Records of the Australian Museum

RECORDS OF THE
AUSTRALIAN MUSEUM

Volume 68 Number 7 7 December 2016

The Tasmanian Mountain Shrimps, Anaspides Thomson, 1894
(Crustacea, Syncarida, Anaspididae)

by

Shane T. Ahyotig

Editorial Board

Dr Shane Ahyong
Dr Don Colgan
Dr Elena Kupriyanova
Dr Andrew Mitchell
Dr Robin Torrence
Editor

Dr Shane McEvey

Journal compilation © 2016 Australian Museum, Sydney

No part of this publication may be reproduced without
permission of the Editor.

Volume 68 Number 7

Published (print and online) 7 December 2016
Price: AU$50.00

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ISSN 0067-1975 (print)

ISSN 2201-4349 (online)

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Cover image —The Tasmanian mountain shrimps of the genus
Anaspides Thomson, 1894 (Syncarida: Anaspididae), are
endemic to Tasmania and often regarded as “living fossils”
owing to the the fact they have changed little since the
Triassic. All live in cold, usually high altitude fresh-waters.
Until recently, only two species were recognized. Australian
Museum carcinologist Dr Shane Ahyong has discovered that
the fauna comprises at least seven species, among which
Anaspides swaini Ahyong, 2015 (cover image and pages
352-361) is frequently encountered in the Weld River, Snowy
Mountains region (Tasmania), Mt Field and Mt Wellington
(North West Bay River catchment) to the Western Arthurs,
throughout the Franklin-Gordon drainages, north to Lake
Rhona and Frenchmans Cap, Mt Rufus and the vicinity of
Lake St Clair; 300-1440 m asl (epigean), 30-1000 m asl
(subterranean).

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© The Author, 2016. Journal compilation © Australian Museum, Sydney, 2016
Records of the Australian Museum (2016) Vol. 68, issue number 7, pp. 313-364.
ISSN 0067-1975 (print), ISSN 2201-4349 (online)
http://dx.doi.Org/10.3853/j.2201-4349.68.2016.1669

The Tasmanian Mountain Shrimps,
Anaspides Thomson, 1894
(Crustacea, Syncarida, Anaspididae)

Shane T. Ahyong

Australian Museum Research Institute,

Australian Museum, 1 William Street, Sydney NSW 2010, Australia, and

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

shane. ahyong@austmus. gov. au

Abstract. The Tasmanian mountain shrimps of the genus Anaspides Thomson, 1894 (Syncarida:
Anaspididae), are endemic to Tasmania and often regarded as “living fossils” owing to the retention
of numerous putatively plesiomorphic eumalacostracan traits and minimal morphological change since
the Triassic. All live in cold, usually high altitude fresh-waters. Until recently, only two species were
recognised: the presumed widespread A. tasmaniae (Thomson, 1893) (type species) and A. spinulae
Williams, 1965a, from Lake St Clair, whose validity was frequently questioned. Independent morphological
and molecular studies revealed previously unrecognized taxonomic diversity, resulting in preliminary
descriptions of three new species in 2015. Anaspides is revised based on extensive collections from
throughout Tasmania. Telson structure and male secondary sexual characters proved taxonomically
instrumental. Seven species are recognized of which two are new to science; all are fully figured and
morphological variation is discussed in detail. Rather than being widespread, Anaspides tasmaniae is
restricted to Mount Wellington; A. spinulae is a valid species known only from Lake St Clair. Two species,

A. clarkei Ahyong, 2015, and A. eherhardi sp. nov. occur only in caves of the Ida Bay-Hastings karst
systems and Junee-Florentine systems, respectively. The three widest ranging species (A.jarmani Ahyong,
2015, A. swaini Ahyong, 2015, and A. richardsoni sp. nov.) are primarily epigean and each contains
several morphological forms that might warrant further taxonomic subdivision. Distributions of species
of Anaspides are largely discrete and broadly correspond to the biogeographical discontinuity known as
Tyler’s Line, dividing the drier eastern from the wetter western parts of Tasmania. Caves are believed to
have acted as oligothermal refuges for Anaspides in the past, and it is notable that the specimens from the
lowest altitudes today are all from caves, including the northernmost record of Anaspides (A. richardsoni ,
Great Western Cave, Gunns Plain, 109 m asl). Given the revised taxonomy of Anaspides , with significantly
altered species distributions, the conservation status of all species of the genus requires review.

Keywords. Crustacea; Anaspidacea; Anaspides ; Tasmania; freshwater; shrimp.

Ahyong, Shane T. 2016. The Tasmanian Mountain Shrimps, Anaspides Thomson, 1894 (Crustacea, Syncarida,
Anaspididae). Records of the Australian Museum 68(7): 313-364. http://dx.doi.Org/10.3853/j.2201-4349.68.2016.1669

314

Records of the Australian Museum (2016) Vol. 68

When G. M. Thomson (1893) described Anaspides tasmaniae
(as Anaspis ) from Mt Wellington, Tasmania, he was not
aware of the significance of his discovery, placing his new
find in the now defunct Schizopoda, alongside mysidacean
and euphausiacean shrimps. It was Caiman (1897), however,
who recognized the affinities of the living Anaspides
with the Palaeozoic fossil syncarids, and would later
recognize a separate eumalacostracan order, Anaspidacea
Caiman, 1904 (Brooks, 1962; Schram, 1984). Anaspides is
remarkable because it retains many putatively plesiomorphic
eumalacostracan traits, such as minimal tagmatisation,
a relatively complete suite of thoracopodal rami, and an
elementary cardioid escape reaction (Schram & Hessler,
1984). As a result, Anaspides has attracted considerable
scientific interest as a potential “living fossil” closely
resembling its Triassic forbears and possibly reflecting the
eumalacostracan stem condition (Coineau & Camacho, 2013).

Until 2015, only two species of Anaspides were recognised,
with A. tasmaniae (Thomson, 1893) (type locality: Mt
Wellington) accorded a wide range throughout most of
central, western and southern Tasmania from both epigean
and subterranean habitats, and A. spinulae Williams, 1965a,
believed to be restricted to central Tasmania from Lake St
Clair (type locality) and immediate environs. Additionally,
Anaspides occurs in subterranean habitats throughout
Tasmania, with some having troglobitic adaptations such as eye
and pigment reduction and unusual telson morphology in which
the posterior telson spines are few, stout and widely spaced,
rather than finely spaced and slender—the “cave type” telson
(Eberhard etal. , 1991; Clarke, 2006). Until recently, taxonomic
questions surrounding Anaspides focussed on whether A.
spinulae was distinct from the presumably widespread A.
tasmaniae , and whether subterranean Anaspides with the
“cave-type” telson constituted a separate species (O’Brien,
1990). Preliminary molecular investigations (Jarman & Elliot,
2000; Andrew, 2005), although not decisive with regards to
previous taxonomic questions, nevertheless identified multiple
divergent lineages within Anaspides. Independent, parallel
morphological and molecular investigations also recognized
significant heterogeneity within the genus, not only among
epigean, but also subterranean forms, leading to preliminary
accounts of three new species of Anaspides (Ahyong, 2015)
in advance of a full taxonomic revision of the genus. The
present study revises Anaspides , restricting A. tasmaniae to
Mt Wellington, recognizes as valid A. spinulae , and five other
species of which two are described as new.

Materials and methods

Measurements of specimens are of total body length,
measured from the apex of the rostrum to the tip of the
telson. Abbreviations: above sea-level (asl); indeterminate
(indet)\ juvenile ( j'm); Mount (Mi); station (stn). Almost 3600
specimens were examined, deposited in the collections of the
Australian Museum, Sydney (AM)-, Haswell Museum, Uni¬
versity of Sydney (HMUS); Museum of Zoology, University
of Sao Paolo, Brazil (MZUSP); Museum of Comparative
Zoology, Harvard University (MCZ); Museum Victoria,
Melbourne (NMV): Otago Museum, Dunedin (OM); Queen
Victoria Museum and Art Gallery, Launceston (QVM)\ South
Australian Museum, Adelaide (SAMA); Tasmanian Museum
and Art Gallery, Hobart (TMAG); Lee Kong Chian Natural
History Museum, National University of Singapore (ZRC);
National Museum of Natural History, Smithsonian Institution,

Washington D.C. (XJSNM% Universidade Federal de Minas
Gerais, Brazil (UFMG); Zoological Collection, Universitat
Rostock (ZRSO); and Yale Peabody Museum, Connecticut
(YPM). Material examined is listed in approximately north-
south and east-west order and grouped according to regions
or localities. In the case of “spinose” A. spinulae-liko forms
ok A. richardsoni , which occur in the Walls of Jerusalem area
and Western Lakes, they are grouped separately or indicated
as such within each entry to distinguish them from the typical
non-spinose forms. Similarly, the three morphological forms
of A. richardsoni from the Mole Creek Karst area are also
grouped for convenience.

Taxonomic characters in Anaspides

Species of Anaspides are morphologically conservative,
exhibiting few differences in major structures. Pleonal
spination, considered diagnostic of A. spinulae by Williams
(1965a), is more variable than previously thought, but when
considered in combination with other characters, remains
taxonomically useful. Several new taxonomic features are
recognized here and given primacy in separating species—
the shape and spination of the telson, and secondary sexual
characters of adult males.

The shape and armature of the posterior margin of the
telson, whether evenly rounded or angular, and whether
armed with stout, well-spaced spines (Figs 8, 12) or lined
with numerous finely-set spines (Figs 21,22), is diagnostic.
These features are taxonomically effective across a wide
size range, noting that the posterior margin of the telson in
the earliest post-hatching juveniles has relatively few spines
and a median cleft or emargination (Figs 33P, 34T), which
rapidly closes with increasing size to form a rounded or
angular spinose margin (Fig. 23B,C).

Two sexually dimorphic features of males have proven
especially useful: the modified inner antennular flagellum and
pleopod 1 endopod. In adult males, the inner (= accessory)
flagellum of the antennule is modified to form a clasping
structure used to hold females, presumably during courtship
(Hickman, 1937). Rather than being straight and flexible as in
females and juvenile males, the proximal portion of the adult
male inner flagellum (articles 1-8) is semi-rigid and double¬
bent into an S-shape, being strongly curled downwardly and
then turning upwards and horizontally in the vicinity of the
sixth and seventh articles (Figs 1, 2D). Each of the proximal
eight articles of the modified inner antennular flagellum has
several short, curled setae on the dorsal and lateral surfaces,
herein called “finger setae”. The proximal 5 articles are as long
as or shorter than wide, articles 6 and 7 are more elongated,
with an obtusely angled inner margin of which the article 7
bears 1-5 slender, prominent, stiff, conical protrusions on the
inner margin, herein called clasping spines (Fig. 2D). The
number of clasping spines is diagnostic and should always
be examined when making identifications. Both left and right
antennules should always be examined in case of damage or
regeneration from damage to these delicate structures. Note
also that the clasping spines usually arise sequentially and
increase in size with successive moults, so juveniles of A.
swaini , for instance, in which only one of the two clasping
spines have appeared, could be mistaken for A. richardsoni ,
which has only one clasping spine in adults. Thus, it is also
important to confirm the maturity of male specimens by
examining the differentiation of the proximal articles of the
inner antennular flagellum and presence of finger setae.

Ahyong: Endemic Tasmanian Mountain Shrimps

315

As in many other malacostracans, the adult male pleopods
1 and 2 are modified as copulatory organs and act together
to facilitate sperm transfer: the petasma. In Anaspides, male
pleopod 1 is broadened and expanded distally, with a small
medial lobe bearing a short row of retinaculae that link the
left and right pleopods to form a functional unit (Fig. 2M).
In lateral view of pleopod 1, whether or not the retinacular
lobe is visible (Fig. 3 I) or obscured (Fig. 71) by the lateral
fold of the pleopod is diagnostic.

In identifying Anaspides , careful consideration should
be given to the sex and maturity of specimens. Adult
males, having a full suite of diagnostic characters, are
best considered first and can be recognized by the fully
modified antennules with well-developed clasping spines,
finger setae and fully curled proximal portion of the inner
antennular flagellum, in addition to the modified pleopod
1-2 endopods. Adult females can be recognized by the well-
developed gonopore on pereonite 8, and the development
of distinctly setose endites on the inner margins of the
pereopod 1-7 coxae, which meet medially. Individuals of
most species are mature by 18-21 mm, but some populations
of A. swaini do not show fully developed secondary sexual
characteristics until 24-33 mm. Several features change
allometrically including the rostrum (becoming more
slender with size), eyes (proportionally larger in smaller
specimens), scaphocerite (more slender in small juveniles),
telson proportions (slightly wider with increasing size),
pleonal setation (shorter with increasing body size), and to
a lesser degree, the proportional width of the uropod rami
(wider with increasing size).

The presence or absence of the pleopodal endopods
in adults is taxonomically important (pleopods 3-5 in A.
darker, pleopod 5 in other species). Pleopodal endopod
development is anamorphic in Anaspides. Endopods first
appear on the anterior pleopods early in development and
progress to the posterior pleopods over successive moults.
In species with a full complement of pleopod endopods in
adults, such as A. richardsoni and A. spinulae, the pleopod
5 endopod may appear at an early or late juvenile stage, and
this may vary between localities.

Systematics

Syncarida Packard, 1885

Anaspidacea Caiman, 1904
Anaspididae Thomson, 1893

Anaspides Thomson, 1894

Anaspis Thomson, 1893: 7 (preoccupied, Anaspis Geoffroy,
1762 [Coleoptera]; type species: Anaspis tasmaniae
Thomson, 1893).

Anaspides Thomson, 1894: 285,286 (replacement name for
Anaspis Thomson, 1893, preoccupied).

Diagnosis. Rostrum prominent, well-developed. Cephalo-
thorax without fenestra dorsalis. Body subcylindrical,
straight or evenly curved, without prominent “bump-like”
flexure at pleonite 1. Free pereonites length subequal,
shorter than pleonites. Pleonite 6 shorter than twice length
of pleonite 5; surface of integument with few setae but no
spines. Telson dorsoventrally compressed; posterior margin
rounded or angular, with spine row. Antennular and antennal

peduncles unarmed. Scaphocerite with small lateral spine.
Thoracopod 1 (maxilliped) with epipods. Thoracopod 7 with
exopod. Uropodal endopod more than three-fourths length
of exopod; exopod with small group of movable spines near
position of diaraesis.

Description. Body subcylindrical, straight or evenly curved,
without prominent “bump-like” flexure at pleonite 1.
Rostrum prominent, apex blunt, rounded, slightly deflected
ventrally; few distal setae, arising submarginally. Head
(cephalothorax) comprised of fused cephalon and pereonite
1; cervical groove distinct; dorsal organ present on dorsal
midline anterior to cervical groove; mid-lateral surface
posterior to cervical groove with shallow diagonal groove.
Pereonites 2-8 length subequal, subparallel, shorter than
pleonites. Pleonites 1-5 length subequal; subparallel; pleura
well-developed, rounded. Pleonite 6 shorter than twice
length of pleonite 5; surface of integument with few setae
but no spines. Telson dorsoventrally compressed, with low,
broad median ridge; posterior margin rounded or angular,
with spine row; surface with low, broad median crest, few
scattered setae. Female gonopore (spermatheca) on pereonite
8 sternum between coxae; bulbous, directed anteriorly,
anterior surface with genital orifice as narrow transverse slit.
Pleonal sternites 3-5 with low median processes between
pleopod bases.

Eyes pedunculate; cornea usually well-developed.

Antennular peduncle 3-articulate, unarmed; article 1
with statocyst; biflagellate, inner (= accessory) flagellum
shorter than outer; adult males with proximal portion of
inner flagellum modified to form clasping structure, with
proximal 8 articles bearing dorsal finger setae and together
semi-rigid, S-shaped, curled, strongly curved downward,
then turning upwards and horizontally in vicinity of articles
5 and 6, with slender clasping spines on obtusely angled
inner margin of article 7.

Antenna uniflagellate; protopod 3-articulate, unarmed;
exopod (scaphocerite) laminar, broadly ovate, small lateral
spine, mesial and distal margin setose to base of lateral spine;
endopod peduncle 2-articlulate, unarmed.

Labrum with shallow proximal constriction, anterior
proximal surface with blunt median ridge; distal margin
convex to slightly concave, finely setose.

Mandibular corpus (apophysis) robust; molar process and
incisor process well-developed; molar with elongate, ovate,
triturating surface, surrounded by spiniform setae; incisor
process diagonal to axis of mandibular corpus; left incisor
process with 7 triangular teeth in sinuous row, proximally
with spine row between proximal incisor tooth and molar
process; right incisor process similar to left except with 6
triangular teeth, proximal tooth usually apically divided;
palp 3-articulate, setose, article 1 short, subquadrate, article
2 slender, almost twice length of article 3.

Paragnaths widely separated by deep V-shaped incision,
without lobes, distal half finely setose, especially mesially.

Maxillule with 2 endites; proximal endite distally setose;
distal endite spinose distally, lateral surface with small
rounded palp.

Maxilla with 4 endites, proximal 2 endites with plumose
setae, distal 2 endites densely arrayed with serrulate setae.

Thoracopods 1-8 protopod with coxa, basis, preischium,
ischium, merus, carpus, propodus and dactylus; flexure at
carpus-merus articulation.

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Records of the Australian Museum (2016) Vol. 68

Thoracopod 1 (maxilliped) coxa inner margin with setose
coxal endites, outer margin with 2 slender, lamellar epipods,
proximal wider than distal; basis with slender, flattened,
strap-like exopod; coxa-basis demarcation often ill-defined;
preischium rectangular, more than twice length of quadrate
ischium; merus as long as preischium, slightly tapering
distally; carpus triangular, longer than high, half length
of merus; propodus slender, slightly shorter than merus;
dactylus short, terminating in slender claw, with 2 slender
movable spines on lateral side, 3 on mesial side.

Thoracopods 2-8 (pereopods) as ambulatory legs.
Thoracopods 2-6 structurally similar, distal 4 articles with
tufts of setae, primarily along flexor margins, dactylus strongly
setose; thoracopods 4-5 longest; coxa outer margin with 2
ovate, lamelliform epipods, inner margin in adult females
with setose endite; basis short, partially fused with preischium;
exopod articulating with outer margin of basis, with elongate
basal article and setose multi-annulate flagellum (< 30);
ischium about as long as basis-preischium; merus elongate,
slightly tapering distally, about twice length of ischium; carpus
triangular, longer than high, about half length of merus or
slightly less; propodus elongate, slender, shorter than merus;
dactylus short, terminating in long, slender claw, with slender
movable spine on lateral side, 2 movable spines on mesial
side. Thoracopod 7 similar to thoracopods 2-6 except epipods
proportionally more slender; exopod a single narrow lamella.
Thoracopod 8 structurally similar to preceding thoracopods
but lacking epipods or exopod; basis and preischium
indistinguishably fused; longer than thoracopod 7.

Pleopods 1-5 exopod long, slender, setose, 25-30-
annulate. Pleopods 1-2 endopod always present, unmodified
endopod ovate, lamellar, length subequal to first exopod
article, variously present on pleopods 3-5 in females and
juvenile males; adult male pleopods 1-2 endopod modified
as copulatory structures (petasma). Adult male pleopod 1
elongate, directed anteriorly, reaching beyond thoracopod
8 coxa; slender proximally, expanded distally forming
cannulate, scoop-like structure, hollowed mesially; distally
rounded, lateral margin thin, lamellate; inner margin with
short row of retinaculae near distal one-third, forming small
rounded lobe; inner proximal surface with scattered setae and
spinules. Male pleopod 2 endopod of 2 articles, longer than
that of pleopod 1, directed anteriorly, reaching to thoracopod
8 coxa; proximal article twice length of distal article, mesial
proximal margin with row of retinaculae; distal article
broadly curved, mesially hollowed, apex blunt.

Telson and uropods forming tail-fan. Uropodal exopod
lateral margin with indistinct, partial diaeresis near distal
one-third; movable spines near position of diaeresis, flanked
by tufts of setae; inner margin and outer margin distal to
diaeresis setose. Uropodal endopod slightly shorter than
exopod, margins setose.

Species composition. Anaspides clarkei Ahyong, 2015, A.
eberhardi sp. nov., A. jarmani Ahyong, 2015 ,A. richardsoni
sp. nov., A. spinulae Williams, 1965a, A. swaini Ahyong,
2015,^4. tasmaniae (Thomson, 1893) (type species).

Remarks. Anaspides is one of three anaspidid genera endemic
to Tasmania, others being Paranaspides Smith, 1908 (with
only the type species, Paranaspides lacnstris Smith, 1908),
and Allanaspides Swain, Wilson, Hickman & Ong, 1970 (with
the type species A. helonomus Swain, Wilson, Hickman &
Ong, 1970, and A. hickmani Swain, Wilson & Ong, 1971)

(Lake et al., 2002). Allanaspides, which uniquely has a
fenestra dorsalis on the cephalothorax, lacks maxillipedal
epipods and lacks the exopod on thoracopod 7, is believed
to be sister to other anaspidids (Jarman & Elliott, 2000).
Paranaspides is most closely related to Anaspides , being
sister to, or possibly nested within, the latter (Jarman &
Elliott, 2000). Few major features separate the two genera,
however. Paranaspides , with a pelagic rather than benthic
habit, is considerably more spinose than Anaspides but the
most important distinguishing feature is the distinct pleonal
flexure of the former as a result of the wedge-shaped pleonite 1.

Phylogenetic relationships inferred from mitochondrial
16S sequences from selected Anaspides populations (Jarman
& Elliott, 2000; Andrew, 2005), although with low resolution,
recognized three broad clades: a southern group corresponding
to A. jarmani and A. clarkei, which diverged from other
Anaspides approximately 25 ma; a southwestern group (A.
swaini ); and a northern-eastern group corresponding here to
A. tasmaniae, A. spinulae, A.richardsoni and A. eberhardi).

The phylogenetic significance of Anaspides for Malacos-
traca has prompted numerous morphological, ultrastructural
and physiological studies, all under the name A. tasmaniae.
Those applicable to A. tasmaniae sensu stricto are as follows:
embryology (Hickman, 1937); giant lateral neurone (Silvey
& Wilson, 1979); organ of Bellonci (Kauri & Lake, 1972);
locomotory function (Macmillan et al, 1981); spermogenesis
(Jespersen, 1983); foregut morphology (Wallis & Macmillan,
1998); and ommatidial structure (Richter, 1999); and cuticular
sclerites (Kutschera et al, 2015). Manton’s (1930) study of
habits and feeding are based on A. tasmaniae and A. swaini.
Studies of functional morphology and excretion (Cannon &
Manton, 1927; Manton, 1929,1931) are probably based on^4.
swaini given the photograph by Sidnie Manton, presented by
William Caiman to the Royal Zoological Society of London
depicting what appears to be A. swaini (MacBride, 1930).
Analyses of ecology and life history (Swain & Reid, 1983)
and mandibular morphology (Richter et al., 2002) are based
on A. richardsoni from Mt Field. Serov (1988) examined
ecology of populations from Browns River near Silver Falls,
which are thus referable to A. tasmaniae. Specimens studied
by Tjonneland et al. (1984) for heart ultrastructure, from
Myrtle Forest Creek, are referrable to A. swaini. Smith’s (1908,
1909b) studies of general morphology may be based several
species given that he accessed material from Mt Wellington
(North West Bay River), Mt Field and the Hartz Mountains,
localities at which A. swaini, A. richardsoni and A. jarmani
occur, respectively. Internal anatomy (gonads and alimentary
canal) reported by Nicholls & Spargo (1932) is probably
based on A. richardsoni given that Nicholls collected widely
in the Great Lake area, many specimens of which are still
extant in the collections of the Western Australian Museum
and Tasmanian Museum. Specimens of Anaspides collected
by Smith were the source material for a parasitic protozoan,
Ganymedes anaspidis described by Huxley (1910). Given the
uncertainty over the identity of Smith’s Anaspides specimens,
however, the host species of the type material of G. anaspidis
likewise remains unclear.

