UF 3282 (alcohol).
Paratypes. Type locality and date, UF 70 (5, alcohol); UF 3283 (1,
dry, dissected); Masirah I., reef slope, under rocks, 1-5 m, G. Paulay,
1 Nov 1999, UF 356 (1, alcohol).
Other material. Oman, Bar al Hikman peninsula, under rocks,
0-6 m, Jan 2005, UF 4142 (10); UF 4148 (1); UF 4144 (1); UF 4192 (2);
Red Sea, Egypt, near Qusier, 2002, NMV F106970 (1); Mauritius,
MNHN EcAsll866 (1); ex MAU 74-6, EcAsll867 (1); ex MAU 74-
20, EcAs 11868 (1); ex MAU 74-23, EcAsll869 (1); ex MAU 74-24,
EcAsll870 (1); ex MAU 74-33, EcAsll871 (2); ex MAU 74-36,
EcAsll872 (1); ex MAU 74-37, EcAsll873 (1); ex MAU 74-38,
EcAsll874 (1); ex MAU 74-40, EcAsll875 (1); Mauritius, Robillard,
NHM [18]89.3. 11.7-9 (3); Cape Malheureux, Coin de Mire I., 24 m, 15
Nov 1999, UF 3281 (1); Rodrigues I., Riviere Banane, in algae, 21 Sep
2001, AM J24290 (1); Trou Malabar, coral rubble, 10 m, 22 Sep 2001,
NHM 2004.2813-2814 (2); Madagascar, Nossi-Bi, littoral reef, 2 Dec
1959, EcAsl 1906(1).
Diagnosis. Non-fissiparous Aquilonastra species; rays 5, broad
basally, tapering, rounded distally, up to R = 19 mm, r = 11
mm; gonopores abactinal; few pedicellariae in abactinal
interradii, each with 2 thick curved pointed differentiated
valves, not evident on small specimens.
At R = 16 mm, r = 9 mm, 0-10 proximal doubly papulate
carinal plates, each with up to 4 spinelet clusters each cluster
with up to 7 spinelets; up to about 20 spinelets on proximal
abactinal plates, in clusters or band across proximal edge of
plate; rare small secondary plates; spinelets small, thin, long,
digitiform to predominantly bluntly conical, sometimes
subsacciform, rarely splay-pointed; up to about 8 in tufts on
distal interradial plates, not overlapping adjacent plate spinelets
if splayed; superomarginal plates smallerthaninferomarginals,
superomarginals with up to about 7 spinelets per plate,
inferomarginals with up to about 14 larger spinelets per plate.
Spines per actinal plate up to: oral 7, suboral 6, furrow 6,
subambulacral 6, actinal interradial 8, predominantly 5-6;
interradial spines thick basally, conical, subsacciform.
Colour (live). Mottled pale brown, red-brown, grey-brown,
blue-grey, off-white (photo by G. Paulay).
Distribution. Arabian Sea, Oman; Red Sea, Egypt; W Indian
Ocean, Mauritius, Rodrigues I.; 0-24 m.
284
P. Mark O’Loughlin and Francis W.E. Rowe
Etymology. Named for Jon Waters, of the University of Otago in
New Zealand, in appreciation of his significant contribution to
our understanding of the molecular phylogeny of Asterinidae.
Remarks. Two of the larger specimens from Mauritius (UF
3281, MNHN EcAsll868) have the characteristic pedicellariae
of the types from Oman. For population studies by Soliman
(1999) see Remarks under A. burtonii above. A. watersi is
distinguished by differentiated thick pedicellariae, and small,
thin, long, subacicular spinelets.
Aquilonastra yairi sp. nov.
Figures 1, 4f, lOf
Asterina wega.— Achituv, 1973a: 333-336 (part. Acre and Akhziz
pool populations).
Asterina burton ii.— Smith, 1927a: 641-645 (part).— Tortonese,
1960: 20-21 (probably part).
Asterina burtoni.— Price, 1983: 47^48, fig. 14 (part). —Achituv and
Sher, 1991: 670 (part).— Mladenov and Achituv, 1999: 152 (part).—
Karako et al., 2002: 139-144 (part, Akhziv, Shikmona, and Mikhmoret
Mediterranean populations).— Waters et al., 2004: 874, 876-877, figs 1,
2, tbl. 1 (non Asterina burtonii Gray, 1840; see above).
Asterina burtoni burtoni.— A.M. Clark and Rowe, 1971: 68, fig.
17c (Acre), tbl. 1 (part).-A.M. Clark, 1993: 207-208 (part; non
Asterina burtonii Gray, 1840).
Aquilonastra burtoni.— O’Loughlin and Waters, 2004: 11, 13
(part), 14, fig. 1.— Byrne, 2006: 245, tbls 1, 2, fig. 1 (non Asterina
burtonii Gray, 1840).
Material examined. Holotype. Mediterranean Sea, Israel, Michmoret,
Y. Achituv, 11 Jun 2005, NMV FI 12174 (alcohol). Paratypes. Type
locality and date, F107434 (18, alcohol).
Other material. Israel, Akhziv, 25 Oct 1966, MNHN EcAsll042
(67, dry); Gulf of Suez, A.P. Dollfus, 25 Dec 1928, EcAsll840 (1,
dry); AM J17892 (2, dry); Red Sea, NHM [18]40.3.23.55 (1, dry); don.
J. Waters, F104975 (22, alcohol); F104974 (2, alcohol).
Diagnosis. Fissiparous Aquilonastra species; rays up to 8,
predominantly 6, form frequently asymmetrical post-fissiparity;
form of larger specimens sometimes symmetrical with 5 equal
rays, most interradii with inconspicuous madreporite; rays
narrow basally, tapering, narrow rounded distally, digitiform;
up to R = 7 mm, r = 4 mm; gonopores not seen.