Anaspides tasmaniae was originally described as the type
species of Anaspis Thomson, 1893. Anaspis Thomson, 1893,
however, being preoccupied by Anaspis Geoffroy, 1762
(Coleoptera), was replaced by Anaspides Thomson, 1894.
Seven species of Anaspides are recognized here of which
two are new to science.

Ahyong: Endemic Tasmanian Mountain Shrimps

317

1

2

3

4

5

6

Key to species of Anaspides

Telson posterior margin with 4-10 (rarely 15), stout, well-spaced

spines. 2

Telson posterior margin lined with more than 17 (usually > 20),

slender, finely-spaced spines . 3

Telson about as long as wide, posterior marginal spines limited
to posterior one-fourth. Pleonite 6 posterior margin and pleura
4-5 unarmed. Adult male with 4 (rarely 5) antennular clasping

spines. Pleopods 4-5 (usually 3-5) without endopod. A. clarkei Ahyong, 2015

Telson distinctly longer than wide, posterior marginal spines
limited to posterior one-third. Pleonite 6 posterior margin
denticulate; pleura usually with 1 or more small spines.

Adult male with 1 antennular clasping spine. Pleopods

3-4 with endopod . A. eberhardi sp. nov.

Adult males with 1 antennular clasping spine. A. richardsoni sp. nov.

Adult males with 2 or more antennular clasping spines. 5

Telson linguiform, elongate, lateral margins seamlessly grading

into evenly rounded, posterior margin . A. tasmaniae (Thomson, 1893)

Telson polygonal, transition between lateral and posterior
margins obtusely angular, blunt; posterior margin angular to

rounded. 5

Adult males with 2 antennular clasping spines. Male pleopod 1

endopod with retinacular lobe visible in lateral view. 6

Adult males with 3-5 (usually 4) antennular clasping spines.

Male pleopod 1 endopod with retinacular lobe obscured, not

visible in lateral view . A.jarmani Ahyong, 2015

Pleonite 6 posterolateral margins blunt, rounded, at most with
a minute spinule. Pleonites 5-6 with or without short spines
on posterior tergal margins, usually absent on tergite 5; pleura
3-5 with or without small spines, usually absent or at most 1 or
2 small spines on pleura 4-5 .

Pleonite 6 posterolateral margins produced to prominent
spine. Pleonites 5-6 with prominent spination on posterior
tergal margins, spines distinctly longer than wide; pleura 3-5
prominently spinose .

A. swaini Ahyong, 2015

A. spinulae Williams, 1965a

Anaspides tasmaniae (Thomson, 1893)

Figs 1-4, 35A, 36

Anaspis tasmaniae Thomson, 1893: 7-10 (type locality: near
The Springs, Mt Wellington).

Anaspides tasmaniae. —Caiman, 1897: 787-794, pi. 1, 2,
fig. 12-14.—Thomson, 1897: 580; 1926: 161.—Manton,
1930: pi. 4. —Hickman, 1937: 2, tab. 1-3, pi. 1-13.
—Hewer, 1967: 1-2. —Kauri & Lake, 1972: 432, figs.
1-17.—Williams, 1974: 80, fig. 4.6, tab. 4.1.— Silvey &
Wilson, 1979: 122. —Jarman & Elliot, 2000: 624, tab. 1
(Mt Wellington). —Jarman, 2001: 201, tab. 1. —Jarman
etal. 2000: 27, tab. 1 —Lake etal., 2002: 11-12. —Serov,
2002: 8, 15. —Camacho, 2006: 4.

Type material. Lectotype: AM G2130, male (23 mm), Mt Wellington, “4000
ft”, per G.M. Thomson. Par ate ctotypes: AM P99315, 1 female (24 mm), 1 juv.

(19 mm), collected with lectotype; OM Iv. 1396, 1 $ (22 mm), Mt Wellington, “4000
ft” [1200 m], coll. G.M. Thomson.

Other material examined, tmag 14370/G114, (14-26 mm), 19

(29 mm). New Town Rivulet, Mt Wellington, 42°52.5'S 147°15.8'E, 1000 ft asl [300
m], coll. J. Pearson, 20 Jun 1937; QVM 10:8079, 1<J (damaged, c. 27 mm), 2??
(26-27 mm), New Town Creek, Mt Wellington, 42°52'S 147°16'E, coll. E. Guiler,

1956; TMAG G6433, 2$$ (22-24 mm), 6§ $ (22-26 mm), Lenah Valley, Newtown
Rivulet, 42°51.6'S 147°16.9'E, 150 m asl, coll. R. Swain, Jul 1969; NMV J42438,1$
(28 mm), 2$$ (21-25 mm), 2 juv. §? (13-14 mm), Organ Pipes, Mt Wellington,
42°53.8'S 147°14.5'S coll. 12 May 1912, pres J. Searle, Feb 1936; TMAG G6383,
59$ (27-32 mm), Picnic Hut, Mt Wellington, 42°53.9'S 147°14.2'E, 1250 m asl,
coll. R. Swain, 4 Nov 1969; TMAG G6404, 2$$ (22-24 mm), 10 (22-27

mm), Picnic Hut, Mt Wellington, 42°53.9'S 147°14.2'E, 1250 m asl, coll. R. Swain,
Apr 1969; TMAG G6365, 2$$ (21-24 mm), 7£$ (24-30 mm), Picnic Hut, Mt
Wellington, 42°53.9'S 147°14.2'E, 1250 m asl, coll. R. Swain, 14 Sep 1969; AM
P14157, 2 specimens (slide preparations), Picnic Point, Mt Wellington, 42°54.9'S
147°14.7'E, stream, 800 m asl, coll. W.D. Williams 29 Jan 1963; AMP99314, (28

mm), The Chalet, Mt Wellington, 42°53.43'S 147°14.04'E, stream, 970 m asl, coll. S.
Jarman, Nov 1997; TMAG G6414, (27-33 mm), 7$? (27-35 mm), Fem Tree,

Mt Wellington, 42°55.5'S 147°15.6'E, 420 m asl, coll. R. Swain, 14 Feb 1971; AM
P97847, 1 juv. $ (12 mm, Silver Falls, Mt Wellington, 42°55.3'S 147°14.9'E, 1500
ft asl [450 m], 28 Feb 1935; TMAG G6431,(26 mm), 3 juv. $$ (19-21 mm),
59$ (19-28 mm), Browns River, above Silver Falls, 42°55.1'S 147°14.7'E, 620 m
asl, coll. I. Wilson & B. Knott, 20 Jan 1971; AM P98089, l<$ (24 mm), 4 juv. SS
(14—16 mm), 2$$ (23-28 mm), 4$$ (10-16 mm), Mt Wellington, creeks, coll. R.
Swain & A. Richardson, Jul 1990; MZUSP 33665,1 juv. $ (16 mm), Mt Wellington,
coll. R. Swain & A. Richardson, Jul 1990; UFMG, 1 $ (19 mm), Mt Wellington, coll.
R. Swain & A. Richardson, Jul 1990; USNM 60111, lA (25 mm), 1 juv. c? (15 mm),
1$ (23 mm), Silver Falls, Mt Wellington, coll. W.M. Tattersall, 1914; HMUS Cr(M)
II/U/1—l(i), \ (14-24 mm), 16?$ (14-28 mm), from creeks on Mt Wellington,

26 Aug 1965; USNM 291481, \$ (24 mm), Mt Wellington, coll. F.R. Schram, 25
May 1980; NMV J42440,1$ (28 mm), Mt Wellington, 4000 ft asl [1200 m], coll. A.
Neboiss, 22 Feb 1967; TMAG, 5$S (24-26 mm), no data; USNM 30578,2 juv. <$<$
(13-14 mm), 1$ (22 mm), 2 juv. ?? (13-14 mm), from G.M. Thomson, no data.

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Records of the Australian Museum (2016) Vol. 68

Description. Eyes with well-developed cornea, pigmented,
subglobular, longer than half length and slightly wider than
stalk, stalk with subparallel margins. Rostrum narrow in
adults, apex blunt.

Pleonites with sparsely setose pleural margins, rounded;
pleuron 1 unarmed; pleura 2-3 with 0-2 small spines;
pleuron 4 with 1-4 small spines and scattered setae. Pleonite
5 pleuron with 1-4 spines and scattered setae; posterior
tergal margin with 3-5 spines either side of midline, setose.
Pleonite 6 posterior margin spinose, setose; posterolateral
margin setose, rounded. Pleonal sternites 3-5 with low,
weakly bilobed median processes between pleopod bases,
widest on sternite 3, narrowest on sternite 5.

Telson longer than wide, linguiform, widest proximally;
lateral margins sinuous in dorsal outline, distally convergent;
transition from lateral to rounded posterior margin evenly
curved, seamless; posterior spine row with 19-37 short,
evenly graded, slender, closely spaced spines, generally
longest medially.

Antennule inner flagellum about 0.2 x body length (19-20
articles in figured 28 mm male); article 7 inner margin
obtusely angled in adult males, with 2 long, slender clasping
spines proximally; outer flagellum 0.4-0.5 x body length
(69 articles in figured 28 mm male). Antennal flagellum
0.3-0.4 x body length (47 articles in figured 28 mm male);
scaphocerite elongate, ovate, lateral spine slightly distal
to midlength; apex reaching almost to midlength of distal
peduncular article.

Right mandibular incisor process with proximal tooth
distally undivided to trifurcate, usually bifid.

Pleopods 1-4 with endopod in adults (rarely on one
side on pleopod 5). Adult male pleopod 1 distally widened,
scoop-like, lateral margins weakly expanded, not obscuring
retinacular lobe in lateral view.

Uropodal protopod dorsally unarmed or with 1 or 2 small
spines; exopod with 2^1 movable spines on outer margin near
position of partial diaeresis; exopod length about 2.5-3 times
width, as wide as endopod, apex rounded, relatively broad.

Measurements. Male (n = 55) 14-33 mm; female (n = 96)
10-35 mm.

Remarks. Anaspides tasmaniae is readily distinguished
from other species of the genus by the combination of well-
developed eyes, the presence of two antennular clasping
spines in adult males (Fig. 2D), the elongated telson with
an evenly rounded posterior margin lined with closely
set spinules (Fig. 2B), and a male pleopod 1 in which the
retinacular lobe is visible in lateral view (Fig. 31). Anaspides
tasmaniae shares the presence of two male antennular
clasping spines with A. swaini and A. spinulae , but differs
by the rounded versus triangular posterior margin of the
telson. Although some specimens of A. swaini may also
have a slightly rounded posterior margin of the telson (Fig.
33L), the transition between the lateral and posterior margins,
marked by the beginning of the spine row, is seamless in A.
tasmaniae rather than bluntly and obtusely angular.

Morphological variation in A. tasmaniae is not marked. The
pleonite 4-5 pleura (sometimes also 2-3) are multidenticulate,
with 1-4 (usually 2 or 3) small pleural spines on pleonites 4
and 5 (Fig. 4). The posterior tergal margin of pleonite 5 is
spinose (usually 2-4 small spines on either side of the midline)
and that of pleonite 6 is spinose along the entire posterior
margin. The extent of pleonal spination of A. tasmaniae ,
like A. swaini , may approach that of A. spinulae. In A.
spinulae , however, the pleonal spines are always considerably
more prominent, even in juveniles (Fig. 28B,E), and the
posterolateral angle of somite 6 is produced to a prominent
spine (Fig. 25C), rather than forming a blunt lobe (Fig.

Ahyong: Endemic Tasmanian Mountain Shrimps

319

Figure 2. Anaspides tasmaniae (Thomson, 1893), male, 28 mm, Fern Tree, TMAG 6414. (A) cephalothorax, dorsal view; (B) pleonite 6,
telson and right uropod; (C) pleonites 3-6 pleura, right lateral view; (D) right antennule; (E) right antenna; (F) labrum, anterior view; (G)
right mandible; (H) right mandible incisor process; (I) left mandible incisor process; (J) paragnaths, anterior view; (K) right maxillule;
(L) right maxilla; (M) pleopods 1-2, in-situ, ventral view. Scale: A-E, M - 1.0 mm; F-E = 0.7 mm.

2B,C). The proximal incisor tooth on the right mandible of
A. tasmaniae is usually distally bifid or trifid (occasionally
undivided) (Fig. 2H). As in other congeners, the rostrum of A
tasmaniae is proportionally broadest in juveniles, becoming
proportionally narrower with increasing body size, and

the proportional length of setae on the pleonal pleura and
posterior margins decreases with increasing body size. The
pleopod 5 endopod is absent in all specimens except for a
male from Lenah Valley (24 mm; TMAG G6433) in which
an endopod is present on the right side. Both sexes exhibit

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Records of the Australian Museum (2016) Vol. 68

Figure 3. Anaspides tasmaniae (Thomson, 1893), Fern Tree, TMAG 6414, male 28 mm (A-N), female 30 mm (O-P). (A) right thoracopod
1 (maxilliped); (B-H) right thoracopods 2-8; (I) right pleopod 1 endopod, lateral view (retinacular lobe indicated by arrow); (J) right
pleopod 2 endopod, lateral view; (K) right pleopod 4, anterior view; (L-N) pleonites 3-5 median sternal processes; (O-P) female gonopore,
right lateral and ventral views. Scale A-H = 1.0 mm; I-P = 0.7 mm.

full secondary sexual characters by 20-21 mm body length.

Anaspides tasmaniae was previously accorded a wide
distribution throughout Tasmania, but is here restricted to
southeastern Tasmania from surface localities on the eastern

and southeastern face of Mt Wellington at altitudes of
150-1250 m above sea-level; it is not yet known from caves.
Anaspides from elsewhere in Tasmania are referrable to other
species (Ahyong, 2015). The locality data accompanying a

Ahyong: Endemic Tasmanian Mountain Shrimps

321

Figure 4 . Anaspides tasmaniae (Thomson, 1893). (A-D) lectotype male, 23 mm, AMG2130, Mt Wellington. (E) juv. male, 20 mm, Silver
Falls, Browns River, TMAG G6431. (F-J) female, 28 mm. Silver Falls, Browns River, TMAG G6431. (K) female, 27 mm, Silver Falls,
Browns River, TMAG G6431. (L-O) female, 32 mm, Picnic Hut, Mt Wellington, TMAG G6383. (P-R) female, 23 mm, Mt Wellington,
AM P98089. A, G, M, P, telson. B, E, H, K, Q, N, right pleonal pleura. C, right antennular clasping spines. D, right pleopod 1 endopod,
right lateral view. F, L, anterior cephalothorax. I, R, O, right scaphocerite. J, right uropodal endopod. Scale 1.0 mm.

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Records of the Australian Museum (2016) Vol. 68

number of specimens of A tasmaniae sensu stricto from Mt
Wellington do not indicate the drainage or precise location,
but those with more specific locality data are all from eastern
drainages on Mt Wellington between the catchments of
New Town Rivulet and Browns River. Previous records of
A. tasmaniae from northern and western localities around
Mt Wellington, such as Myrtle Forest (Collinsvale) and the
catchment of the Northwest Bay River (O’Brien, 1990) are
referrable to A. swaini , having an angular rather than evenly
rounded posterior margin of the telson and minimal pleonal
spination (at most with scattered denticles on pleonite 6 and
occasionally one or two denticles on the pleonite 5 pleuron)
(Fig. 33 A-J). A series from the Huonville area slightly west
of Mt Wellington (presumably from the catchment of the
Mountain River) is also referable to A. swaini. Specimens
corresponding to A. swaini , but labelled only as coming
from Mt Wellington, were presumably collected from a
westerly locality in the North West Bay River catchment.
Two anomalous lots with specific locality data, however,
were collected in 1928 from Fem Tree Glen and Wishing
Well on the southeastern face of Mt Wellington (AM P9217,
9218). The location of Wishing Well is well known but “Fem
Tree Glen” is probably an error for Fern Tree Bower on
the lower Browns River, well downstream of Silver Falls.
Either way, the presence of A. swaini in the Browns River
catchment, an eastern locale, is unexpected. Other specimens
from the Browns River catchment (Silver Falls and Fern
Tree) represent A. tasmaniae sensu stricto but those from
“Fern Tree Glen” and Wishing Well agree with those from
St Crispins Well (Fig. 33A-F), in the catchment of the North
West Bay River. The North West Bay and Browns rivers,
however, do not share catchments, making natural dispersal
between these drainages unlikely. Fern Tree Bower and
Wishing Well, however, are notable as two key points along
the aqueduct that drew water from the western side of Mt
Wellington (starting at St Crispins Well) to Hobart since at
least 1881. As a result, the records of A. swaini in the lower
Browns River from 1928 could be the result of accidental
translocation from the western side of Mt Wellington; it
remains to be determined whether A. swaini is still present
there. The western and northern Mt Wellington specimens
here referred to A. swaini differ slightly from topotypic
material (see Remarks under account of A. swaini).

Anaspides was previously recorded from the plateau near
the summit of Mt Wellington in tarns and creeks, most of
which drain into the Northwest Bay River (Manton, 1930).
It apparently no longer occurs there, however, probably as
a result of major bushfires that swept across the top of Mt
Wellington in 1930s (Swain pers. com. in O’Brien, 1990).
Anaspides populations are now restricted to catchment creeks
to the periphery of the summit— A. tasmaniae to the east and
south-east, and A swaini to the west and northwest. Material
from the uppermost parts of Mt Wellington collected during
or before the early decades of the 20th century includes
both A. swaini and A tasmaniae , so it is possible they were
sympatric or lived in close proximity atop Mt Wellington
prior to the 1930s bushfires. Manton (1930) observed two
colour forms in Anaspides on Mt Wellington: a dark brown
to olive-green form found on the Mt Wellington plateau, and
a light brown form occurring on the slopes. Notwithstanding
likely habitat related colour variation, the distribution of
Anaspides on Mt Wellington as determined herein suggests
that Manton’s dark form is referrable to A. swaini , and the

light brown form, to A. tasmaniae (Manton, 1930: pi. 2-3,
4). Nicholls’ (1947) remark that the “dark coloration prevails
almost everywhere the species is taken (many quite remote
from Mt Wellington)” is consistent with the much wider
range of A. swaini relative to A. tasmaniae.

The type series of A. tasmaniae was collected from Mt
Wellington on two occasions in 1892. Specimens were first
collected in January 1892 by G.M. Thomson during a visit to
Hobart for the Congress of the Australasian Association for
the Advancement of Science (Morton, 1893; Thomson, 1926)
and by Leonard Rodway (Tasmanian Government botanist)
on 24 May 1892 from the original locality (Thomson, 1893,
1894). Thomson’s (1893, 1894) early accounts suggest
that the specimens were collected near the summit of Mt
Wellington at an elevation of “over 4,000 ft” (= 1200 m).
Thomson’s (1926: 161) more detailed account of events,
however, indicates the specimens were collected “about
three-fourths of the way to the summit, where the water
issues from a few deep pools among the rocks” near “The
Springs”. Thus, rather than the summit, the type series would
have originated from the eastern face of Mt Wellington,
nearer to 700 m asl. These specimens of A. tasmaniae
are now in poor condition, having at one time been dried,
but clearly preserve the primary diagnostic features —an
elongated telson with rounded posterior margin, two male
antennular clasping spines, spinose pleural margins of
pleonites 2-5, spinose tergal margins of pleonites 5-6, and
absence of the pleopod 5 endopod. The male (Fig. 4A-D) is
herein designated as the lectotype to fix the identity of the
species. Other type specimens become paralectotypes. The
female paralectotype in the Otago Museum (OM Iv.1396)
was collected by Thomson, corresponding to a January 1892
collection date.

Being confined to catchments on the eastern face of Mt
Wellington, Anaspides tasmaniae as currently understood
has a considerably narrower distribution than previously
thought. Although A. tasmaniae resides within the relative
protection of the Wellington Park reserve its conservation
status requires reassessment along with detailed delimitation
surveys to precisely determine its current distribution.

Distribution. Eastern Mt Wellington, from the catchments of
the Newtown Rivulet to the Browns River; 150-1250 m asl.

Anaspides clarkei Ahyong, 2015

Figs 5-8, 35B-C, 36

Anaspides tasmaniae. —Lake & Coleman, 1977: 12-13,
pi. 2. —Jarman & Elliot, 2000: fig. 4 (Wolf Hole), tab. 1
(part, Wolf Hole). —Clarke, 2000: 30; 2006, fig. 5.13-15.

—Eberhard, 2001: 97. —Boulton etal ., 2003: 48.

Anaspides sp. (telson ‘cave’ type). —Eberhard et ah, 1991:

48 (Hastings and Ida Bay systems only).

Anaspides. —Gooderham & Tsyrlin, 2002: 73, unnumbered
colour figure (Exit Cave).

Anaspides clarkei Ahyong, 2015: 596-597, fig. 1A-D(type
locality: Exit Cave, Ida Bay).

Type material. Holotype: SAMAC6301,c? (29 mm), Exit Cave, Ida Bay, 70 m asl,
BS1848, coll. E. Hamilton Smith, 24 May 1969. Paratypes: SAMAC6302, 1$ (32
mm), Exit Cave, Ida Bay, 70 m asl, BS1848, coll. E. Hamilton Smith, 24 May 1969;
AM P73045, 1 juv. £ (18 mm). Base Camp Tributary, Exit Cave, Ida Bay, 43°28.2'S
146°51'E, coll. S. Gersbach (#64631); TMAGG6033,1 ? (38 mm), Exit Cave (IB-120),
Ida Bay Karst, rock pool in Skeleton Creek, coll. A. Clarke, 20 Jan 1998.

Ahyong: Endemic Tasmanian Mountain Shrimps

323

Other material examined. Hastings Karsf. AM P73047, 1<J (28 mm),
far end of Lake Pluto, Wolf Hole, above mud, 43°23.3'S 146°51.4'E, 998-23, coll. A.
Clarke, 20 Sep 1998; AM P73046, 1? (31 mm), Lake Pluto, Wolf Hole, 43°23.3'S
146°51.4'E, above muddy substrate, 200 m asl, 998-22, coll. A. Clarke, 20 Sep 1998;
QVM: 10:47694, 1$ (28 mm), Lake Pluto, Wolf Hole, 43°23.3'S 146°51.4'E, above
muddy substrate, 200 m asl, 998-22, coll. A. Clarke, 20 Sep 1998; AM P99289, \$
(22 mm), 1$ (23 mm), 1 juvenile? (12 mm), Newdegate Cave (H-X7), Hastings
Karst, 43°23.0'S 146°50.5'E, from pools in Mystery Creek streamway beyond
Binney Chamber, 1298-02, coll. A. Clarke, 1 Dec 1998; AMP99290,1 indeterminate
juvenile (shrivelled, poor condition, >8 mm), Hell’s Half Acre streamway, 1.6 km into
Newdegate Cave (H-X7), Hastings Karst, 43°23.0'S 146°50.5E, from pools in Mystery
Creek streamway beyond Binney Chamber, 1298-07, coll. A. Clarke, 1 Dec 1998.