At R = 7 mm, lacking carinal plates; lacking large irregular
proximal abactinal plates; 2 irregular longitudinal series of
papulae on each side of rays; spinelets subgranuliform, short
thick conical to columnar, splay-pointed, cover plates closely,
frequently clustered in groups of up to 8 per group, up to about
16 spinelets on proximal plates, up to about 14 on mid-
interradial plates; superomarginal plates each with up to about
7 spinelets, inferomarginal plates each with up to about 14
taller spinelets.
Spines per actinal plate up to: oral 5, suboral 4 (frequently
3), furrow 4, subambulacral 4, actinal interradial 4
(predominantly 3); interradial spines conical, thin, pointed.
Colour (live). Specimens from Shikmona were all “mottled
brown and orange” (pers. comm. Y. Achituv).
Distribution. Eastern Mediterranean, Acre, Akhziz pool
populations, Michmoret, Shikmona; Red Sea, Gulf of Suez.
Etymology. Named for Yair Achituv (Bar-Ilan University,
Israel), with appreciation of his contribution of material for this
study and his research on the asterinids of the eastern
Mediterranean and Red Sea.
Remarks. At R = 7 mm, A. yairi is distinguished from A.
burtonii by: having more actinal spines per plate (up to four not
two suboral; up to four not two interradial); close cover of
spinelets on abactinal plates, frequently clustered into groups
(not covering plates sparsely); smaller and more regular
abactinal plates; more numerous papulae.
Achituv (1969) studied “pentaradiate and pluriradiate”
forms of an asterinid from Elat in the Gulf of Aqaba, and
referred the pentaradiate form to Asterina burtoni and the
pluriradiate form to Asterina wega. We support his conclusion
that there are two discrete species. But we refer the non-
fissiparous pentaradiate form to Aquilonastra marshae sp. nov.
(above). The fissiparous pluriradiate form had R up to 12 mm,
and is judged here to be Aquilonastra burtonii (Gray, 1840)
and not conspecific with the smaller fissiparous species A.
yairi. We have seen no evidence that A. yairi occurs in the Gulf
of Aqaba. The use of “pentaradiate” to determine non-
fissiparous specimens can be misleading as the larger fissiparous
specimens frequently have five equal rays but continue to have
more than one inconspicuous interradial madreporite.
For evidence from Achituv (1973) that both A. burtonii
and A. yairi occur in the eastern Mediterranean, see Remarks
under A. burtonii above.
Specimens (NMV F104974, F104975) were assumed to be
A. burtoni and reported in the molecular phylogenetic work of
Waters et al. (2004), and represented in the phylogenetic tree
of O’Loughlin and Waters (2004). Both lots are redetermined
in this work as A. yairi. In O’Loughlin and Waters (2004), the
material AM J17892 (2) was assigned to A. burtoni. It is
redetermined here as A. yairi.
The discovery of two type specimens of Asterina burtonii
by Smith (1927a) is discussed above under A. burtonii. The
smaller of these two types is assigned here to A. yairi.
Acknowledgments
We acknowledge with gratitude the generous availability
and loans and donations of an abundant array of asterinid
specimens and photos from the Indo-Pacific, Red Sea and
Mediterranean regions by: Yair Achituv (Bar-Ilan University,
Israel); Leon Altoff and Audrey Falconer (Marine Research
Group, Victoria); Nadia Ameziane and Marc Eleaume
(MNHN); Nechama Ben-Eliahu (HUJ); Prof. Benayhu and
Alex Shlagman (TAU); Penny Berents and Roger Springthorpe
(AM); Maria Byrne (University of Sydney, Australia); Andrew
Cabrinovic (NHM); Neville Coleman (Australia); Chantal
Conand (University of La Reunion); Toshihiko Fujita (NSMT);
Johann Hinterkircher (photographs); Mieko Komatsu (Toyama
University); Loisette Marsh (WAM); Horst Moosleitner
(Zoologisches Institut der Universitat, Salzburg); Gustav
Paulay (UF); Matthew Richmond (Tanzania); Yves Samyn
(Royal Belgian Institute of Natural Sciences); Quaseem Tahera
(University of Karachi, Pakistan); M. Tsurnamal (Israel);
Elizabeth Turner (TM); Sven Uthicke (photographs); Jon
A systematic revision of the asterinid genus Aquilonastra O’Loughlin, 2004 (Echinodermata: Asteroidea).
285
Waters (University of Otago, New Zealand). We are grateful
for the assistance of: Cynthia Ahearn (Smithsonian Institution,
provision of references), Ben Boonen (preparation and format
of figures), Liz Dane, Anna McCallum, Mark Norman, Chris
Rowley, David Staples (NMV, assistance with photographic
images), Ruth Doran (NMV, map), Tim O’Hara (NMV,
molecular phylogeny discussion), Sarah Thompson (NMV,
assistance with data). We are grateful to Alan Dartnall and
Chris Mah for their helpful reviews.
Specimens from Mnazi Bay, southern Tanzania, were collected
during biodiversity survey work for the Mnazi Bay-Ruvuma
Estuary Marine Park, as part of the IUCN coordinated
biodiversity and mapping programme supported by GEF and
the government of France. Specimens from PNG and Mauritius
were collected during field work sponsored by the Coral Reef
Research Foundation under contract to the US National Cancer
Institute contract no. N01-CM-27704. We gratefully
acknowledge that the map incorporates software data that are
© copyright ESRI 2004.
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