Ida Bay Karst QVM 10:49168,3? ? (21-24 mm), Eastern Passage streamway of Little
Grunt Cave and Exit Cave, 43°28.3'S 146°51.7'E, coll. S. Eberhard, Aug 1993; QVM
10:13261,3?? (c. 19-31 mm). Little Grunt Cave (IB29), 43°28.3’S 146°5L7'E, coll.
S. Eberhard, 17 Feb 1992; QVM 10:13254, 1 ? (19 mm), 1 juvenile? (14 mm). Exit
Cave, Eastern Passage, 43°28.3'S 146°51.3'E, coll. S. Eberhard, 15 Feb 1992; QVM
10:12248,3 juvenile c?c? (9-19 mm), Loons Cave (IB2-7), Ida Bay Karst, 43°27.4'S
146°52.1'E, deep stream, 80 m asl, coll. S. M. Eberhard & J. Jackson, 10 May 1989;
QVM 10:12177, 2?? (23-25 mm), 1 indet juv. (10 mm), Milkrun Cave (IB38-5),
43°28.3'S 146°51.3'E, 360 m asl, about 200 m from entrance, coll. S. Eberhard, 22
August 1985; TMAGG6487,1? (24 mm), Arthurs Folly (IB10),43°27.3'S 146°52.3'E,
stream, dark zone, 690-05, coll. A. Clarke, 24 Jun 1990; TMAG G6488, 1 $ (23 mm),
Cyclops Pot (IB57), pool at bottom of lower pitch, 190 m depth, 290-01, coll. D.
Morgan & J. Butt for A. Clarke, 18 Feb 1990; TMAG G6486, 1 juvenile;? (18 mm),
1? (21 mm), Revelation Cave (IB1), 43°27.8'S 146°50.3'E, from deep pool, base of
underground shaft, 240 m asl, CV49, coll. A. Goede, 14 Jun 1969; QVM 10:13230,1 $
(24 mm), 2? ? (22-23 mm). Exit Cave, 43°24'S 146°52'E, tributary, coll. S. Eberhard,
15 Feb 1992; AM P82857, 1 juvenile? (14 mm), Exit Cave, 43°28.2'S 146°51.0'E,
trickle at side entrance from IB 161, Bobs Hole, in twilight zone, JHB T0701, coll.
J.H. Bradbury, 7 Mar 1997; AM P99291, 1 ? (21 mm). Exit Cave (IB-14), pool near
Ballroom passage, 700 m into cave, 43°28.6'S 146°51.3'E, 202-11, 14 Feb 2002; AM
P99292, \S (20 mm), 1 juv. ? (21 mm), Exit Cave, riffle zone near Ballroom Passage
junction, 43°28.6'S 146°51.3'E, dark zone, 397-63, coll. A. Clarke, 7 Mar 1997; AM
P99293, 1 juvenile;? (18 mm), 2 juvenile ?? (18-19 mm), Exit Cave (IB-120), Lost
Squeeze Passage, pools, 194-04, coll. A. Clarke, 20 Jan 1998; AM P99294,2?? (24
mm), Exit Cave (IB 14), Base Camp Tributary, c. 1.75 km into Exit Cave, 202-19, coll.
A. Clarke, 14 Feb 2002; TMAG G6489, 3 juvenile $$ (15-22 mm), 1 ? (21 mm),
Base Camp Tributary, c. 1.75 km into Exit Cave (IB-14), 193-112, coll. A. Clarke, 29
Jan 1993; TMAG G6485, (21-24 mm), 3?? (20-29 mm), Exit Cave, Devil’s

Stovepipe, pool at bottom of shaft, dribble system, at rock base with pools, “no pigment

regeneration after 4 weeks in lab”, coll. A. Goede & B. Collins, 2 Mar 1969; TMAG
G6484, 1 ? (32 mm), Exit Cave, 50 m downstream of “Waddle ‘n’ Splosh”, coll.
Laimonis Kavalieris, 23 January 1973; TMAG G6494, 3 juvenile $$ (22-25 mm),
5? ? (23-32 mm). Exit Cave, pool at base of shaft near Keller’s Squeeze, 43°28. l'S
146°50.7'E, CV42, coll. A. Goede & A. Keller, 29 Mar 1969; TMAG G6490, 1 ? (20
mm), Exit Cave (IB 14), 20 m upstream from site of monitoring probe, dark zone,
996-05, coll. L. Gardner & J. Hammond for A. Clarke, 7 Sep 1996; TMAG G1287,
2?? (20-23 mm), Exit Cave, coll. W.D. Williams, 1 Jun 1968;ZSR0385,1 juvenile?
(15 mm), Bradley-Chesterman Cave (IB6), c. 75 m upstream from entrance, 43°27.7'S
146°51.8'E, coll. A. Clarke, 5 Mar 2006.

Description. Eyes with cornea pigmented or unpigmented,
strongly reduced, narrower than stalk, shorter than one-fourth
length of stalk; stalk with subparallel or slightly convergent
margins.

Rostrum broadly triangular, almost equilateral, apex blunt.

Pleonites 1-5 unarmed, with sparsely setose pleural
margins, rounded. Pleonite 6 posterior and posterolateral
margins unarmed, setose. Pleonal sternites 3-5 with low,
median processes between pleopod bases, bilobed and widest
on sternite 3, bilobed on sternite 4, unilobate on sternite 5.

Telson length and width subequal to slightly longer
than wide, widest proximally; lateral margins sinuous in
dorsal outline, distally convergent; transition from lateral to
posterior margin obtusely and bluntly angular to rounded;
posterior margin bluntly angular to broadly rounded;
posterior spine row with 4-15 (usually 6-8) stout, well¬
spaced spines, directed posteriorly, arrangement usually
subsymmetrical, though frequently distinctly asymmetrical;
proximalmost spines near posterior 0.25 of telson length.

Antennule inner flagellum about 0.2 x body length (26-28
articles in holotype); article 7 inner margin obtusely angled
in adult males, with 4 (rarely 5) relatively short, slender,

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Records of the Australian Museum (2016) Vol. 68

Figure 6. Anaspides clarkei Ahyong, 2015, holotype male, 29 mm, Exit Cave, Ida Bay, SAMA C6301. (A) cephalothorax, dorsal view;
(B) pleonite 6, telson and right uropod; (C) pleonites 4-6 pleura, right lateral view; (D) right antennule; (E) right antenna; (F) labrum,
anterior view; (G) right mandible; (H) right mandible incisor process; (I) paragnaths, anterior view; (J) right maxillule; (K) right maxilla;
(L) pleopods 1-2, in-situ, ventral view. Scale: A-E, L = 2.0 mm; F-K = 0.7 mm.

closely spaced clasping spines; outer flagellum 0.6-0.8
x body length (103-108 articles in holotype). Antennal
flagellum 0.4-0.6 x body length (98 articles in holotype);
scaphocerite ovate, lateral spine near distal one-fourth; apex
slightly overreaching penultimate peduncular article.

Right mandibular incisor process with proximal tooth
distally trifurcate.

Pleopods 1-2 (usually) or 3 with endopod in adults
(occasionally with pleopod 3 endopod present on one side
only). Adult male pleopod 1 distally widened, scoop-like,
lateral margins expanded, obscuring retinacular lobe in
lateral view.

Uropodal protopod dorsally unarmed; exopod with 2 or
3 movable spines on outer margin near position of partial
diaeresis; exopod length about 3.5-4 times width, slightly
wider than endopod, apex rounded, relatively narrow.

Measurements. Male (n = 20) 15-29 mm, female (n = 47)
9-38 mm, sex indeterminate (n = 2) 8-10 mm.

Remarks. The cave dwelling Anaspides clarkei is highly
distinctive in having the posterior margin of the telson armed
with few (4-15, usually 6-8) stout, well-spaced spines
(versus a row of fine, closely-spaced spines), the strongly
reduced cornea, and long outer antennular flagella that

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Figure 7. Anaspides clarkei Ahyong, 2015. (A-O) Holotype male, 29 mm, Exit Cave, Ida Bay, SAMAC6301; (P-Q) paratype female, 32
mm, Exit Cave, SAMA C6302. (A) right thoracopod 1 (maxilliped); (B-H) right thoracopods 2-8; (I-J) right pleopod 1 endopod, lateral
and ventral views; (K) right pleopod 2 endopod, lateral view; (L) right pleopod 3, anterior view; (M-O) pleonites 3-5 median sternal
processes; (P-Q) female gonopore, right lateral and ventral views. Scale A-H, L-0 = 2.0 mm; I-K = 1.0 mm.

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Figure 8. Anaspides clarkei Ahyong, 2015. (A-C) juv. female, 15 mm, Bradley Chesterman Cave, Ida Bay, ZSRO; (D-F) female, 24
mm, Arthur’s Folly, Ida Bay, TMAG G6487; (G) female, 18 mm, Exit Cave, Ida Bay, AM P73045; (H-I) juv. female, 14 mm, Exit Cave,
AM P82857; (J-L) male, 28 mm, Wolf Hole, Hastings, AM P73047; (M) female, 28 mm, Wolf Hole, QVM 10:47694; (N) female, 31
mm, Wolf Hole, AM P73046; (O) paratype female, 32 mm, Exit Cave, SAMA C6302; (P) paratype female, 38 mm, Exit Cave, TMAG
G6033; (Q) female, 23 mm, Newdegate Cave, Hastings, AM P99289; (R) male, 22 mm, Newdegate Cave, AM P99289; (S) female, 19
mm, Revelation Cave, Ida Bay, TMAG G6486; (T-U) indet juv. (10 mm), female (25 mm), Milkrun Cave, Ida Bay, QVM 10:12177; (V)
female, 31 mm, Little Grunt Cave, Ida Bay, QVM 10:13261; (W-Y) female, 32 mm, “Waddle ‘n splosh”, Exit Cave, TMAG G6484; (Z)
female, 24 mm, Exit Cave, AM P99294. A, D, H, J, W, anterior cephalothorax; B, E, G, I, L, M-V, X, Z, telson; C, F, K, Y, scaphocerite.
Scale: A-C, H-I, T = 0.5 mm; D-G, J-S, U-Z = 1.0 mm.

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327

are distinctly longer than half the body length (Figs 5, 6).
Anaspides clarkei also differs from adults of other congeners
in lacking the endopod on pleopods 4 and 5, usually also
pleopod 3. The endopod of pleopods 1-4, and usually also
on pleopod 5, is present in adults of all other species of
Anaspides. Although pleopodal endopod development is
anamorphic in Anaspides , but in A. clarkei , the endopods
do not develop beyond pleonite 3, usually pleonite 2.
Similarly, the rostrum of other species of Anaspides becomes
increasingly slender with age, but the rostrum of A. clarkei
remains broad into maturity. Thus, the reduction in pleopodal
endopods and broad rostrum in adults suggests that A clarkei
is, in some respects, paedomorphic.

The “cave-type” telson of A clarkei , with few, stout well¬
spaced spines, superficially resembles that of A. eberhardi
from caves in the Junee-Florentine karst system. Distinctions
between the two species are outlined under the account of A.
eberhardi. Despite the superficially similar telson spination
to A. eberhardi , A. clarkei is most closely related to A.
jarmani from the neighbouring surface localities, uniquely
sharing similar male pleopod 1 morphology in which the
retinacular lobe is obscured in lateral view, the presence
of 4 (rarely 5) antennular clasping spines in adult males,
and in the absence of pleonal spines. Anaspides clarkei and
A. jarmani differ chiefly in the well-developed eyes in the
latter, different telson spination (spines widely spaced in A.
clarkei ), and absence of endopods on pleopods 4-5 (usually
also absent on 3) in A. clarkei. Other distinctions between
the two species are outlined under the account of A.jarmani.
As in other species of the genus, the uropodal endopod in
A. clarkei exceeds three-fourths the length of the exopod.
The right uropodal endopod of the holotype of A. clarkei ,
however, is abnormally shortened (Fig. 6B), possibly as a
result of damage or moult irregularities.

The arrangement of posterior telson spines of A. clarkei
is usually only approximately symmetrical, contrasting
with the highly degree of symmetry seen in A. eberhardi.
Several specimens of A. clarkei have aberrant, distinctly
asymmetrical telson spination, probably as a result of
injury or moult irregularities (e.g., Fig. 8N,P,S). A specimen
(female, 32 mm, TMAG G6484; Fig. 8W-Y) from “Waddle
‘n Splosh”, Exit Cave, is aberrant in having more numerous,
somewhat asymmetrically developed, less widely spaced
telson spines, and much more strongly reduced, almost
flattened corneas; in other respects, including pleopodal
endopod reduction, the specimen agrees with typical A.
clarkei. Development of the male antennular clasping spines
is sequential, with two or three present in juveniles, reaching
the full complement of four spines in adults. Both sexes of
A. clarkei appear to be mature by 19-25 mm.

Anaspides clarkei is known only from the Ida Bay and
Hastings karst systems, exhibiting troglobitic adaptations in
the significantly reduced cornea with degenerate ommatidial
facets, reduction or absence of comeal and body pigmentation,
and proportionally much longer antennular and antennal
flagella, measuring distinctly more than half body length.
General setal development in A. clarkei is uniform and
resembles that of epigean forms, but body pigmentation
differs between localities. Those from Ida Bay and
Newdegate Cave (Hastings) are unpigmented or significantly
de-pigmented, at most with diffuse body pigmentation; all
have corneal pigmentation (Fig. 35B). As observed by Goede
(1972) and Lake & Coleman (1977), however, specimens

from Wolf Hole lack all body and corneal pigmentation
(Fig. 35C). In addition to the complete loss of pigmentation,
adults from Wolf Hole also differ slightly from Newdegate
Cave and most Ida Bay specimens in the noticeably blunter,
broader and more flattened posterior margin of the telson
(Fig. 8L-N); additional material from Wolf Hole is required
to determine the stability of this feature. The telson in Ida Bay
and Newdegate specimens generally tapers more strongly
and is more rounded distally, although some specimens
from Milkrun Cave and Little Grunt Cave (Fig. 8U,V)
approach those from Wolf Hole. In other respects, the Wolf
Hole specimens agree well with those from other localities.
The 28 mm male from Wolf Hole (AM P73047; Fig. 8J-L)
has five clasping spines on the left antennule, four on the
right; other adult males have four clasping spines on both
antennules. Wolf Hole is the most isolated locality at which
A. clarkei occurs, being hydrologically separated from the
Ida Bay system and neighbouring Newdegate Cave, as well
as from epigean Anaspides. Some degree of subspecific or
possibly even specific differentiation is plausible between
Hastings and Ida Bay populations given the isolation of the
respective karst systems. Anaspides listed from the Hastings
and Ida Bay systems as “telson ‘cave’ type” by Eberhard et
al. (1991) are referrable to A. clarkei.

In both Hastings and Ida Bay systems, the closely related
A. jarmani has entered subterranean waters (Newdegate
Cave and Mystery Creek Cave, respectively). Both A. clarkei
and A. jarmani are present in Newdegate Cave, but it is not
presently known whether they are sympatric there or occur
in different parts of the system.

Quarrying at Ida Bay since the Second World War, with
attendant severe sedimentation and degraded water quality
led to extinction of Anaspides from Bradley-Chesterman
Cave. Closure of the quarry in 1992, however, followed
by catchment rehabilitation saw Anaspides recolonize
Bradley-Chesterman by December 1998 (Eberhard, 1999,
2001), presumably from populations in other parts of the
drainage such as Loon’s, Little Grunt, Arthurs Folly caves.
The re-appearance of A. clarkei in Bradley-Chesterman
Cave corroborates the supposition of subterranean continuity
between Ida Bay cave subsystems (Kieman, 1993).

Distribution. Presently known only from Ida Bay and
Hastings karst systems; 70-360 m asl.

Anaspides eberhardi sp. nov.

Ligs 9-12, 36

Anaspides sp. (telson ‘cave’ type). —Eberhard et al ., 1991:

48 (Junee-Florentine records only).

Type material. Holotype: QVM 10:49169A (32 mm), Niggly Cave (JF-2),
near Maydena, Junee-Florentine Karst, 42°42'S 146°31'E, coll. S. Eberhard, Aug
1993. Paratypes: QVM:2016:10:0001, (30 mm), Niggly Cave (JF-2), collected
with holotype; QVM:2016:10:0002,1 $ (32 mm), Niggly Cave (JF-2), collected with
holotype; TMAG G6479, 1 S (32 mm), 2£$ (28-34 mm), Niggly Cave (JF2-1),
42°42'S 146°31'E; QVM 10:12917, \S (31 mm), 2$£ (c. 25-33 mm), Cauldron Pot
(JF-2), Maydena, Junee-Florentine Karst, 42°42.9'S 146°34.4'E, 720 m asl. Brew Ch,
E series streamway, coll. S.M. Eberhard, 28 Jan 1990; TMAG G6492, 1 $ (30 mm),
Cauldron Pot (JF-2), Maydena, Junee-Florentine Karst, 42°42.9'S 146°34.4'E, 720 m
asl, coll. R. Eberhard, 1 Feb 1985.

Other material examined. (Junee-Florentine Karst). TMAG G6467, 1
juv. cj (19 mm), 1§ (43 mm), cave JF-104, at foot of Tiger Range, Florentine Valley,
42°33'S 146°25'E, underground stream, A. Goede, 9 Oct 1976; AM P99295, 1 juv. S
(shrivelled, poor condition, c. 13 mm), Welcome Stranger Cave (JF-229), 42°37.8'S

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Figure 9. Anaspides eberhardi sp. nov., male holotype, 32 mm, Niggly
Cave, QVM 10:49169. Habitus, right lateral view. Scale 2.0 mm.

146°30.4'E, from sump pool at end of cave, 398-03, coll. P. Verwey, 8 Mar 1998;
TMAG G6469, 1? (22 mm), Welcome Stranger Cave, 42°37.8'S 146°30.4'E, in
stream, CV50, coll. D. Maloney, 3 Apr 1971; TMAG G6470, \<$ (24 mm), 1 juv. $
(18 mm), Welcome Stranger Cave, 42°37.8’S 146°30.4'E, in stream, CV39, coll. S.
Eberhard,A. Richardson & R. Swain, 28 Oct 1984; TMAG G6471,2$$ (21-27 mm),
Welcome Stranger Cave, 42°37.8'S 146°30.4'E, coll. R. Eberhard, 6Aug 1990; TMAG
G6482,1$ (35 mm), Porcupine Pot(JF387), Florentine Valley, 42°40.9'S 146°30.2'E,
640 m asl, CV1, coll. S. Eberhard, 10 Nov 1985; TMAG G6480, (29-32 mm),

2$£ (31-37 mm), Gormenghast Cave (JF35-8), Florentine Valley, 42°41'S 146°30'E;
TMAG G6483, 1$ (c. 26 mm), Pendant Pot (JF37), Florentine Valley, 42°41.4'S
146°30.0'E, 550 m asl, CV36, coll. R. Eberhard, Apr 1984; TMAG G6472, 1$ (24
m m ), Growling Swallet (JF-36), 42°41.4'S 146°30.0’E, 570 m asl, CV25, coll. S.
Eberhard, 29 Sep 1985; TMAG G6474,1 juv. $ (10 mm), Growling Swallet (JF-36),
42°41.4'S 146°30.0'E, at least 400 m depth, in stream, 590 m asl, coll. S. Eberhard, 11
Dec 1983; TMAG G6475,5 $ § (27-47 mm), Junee Cave, Florentine Valley, 42°44.3'S
146°35.7'E, 340 m asl, coll. S. Eberhard, 3 Jan 1985; TMAG G6468, 1$ (31 mm),
Settlement Cave, Florentine Valley, CV45, coll. D. Maloney, 17 Apr 1971; TMAG
G6477,1 £ (45 mm), Florentine Valley, cave, no other data, CV48, Jun 1972; TMAG
G6476, 1 $ (c. 20 mm, very poor condition), Florentine Valley, cave, no other data,
CV47, Jun 1972; TMAG G6478, 26<$ (17-18 mm), 2££ (17-20 mm), Florentine
Valley, unnamed cave, found in pool of standing water in cave passage which acts as
a flood overflow, CV43, coll. A. Goede, 2 Nov 1969.

Description. Eyes with cornea pigmented, reduced,
narrower than stalk, shorter than half length of stalk; stalk
with subparallel or slightly convergent margins. Rostrum
triangular, apex narrow, blunt.

Pleonites with sparsely setose pleural margins, rounded;
Pleonites 1-3 unarmed. Pleonite 4-5 posterior tergal margins
unarmed; pleura rounded, with 0-3 (usually 1) small spines
and scattered setae. Pleonite 6 posterior margin with row of

small spines, setose; posterolateral margin setose, rounded.
Pleonal sternites 3-5 with low, median processes between
pleopod bases, bilobed and widest on sternite 3, bilobed on
stemite 4, unilobate on sternite 5.

Telson longer than wide, slender, triangular, widest
proximally, lateral margins slightly sinuous in dorsal outline,
distally strongly convergent on narrow apex; normally with
4 pairs of stout, graded, symmetrically arranged, well¬
spaced spines, anterior 3 pairs directed posterolaterally,
distal pair directed posteriorly; occasionally with abnormal
asymmetrical spination (3/4,3/5 or 4/5 spines on either side);
proximalmost spines near posterior 0.3-0.4 of telson length;
occasionally with stout posteromedian spine and small dorsal
median spine above posterior margin.

Antennule inner flagellum about 0.2 x body length
(24-29 articles in holotype); article 7 inner margin obtusely
angled in adult males, with 1 long, slender, clasping spine at
proximal comer; outer flagellum 0.6-1.1 (usually 0.7-0.8)
x body length (139 articles in holotype). Antennal flagellum
0.5-0.6 x body length (111 articles in holotype); scaphocerite
elongate, ovate, lateral spine at distal one-third; apex not
reaching midlength of distal peduncular article.

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329

Figure 10. Anaspides eberhardi sp. nov., male holotype, 32 mm, Niggly Cave, QVM 10:49169. (A) cephalothorax, dorsal view; (B) pleonite
6, telson and right uropod; (C) pleonites 4-6 pleura, right lateral view; (D) right antennule; (E) right antenna; (F) labrum, anterior view;
(G) right mandible; (H) right mandible incisor process; (I) paragnaths, anterior view; (J) right maxillule; (K) right maxilla. Scale: A-E
= 2.0 mm; F-K = 1.3 mm.

Right mandibular incisor process with proximal tooth
distally trifurcate.

Pleopods 1-4 with endopod. Adult male pleopod 1 distally
widened, scoop-like, lateral margins weakly expanded, not
obscuring retinacular lobe in lateral view.

Uropodal protopod with 1-3 small dorsal spines; exopod

with 1-3 movable spines on outer margin near position of
partial diaeresis; exopod length 3.5-4 times width, slightly
wider than endopod, apex rounded, relatively narrow.

Etymology. Named in honour of Stefan Eberhard, who
collected the majority of specimens of this new species.

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Figure 11. Anaspides eberhardi sp. nov. (A-O) male holotype, 32 mm, Niggly Cave, QYM 10:49169; (P-Q) paratype female, 32 mm,
QVM:2016:10:0002; (A) right thoracopod 1 (maxilliped); (B-G) right thoracopods 2-7; (H) leftthoracopod8; (I-J) rightpleopod 1 endopod,
lateral and ventral views; (K) right pleopod 2 endopod, lateral view; (L) right pleopod 5, anterior view; (M-O) pleonites 3-5 median
sternal processes; (P-Q) female gonopore, right lateral and ventral views. Scale A-H, L = 2.0 mm; I-K, M-0 = 1.0 mm; P-Q = 1.4 mm.

Measurements. Male (n = 12) 13-32 mm, female (n = 26) of the genus with a “cave-type” telson, is most closely related
10-47 mm. to A. richardsoni , uniquely sharing the single antennular

clasping spine in adult males (Fig. 9D). The two species
Remarks. Anaspides eberhardi sp. nov., the second species are readily separated by the armature of the telson, with

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331

Figure 12. Anaspides eberhardi sp. nov., selected features. (A) juv. female, 10 mm, Growling Swallet, TMAG G6474; (B) juv. male, 18
mm, foot of Tiger Range, TMAG G6467; (C) female, 20 mm, unnamed cave CV43, TMAG G6478; (D) female, 31 mm, Settlement Cave,
TMAG G6468; (E) paratype female, 30 mm, Cauldron Pot, TMAG G6492; (F) female, 42 mm, foot of Tiger Range, TMAG G6467;
(G) female, 47 mm, Junee Cave, TMAG G6475; (H) female, 33 mm, Junee Cave, TMAG G6475; (I) male, 24 mm, Welcome Stranger,
TMAG G6470; (J) paratype male, 31 mm, Cauldron Pot, QVM 10:12917; (K) paratype male, 30 mm, Niggly Cave, QVM:2016:10:0001;
(L) paratype female, 32 mm, Niggly Cave, QVM:2016:10:0001. Scale A = 0.5 mm; B-D = 2.0 mm, F-G = 2.0 mm; H-F = 1.0 mm.

few, well-spaced spines in the new species (compared to the
closely packed posterior spine row in A. richardsoni ), and
absence of the pleopod 5 endopod in adults of A eberhardi
(rarely absent in A. richardsoni). Moreover, the telson of A.
eberhardi is distinctly more elongate and posteriorly tapering
than any other species of the genus. Anaspides eberhardi
also shows cave adaptations in corneal reduction, more
elongate antennular flagella and body depigmentation. The

eyes, however, remain pigmented and the corneas, although
reduced compared to epigean forms, are comparatively larger
than in the other obligate troglobite, A. clarkei. The telson
of A. eberhardi is like that of A. clarkei , however, in having
few, stout, widely spaced spines, rather than the fine, closely
packed spine row of epigean forms. Anaspides eberhardi is
readily distinguished from A. clarkei by the presence of one
instead four (rarely five) antennular clasping spines in adult

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males, posterior denticles on pleonal pleura 5-6 (unarmed
in A. clarkei ), a distinctly more elongate and more distally
tapering telson in which the posterior spines extend anteriorly
to the posterior one-third of the telson, rather than posterior
one-fourth in A. clarkei ; and presence of the pleopod 4-5
endopod (absent in A. clarkei).

Variation in the A. eberhardi is slight. The posterior tergal
margin of pleonite 6 always has a series of small denticles,
and 1-3 small denticles usually on pleuron 5, often also on
pleuron 4 (Fig. 12). The outer margin of the uropodal exopod
has 1-3 movable spines, and the uropodal protopod bears 1-3
small dorsal spines. The outer antennular flagellum ranges
in length from 0.6-1.1 times body length, generally being
proportionally longer in smaller specimens. The arrangement
of telson spines is highly consistent in the present series
with four pairs of well-spaced marginal spines present from
the smallest through largest specimen examined (10-46
mm). Only six specimens differ from the typical pattern of
spination. Two specimens (paratype female, 30 mm, TMAG
G6492; female, 43 mm, TMAG G6467) have, in addition
to the four pairs of marginal spines, a short median spine
above the posterior margin, the smaller of which also has an
additional marginal median spine (Fig. 12E,F). The telsons of
the other four specimens have asymmetrical arrangements,
with unequal numbers of spines on either side of the
midline (3/4,3/4,3/5) (Fig. 12C,D), or distinctly irregularly
arranged spines (Fig. 12H). Although normally remarkably
consistent in arrangement, the length of the telson spines
varies allometrically, becoming proportionally shorter with
increasing body size.

Anaspides listed from the Junee-Florentine system by
Eberhard et al. (1991) as “telson ‘cave’ type” are referrable
to A. eberhardi. Several specimens appear to have clusters
of fungal hyphae on various parts of the body including the
cephalothorax, pleon, uropods and bases of the pereopods.

Given the secondary sexual features of adult males,
particularly the single antennular clasping spine and similar
pleopod 1, a close relationship between H. eberhardi and A
richardsoni is likely. The caves occupied by A. eberhardi
are at the southern end of the wide geographic range of A.
richardsoni. Anaspides richardsoni occurs throughout the
caves in the Mole Creek area, but in caves around Mt Field,
it has been recorded only from Rift and Growling Swallet
caves, the latter of which is also occupied by A. eberhardi.

Distribution. Presently known only from caves in the
Junee-Florentine Karst area: base of the Tiger Range to
Cauldron Pot Cave, Niggly Cave and Welcome Stranger;
340-720 m asl.

Anaspides jarmani Ahyong, 2015

Figs 13-17, 36

Anaspides tasmaniae. — Smith, 1909a: 64, 70 (Harz [sic]
Mountains). Thomson, 1926: 161 (Hartz Lake only).

—Knott etal., 1978: 703, 705, tab. 5. —Eberhard etal.,
1992: tab. 2. —Jarman & Elliot, 2000: fig. 4 (clade D,
part), tab. 1 (part, Adamsons Peak, Hartz Mtns).

Anaspides sp. —Eberhard et al., 1992: tab. 2.

Anaspides sp. (telson ‘normal’ type). —Eberhard et al., 1991:

48 (Judds Cavern, Newdegate Cave).

Anaspides sp. (telson type intermediate). —Eberhard et al.,
1991: 48 (Mystery Creek Cave).

Anaspides jarmani Ahyong, 2015: 598: fig. 1E-H (type
locality: Adamsons Peak).

Type material. Holotype: AM P73039,6' (24 mm), Adamsons Peak,
43°20.94'S 146°49.94'E, stream, 1200 m asl, coll. S. Jarman. Paratypes: AMP73040,
1 (j (26 mm), 1 juv. 8 (18 mm), 3 9$ (27-30 mm), 5 juv. (7-19 mm), type locality.

Other material examined. Adamsons Peat tmag G6395,5 juv. <38

(6-12 mm), 11 juv. 9 $ (7-11 mm), Adamsons Peak, 50 yards from track signposted
“water”, 43°21.2'S 146°49.3'E, coll. R. Swain & J. Ong, 8 Feb 1970.

Hartz Mountains'. AM P73041, 388 (20-23 mm), 1 juv. 8 (18 mm), 3^£ (19-24
mm), 6 juv. $$ (8-18 mm), Ladies Tam, 43°14.83'S 146°46.18'E, 955 m asl, coll. S.
Jarman; ZSRO 375, 388 (19-24 mm), 1 juv. 8 (20 mm), 4^ (22-24 mm), Ladies
Tarn, 43°14.83'S 146°46.18'E, 24 Feb 2006; QVM 10:49055, 1<J (19 mm), 1 juv. 8
(16 mm), 1 juv. $ (18 mm), 2 indetjuv. (5 mm), Ladies Tam, 43°14.34'S 146°46.03'E,
980 m, rock fauna, coll. S. Chilcott, Inland Fisheries Commission, 14 Jan 1988;
QVM 10:49056, 1 juv. 8 (21 mm), 6 juv. (12-16 mm), Hartz Lake, 43°14.55'S
146°45.15'E, rock fauna, 940 m asl, coll. S. Chilcott, Inland Fisheries Commission,
14 Jan 1988; TMAG G6399, 3 juv. 88 (16-25 mm), 7 juv. (16-25 mm), Hartz
Lake, 43°14.55'S 146°45.15'E, 940 m asl, coll. D. Coleman, 9 Jan 1974.

Ida Bay Karst. AMP99299,3 juv. $$ (c. 18-25 mm, poor condition), Mystery Creek
Cave (IB 10), Cephalopod Creek side passage, Ida Bay karst, 43°27.8'S 146°50.9'E,
1196-14, coll. A. Clarke, 2 Nov 1996; AM P99300, 3 damaged juveniles (c. 12-13
mm), Mystery Creek Cave (IB10), IdaBay Karst, 43°27.8'S 146°50.9'E, 1196-15, coll.
A. Clarke, 2 Nov 1996; AM P99301,1 juv. 8 (18 mm), Mystery Creek Cave (IB10),
Ida Bay karst, 43°27.8'S 146°50.9'E, 105-04, coll. A. Clarke & T. Murakami, 5 Jan
2005; AM P99302, 1 juv. $ (c. 16 mm, shrivelled, poor condition), Mystery Creek
Cave (IB 10), Cephalopod Creek passage, Ida Bay Karst, 43°27.8'S 146°50,9'E, pool,
998-28, coll. A. Clarke, 21 Sep 1998; AM P99303, 2 juv. 88 (12-16 mm), Mystery
Creek Cave (IB10), Cephalopod Creek side passage, 43°27.8'S 146°50.9'E, plunge pool
& streamlet, 1004-04, coll. A. Clarke, 21 Oct 2004; AM P99304,1 juv. 8 (shrivelled,
poor condition, c. 17 mm), 1 juv. $ (shrivelled, poor condition, c. 16 mm), Mystery
Creek Cave (IB10), 43°27.8’S 146°50.9’E, pool on side of main streamway, 400 m into
cave, dark, 1196-09, coll. A. Clarke, 2 Nov 1996; QVM 10:12175,1 juv. £ (10 mm).
Entrance Cave [= Mystery Creek Cave] (IB 10-4), Ida Bay, 43°27.8'S 146°50.9'E, cave
stream, coll. S. Eberhard, 11 Nov 1986; USNM 1277680,2 88 (20-23 mm), 1$ (24
mm), 1 juv. ^ (14 mm), 1 indetjuv. (6 mm). Mystery Creek Cave, Ida Bay, 43°27.7'S
146°50.8'E, 0-0.2 m, stn 87-254, coll. T. Iliffe, 29 Dec 1987.

Hastings Karst TMAG G6493, 1^ (21 mm), Hell’s Half Acre, Newdegate Cave,
43°23.0'S 146°50.5'E, small creek, coll. A. Goede, 1 Nov 1970.

Vanishing Falls karst, Salisbury River. QVM 10:13005,1 8 (26 mm), Salisbury River
Cave, 43°22.8'S 146°37.5'E, VF-X2, coll. S. Eberhard, 25 Apr 1992; QVM 10:13014,
2 88 (21-22 mm),2^ (c. 24-27 mm), Salisbury River Cave, 43°22.8'S 146°37.5'E,
VF6, flood overflow passage, coll. S.M. Eberhard & V. Wong, 21 Apr 1992.

Cracroft : QVM 10:12326,1 8 (28 mm), Wargata Mina, Judds Cavern, in main stream,
1.5 km from entrance, 43°15.3'S 146°35.0'E, 380 m asl, Cl-8, coll. S. Eberhard, 4
Apr 1989; QVM 10:12327,1 $ (22 mm), Wargata Mina (C-001), Judds Cavern, main
stream, 43°15.3’S 146°35.0'E,C1-19,380 m asl, coll. J. Jackson, 25 Nov 1989; TMAG
G6495,1 $ (31 mm), Judds Cavern, 43°15.3'S 146°35.0'E, Cl -28, route 66,2 Mar 1990.

Precipitous Bluff'. QVM 10:13279, 1 8 (26 mm), 1£ (27 mm), Bauhaus Cave (PB6),
Persephone Stream, Precipitous Bluff, 43°29.0'S 146°37.0'E, Screaming Stals
streamway, stn4, coll. S. Eberhard, 23 Dec 1991; QVM 10:12322,1 indetjuv. (7 mm),
Persephone Cave, Precipitous Bluff, 43°28.9'S 146°35.2'E, deep stream, PB17-6, coll.
S.M. Eberhard, 3 Jan 1990.

Southern Ranges'. TMAG G6366,4££ (23-28 mm), Ooze Lake, 43°30.2'S 146°42.0'E,
900 m asl, coll. P. Davies, Oct 1985.

Description. Eyes with well-developed cornea, pigmented,
wider than and longer than half length of stalk (epigean
specimens) to narrower than stalk, strongly reduced, shorter
than half length of stalk (in some subterranean forms); stalk
with subparallel margins.

Rostrum narrow in adults, apex blunt.

Pleonites 1-5 unarmed, with sparsely setose pleural
margins, rounded. Pleonite 6 posterior and posterolateral
margins unarmed, setose. Pleonal sternites 3-5 with low,
median processes between pleopod bases, bilobed and widest
on sternite 3, bilobed on sternite 4, unilobate on sternite
5. Pleonal sternites 3-4 with distinctly bilobed median
processes between pleopod bases, widest on sternite 3;
sternite 5 with narrow rounded lobe.

Telson length and width subequal or slightly longer than
wide, pentagonal, widest proximally; lateral margins sinuous
in dorsal outline, distally subparallel to convergent; transition
from lateral to posterior margin obtusely angular; posterior
margin acutely to obtusely angular, blunt medially; posterior
spine row with 17-37 slender, evenly graded, closely spaced
spines, longest medially.

Antennule inner flagellum about 0.2 x body length (20

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333

articles in holotype); article 7 inner margin obtusely angled
in adult males, with 3-5 (usually 4) relatively short, slender,
closely spaced clasping spines; outer flagellum 0.4-0.8
x body length (81-84 articles in holotype) in epigean
specimens, 0.4-1.1 x body length in subterranean specimens.
Antennal flagellum 0.4-0.5 x body length (57-62 articles
in holotype) in epigean specimens, 0.5-0.8 x body length
in subterranean specimens; scaphocerite ovate, lateral spine
usually near distal one-fourth, slightly distal to midlength in
specimens from Hartz Mountains; apex slightly overreaching
penultimate peduncular article. Right mandibular incisor
process with proximal tooth distally bifid to quadrifid.

Pleopods 1-4 or 5 with endopod in adults. Adult male
pleopod 1 distally widened, scoop-like, lateral margins
expanded, obscuring retinacular lobe in lateral view.

Uropodal protopod dorsally unarmed; exopod with 2 or
3 movable spines on outer margin near position of partial
diaeresis; exopod length 3-4 times width, slightly wider than
endopod, apex rounded, relatively narrow.

Measurements. Male (n = 32) 16-28 mm, female (n = 66)
7-31 mm, indet (n = 4) 5-7 mm.

Remarks. Anaspides jarmani and A. clarkei are unique in
the genus in their male pleopod 1 morphology in which
the lateral margins obscure the retinacular lobe in lateral
view (Fig. 151), the presence of 3-5 (usually 4) closely set
antennular clasping spines (Fig. 14D) and complete absence
of spines or denticles on the pleonites, unlike other species
of the genus in which one or more pleonites have some
degree of spination. Anaspides jarmani differs from A.
clarkei in the spination of the posterior margin of the tel son
(lined with numerous, fine, closely set spines rather than
stout, well-spaced spines), in having well-developed or only
slightly reduced eyes (rather than strongly reduced), and in
the presence in adults of endopods on pleopods 3-4 or 3-5
depending on locality. In adult A. clarkei , the endopod of
pleopod 4-5 (usually 3-5) is absent. Note that in the smallest

juveniles (7-10 mm) of A. jarmani from all localities as well
as larger juveniles from Mystery Creek Cave (Ida Bay), the
endopods of pleopods 4-5 are as yet undeveloped. In A.
jarmani , the posterior margin of the telson becomes angular
by 8 mm body length, more or less attaining its adult shape
by about 17 mm. Maturity is reached by 19-25 mm. The
angle of the posterior margin of the telson in adults is slightly
acute to approximately right angled in eastern specimens
(Adamsons Peak, Ida Bay, Hartz and Hastings), and obtuse in
westerly specimens, most of which are from caves (Cracroft,
Vanishing Falls, Precipitous Bluff, Southern Ranges).

Anaspides jarmani has a narrow distribution in southern
Tasmania, constrained in the northeast by the Hartz
Mountains, the southeast by Ida Bay and in the west by
the New River. Adult A. jarmani from the southeastern
part of its range (Adamson’s Peak and Newdegate cave,
Hastings) differ from those from other localities in the
presence of the endopod on pleopod 5 and a proportionally
broader scaphocerite in adults (noting that the scaphocerite
is typically more slender in juveniles than adults). Adults of
A. jarmani from other localities lack the pleopod 5 endopod
except for three specimens from the Hartz Mountains (18
mm female, AM P73041; 16 mm juvenile female, QVM
10:49056; 21 mm juvenile female, TMAG G6399) in which
the endopod is present on the right side, absent on the left.
These differences in pleopod 5 endopod condition and subtle
proportional differences in the scaphocerite might reflect
significant population differences, but all are presently
considered to represent a single species, A. jarmani, pending
further study.

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Figure 14. Anaspides jarmani Ahyong, 2015, holotype male, 24 mm, Adamsons Peak, AM P73039. (A) cephalothorax, dorsal view; (B)
pleonite 6, telson and right uropod; (C) pleonites 4-6 pleura, right lateral view; (D) right antennule; (E) right antenna; (F) labrum, anterior
view; (G) right mandible; (H) right mandible incisor process; (I) paragnaths, anterior view; (J) right maxillule; (K) right maxilla. Scale:
A-E = 1.0 mm; F-K - 0.7 mm.

Although normally epigean, Anaspides jarmani has
entered caves throughout its range: Hastings (Newdegate
Cave), Ida Bay (Mystery Creek Cave), Cracroft (Judds
Cavern), Vanishing Falls Karst (Salisbury River Cave) and
Precipitous Bluff (Bauhaus Cave, Persephone) (Fig. 17).
Specimens from Newdegate Cave (Fig. 17E) agree well
with epigean forms, including pigmented eyes, differing
only in body depigmentation and longer antennular flagella;
they are readily distinguished from A. clarkei by the much

better developed cornea, telson spination and presence of
the pleopod 5 endopod. Distinct comeal reduction is evident
in unpigmented specimens from Judds Cavern and some
specimens from Salisbury River Cave (Fig. 17A,B) as in
cave forms of A. richardsoni from the Honeycomb and Wet
Caves, Mole Creek. In addition to the cave-adapted form in
Salisbury River Cave (Fig. 17B), pigmented epigean forms
(Fig. 17C) are also present, the former from streamways and
the latter from seeps (Eberhard etal., 1991,1992). Specimens

Ahyong: Endemic Tasmanian Mountain Shrimps 335

Figure 15. Anaspides jar mam Ahyong, 2015. (A-P) holotype male, 24 mm, Adamsons Peak, AM P73039; (Q-R) paratype female 27 mm,
Adamson’s Peak, AM P73040; (A) right thoracopod 1 (maxilliped); (B-H) right thoracopods 2-8; (I-J) right pleopod 1 endopod, lateral
and ventral views; (K) right pleopod 2 endopod, lateral view; (L) pleopods 1-2, in-situ, ventral view; (M) right pleopod 3, anterior view;
(N-P) pleonites 3-5 median sternal processes; (Q-R) female gonopore, right lateral and ventral views. Scale = 1.0 mm.

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Figure 16. Anaspides jarmani Ahyong, 2015, anterior cephalothorax, telson and scaphocerite. (A) paratype juv. female, 8 mm, Adamsons
Peak, AM P73043; (B) paratype juv. male, 18 mm, Adamsons Peak, AM P73043; (C) female, 25 mm, Ooze Lake, TMAG G6366; (D)
male, 23 mm, Ladies Tarn, Hartz Mountains, AM P73041. Scale: A = 0.25 mm; B = 0.5 mm; C-D = 1.0 mm.

from Precipitous Bluff (Fig. 17D) are unusual, with the
seemingly mature male from Bauhaus Cave having two
antennular clasping spines, but atypical pleopod 1 endopod;
they may represent a separate species but tentatively assigned
to A. jarmani pending further study. As in epigean specimens
from Ooze Lake (Fig. 16C) and the Hartz Mountains
(Fig. 16D), cave specimens from Cracroft, Salisbury and
Precipitous Bluff lack the endopod on pleopod 5, whereas
the endopod is present in the Newdegate Cave specimen as
in the adjacent epigean specimens from Adamsons Peak.
Specimens of A. jarmani from Mystery Creek Cave, Ida
Bay, are juveniles and apart from depigmentation and an
incomplete complement of pleopod endopods, agree well
with surface forms.

Distribution. Southern Tasmania in epigean habitats from
the Hartz Mountains to Adamson’s Peak and Ooze Lake, and
from caves in the Hastings, Ida Bay, Cracroft, Salisbury karst
systems and Precipitous Bluff; 160-380 m asl (subterranean),
900-1340 m asl (epigean).

Anaspides richardsoni sp. nov.

Figs 18-23, 35D-F, 36

Anaspides tasmaniae. —Caiman, 1897: 802 [“Lake Field” (=

Mt Field) specimens only (OM Iv. 1394, below)]. —Smith,
1909a: 64, 70 (Mt Read, Mt Field). —Williams, 1965b:
333-334, fig. 1, 2; 1965a: 106, tab. 6. —Swain & Reid,
1983: 163-171. —Richardson & Swain, 1989: 277, tab.

1, app. 1. —Jarman & Elliot, 2000: fig. 4 (clade E, part),
tab. 1 (part, Central Plateau). —Gooderham & Tsyrlin,
2002: 73 (unnumbered colour figure). —Richter et al.,
2002: 341,347, fig. 31-33.

Anaspides spinulae. —O’Brien, 1990: frontispiece, pi. 1.
Anaspides sp. (telson ‘normal’ type). —Eberhard etal ., 1991:

48 (Mole Creek caves).

Type material. (Mt Field). Holotype: AM P72839,$ (33 mm), between
Newdegate Pass and Mawson Plateau, coll. J. Kunze, 27 Feb 1974. Paratypes: AM
P72840, 1 $ (TL 32 mm), between Newdegate Pass and Mawson Plateau, 42°40.2'S
146°33.6'E, coll. J. Kunze, 27 Feb 1974; AM P72841, { 23-21 mm), 5 juv.

(17-21 mm), 4^ (24-31 mm), 5 juv. ^ $ (14-20 mm), between Newdegate Pass and
Mawson Plateau, 42°40.2'S 146°33.6'E, coll. J. Kunze, 27 Feb 1974; ZRC 2016.0491,
1V (30 mm), 1 $ (31 mm), between Newdegate Pass and Mawson Plateau, 42°40.2'S

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337

Figure 17. Anaspides jar man i Ahyong, 2015, anterior cephalothorax, telson and scaphocerite. (A) male, 28 mm, Judds Cavern, QVM
10:12326; (B) female, c. 24 mm, Salisbury River Cave, QVM 10:13014; (C) male, 26 mm, Salisbury River Cave, QVM 10:13005; (D)
female, 27 mm, Bauhaus Cave, Precipitous Bluff, QVM 10:13279; (E) female, 21 mm, Hell’s Half Acre, Newdegate Cave, TMAG G6493;
(F) male, 23 mm, Mystery Creek Cave, USNM 1277680. Scale = 1.0 mm.

146°33.6'E, coll. J. Kunze, 27 Feb 1974; TMAG G6174, (27 mm), 2 juv. SS

(14-16 mm), 19 (34 mm), Mt Field, small tarn above University ski lodge, 42°40.134'S
146°34.152'E, tam in herbfield with dolerite rocky outcrops, dipnet, 1240 m asl, coll.
R. Mollison, 27 Apr 2011; TMAGG6363, 1? (45 mm), midway between Newdegate
Pass and Newdegate Tam, 42°39.6'S 146°33.6'E, in small tam, 1200 m asl, coll. I.
Wilson & J. Ong, 25 Jan 1970; TMAG G6419, 2$$ (25-27 mm), 21 juv. $$ (11-21
mm), 1? (22 mm), 37 juv. ?? (11-21 mm), Newdegate Pass, 42°39.6'S 146°33.0'E,
sample 2, coll. I. Wilson & J. Ong, 25 Jan 1970; TMAG G6445, (20-21 mm),

6 juv. # 03-19 mm), 2?? (24-29 mm), 5 juv. ?? (13-15 mm), Newdegate Pass,
42°39.6'S 146°33.0'E, sample 1, coll. I. Wilson & J. Ong, 25 Jan 1970; TMAG G6429,
11 SS (22-32 mm), 3 juv. $$ (18-21 mm), 25?$ (22-34 mm), 5 juv. ?? (14-21
mm), near Sitzmark Lodge, Mt Field, 42°40.8'S 146°34.8'E, tarn, coll. C. Reid, 15-18
Oct 1974; USNM 1277685, \<$ (26 mm), 1 juv. S (17 mm), 3?? (25-30 mm), near
Sitzmark Lodge, Mt Field, 42°40.8'S 146°34.8'E, tarn, coll. C. Reid, 15-180ct 1974.

Other material examined. Gunns Plain to Black Bluff. AM P99296, 1
juv. C?(12 mm), Great Western Cave (GP27), G u nn s Plains karst area, NW Tasmania,
41°17.8'S 146°00.3'E, from riffle pool 250 m upstream, 1296-24, 109 m asl, coll. A.
Clarke, 29 Dec 1996; QVM 10:13975,1 ? (23 mm), Paddy’s Lake, below Black Bluff,
near Loongana, Nietta South, 41 °27.2'S 145°57.6'E, 1070 m asl, coll. T. Hume, 5 May
1972; TMAG G6398, 14# (20-24 mm), 1 juv. <J (16 mm), 32?? (19-30 mm), 3
juv. ? ? (16-17 mm), Paddy’s Lake, Black Bluff, 41°27.2'S 145°57.6'E, 1070 m asl,

coll. C. Binks, 25 Apr 1979; TMAG G6169,1 juv. $ (13 mm), 6 juv. ?? (10-17 mm),
Vale of Belvoir, 41°32.95'S 145°53.54'E, coll. R. Mollison, 16 Mar 2010.

Deloraine. TMAG 14391/G135,2<?<? (22-24 mm), Deloraine, coll. C. King, Nov 1937.

Mole Creek (Form 1)\ SAMAC8481,2 immature c?<3 (16-21 mm), 2?? (30-39 mm),
1 indet juv. (9 mm), about 200 m from cave entrance, Marakoopa Cave, 41°34.9'S
146°17.3'E, 490 m asl, small stream, BS0464, coll. E. Hamilton Smith, 19 Nov 1963;
USNM 1277682,3 $$ (29-34 mm), 3 juv. # (12-16 mm), 3? ? (21-30 mm), 2 juv.
?? (16-17 mm), Marakoopa Cave, 0.3 m, stn 87-253, coll. T. Iliffe, 27 Dec 1987;
TMAG G646, \S (34 mm), 1 ? (45 mm), Long Creek, Marakoopa Cave (MCI20),
4 Dec 1991; QVM 10:13973, 1? (33 mm), Marakoopa Cave II (MC-015), 41°34.9'S
146°17.3'E, coll. K. Crocker, 1 Jul 1981; TMAG G6458, 2# (31-32 mm), 2??
(32-33 mm), Lake Passage, Marakoopa Cave II (MC15-1), 41 °34.9'S 146° 17.3'E, 490
m asl, coll. A. Goede, 21 Aug 1982; TMAG G6459, 2$$ (26-28 mm), 3?? (21-32
mm), Lake Passage, Marakoopa Cave II (MCI 5-1), 41°34.9'S 146°17.3'E, 490 m asl,
coll. A. Goede, 21 Aug 1982; TMAG G6462, 2?? (36-41 mm), Prohibition Cave,
41°35.6'S 146°19.5’E, coll. R. Eberhard, 19 Apr 1991.

Mole Creek (Form 2): AM P73038, 1 j (37 mm), Honeycomb Cave (MC107), lower
pool, dark zone, 41°36.0'S 146°24.4'E, 1096.03, coll. S. Bunton, 27 Oct 1996; ZSRO
863a, 1 ? (44 mm), Honeycomb Cave, KS34, Anal6A011,27 Feb 2013; ZSRO 863b,
1? (46 mm), Honeycomb Cave, KS34, Anal7 A011,27 Feb 2013; USNM 1277681,
\S (33 mm), 1 juv. S (22 mm), 7?? (32-55 mm), Honeycomb Cave, 0-0.5 m,

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Records of the Australian Museum (2016) Vol. 68

mm), 222 (30-37 mm), Kellys Pot (MC207-13), 41°36.6'S 146°22.5'E, 520 m asl;
TMAG G6460,1 $ (37 mm), Mole Creek caves, in water pool, coll. R.A. Rafferty, 28
Apr 1938; TMAG G6457, 1<J (36 mm), 222 (37-46 mm), Herbert’s Pot (MC202),
41°36.9'S 146°23.3'E, 500 m asl, CV40, coll. S. Eberhard, 6 Apr 1985; AMP56373,
1 S (TL 35 mm), 1 $ (TL 35 mm). Wet Cave near Caveside, 41 °36'S 146°25’E, C.92T,
moderate flowing stream, disappears before cave entrance, cave with few pools, coll.
W. Ponder et al., 18 Jan 1982.

Mole Creek (Form 3): SAMAC8482, 1 2 (24 mm), 2 juv. (7-9 mm), unnamed cave,
Sassafras Creek, about 50 m from entrance, 41°33.7'S 146°21,9'E,270 m asl, clear pool,
mud bottom, BS0457,18Nov 1963; TMAG G6454, 3$$ (23-32) mm), 1 $ (23 mm),
5juv. (13-20 mm), Kubla Khan Cave, 41°33.2'S 146°17.6'E,coll. S. Eberhard, R.
Swain & A. Richardson, 23 Sep 1981; TMAG G6455, IS (26 mm), 1 juv. 3 (20 mm),
2$ $ (26-28 mm), 3 juv. $ $ (10-21 mm), Kubla Khan Cave, 41°33.2'S 146°17.6'E,
coll. S. Eberhard, A. Richardson & R. Swain, 25 May 1986; TMAG G6456, \S (29
mm), 222 (23-39 mm), Kubla Khan Cave, 41°33.2'S 146°17.6'E, in river, c. 500 m
into cave, 320 m asl, coll. S. Eberhard, 12 Nov 1983; TMAG G6466, 3$$ (25-30
mm), 3$ ? (23-28 mm), Kubla Khan Cave (MC1-30), 41 °33.2'S 146°17.6'E; USNM
1277679, Id' (25 mm), Kubla Khan Cave, 0-0.5 m, stn 87-250, coll. T. Ilifle, 26 Dec
1987; USNM 1277683, 3 S3 (23-26 mm), 522 (28-40 mm), Mayberry area, Mole
Creek, water cave, from sinkhole near farmhouse, baited traps, 0.2m, stn 87-251,
coll. T. Iliffe, 27 Dec 1987.

West Coast Range. TMAG G6411, \S (26 mm), 3 juv. SS (16-20 mm ) 2 juv. 92
(20-21 mm), tam II, S of Lake Tyndall, 41°57.1'S 145°35.2'E, 980 m asl, coll. C.J.
Binks & B. Knott, 16 Jan 1973; TMAGG6312, %SS (23-25 mm), 3 juv. SS (16-21
mm), 822 (23-30 mm), 3 juv. 2$ (15-18 mm), tam S of Lake Tyndall, 41°571’S
145°35.2,980 m asl, coll. C.J. Binks & B. Knott, 16 Jan 1973; TMAG G6362,1 juv. $
(18 mm), 2 juv. 2? (16-17 mm). Lake Sandra, Mt Murchison, 41°49.9'S 145°35.8'E,
940 m asl, 10 Dec 1973; TMAG G6385, 12 (damaged, c. 26 mm), Lake Sandra, Mt
Murchison, 41 °49.9'S 145°35.8'E, 940 m asl, coll. W. Fulton,Nov 1983; TMAG G6392,
12 (27 mm), N of Geikie, Tyndall Range, alpine plateau above climbing route and
camp, 41°57.4'S 145°35.0’E, 1000 m asl, coll. C.J. Binks & B. Knott, 16 Jan 1973.

Cradle Mountain Lake St Clair National Park. TMAG G6352, 1 d (21 mm), 12 (23
mm). Twisted Lake, Cradle Mountain, 41°40.22'S 145°58.09'E, 1116 m asl, coll. D.
O’Brien, 8 Mar 1990; SAMA C6303, 1 juv. S (18 mm), top of Cradle Mountain,
41°40.8'S 145°57.0'E, small pool in creek, 4850 ft asl [1455 m], coll. A. Kowanko, Dec
1967; SAM C6304,2 2 2 (21-32 mm),top of Cradle Mountain,41°40.8'S 145°57.0'E,
small pool in creek, 4850 ft asl [1455 m], coll. A. Kowanko, Dec 1967; QVM 10:13835,
\S (20 mm), Sutton’s Tarn, Cradle Mountain, 41°41.01'S 145°55.98'E, 1089 m asl,
coll. Kingston, 1991; TMAGG6350, 1 S (26 mm), 12 (24 mm), Sutton’s Tarn, near
Kitchen Hut, Cradle Mountain, 41°41,0'S 145°56.0'E, 1089 m asl, coll. D. O’Brien, 8
Mar 1990; TMAG G6351,222 (30-31 mm), Sutton’s Tarn, near Kitchen Hut, Cradle
Mountain, 41°41.0'S 145°56.0'E, 1089 m asl, coll. D. O’Brien, 8 Mar 1990; TMAG
G6381, 2.SS (27-31 mm), 1222 (21-32 mm), Mt Doris, tarn on S side, 41°52.6'S
146°02.7'E, 1170 m asl, coll. B. Knott, 12 May 1970; AMP72843,1 juv. S (21 mm),
12 (23 mm), Mt Ossa-Mt Doris Ridge, Mt Ossa, 41°52.2'S 146°04.8'E, 1255 m asl,
coll. C. Sands; AM P82858, 2$S (23-24 mm), 4 juv. 22 (14-21 mm), 822 (23-30
mm), 5 juv. 22 (13-18 mm), S of Mt Doris, runnel crossing Mt Ossa track, 41 °52.2'S
146°01,8'E, deep runnel in grassy lawn in alpine shrubbery, FW18, coll. A. Richardson
& PA. Serov, 28 Jan 1990; TMAG G6413, 6 SS (24-29 mm), 622 (26-34 mm), 1
juv. 2(16 mm), near summit of Cathedral (NE of summit), 41°53.4'S 146°07.2'E, c.
1350 m asl, tam, small, clear & with rocky bottom, sample 6, coll. C.J. Binks & B.
Knott, 31 Jan 1972; WAM Cl 1772, lc? (21 mm), 322 (21-23 mm). Cradle Mountain
district, coll. A. Connell, 1939; WAM C58160, 1 juv. d 1 (16 mm), 422 (25-39 mm),
Cradle Mountain-Lake St Clair, National Park, 16 Oct 1947.

Walls of Jerusalem National Park (non- “spinulae ’’form) : TMAG G6448, 28 juv. SS
(13-18 mm), 49 juv. 2 2 (12-19 mm), 1 indet juv. (5 mm), Jaffa Vale, at Dixon’s Hut,
Walls of Jerusalem, 41°40 77'S 146°06.37'E, 1250 m asl, coll. ?R. Swain, 1969; TMAG
G6449,2 SS (25-28 mm), 2juv. SS (18-19 mm), 4 juv. 22 (15-17 mm), Jaffa Vale,
at Dixon’s Hut, Walls of Jerusalem, 41°40.77'S 146°06.37'E, 1250 m asl, coll. ?R.
Swain, 1969; TMAG G6319,2 SS (23-29 mm), 12 (20 mm), 1 juv. 2 (13 mm), pool
50 m W of Herod’s Gate Pool, 41 °48.7'S 146°16.8'E, 1220 m asl, coll. S. Smith, 16 Apr
1990; TMAG G6380, 6 juv. SS (13-16 mm), 8 juv. 22 (12-19 mm), Lake Adelaide
track between Fish Rock and Herod’s Gate, Walls of Jerusalem, 41°48.7'S 146°16.8'E,
runnel, 1200 m asl, coll. B. Knott, 18 Nov 1971; TMAG G6368, 5d'd' (24-26 mm), 10
juv. SS (14-21 mm), 822 (25-35 mm), 9 juv. 22 (H~20 mm), near Herod’s Gate,
Walls of Jerusalem, 41°48.7’S 146°17.4'E, small creek, 1200 m asl, coll. B. Knott, 17
Nov 1971; TMAG G6446, 2 SS (25-27 mm), 1 juv. 3 (17 mm), 222 (26-30 mm),
10 juv. 22 (14-18 mm), Zion Gate west, Walls of Jerusalem, 41°48.9'S 146°19.6'E,
1280 m asl, coll. ?R. Swain, 1969; TMAG G6393, 1 S (24 mm), 10 juv. SS (11-18
mm), 22 2 (24-33 mm), 13juv. 22 (15-18 mm), Saddle through to east wall, Walls of
Jerusalem, 41°48.9'S 146°19.2’E, 1230 m asl, coll. J. Bludhorn, 18 Nov 1971; TMAG
G6418, ASS (29-32mm),4juv. SS (16-23 mm),222 (26-29 mm), 2juv. 2$ (13-18
mm), E side of Zion Gate, Walls of Jerusalem, 41°49.0'S 146°19.6'E, 1240 m asl, coll.
?R. Swain, 1969; TMAG G6345,3 SS (21-22 mm), 12 (25 mm), 20 m below Gate of
Chain, Walls of Jerusalem, 41°49.12'S 146°18.39'E, pool in pencil pines, 1300 m asl,
coll. S. Smith, 16Apr 1990; TMAG G6346, 1 S (21 mm), 322 (20-22 mm), 1 juv. 2
(14 mm). Gate of Chain, 50 m belowridge. Walls of Jerusalem, 41°49.13'S 146°18.53'E,
1280 m asl, coll. S. Smith, 16 Apr 1990; TMAG G6318, 2 SS (24-27 mm), 1 juv. 3
(13 mm), 22 2 (26-30 mm), 100 m E of ridge, Gate of Chains, Walls of Jerusalem,
41°49.13'S 146°18.60'E, 1260 m asl, coll. S. Smith, 16 Apr 1990; TMAG G6440, 1 S
(23 mm), 1 juv. S (16 mm), 12 (23 mm), 6 juv. 2,2 (12-16 mm), near Lake Ball, Walls
of Jerusalem, 41°49.23'S 146°17.95'E, 1240 m asl, coll. V. Thorp, 3 Apr 1972; TMAG
06320,6* (26 mm), 1 juv. S (15 mm), 22 2 (25-29 mm), 1 juv. 2 (16 mm), 200 m
below Pool of Bethesda, Walls of Jerusalem, 41°49.34'S 146°17.81'E, runnel, #1,1260
m asl, coll. S. Smith, 14 Apr 1990; TMAG G6348, 6 SS (25-32 mm), 12 (32 mm),
50 m SE of Damascus Gate, 41°49.61'S 146°17.59'E, pool in grassland, coll. R. Smith,
1350 m asl, 15 Apr 1990; TMAG G6376, 3 juv. S3 (15-16 mm), 11 juv. 22 (12-17
mm) Damascus Gate, 41°49.61'S 146°17.95'E, coll. R. Swain, 1969; TMAG G6422,
IS (29 mm), 1 juv. 33% mm),222 (26-28 mm), 6juv. 22 (12-15 mm), Damascus
Gate, Walls of Jerusalem, runnel draining into Lake Calvine, 41°49.99'S 146°18.16'E,
1340 m asl, coll. ?R. Swain, 1969; TMAG G6394, 9 juv. SS (15-20 mm), 12 (24
mm), 8 juv. 22 (13-18 mm), near Junction Lake immediately under Moraine retaining
Lake Meston, Walls of Jerusalem, 41°55.35'S 146°11.52'E, bog drainage pool (chiefly
Gleichenia), “sample 2”, 950 m asl, coll. C. Binks & B. Knott, 31 Jan 1972; TMAG
G6397, 7 juv. S3 (10-22 mm), 12 (29 mm), 5 juv. 22 (17-21 mm), creek draining
into Junction Lake, second of four or five drainage systems on broad plain from Lake
Meston, Walls of Jerusalem, 41°55.35'S 146°11,52'E, clear pools under Nothofagus lined
creek, “sample 3”, 950 m asl, coll. C. Binks & B. Knott, 31 Jan 1972.

Walls of Jerusalem National Park (mixed “spinulae" and non-“spinulae” forms)'.
TMAG G6396, ASS (21-22 mm), 6 juv. 33 (15-18 mm), 12 (24 mm), 2 juv. 2 2
(17-20 mm), Zion Vale, Walls of Jerusalem, 41°48.64'S 146°18.83’E, pool, 1200 m
asl, coll. R. Swain, 1969; TMAG G6379,1 S (25 mm), 2 juv. 33 (18-22 mm), 222
(24-25 mm), Maximal Creek, W of Herod’s Gate, Walls of Jerusalem, 41°48.62'S
146°15.94'E, 1150 m asl, coll. B. Knott, 16 Nov 1971.

Walls of Jerusalem National Park (“spinulae" form): WAM Cl 1771, 4 juv. SS
(shrivelled, c. 13-16 mm), 12 (shrivelled, c. 19 mm), Jones Tam, at foot of Western
Wall, Walls of Jerusalem, 41°49.02'S 146°18.46'E, approx. 4200 feet [1260 m], 26

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Figure 19. Anaspides richardsoni sp. nov., holotype male, 33 mm, Mt Field, AM P72839. (A) cephalothorax, dorsal view; (B) pleonite
6, telson and left uropod; (C) pleonites 4-6 pleura, right lateral view; (D) right antennule; (E) left antenna; (F) labrum, anterior view;
(G) right mandible; (H) right mandible incisor process; (I) paragnaths, anterior view; (J) right maxillule; (K) right maxilla. Scale: A-E
= 1.0 mm; F-K = 0.7 mm.

Apr 1935; TMAG G6347, 4 juv. (9-12 mm), 7 juv. 99 (11-15 mm), 100 m E
of ridge, Gate of Chain, Walls of Jerusalem, 41°49.13'S 146°18.53'E, 16 Apr 1990;
TMAG G6391,9juv. $$ (14-18 mm), 19 (27 mm), 11 juv. 99 (13-17 mm), SE end
Cloister Lagoon, 41 °54.26'S 146° 10.66'E, from small drainage pools in predominantly
Gleichenia covered bog, sample 4, 1100 m asl, coll. C.J. Binks & B. Knott, 31 Jan
1972; TMAG G6434,(23-30 mm, 5 9 9 (24-31 mm). Pool of Bethesda, Walls of
Jerusalem, 41°49.28'S 146°17.95’E, 1270 m asl, coll. R. Swain, 1969; TMAG G6349,
2 SS (21-23 mm), 1 juv. S (15 mm), 399 (26-27 mm), E edge Pool of Bethesda,
41°49.28'S 146°17.88'E, 1270 m asl, coll. S. Smith, 16 Apr 1990; TMAG G6427,1<J
(24 mm), 19 juv. S6 (12-18 mm), 11 juv. 9$ (12-20 mm), 6 indet juv. (6-7 mm),
41 °53.01'S 146°09.23'E, from dirty, deep pool system draining through pineapple grass
into Lake Chalice, 1020 m asl, “sample 5”, coll. C. Binks & B. Knott, 31 Jan 1972;
TMAG G6330, 3<?<? (22-23 mm), 19 (24 mm), [no label] Lake Chalice, 41°52.84'S
146°08.87'E, coll. W. Fulton, 2 Feb 1988.

Western Lakes—Great Western Tiers'. AMP57906,19 (21 mm), Western Bluff, 41 °37'S
146°17'E, 26 Jan 1964; QVM 10:13977,19 (36 mm), 1 juv. 9 08 mm), Jacks Lagoon,
5 km SW of Lake Mackenzie, 41 °42. l'S 146°20.6’E, “abundant”, “no fish”, 1260 m asl,
coll. E.V. Terry, 7 Apr 1989; WAM C58161, 3$$ (26-29 mm), 299 (25-26 mm),
Ironstone Mountain, Northwestern Tiers, 41 °42.8'S 146°28.5'E, from temporary creek,
100 m asl, coll. I. Gooch, 1946; QVM 10:13978, 1<J (23 mm), 299 (23-31 mm),
Meander Falls, 41°44. l'S 146°30.3'E,coll. S. Merry, 1957; QVM 10:49162 (“spinulae”
form), 1 S (23 mm), 599 (23-28 mm), Lake Johnny, 41°43.6'S 146°25.2'E, 1210 m
asl, coll. W. Fulton, 9 Dec 1987; QVM 10:49163 (“spinulae” form), 2$$ (18-20 mm),
599 (23-26 mm), Lake Halkyard, 41°44.2'S 146°18.9'E, 1210 m asl, coll. W. Fulton,
9 Dec 1987; QVM 10:49164 (“spinulae” form), 2t?c? (22-23 mm), 59 9 (20-30 mm),
Lake Fox, 41°43.9'S 146°24.8'E, 1230 m asl, coll. W. Fulton, 9 Dec 1987; QVM
10:49165, (28-31 mm), 8 9 9 (24-29 mm), 3 juv. (20-21 mm), 15 juv. 99

(11-19 mm), 300 m Wof Lake Fox, 41°43.9’S 146°24.00'E, pool, 1210 m asl, coll. W.

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Figure 20. Anaspides richardsoni sp. nov. (A—P) holotype male, 33 mm, Mt Field, AM P72839; (Q-R) paratype female, 32 mm, Mt
Field, AM P72840; (A) right thoracopod 1 (maxilliped); (B-H) right thoracopods 2-8; (I-J) right pleopod 1 endopod, lateral and ventral
views; (K-L) right pleopod 2 endopod, lateral and ventral views; (M) right pleopod 5, anterior view; (N-P) pleonites 3-5 median sternal
processes; (Q-R) female gonopore, right lateral and ventral views. Scale A-M = 2.0 mm; N-R = 1.0 mm.

Fulton, 9 Nov 1987; QVM 10:13980,2juv. 33 (17-18 mm), 1 juv. $ (19 mm), stream
on top of Western Tier near Lake Lucy Long, 41°42.2'S 146°26.0'E, 1190 m asl, coll.
J. Simmons, 3 Mar 1963; TMAG G6438, 3 33 (25-29 mm), 4 juv. 33 (17-22 mm),
1$ (28 mm), 7 juv. 99 (14-21 mm), edge of Great Western Tiers near Pine Lake,
41°44.7'S 146°43.0'E, runnel draining opposite way, 1220 m asl, coll. B. Knott, 1970;
WAM C58166,1^ (31 mm), creek north of Pine Lake, coll. G.E. Nicholls, 27 Jan 1947;
WAM C58165, 7 33 (19-23 mm), 4 juv. 33 (17-17 mm), 799 (19-27 mm), 4 juv.
$9 (16-18 mm), N end of Pine Lake, 41°44.2'S 146°42.2'E, from linked puddle, coll.
G.E. Nicholls, 27 Jan 1947; TMAG G6443, lc^ (21 mm), 16 juv. 33 (16-20 mm), 8$ $

(19-26 mm), 10 juv. f f (14-19 mm). Pine Lake, 41°44.6'S 146°42.0'E, 1190 m asl,
coll. R. Swain, 22 Nov 1969; TMAG G6420,19 33 (20-24 mm), 9 juv. 33 (9-18 mm),
1999 (18-22 mm), 6 juv. 99 (14-19 mm), creeks entering Pine Lake, 41°44.6'S
146°42.0'E, coll. R. Swain, 3 Feb 1969; AMP97845,3 juv. 33 (7-12mm), 15 juv. 99
(6-11 mm), Halfmoon Creek, roadcrossing below Pine Lake, 41°45.01'S 146°42.75'E,
from rocky pools, hand sieves, 1159 m asl, TAS-516, coll. G.D.F. Wilson & S.J. Keable,
9 Mar 2001; TMAG 14371/G115,19 (29 mm), stream, 1 mile N of Rainbow Chalet,
Great Lake, 41°46.0'S 146°33.7'E, 3000 ft asl [914 m], coll. J. Pearson, Apr 1939; TMAG
G1309, numerous juv, Breona, 41 °47’S 146°42’E, 1060 m, from well, coll. G.E. Nicholls,

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Figure 21. Anaspides richardsoni sp. nov., selected features. (A) male, 23 mm, S of Lake Tyndall, TMAG G6312; (B) female, 23 mm,
Loongana, QVM 10:13975; (C) male, 22 mm, Deloraine, TMAG G135; (D) female, 26 mm, Little Pine Lagoon, QVM 10:13981; (E)
juv. male, 18 mm, Cradle Mountain, SAM C6303; (F) female, 28 mm, W of Lake Fox, QVM 10:49165; (G) female, 27 mm, Mt Ossa,
AM P82858; (H) female, 36 mm, Oatlands, TMAG G6360. Scale = 1.0 mm.

6 Feb 1945; TMAG G1310, 8 mature 33 (18-24 mm), (21-35 mm), Breona,
from well, coll. G.E. Nicholls, 6 Feb 1945; TMAG G1314, \3 (28 mm), 3 juv. 33
(12-17 mm), 1$ (26 mm), 2 juv. (11-15 mm), Breona, Great Lake, north-east, in
creek supply hotel, coll. G.E. Nicholls, 23 Dec 1943; TMAG G1311, 1$ (19 mm) and
numerous juv, Breona, from well behind Stewart’s house, coll. G.E. Nicholls, 6 Feb
1945; TMAG G1312,1 3 (20 mm) and numerous juv, Breona, from waterhole draining
creek, coll. G.E. Nicholls, 6 Feb 1945; WAMC11773,1 3 (c. 18 mm), 1 juv. 3 (c. mm),
T$ (22 mm), 3 juv. $ 9 (c. 10-16 mm), Stewarts Water Hole, Breona, coll. G.E. Nicholls,
8 Jan 1946; WAM C58164, Id' (18 mm), 3 juv. 33 (12-14 mm), 1$ (21 mm), 7 juv.
y V (6-15 mm), Breona, log behind (north) Stewart’s Cottage, 30 Jan 1947; WAM

Cl 1769,2 33 (18-22 mm), 2 juv. 33 (13-16 mm), 1 $ (21 mm), 5 juv. (7-20 mm),
creeks at Breona, Great Lake, 41°47'S 146°42'E, coll. Mr Stewart et al., 25 Jan 1947;
WAM Cl 1774, 233 (21-23 mm), 1 juv. 3 (17 mm), 1$ (18 mm), 2 juv. (9-10
mm) Breona, wells, coll. G.E. Nicholls, 6 Feb 1945; WAM C58158,27juv. 33 (11-18
mm), 43 juv. (8-17 mm), indet juv. (6-7 mm), Breona, Great Lake, puddles near
haulage, coll. G.E. Nicholls, 28 Jan 1947; WAM C11770, 8 33 (24-31 mm), 8$$
(27-42 mm), Brandons [=Brandum], Great Lake, 41°49.6'S 146°40.5'E, 1034 m asl,
Easter 1946; WAM C11775,1 juv. 3 (16 mm, 1 juv. $ (16 mm), Reynolds Neck, Great
Lake,41 0 51.TS 146°41.4'E, 1034 m asl, coll. A. Pike, 1935; TMAG G1630 ,433 (22-23
mm), 19 (23 mm), near Sh a nnon Lagoon, 41°59.6'S 146°44.1'E, pool in swamp, 1040

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Figure 22. Anaspides richardsoni sp. nov., selected features. (A-D) female, 30 mm, Clarence Lagoon, TMAG G6359; (E) juv. female,
13 mm, Clarence Lagoon, TMAG G6359; (F-I) male, 23 mm, Clarence Lagoon, TMAG G6359; (J-M) male, 30 mm, Pool of Bethesda,
TMAG G6434; (N-P) male, 23 mm, Lake Johnny, QVM 10:49162; (Q-S) Lake Fox, QVM 10:49164, female (29 mm), female (20 mm),
female (30 mm); (T-U) Lake Halkyard, QVM 10:49163, male (18 mm), female (27 mm). Scale: A-D, F-U =1.0 mm; E - 0.5 mm.

m asl, coll. J.W. Evans, 13 Dec 1936; TMAG G131,2 juv. ff (15-21 mm), mouth of
creek, Great Lake, coll. A.W.G. Powell, 29 Mar 1937; NMV J42435, 20 0 (25.0-36.0
mm), Great Lake, coll. F.E. Burbury, 1942; NMV J42443, 1$ (24.0 mm), Great Lake,
coll. F.F. Wilson, Jan 1933; AMP99298,2juv. SS (10-11 mm), 12juv. (8-14mm)
Sandbanks Tier, 41°50'28.69"S 146°51'10.95"E, 1150 m asl, stream from under
boulderfield, coll. S. Jarman; QVM 10:22606,1$ (34 mm), Lake River Valley, coll. C.

Spencer, 1997; WAM C58156, 33SS (18-31 mm), 48 juv. AS (12-18 mm), 23$$
(19-34 mm), 174 juv. ?? (9-18 mm), 5 miles N of Breona, 41°42.2'S 146°43.4'E,
creek, coll. G.E. Nicholls, 7 Feb 1945; NMV J42444,4juv. OO (12.0-16.0 mm), Ouse
River, 5 miles W of Miena, coll. A. Neboiss, 28 Feb 1967; USNM 1277684, 1 juv. S
(15 mm), Ouse River, 5 miles W of Miena, coll. A. Neboiss, 28 Feb 1967; TMAG G6369
(“spinulae” and non-“spinulae” forms), 8SS (19-22 mm), 4 juv. SS (15-17 mm), 10

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Figure 23. Anaspides richardsoni sp. nov., selected features. (A-C) Marakoopa Cave, SAM BS0464, female (31 mm), juv. female (21
mm), juv. female (9 mm); (D) male, 37 mm, Honeycomb Cave, AM P73038; (E) male, 25 mm, Kubla Khan Cave, USNM 1277679; (F)
female, 28 mm, Mayberry area, sink hole, Mole Creek, USNM 1277683; (G-H) Growling Swallet, female 41 mm (TMAG G6473), juv.
female 12 mm (TMAG G6481); (I) juv. female, 20 mm. Rift Cave, QVM 10:12158; (J) juv. female, 12 mm, Great Western Cave, Gunns
Plain, AM P99296. Scale: A, B, D-G = 1.0 mm; C, H-J = 0.5 mm.

$ $ (18-27 mm), 1 juv. $ (15 mm), Ouse River, semi creek, 22 Nov 1969; AM PI 1873,
1 $ (24 mm), 4-4 !4 miles S of Miena, in spring off road between Miena and Bothwell,
about 20 yards to left of road near old notice on tree, marked “water”, 41°59.8'S
146°47.2'E, coll. J. Waterhouse, c. 1930; TMAG G6428, 16 <$<$ (23-33 mm), 7$$
(32-35 mm), small stream running into Ouse River on Lake Auga Road, 41°52.61'S
146°36.28'E, drift sample, 1150 m asl, coll. P. Davies, Nov, 1985; AM P14158, 6 juv.

(slide preparations), near Little Pine Lagoon, 42°00.0'S 146°35.7'E, 1000 m asl, stream,
coll. W.D. Williams, 5 Feb 1963; QVM 10:13981, (19-25 mm), 1 juv. $ (18

mm), 79$ (22-26 mm), 5 juv. 9? (10-18 mm), about 10 miles from Great Lake on
Missing Link Road, between Great Lake and Bronte Park, from little stream on S side
of McKenzies Tier leading into Little Pine River, 42°02.2'S 146°33.0'E, coll. Sgt.
McIntyre, 1958 via Arthur Fleming; SAMAC8446,4juv. <$<$ (13-18 mm), 2$$ (36-41

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mm), 9 juv. $$ (9-21 mm), about 10 km N of Bronte Park, 42°04'S 146°29'E, from
small stream above Pine Tier Dam, coll. W. Zeidler, 16 Jul2001; AMP99297, \$ (21
mm), Pine Tier Lagoon, near shore, 42°04.38'S 146°29.30'E, coll. S. Richter & C.
Wirkner, 28 Feb 2006; ZSRO 372, 1 juv. 8 (11 mm), 7 juv. $ $ (6-12 mm), Pine Tier
Lagoon, near shore, 42°04.38'S 146°29.30'E, coll. S. Richter & C. Wirkner, 28 Feb
2006; ZSRO, 1$ (38 mm), tributary of Travellers Rest River, coll. A. Richardson, 4
Aug 2016; TMAG G6321,2 $ $ (21-28 mm), Clarence Lagoon, 40-50 m from outflow,
42°05.12'S 146°18.87'E, 980 m asl, coll. D. O’Brien & R. Kirkwood, 30 Mar 1990;
TMAG G6439, 288 (24-29 mm), 5$ $ (21-36 mm), 6 juv. $ $ (11-12 mm), Clarence
Lagoon, 42°05.2'S 146°19.2'E, on scuba, 980 m asl, coll. R. Mawbey 11 Jan 1984;
TMAG G6442, 388 (21-23 mm), 2$$ (26-29 mm), Clarence Lagoon, 42°05.12'S
146°18.87'E, stomach of brook trout, 980 m asl, coll. PA. Tyler, 5 Aug 1970; TMAG
G6355, 1$ (32 mm), Clarence Lagoon, 42°05.2'S 146°19.2'E, outflow creek, 980 m
asl, in FBA net, coll. R. Mawbey, 13 May 1985; TMAG G6359, 388 (22-24 mm), 1
juv. 8 (12 mm), 1 $ (30 mm; non-“spinulae” form), 1 juv. $(13 mm), Clarence Lagoon,
near start of Clarence River, 42°05.1'S 146°18.9'E, rocky shoreline, 980 m asl, coll. W.
Fulton, 22 May 1976; TMAG G6367(non-“spinulae” form), \88 (22-26 mm), 6 juv.
88 (18-21 mm), 2$$ (23-24 mm), 4 juv. $$ (10-21 mm), 2 indet juv. (7 mm),
moorland pools draining into Clarence Lagoon, 42°04.42'S 146°18.80'E, 980 m asl,
coll. R. Swain,21 Dec 1982; TMAGG6353,15juv. 88 (12-15 mm), 13juv. $$ (11-14
mm), Silver Plains Creek, Alma Tier, 42°07.19'S 147°04.57'E, 920 m asl, coll. R.
Mawbey, 11 Mar 1979; TMAG G6360, \$ (30 mm), 1$ (36 mm), “The Groves”,
Oatlands, 2 Mar 1969;AMP14160,1 8 (slide preparation), 1$ (slide preparation), near
Tarraleah, 42°17.2'S 146°26.3'E, 470 m, stream, coll. W.D. Williams, 7 Feb 1963.

Mt Field National Part OM Iv.1394, A88 (21-30 mm), 3$$ (21-32 mm), Mt Field,
4000 ft asl [1200 m]; AM P97846,1 $ (38 mm), 1 juv. $ (16 mm), Tam Shelf, Rodway
Range, coll. M.S. Moulds, 3 Feb 1992; AM P14159,4 specimens (slide preparations c.
16-18 mm), near Lake Dobson, 42°41'S 146°36'E, stream, coll. WD. Williams, 28 Jan
1963; AM P73058, \<$ (28 mm), Wombat Moor, Mt Field, spring head of small spring
fed sphagnum swamp on hillside, 42°40.98'S 146°36.48'E, TAS-484, coll. G. Wilson &

S. Keable, 4 Mar 2001; AM P73059, Id? (25 mm), 3$$ (19-25 mm), Wombat Moor,
Mt Field, spring head of small spring fed sphagnum swamp on hillside, 42°40.98'S
146°36.48'E, TAS-484, coll. G. Wilson & S. Keable, 4 Mar 2001; AM P97844,1 juv. 8
(15 mm), 9 juv, $ $ (6-15 mm), spring flowing out of hillside near ski lodge at bottom
of walking track skirting Lake Dobson and continuing to Mt Field West, 42°41.11'S
146°35.44'E, pool among water mosses, with phreatoicidean isopods, hand sieves, 1037
m asl, TAS-483, coll. G.D.F. Wilson & S. J. Keable, 4 Mar 2001; TMAG G6405,10 88
(25-33 mm), 5$ $ (28-38 mm), Mawson Plateau, Mt Field, 42°41,4'S 146°35. PE, 1270
m asl, coll. D. Hamilton, Apr 1970; TMAG G6371, 1 juv. c? 0 8 mm), Mt Field West,
42°39.5'S 146°30.7'E, 1400 m asl, coll. M. Fenton, 20 Jan 1971; TMAG G6401, 288
(27-29 mm), 5$ $ (26-38 mm), Newdegate Pass, Mt Field, 42°39.5'S 146°33.3'E, coll.
C. Reid, 23 May 1974; TMAG G6441, 6<SS (24-29 mm), 2 juv. 88 (9-10 mm), 6$$
(24—33 mm), between Mackenzie tarn and James Tam, 42°40.2'S 146°33.7'E, small
tarn, 1180 m asl, coll. T. Walker, Mar 1972; TMAG G133, 1$ (29 mm), Lake Fenton,
42°40.4'S 146°36.9'E, 1006 m asl, coll. Pott, Apr 1936; TMAG G6375, 388 (29-31
mm), 2 juv. 88 (15-25 mm), 1$ (33 mm), Mt Mawson, 42°41.7'S 146°35.8'E, 1280
m asl, coll. D. Hamilton, Apr 1970; TMAG G6444, 588 (25-29 mm), 1 juv. 8 (19
mm), 5$ $ (26-42 mm), 5 juv. $$ (8-22 mm), Mawson Plateau, 42°41.4'S 146 0 35.FE,
pool with mud bottom, 1267 m asl, coll. T. Walker, Mar 1972; TMAG G6415, 2 juv.
$ $ (13 mm), Mawson Plateau, 42°41,4'S 146°35.0'E, pool with rocky bottom, coll. T.
Walker, Mar 1972; TMAG G6384, A88 (25-30 mm), 7$$ (26-36 mm), 8 juv. OO
(12-20 mm), Mawson Plateau, 42°41.4'S 146°35.0'E, pool with rocky bottom, coll. T.
Walker, Mar 1972; TMAG G6432,1 8 (25 mm), 1 juv. 8 (20 mm), 1$ (31 mm), 6 juv.
$$ (8-23 mm), 1 indet juv. (7 mm), W side of Rodway Range, 42°40.2'S 146°32.4'E,
pool with muddy bottom, coll. T. Walker, Mar 1972; TMAG G6373, 3 indet juv. (7
mm), W side of Rodway Range, 42°40.2'S 146°32.4'E, pool with muddy bottom, coll.

T. Walker, Mar 1972; QVM 1043547, 2%88 (20-32 mm), 18$$ (20-41 mm), 1 juv.
$ (13 mm), Mawson Plateau, Mt Field, 42°41.4'S 146°35.TE, 1270masl, coll. C. Reid,
1975; TMAG G6314, 7$$ (25-36 mm), Robert Tam, Tam Shelf, Mt Field, 42°40.7'S
146°34.2'E, 1200 masl, coll. T. Walker, Mar 1972; TMAGG6372,10 88 (25-32 mm),
8 juv. 88 (16-21 mm), 9$$ (20-32 mm), 11 juv. $$ (10-22 mm), Robert Tam, Tarn
Shelf, Mt Field, 42°40.7'S 146°34.2'E, 1200 m asl, sample 1, tube 1(2), coll. I. Wilson
& J. Ong, 25 Jan 1970; TMAG G6408, 1 8 (31 mm), 1$ (38 mm), Mt Field, midway
between Newdegate Pass and Newdegate Tam, 42°39.6'S 146°33.6'E, in small tarn,
1200 m asl, coll. I. Wilson & J. Ong, 25 Jan 1970; TMAG G6423, 1 8 (27 mm), Tarn
Shelf, Mt Field, pool on lodge side, 42°40.1'S 146°33.6’E, 1260 m asl, coll. T. Walker,
Mar 1972; TMAGG6386,3 juv. 88 (15-18 mm), 2$$ (20-26 mm), 6juv. $$ (9-17
mm), Lake Dobson Road, 3 miles before Lake, 42°40.8'S 146°37.7'E, 1000 m asl, coll.
R. Swain, 18 Dec 1969; TMAG G6410, 4 88 (20-29 mm), 6 juv. 88 (10-17 mm),
12$$ (19-28 mm), 10 juv. $$ (11-14 mm), Tam Shelf, near James Tam, 42°40.3’S
146°33.8'E, small tarn with stream, sample 4, coll. I. Wilson & J. Ong, 25 Jan 1970;
TMAG G6421, 588 (22-26 mm), 4 juv. 88 (13-17 mm), 3$$ (24-30 mm), 5 juv.
$$ (12-18 mm), Tam Shelf, near Backhouse Tam, 42°40.1'S 146°33.6'E, sm a ll tarn,
sample 5, coll. I. Wilson & J. Ong, 25 Jan 1970; TMAG G6445, 588 (20-21 mm),
6 juv. 88 (13-19 mm), 2$$ (24-29 mm), 5 juv. $$ (13-15 mm), Newdegate Pass,
42°39.6'S 146°33.0'E, sample 1, coll. I. Wilson & J. Ong, 25 Jan 1970.
Junee-Florentine Karst. QVM 10:12158,1 juv. $ (20 mm), Rift Cave (JF34-5), Junee,
Florentine Valley, 42°44.3'S 146°35.6'E, 680 m asl, coll. S.M. Eberhard, 4 Jan 1985;
TMAG G6481,1 juv. 8 (12 mm), Growling Swallet Cave, Florentine Valley, 42°41,4'S
146°30.0'E, JF36-39,570 m asl, CV10, coll. S. Eberhard, 2 Jun 1985; TMAG G6473,
1$ (41 mm), Growling Swallet Cave (JF36), Florentine Valley, 42°41.4'S 146°30.0'E,
550 m asl, CV41, coll. S. Eberhard, 14 Apr 1985.

No data : AM PI 1897,2$ $ (31-39 mm), Tasmania, coll. B. Plomley via J. Waterhouse,
pre 1949;QVM 10:13484,1 8 (24 mm), 2$$ (21-24mm), unidentified cave, coll. S.
Eberhard, 71993; QVM 10:13976, 1 $ (31 mm), coll. E.O.G. Scott; USNM 1277686,
1 $ (31 mm), from G.M. Thomson, no data.

Description. Eyes with well-developed cornea, pigmented,
wider than stalk (epigean forms) to narrower than stalk in
some subterranean forms, longer than half length of stalk;
stalk with subparallel margins (except in some “spinose”
specimens). Rostrum narrow in adults, apex blunt.

Pleonites with sparsely setose pleural margins, rounded;
posterior margin of tergites 5-6 setose. Pleuron 1 unarmed.
Pleonites 2-6 usually with posterior margin of tergite 5
unarmed and pleura 2-5 unarmed or with 1 or 2 small spines
on pleuron 5; pleonite 6 posterior margin unarmed or minutely
spinose, posterolateral margin rounded, with or without minute
denticle. Pleonites 2-6 of “spinose” specimens with pleura
3-5 (sometimes also pleuron 2) strongly spinose; posterior
margin of tergites 5-6 strongly spinose; posterolateral margin
produced to distinct spine. Pleonal stemites 3-5 with low,
median processes between pleopod bases, distinctly bilobed
and widest on stemite 3, bilobed on sternite 4, weakly bilobed
and narrowest on sternite 5.

Telson slightly wider than long to longer than wide
(usually slightly longer than wide), pentagonal (usually) to
sub-linguiform, widest proximally; lateral margins sinuous
in dorsal outline, distally subparallel to convergent; transition
from lateral to posterior margin obtusely angular; posterior
margin angular to rounded, blunt medially; posterior spine
row with 19-50 slender, closely spaced spines, (usually)
short, evenly graded to distinctly uneven in some “spinose”
specimens.

Antennule inner flagellum about 0.2 x body length (21
articles in holotype); article 7 inner margin obtusely angled in
adult males, with 1 long, slender clasping spine at proximal
corner; outer flagellum 0.4-0.6 x body length (83 articles
in holotype) in epigean specimens, 0.4-0.9 x body length in
subterranean specimens. Antennal flagellum 0.4-0.6 x body
length (62-65 articles in holotype), 0.5-0.9 in subterranean
specimens; scaphocerite elongate, ovate, lateral spine slightly
distal to midlength; apex reaching almost to midlength of
distal peduncular article.

Right mandibular incisor process with proximal tooth
distally undivided to trifurcate, usually bifid.

Pleopods 1-5 (rarely 1^1) with endopod in adults. Adult
male pleopod 1 distally widened, scoop-like, lateral margins
weakly expanded, not obscuring retinacular lobe in lateral
view.

Uropodal protopod dorsally unarmed or with 1-3 spines;
exopod with 2-4 (usually 3) movable spines on outer margin
near position of partial diaeresis; exopod length about
2.5-3.5 times width, as wide as endopod, apex rounded,
narrow to relatively broad.

Etymology. Named in honour of Alastair Richardson, for his
many contributions to Tasmanian carcinology and limnology.

Measurements. Male (n = 756) 7-37, female (n = 1151)
6-55 mm, indet (n = 16) 5-7 mm.

Remarks. Anaspides richardsoni sp. nov. is characterized
by the combination of a single antennular clasping spine
in adult males (Fig. 19D), the telson posterior margin with
a close-set spine row (Fig. 19B) and the male pleopod 1
endopod having the retinacular lobe visible in lateral view
(Fig. 201). Additionally, the pleopod 5 endopod is present in
adults (except in some cave specimens from parts of the Mole
Creek and Junee-Florentine systems; see below). Anaspides
richardsoni is widespread across central Tasmania (Fig. 36)
in a wide arc stretching from the West Coast Range in the

Ahyong: Endemic Tasmanian Mountain Shrimps

345

northwest, eastwards to Cradle Mountain and as far north as
Gunns Plains and Mole Creek, across the Central Plateau, and
south to Mt Field. It is primarily an epigean species occurring
in springs, creeks and lakes, but also caves.

As presently understood, Anaspides richardsoni is the
most morphologically variable species of the genus. Through
the central-eastern (Mt Field east to Oatlands and north
to Deloraine) and north-western part of its distributions
(West Coast Range to Cradle Mountain and Mt Ossa
to Gunns Plains), however, A. richardsoni is relatively
uniform morphologically. The pleonites are almost always
unarmed (or with few spinules on the posterior margin
of pleonite 6), and in specimens from Mount Field to the
Great Lake-Deloraine area, the telson is about as long as or
little longer than wide, with the posterior margin distinctly
angular, approximately right-angled (Fig. 21). The telson
in specimens from the far north (Gunns Plains, Loongana,
Vale of Belvoir, Mole Creek) and western Central Plateau
(Walls of Jerusalem area, Western Lakes) tend to be more
elongate and often more acutely angular posteriorly (Figs
21B, F; 22). Specimens from the Mt Field area at the southern
extremity of its range generally have more slender uropods
than those from the northeast (most evident in large adults),
and a usually bifid proximal tooth on the right mandibular
incisor process (Fig. 19H) compared to the typically trifid
condition found in other areas. Populations of A. richardsoni
from the peripheries of the main distribution, however, show
additional notable variations, particularly those from the far
north, the western Central Plateau and the Mole Creek Caves.

Specimens from parts of Cradle Mountain and further
north (Black Bluff, Vale of Belvoir, Gunns Plains) have
a less angular telson than usual, sometimes being almost
rounded. At a relatively small size (18 mm) some Cradle
Mountain specimens already have a noticeably shortened
telson (slightly wider than long; Fig. 2IE), broad uropods
and broad scaphocerite compared to size-matched specimens
from elsewhere.

Few specimens are known from the West Coast Range (Mt
Murchison), and are the most westerly specimens examined
(Fig. 21 A). All have minimal pleonal spination (at most with
few small spines on the posterior margin of pleonite 6). Most
known males are juveniles with a single antennular clasping
spine, although the largest male has (TMAG G6411) has one
antennular clasping spine on the left and two on the right.
The right antennule (peduncle and flagellum), however, is
shortened overall and appears abnormal, possibly as a result
of developmental irregularities. Smith’s (1909a) report of A
tasmaniae from Mt Read (near Mt Murchison, West Coast
Range) is probably referable to A. richardsoni.

A number of specimens of A. richardsoni from the
northwestern Central Plateau encompassing the Walls of
Jerusalem and Clarence Lagoon east to the Ouse River have
strongly spinose pereonites as in A. spinulae from Lake St
Clair —the A. “spinulae” of O’Brien (1990) (Fig. 22). In
these specimens, pleonite 4-5 pleura (usually also pleonite
3) and the posterodorsal margins of pleonites 5 and 6 (5
less often so) are prominently spinose and the posterolateral
margin of pleonite 6 is usually produced to a prominent
spine. The proximal tooth on the right mandibular incisor
process is usually trifid. In addition, the telson is often more
elongate than usual, with a sometimes somewhat rounded
margin on which the posterior spines vary from evenly
graded to distinctly uneven in length. Telson spination differs

slightly between sampled lakes and the Walls of Jerusalem.
Specimens from Lakes Fox and Johnny (Fig. 22N-S)
have relatively uneven telson spination compared to those
from Clarence Lagoon (Fig. 22A-I), Lake Halkyard (Fig.
22T,U), Jacks Lagoon and most specimens from the Walls
of Jerusalem (Fig. 22J-M), which have more evenly graded
telson spines like those of typical epigean forms. It should
be noted, however, that most specimens found throughout
this area are non-spiny or less-spiny than “ spinulae ” (in
which only pleura 4-5 and the dorsal margin of pleonite 6
may be spinose, e.g., Jacks Lagoon, QVM 10:13977). Both
spiny and non-spiny forms were found together in tarns and
runnels in the Walls of Jerusalem (TMAG G6396, TMAG
G6379), the vicinity of Lake Fox (Fig. 21F), the Ouse River
(TMAG G6359) and Clarence Lagoon (TMAG G6359) (Fig.
22A-I). The strong similarity in pleonal spination of the
most spinose specimens of A. richardsoni to A. spinulae is
remarkable, suggesting similar underlying developmental
processes. O’Brien (1990), noting a correlation between
spininess and lake habitats, suggested that A. spimdae might
be a separate lacustrine species or lacustrine ecomorph,
given that non-spiny specimens seemed to be from creeks,
runnels and tarns. If so, however, it would remain to be
explained why A richardsoni from other lakes, such as Pine
Tier Lagoon and Pine Lake are non-spiny, and why spiny
forms can also be found together with non-spiny forms in
tarns, small creeks, and pools. Both non-spiny and spiny
forms are recorded from Clarence Lagoon (Fig. 22A-I).
Additionally, the “ spinulae ” form of A. richardsoni occurs
only on the Central Plateau to the east and northeast of the
similarly spinose A. spinulae from Lake St Clair, but not
in other lake populations of A. richardsoni (e.g., lakes at
Mt Field and the West Coast Range such as Lakes Tyndall
and Sandra), let alone lake populations of other species of
Anaspides such as A.jarmani and A. swaini. The presence of
predatory fish, such as trout and Galaxias might be posited
to induce a spiny “defensive” phenotype in Anaspides as
has been observed in Daphnia (Adler & Harvell, 1990).
Trout are certainly present in Lake St Clair (A. spinulae
sensu stricto), Clarence Lagoon, Lakes Fox, Halkyard and
Johnny, but they are also present in more easterly water
bodies such as Pine Tier Lagoon, Little Pine Lagoon and
Pine Lake in which A. richardsoni shows no such defensive
spination. Instead, the distribution of the “ spimdae ” form
of A. richardsoni correlates more strongly with geography
than ecology, and might reflect phylogeographic genotypic
variation. In particular, the relationship between these
spinose A. richardsoni and A. spinulae from Lake St Clair
warrants further investigation, with A. spinulae differing
most fundamentally in having two rather than one antennular
clasping spines. Curiously, specimens to the immediate
south and west of Lake St Clair, all referrable to A. swaini ,
do not have the spinose pleon of A spinulae. Specimens to
the immediate east of Lake St Clair from the eastern side
of the Traveller Range are non- or minimally spinose as in
typical specimens of A richardsoni. In addition, the telsons
of both spiny and non-spiny specimens from the Walls of
Jerusalem to the northern tiers (as well as Mole Creek) tend
to be more elongated than specimens from elsewhere. More
detailed population level studies, currently underway in
collaboration with S. Richter and colleagues, are required to
further understand these morphological patterns.

Anaspides richardsoni has entered subterranean habitats in

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Records of the Australian Museum (2016) Vol. 68

at least three parts of the periphery of its range (Fig. 23): Mole
Creek karsts and Great Western Cave (Gunns Plain) in the
north, and the Junee-Florentine system in the south. The single
known specimen from Great Western Cave (AM P99296; Fig.
23 J) is a juvenile male that agrees well with epigean juveniles
(e.g., TMAG G6169); it also represents the northernmost
record of Anaspides. Aside from loss of body pigmentation,
the specimen exhibits no obvious troglobitic adaptations and
has well-developed, pigmented eyes. At Mole Creek, three
subterranean forms occur, differing between cave systems.

• Form 1 occurs in the Sassafras (Prohibition Cave) and
Marakoopa systems (Marakoopa I and Marakoopa II
caves) (Figs 23A-C, 35F). Apart from reduced body
pigmentation, it closely resembles epigean specimens,
having well-developed eyes, the outer antennular
flagellum 0.4-0.6 (usually 0.5-0.6) body length, unarmed
pleonites except for scattered denticles along the upper
posterior margin of pleonite 6, and in having a finely-
graded spine row on the posterior margin of the telson.

• Form 2 is found in caves in the Mole-Lobster system
(Honeycomb Cave, Wet Cave, Herberts Pot and Kellys
Pot) (Figs 23D, 35E). It resembles Form 1 in telson
structure, but has reduced eyes, outer antennular flagella
0.5-0.8 (usually 0.6) body length, and more extensive
pleonal spination, with denticles on the pleura of pleonites
4-5 as well as the dorsal posterior margin of pleonite
6. Form 2, reaches the greatest size of the three forms
(>50 mm), the largest being from Honeycomb Cave.

• Form 3 occurs in the Kubla Khan system (Kubla Khan
Cave) and an unnamed cave in the Sassafras system (Fig.
23E,F). It resembles Form 1 in eye development and
minimal pleonal spination, but differs from both Forms
1 and 2 in consistently having much more elongate
antennular flagella (0.7-0.9, usually 0.8) and a more
slender telson with longer, more robust spines, somewhat
approaching that of A. richardsoni from the Walls of
Jerusalem and localities close to Mole Creek such as
Western Bluff, and Ironstone Mountain. Additionally,
the pleopod 5 endopod is frequently absent in specimens
from the Kubla Khan system. One series corresponding
to Form 3 (USNM 1277683; Fig. 23F) was collected
from a deep, water filled sink hole in the Mayberry
area (near Mole creek), believed to be hydrologically
connected to the Kubla Khan system (Iliffe, 1988);
the specimens differ from the Kubla Khan specimens
only in having more extensive body pigmentation
typical of epigean forms, and a slightly wider cornea.

Although most common on the surface around Mt Field, A.
richardsoni is recorded from the Junee-Florentine system
(Rift Cave and Growling Swallet; Fig. 23G-I) on the basis
of two juveniles and an adult female. These specimens agree
well with epigean A. richardsoni , including pigmentation
(albeit seemingly somewhat reduced) and well-developed
eyes, but differ in lacking the pleopod 5 endopod. The
right eye of the Rift Cave specimen is deformed, seemingly
having been damaged (Fig. 231). Two other species of
Anaspides occur in caves in the Mt Field area: A. eberhardi
(Junee-Florentine karsts including Growling Swallet), which
is closely related to A. richardsoni , and A. swaini (Junee-
Florentine and Risby’s Basin karsts), which also sometimes
occurs on the surface at Mt Field, but is the dominant epigean
species to the south and west of the area.

Anaspides richardsoni features well-developed secondary
sexual characteristics by 18-25 mm (usually 20-22 mm).
The pleopod 5 endopod is the last of the endopods to
develop, appearing in juveniles as small as 11 mm to as
large as 22 mm, but usually by 15-18 mm. The presence of
a single male antennular clasping spine is highly consistent,
with, in rare cases, a second spine developing on one side,
often associated with possible regeneration after damage.
The male from Wet Cave, however, is highly aberrant in
having two clasping spines on one side, three on the other.
In both cases, the normal single clasping spine is present
on the proximomedial corner of flagellar article 7, but the
additional abnormally and asymmetrically developed spines
arise on the anteromedial comer. The abnormal positions and
asymmetrical development of the clasping spines in the Wet
Cave male indicate that it is aberrant.

Anaspides richardsoni is the most taxonomically
challenging species of the genus given the range of forms
encountered, especially those from the peripheries of its
range with distinctive morphologies. These may carry
the signature of populations from areas untouched by
the Pleistocene glaciations, which otherwise dominated
the Central Plateau at the time, or may indicate ongoing
differentiation (Andrew, 2005). Further sampling is required
to better understand these peripheral populations, especially
since some localities, such as Gunns Plain, the West Coast
Range and Sandbanks Tier are represented by juveniles or
only few specimens. Further sampling is also required in
the eastern Great Western Tiers in the vicinity of Deloraine,
Great Lake, and Lake Sorell down to the Oatlands area to
determine the current extent of occurrence. The specimens
from the Oatlands area (Fig. 21H) are the easternmost
records of the genus, although O’Brien (1990) speculated
their provenance to be from surrounding hills slightly to the
west. Likewise, specimens from Deloraine were speculated
to originate from hills slightly to the south (O’Brien, 1990).
The known distribution A. richardsoni is discontinuous
between the West Coast Range and Cradle Mountain area,
and between the Western Lakes and Mt Field. Whether
these “gaps” are real or owe to lack of sampling remains to
be determined. Despite the wide morphological diversity,
A. richardsoni is readily diagnosed by the combination of
the single antennular clasping spine in adult males and the
close-set spine row on the posterior margin of the telson.

Distribution. Wide ranging in central Tasmania, from the
West Coast Range and Cradle Mountain in the west to the
Central Plateau from Mt Ossa and localities east of Lake St
Clair, to Mole Creek and the vicinity of Great Lake, south
to Mt Field and as far east as the Oatlands area; 470-1455
m asl (epigean), 109-520 m asl (subterranean).

Anaspides spinulae Williams, 1965

Figs 24-28, 36

Anaspides tasmaniae. —Powell, 1946: 84.

Anaspides spinulae Williams, 1965a: 117-123, fig. 5 (type
locality: Lake St Clair, S of pumping station). —Williams,
1974 : 84-85, tab. 4.1. —Knott, 1975: 157 (Lake St
Clair specimen only), 173. —Michaelis, 1985: tab. 2.

—Richardson, 1985: 3. —O’Brien, 1990: 11-18, tab.

2.2, 2.4, pi. 2. —Jarman & Elliot, 2000: fig. 4, tab. 1. —
Camacho et al., 2002: fig. 1, tab. 1. —Lake et al ., 2002:

11. —Serov, 2002: 8, 15. —Camacho, 2006: 4.

Ahyong: Endemic Tasmanian Mountain Shrimps

347

Figure 24. Anaspides spinulae Williams, 1965, holotype female, 25 mm,
Lake St Clair, AM P14146. Habitus, right lateral view. Scale 2.0 mm.

Type material. Holotype: AM P14146,? (25 mm), Lake St Clair, S of pumping
station, 42°06.6'S 146°12.0'E, 3^4.5 m, coll. J.H. Wilson & W.D. Williams, 8 Feb
1963. Paratypes: AMP14147,1 juv. S (12 mm), type locality; AMP14148,1^ (slide
preparation), type locality; AM PI4149,1 specimen (slide preparation), type locality;
AMP14150, (slide preparation); AM P14151,1 specimen (slide preparation), type
locality; AM PI4152,4 specimens (slide preparation), type locality; AM P14153,1
(slide preparation), type locality; AM P14154,4 specimens (slide preparation), type
locality; AM P14155, 1 juv. $ (c. 11 mm), type locality; AM PI4156, l<j> (21 mm),
1 juv. S (c. 11 mm), type locality.

Other material examined, ypm 9194, 2 $$ (13-14 mm), 1$ (is mm),

Lake St Clair, coll. J.H. Wilson, Mar 1961; TMAG G124, 2 juv. SS (11-13 mm),
599 (9-18 mm), Lake St Clair, bottom drag at midnight, coll. A.W.G. Powell, 24 Jul
1937; TMAG G134, 2 juv. <$<$ (10-12 mm), Cynthia Bay, Lake St Clair, 42°06.6'S
146°10.TE, bottom drag, coll. A.W.G. Powell, 26 Aug 1937; AMP99312,2 juv. <S<3
(8-9 mm), Lake St Clair, 42°06.75'S 146°11,81'E, 5 m depth, Lake bottom, towed
net, 730 m asl, coll. S. Jarman; AM P99313, 16 juv. $$ (7-9 mm), 11 juv. $ $ (7-10
mm), Lake St Clair, S of Pumphouse Point, 42°06.38'S 146°12.10'E, 2 m, weed bed,
on scuba, 730 m asl, coll. M. Driessen & J. Andrew; MCZ IZ:68029,1 juv. $ (9 mm),
1 juv. ^ (8 mm), Lake St Clair, S of Pumphouse Point, 42°06.38'S 146°12.10'E, 2
m, weed bed, on scuba, 730 m asl, coll. M. Driessen & J. Andrew, late 1990s to pre
2005; TMAG G6325, 1<? (23 mm), 1$ (23 mm), Ida Bay, Lake St Clair, 42°01.7'S
146°08.5'E, 3-7 m, sand & under rocks, stones, plentiful to 7 m, 737 m asl, coll. R.
Holmes, 20 Mar 1990; TMAGG6327, 1$ (22 mm), 2 juv. £$ (10-11 mm). Lake St
Clair, 42°01.75'S 146°06.94'E, 6 m, between rocks & stones, “No. 4”, coll. D. O’Brien,
20Mar 1990; TMAGG6328 ,\$ (14mm), 1$ (20 mm),2juv. g$ (9-11 mm), Lake
St Clair, 42°04.63'S 146°10.02'E, under logs, 6-12 m, coll. D. O’Brien & M. Driessen,
20 Mar 1990; TMAGG6326,1 juv. <J (11 mm), Lake St Clair, 42°05.08'S 146°11,76'E,
6 m, pebbles and stone outcrops, coll. R. Holmes, 20 Mar 1990; TMAG G6324,3 juv.
$$ (9-10 mm), 5 juv. £ $ (8-9 mm), Lake St Clair, 42°06.70'S 146°11,74'E, 3-4 m,
amongst Isoetes, under stones, rocks, site 9, coll. D. O’Brien, 20 Mar 1990; TMAG
G6323, 2 juv. SS (10-11 mm), 2 juv. (11-12 mm), Lake St Clair, 42°01.04'S
146°06.74’E,4-6m, stones and pebbles, coll. R. Holmes, 20 Mar 1990; TMAGG6329,
1 juv. $ (8 mm), l<j> (20 mm), no locality data.

spaced spines, with several longer spines, approximately
evenly spaced among shorter spines; rarely with dorsomedian
spine above posterior margin.

Antennule inner flagellum about 0.2 x body length (22
articles in holotype); article 7 inner margin obtusely angled in
adult males, with 2 long, slender clasping spines proximally;
outer flagellum 0.5-0.6 x body length (67-71 articles in
holotype). Antennal flagellum 0.3-0.5 x body length (49-50
articles in holotype); scaphocerite elongate, ovate, lateral
spine slightly distal to midlength; apex almost reaching or
slightly overreaching apex of distal peduncular article.

Right mandibular incisor process with proximal tooth
distally bifid or trifid.

Pleopods 1-5 with endopod in adults. Adult male pleopod
1 distally widened, scoop-like, lateral margins weakly
expanded, not obscuring retinacular lobe in lateral view.

Uropodal protopod dorsally with 1 or 2 small spines;
exopod with 2-4 movable spines on outer margin near
position of partial diaeresis; exopod length about 3 times
width, as wide as endopod, apex rounded, relatively narrow.

Measurements. Male (n = 35) 7-23 mm, female (n = 36)

Description. Eyes with large well-developed cornea,
pigmented, distinctly wider than stalk, subglobular, longer
than half length of stalk; stalk with divergent margins.
Rostrum narrow in adults, apex blunt.

Pleonites with sparsely setose pleural margins, rounded;
pleuron 1-2 with 0-3 small spines; pleura 3-5 prominently
spinose. Pleonites 5-6 posterior tergal margins prominently
spinose, setose. Pleonite 6 posterolateral margin produced to
prominent spine, occasionally with secondary spine. Pleonal
sternites 3-5 with low, median processes between pleopod
bases, distinctly bilobed and widest on sternite 3, bilobed
on sternite 4, weakly bilobed and narrowest on sternite 5.

Telson longer than wide, widest proximally; lateral
margins sinuous in dorsal outline, distally convergent;
transition from lateral to posterior margin obtusely angular;
posterior margin angular to slightly rounded, blunt medially;
posterior spine row with 17-32 slender, uneven, closely

7-25 mm.

Remarks. Williams (1965a) distinguished A. spinulae
from A. tasmaniae on the basis of its pronounced pleonal
spination, particularly of pleura 3-5 and tergites 5-6, uneven
telson spine row, and apparently unusual ecology—the
lacustrine habitat of Lake St Clair. Anaspides tasmaniae
(now known to comprise several species) was thought, at
the time Williams wrote, to occupy only surface creeks,
tarns and streams, but subsequent discoveries from lakes
throughout Tasmania refuted that assumption (Williams,
1974). The validity of A spinulae has been questioned ever
since (Williams, 1974; O’Brien, 1990; Jarman & Elliott,
2000). In particular, the observation of a possible “gradation
of increasing spination in a southerly direction across the
western portion of the Great Western Tiers” down to Lake St
Clair cast doubt on the validity of A. spinulae (Knott, 1975;
O’Brien, 1990). Moreover, Anaspides throughout much of

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Records of the Australian Museum (2016) Vol. 68

Figure 25. Anaspides spinulae Williams, 1965, holotype female, 25 mm, Lake St Clair, AM P14146. (A) cephalothorax, dorsal view;
(B) pleonite 6, telson and right uropod; (C) pleonites 1-6 pleura, right lateral view; (D) right antennule; (E) right antenna; (F) labrum,
anterior view; (G) right mandible; (H) right mandible incisor process; (I) paragnaths, anterior view; (J) right maxillule; (K) right maxilla.
Scale: A-E = 1.0 mm; F-K = 0.7 mm.

Tasmania show some degree of pleural and tergal spination,
including d. tasmaniae from Mt Wellington, further eroding
the distinctiveness of A spinulae (see O’Brien, 1990). Others
have accepted a limited range ford, spinulae , from Lake St
Clair to Clarence Lagoon (Richardson, 1985; Swain, 2000).
The status ofd. spinulae has remained debated, partly owing
to the seemingly unreliable diagnostic features identified by
Williams (1965a), and partly owing to the inadequate and

perhaps misleading original account of the species, in which
the holotype was not figured.

Review of all available material from Lake St Clair,
as well as western Central Plateau specimens reported by
Knott (1975) and O’Brien (1990) as A. “ spinulae ” (= A.
richardsoni sp. nov.; Fig. 22), indicates thatd. spinulae is a
valid species. The present concept ofd. spinulae emphasizes
the male secondary sexual characters in addition to the

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349

Figure 26. Anaspides spinulae Williams, 1965, holotype female, 25 mm, Lake St Clair, AM P14146. (A) right thoracopod 1 (maxilliped);
(B-H) right thoracopods 2-8; (I) right pleopod 5, anterior view; (J-L) pleonites 3-5 median sternal processes; (M-N) female gonopore,
right lateral and ventral views. Scale =1.0 mm.

pleonal spination. Thus, A. spinulae can be distinguished
from all other species of the genus by the combination of
two antennular clasping spines in adult males (Fig. 27D),
prominently spinose pleural margins of pleonites 2-5 and
posterior tergal margins of pleonites 5-6, the prominently
spiniform posterolateral angle of pleonite 6 and angular
posterior margin of the telson (Figs 25, 27, 28). The cornea
of A. spinulae is also proportionally more inflated than any

of its congeners at a similar size, being noticeably wider than
the stalk (Figs 25A, 27A, 28A,D).

Despite A. spinulae being a valid species clearly separated
from/l. tasmaniae , its taxonomic boundaries remain to be fully
circumscribed. The secondary sexual modifications of the male
antennule and presence of the pleopod 5 endopod suggest a
close relationship to the northern form of A. swaini occurring
to the immediate west and south of Lake St Clair. Although

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Records of the Australian Museum (2016) Vol. 68

Figure 27. Anaspides spinulae Williams, 1965, male, 14 mm, Lake St. Clair, YPM 9194. (A) cephalothorax, dorsal view; (B) pleonite
6, telson and left uropod; (C) pleonites 2-6 pleura, right lateral view; (D) right antennule; (E) right scaphocerite; (F-G) right pleopod 1
endopod, lateral and ventral views; (H-I) right pleopod 2 endopod, lateral and ventral views. Scale = 0.5 mm.

Williams (1965a) hypothesized that A spinulae survived the
Pleistocene glaciation of Lake St Clair in adjacent periglacial
lakes or melt-waters, areas currently occupied by A swaini, A.
spinulae might equally have persisted in-situ in deeper parts
of the lake during that time. This more easily accounts for
the very limited range of the species today, nested between
the ranges of A. swaini and A. richardsoni. Knott (1975)
alternatively hypothesized A spinulae to be only a temporary
resident in Lake St Clair, being periodically flushed into the
lake from adjacent creeks and streams, with its primary habitat
being the western Central Plateau. Present results, however,
indicate otherwise. The nearest neighbouring Anaspides
populations to the immediate south and west (A swaini ) and
immediate east (A richardsoni , eastern side of the Travellers
Rest Range) of Lake St Clair have “normal”, non-spiny pleonal
ornamentation. The strongly spinose forms (A. “spinulae”
of O’Brien, 1990) occur further afield on the western Central
Plateau to the east and northeast of Lake St Clair, and although
closely resembling A spinulae , are referable to A. richardsoni
having one instead of two male antennular clasping spines. It
is enigmatic that the strongly spinose “spinulae” morphology
is known only in Anaspides from Lake St Clair and the
western Central Plateau, whereas the nearest neighbours to
the lake all have “normal” pleonal ornamentation. As noted
under the account of A. richardsoni , the occurrence of the
spiny body form does not have an immediate ecological
basis, relating neither to a lacustrine habitat nor presence
of fish predators. The observed morphological patterns in
Anaspides from west to east of Lake St Clair are presently

difficult to interpret, and the limited currently available
molecular data are equivocal (Jarman & Elliott, 2000; Andrew,
2005). Thus, the relationship of A. spinulae to neighbouring
Anaspides populations, currently assigned to A. swaini and
A. richardsoni, respectively, requires more detailed analysis
beyond the scope of the present study. Although conceivably
more wide ranging, A. spinulae is presently known with
certainty only from Lake St Clair.

Previous records of A. spinulae from Clarence Lagoon
and elsewhere on the Central Plateau are referable to A.
richardsoni. Although resembling A. spinulae in pleonal
spination, adult males from Clarence Lagoon, like other
spiny Central Plateau forms have a single antennular clasping
spine, diagnostic of A. richardsoni. It is notable though that
the corneas of spiny A. richardsoni from Clarence Lagoon
(Fig. 22F), like true A. spinulae , are proportionally larger
and more expanded than size-matched A. richardsoni from
elsewhere on the Central Plateau (including a non-spiny
specimen from Clarence Lagoon; Fig. 22A).

Anaspides spinulae apparently matures at a smaller size
than congeners. Development of antennular modifications
begins in juvenile males at 11-13 mm, with clasping spines
appearing sequentially and with increasing curvature of the
proximal portion of the flagellum. By 14—15 mm body length,
the male secondary sexual characters of A. spinulae are
well-developed suggesting sexual maturity. Males of other
species of Anaspides do not attain similar development until
18 mm or larger. Possibly also significant is that the largest
known specimen of A. spinulae , at 25 mm, is considerably

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351

Figure 28. Anaspides spinulae Williams, 1965, selected features, Lake St. Clair. (A-C) juv. female, 10 mm, AMP99313; (D-F) paratype
female, 21 mm, AM P14156; (G) paratype, sex indet, c. 10 mm, on slide, AM P14152; (H) paratype male, c. 13 mm, AM P14147; (I)
paratype female, +20 mm, on slide, AM P14148. Scale: A-C, G-H= 0.5 mm; D-F, 1=1.0 mm.

smaller than the largest specimens of congeners (35 mm,
A. tasmaniae ; 38 mm, A. darker, 47 mm, A. eberhardi ; 31
mm,A.jarmani; 55 mm, A. richardsoni, 40 mm, A. swaini).

Specimens of A. spinulae are in most respects morpho¬
logically uniform apart from typical allometric variation in
the slenderness of the rostrum (increasingly slender with
increasing size), eye size (proportionally larger in smaller
specimens), and pleonal spine length (proportionally longest
in the smaller specimens). The endopod of pleopod 5 is
absent or rudimentary in juveniles up to 10 mm body length;
present in all others. Pleonal spination is pronounced, even
in the smallest juveniles, with pleural spines always present
on pleonites 3-5, and those on pleonite 2 appearing at about
20 mm body length. The most significant allometric changes
are in the spination of the posterior margin of the telson.
The telson spination in the smallest specimens is markedly
uneven, with 4-6 long spines evenly distributed among
the shorter remaining spines (Figs 27B, 28C,G,H). With
increasing body size, the longer telson spines become shorter
and more similar to surrounding spines, as in the holotype,
somewhat approaching adults of other epigean species (Figs

25B, 28F,I). Similar, albeit less marked changes, are also
evident in other species, such as A. richardsoni and A. swaini,
indicating a general developmental pattern. Apart from
allometric changes,^, spinulae is morphologically uniform,
with few other observed variations. The posterolateral spine
on pleonite 6 may be accompanied by a secondary spine, and
the uropodal protopod and the outer margin of the uropodal
exopod have 1 or 2 and 2-4 spines, respectively.

Unfortunately, Williams’ original characterization of
A. spinulae as having a strongly uneven spine row on
the telson was misleading, because his accompanying
unsealed illustration (Williams, 1965a: fig. 5E) was of
a small dissected juvenile (estimated c. 10 mm body
length; AM P14152; Fig. 28G) in which the differences
in spine length are most pronounced. At any given size,
the posterior telson spines in A. spinulae are less regular
in length than in other epigean Anaspides of similar size,
but not nearly as pronounced as would be assumed from
Williams’ figure. Subsequent reports (e.g., O’Brien, 1990)
of telsons “intermediate” between A. spinulae and typical
A. tasmaniae were based on the implication from Williams’

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Records of the Australian Museum (2016) Vol. 68

(1965a) figure that an adult was depicted. Further, Williams’
(1965a) description of the pleonal setae of both A. tasmaniae
and A. spinulae as spines further clouded distinguishing the
two forms. This possibly contributed to some of O’Brien’s
(1990) difficulties in distinguishing between what he called
A. spinulae , A. “spinulae ” and A. tasmaniae leading to
the strong suggestion that all represent the same species.
O’Brien (1990) also reported Central Plateau specimens
with “longer setae... ”, including specimens from Butlers
Gorge. As already noted, however, the length of pleonal setae
varies allometrically in Anaspides and is shortest in adults,
as corroborated in specimens from Butlers Gorge examined
herein (referrable to A. swaini). Whereas the posterior telson
ornamentation of adult A. spinulae does differ from most
other congeners in length and regularity, it is not nearly as
marked as originally implied by Williams (1965a).

O’Brien (1990) recorded A. spinulae from around the
margins of Lake St Clair at about 1.5-15 m depth on
multiple substrate types. It apparently does not occur on
silted substrates, but is common on stony or pebbled outcrops
and among exfoliating sheets of rock on weathering dolerite
boulders, with highest densities under rocks, fallen logs and
branches, or in Isoetes algal beds.

Distribution. Presently known only from Lake St Clair;

l. 5-15 m depth; 737 m asl.

Anaspides swaini Ahyong, 2015

Figs 29-36

Anaspides tasmaniae. —Smith, 1909a: 64, 70 (Northwest
Bay River). —Manton, 1930: pi. 2-3. —MacBride,
1930: 1079, unnumbered fig.—Tjonneland etal, 1984:
226, figs. 1-10 (heart ultrastructure). —Desmarchelier
& Clarke, 1998: 14. —Jarman & Elliot, 2000: fig. 4
(clade F), tab. 1 (part, Mt Anne, Snowy North, Weld
River). —Clarke, 2000: 33. —Doran et ah, 2001, tab. 2
(Bill Nielson Cave). —Camacho et ah, 2002: fig. 1, tab.
1—Clarke, 2006: fig. 1.16.

Anaspides sp. (telson ‘normal 5 type). —Eberhard etal., 1991:

48 (Junee-Florentine).

Anaspides sp. (telson type intermediate). —Eberhard et al.,
1991: 48 (Deep Thought Cave, Capricorn Cave).

Anaspides swaini Ahyong, 2015: 598: fig. II—L.

Type material. Holotype: AM P73042,c 9 (27 mm), Weld River, 42°48.78'S
146°27.49'E,460masl, coll. S. Jarman. Paratypes: AMP73043, 299 (22-28 mm),
12 juv. $ 9 (8-16 mm), 4 indet juv. (5-7 mm), type locality.

Other material examined Cradle Mountain Lake St Clair National Park.
TMAG G6370, 1<? (22 mm), 399 (24-34 mm), 2 juv. 9f (13-17 mm), Mt Byron,
rainforest creek flowing from Byron Gap into Lake Petrarch, 42°02.4'S 146°03.9'E,
coll. R. Mawbey et al., 920 m asl, 1970; TMAG G6378,4 juv. f9 (12-14 mm), just
downstream from Lake Petrarch, 42°03.4'S 146°06.1'E, 880 m, coll. R. Mawbey et
al, 29 Sep 1973; TMAG G6377, 6 juv. A A (11—43 mm), 5 juv. 99 (10-12 mm),
Cuvier Valley below Mt Olympus, 42°03.8'S 146°06.8'E, subterranean pools in
rainforest floor, 875 m asl, coll. R. Mawbey etal., 29 Sep 1973; TMAG G6426, 2SS
(23-28 mm), 6 juv. A A (14-18 mm), 399 (25-29 mm), 6 juv. 99 (15-18 mm),
Cuvier Valley, below Lake Petrarch, 42°04,8'S 146°08.6'E, small creek in buttongrass
plain, 800 m asl coll. R. Mawbey etal, 28 Sep 1973; TMAG 14388/G132, \A (22
mm), (21-24 mm), “Lake St. Clair?”, coll. D. Handley, 1937; TMAG G126,
SAA (14-23) mm), 599 (12-32 mm), Lake St Clair, coll. D. Turner, Feb 1941; AM
P72844, 1 juv. A (22 mm), 1 juv. 9 (13 mm), Mt Rufus, from tarn below summit,
42°07.18'S 146°06.42'E, DP265344(8113), 292-23, coll. A. Terauds, 16 Feb 1992;
AM P72842, 1 juv. <3* (18 mm), 4 juv. AS (6—7 mm), 1? (30 mm), 6 juv. 99 (6-16
mm), Mt Rufus, 42°07.20'S 146°06.43'E, 1170 m, stream flowing into Lake St Clair,
coll. S. Jarman; AM P82856, \A (19 mm), 1$ (21 mm), Mt Rufus, canal near wooden
bridge, 42°09.0'S 146°07.2'E, JHB T0201, coll. J.H. Bradbury, 2 Mar 1997; QVM
10:13979, If (38 mm), Mt Rufus, 42°07.6'S 146°06.0'E, tarn plateau, 1380 m asl,
coll. L.D. Crawford, 23 Sep 1951; TMAG G6390, 3AS (20-21 mm), 1 juv. A (17
mm) 699 (20-32 mm), 1 juv. $ (16 mm), Mt Rufus, 42°07.6'S 146°05.9'E, 1400

m, coll. B. Knott, 13 Nov 1971.

Wentworth Hills'. QVM 10:49166, 1 A (23 mm), 399 (21-33 mm), 1 juv. A (13 mm),
8 juv. 99 (9-12 mm), Wentworth Hills, 42°12'S 146°19'E, inflow, coll. B. Mawbey,
28 Mar 1990; QVM 10:49167, 3SS (21-25 mm), l A juv. (18 mm), 7$ $ (20-38 mm),
8 juv. 9 9 (9-19 mm), Wentworth Hills, 42° 12'S 146° 19'E, lower soaks, in flow, coll.

B. Mawbey, 28 Mar 1990; TMAG G6311, 2 AS (22-24 mm), 5ff (22-30 mm), 4
juv. A A (10-18 mm), 18 juv. 99 (8-19 mm), Wentworth Hills, stream draining into
Laughing Jack Lagoon, 42°12.47'S 146°19.15'E, 1030 m asl, coll. D. O’Brien & B.
Mawbey,28Mar 1990; TMAGG6313, ISA (21-25 mm), 699 (20-32mm),4juv. AS
(11-14 mm), 6 juv. 9$ (8-11 mm), Wentworth Hills, plateau, 42° 12.36'S 146°18.14'E,
inflow stream to lagoon, 1100 m asl, coll. D. O’Brien & B. Mawbey, 28 Mar 1990.

Butlers Gorge: YPM 9195, 4 A A (22-24 mm), 2 juv. AS (12-19 mm), 329 (25-36
mm), 4 juv. $9 (15-18 mm), near Butlers Gorge, 42°16.6'S 146°16.3'E, 680 m asl,
coll. J.H. Wilson, 20 Oct 1964; TMAG G6406, 5AS (18-22 mm), 2 juv. A A (12-15
mm), 29 9 (21-22 mm), 1 juv. 9 (13 mm), near Butlers Gorge, 42°16.6'S 146°16.3'E,
680 m asl, coll. P. Tyler, 26 Nov 1963.

Franklin-Gordon Wild Rivers National Park: TMAGG6356, 4- A A (22-23 mm), 1 juv.
AS (15-18 mm), 299 (23-27 mm), ljuv. 9 (12 mm), EofMtArrowsmith,42°13.TS
146°05.8'E, underground part of stream in rainforest, air temp 8.3°C, water temp 6.7°C,
800 m asl, coll. A. Richardson, R. Mawbey, B. Knott & P. Suter, 10 Nov 1974; TMAG
G6498, ljuv. 9 (8mm), Capricorn Cave (MR204-24), Mt Ronald Cross karst, 42°13.2'S
146°03.7'E, stream, dark zone, coll. S. Eberhard, 540 m asl, 27 Jan 1989; TMAG 14396/
G140, 1 A (22 mm), 2 juv. 99 (9-11 mm), Lake Tahune, Frenchmans Cap, 42°16'S
145°50'E, 1000 m asl, coll. W.J. Fairbridge, Jan 1945; QVM 10:49053,2 indet juv. (5-6
mm), Lake Tahune, Frenchmans Cap, 42°16'S 145°50'E, outflow creek, drift sample,
1000m asl, coll. S. Chilcott, 28 Jan 1988; AMP99165, lAA (23-29 mm), 999 (23-30
mm), Lake Tahune, Frenchmans Cap, 42°16'S 145°50'E, 1000 m asl, coll. B.V. Timms,
Jan 1989; QVM 10:13972, \A (23 mm), 19 (25 mm), 1 juv. A (damaged, c. 15 mm),
Frenchmans Cap, 42°16.1'S 145°49.6'E, 1440 m, coll. B. McCausland, 15 Mar 1980;
TMAG G6357,19 (27 mm), creek draining into Lake Richmond outflow. King William
Range, 42°18.85'S 146°11.15'E, 740 m asl, coll. A. Richardson & G. French, 28 Jan
1989; TMAG G6497,1 juv. 9(19 mm), 4 indet juv. (5-7 mm), Kutikina Cave (F34-34),
Franklin River karst, 42°31.8'S 145°46.1'E, 60 m asl, coll. S. Eberhard, 23 Mar 1988;
TMAG G6499, 1 A (21 mm), Kutikina Cave (F34-4), 42°31.8'S 145°46.TE, stream,
60 m asl, coll. S. Eberhard, 21 Mar 1989; AM P99306,1$ (damaged), 19 (damaged).
Lake Rhona, 42°33.2’S 146°17.2'E, 860 m asl, amongst cobbles near shore, coll. S.
Jarman; TMAG G1360,4juv. A A (18-30111m), 399 (33-36 mm). Lake Rhona, Upper
Gordon River, below Reed’s Peak, 42°33'S 146°17'E, 860 m asl, coll. A.P. Andrews &
H.D. Barker, 20 Mar 1972; QVM 10:12144,19 (24 mm), Bill Nielson (Rotuli) Cave,
Nicholls Range karst, 42°42.3'S 145°49.3'E, small rocky side stream, 30 m asl, NR1-2,
coll. S. Eberhard, 19 Feb 1987; TMAG G6400,19 (25 mm), 13 juv. AS (8-13 mm),
11 juv. 99 (8-13 mm). Vale of Rasselas, E of “The Thumbs”, 42°40.2'S 146°23.0'E,
buttongrass hole, coll. R. Swain & J. Ong, 17 Feb 1970; TMAG G6409,19 (24 mm),
10 juv. AS (10-13 mm), 17 juv. 99 (8-14 mm), Vale of Rasselas, E of “The Thumbs”,
42°40.2'S 146°23.0'E, buttongrass hole, coll. R. Swain & J. Ong, 17 Feb 1970; TMAG
G6412,1 A (21 mm), 2 juv. AS (11-13 mm), 4 juv. 9? (7-10 mm), Vale ofRasselas,
creekrunning parallel to track from Florentine, 42°41,7'S 146°23.8'E, buttongrass hole,
coll. R. Swain & J. Ong, 17 Feb 1970.

Southwest National Park: ZSRO 859,4juv. AA (9-11-13 mm), 11 juv. 99 (9-15 mm),
Mueller Road, 2 km behind gate, Weld River, creeks, 42°48.76'S 146°24.54'E, 1WP6,
500 m asl, 24 Feb 2013; TMAG G6389, 10 juv. 99 (8-17 mm), SW of Mt Mueller,
Port Davey track, between Scotts Peak Road and “Damper Inn”, 42°49.5'S 146°23.7'E,
first patch of rainforest, 580 m asl, coll. R. Swain, I. Wilson & J. Ong, 18 Feb 1970;
TMAG G6364, \6SS (19-25 mm), 3 juv. AS (11-16 mm), 1799 (19-26 mm), 17
juv. 99 (9-12 mm), S of Mt Mueller, stream at “Damper Inn”, 42°49.8’S 146°27.5'E,
470 m, coll. R. Swain, I. Wilson & J. Ong, 18 Feb 1970; OM Ivl2886, 2AS (20-21
mm), 1 juv. 9 (14 mm), Huon River, 2000 ft asl [600 m]; TMAG G6388,1 A (27 mm),
19 juv. A A (8-13 mm), 61 juv. 99 (7-15 mm), 10 indet juv. (6-7 mm), Port Davey
track N of Mt Bowes, tributary of Weld River, 42°50.3'S 146°24.6'E, 530 m asl, coll. R.
Swain, I. Wilson & J. Ong, 18 Feb 1970; TMAG G128,399 (11-24 mm), Mt Bowes,
42°51.6'S 146°24.6'E, 956 m asl, Apr 1939; TMAG 14369/G113, 4 A A (24-29 mm),
69 9 (23-34 mm), Snowy Mountains, 42°53.4'S 146°39'E, 2000 ft asl [600 m], coll.
CD. King, 20 Feb 1939; TMAG 14373/G117, 4 A A (25-32 mm), 699 (19-33 mm),
Snowy Mountains, 42°55.5'S 146°40.5'E, 3000 ft asl [900 m], small Lake, coll. C.D.
King, Feb 1939; AM P56374,2juv. AA (22-29 mm). Lake Skinner, Snowy Mountains,
42°57'S 146°41'E,C.62T, small stream at end of track to Lake, coll. W. Ponder etal, 15
Jan 1982; AM P73044,2 juv. AA (17-28 mm), 3 juv. 99 (10-25 mm), Snowy North,
stream flowing into Styx River, 42°53.26'S 146°39.30'E, 590 m asl, coll. S. Jarman;
TMAG G397,19 (40 mm), plateau on summit of Mt Snowy, small pool 3 inches deep,
3500 ft asl [1050 m], coll. J.F. Thompson, 31 Jan 1962; TMAG G3432, 59 9 (24-33
mm), Lake Skinner, Snowy Mountains, 42°56.8'S 146°40.5'E, 970 m asl], coll. D. Famell,
28 Jan 1962; TMAG G6387, 28 juv. A A (20-27 mm), 899 (25-33 mm), 18 juv. 99
(18-25 mm), Lake Skinner, Snowy Mountains, 42°56.8'S 146°40.5'E, 970 m asl, coll.
M. Fenton, 29 Nov 1971; TMAG G130, 2 AS (32-34 mm), 699 (17-35 mm), Lake
Denison, 42°57.4'S 146°41.0’E, 1900 ft asl [570 m], coll. C. King, Feb 1936; TMAG
G127,1 A (25 mm), 5 juv. 99 (13-20 mm), Lake Denison, 42°57.4'S 146°41.0'E, 1900
ft asl [570 m], coll. C. King, Feb 1939; TMAG G112,3 A A (23-27 mm), 999 (22-31
mm), Lake Skinner, Snowy Mountains, 42°56.8'S 146°40.1'E,c. 3000 ft asl [970m], coll.

C. D. King, Feb 1939; TMAG G6374, 19 (53 mm), Lake Skinner, Snowy Mountains,
42°56.8'S 146°40.5'E, 970 m asl, coll. D. Farnell, 28 Jan 1962; TMAG G6407, 2 AS
(28-29 mm), 6 juv. AA (20-22 mm), 10 9 9 (23-26 mm), 9 juv. 99 (8—14 mm), Snowy
Mountains, 42°54'S 146°42'E, tarn, coll. M. Fenton, 19 Nov 1970; TMAG G6361, 1
juv. A (12 mm), Snowy South, 42°56.7'S 146°39.4'E, small Lake on slope, 1340 m asl,
coll. P. Davies, Apr 1986; NMV J42439,1 A (25 mm), 19 (26 mm), Gordon River, Capt.
Sutton, Jan 1949; TMAG G136,1 juv. 9 (18 mm), Lake Pedder [probably near Maria
Creek, 42°53.8'S 146°17.3'E, coll. C. King, Apr 1939; AM P99307, 299 (damaged).
Coronation Peak, 42°54.81'S 146°00.68'E, tarn, coll. S. Jarman; TMAG G6316,1 A (23
mm), 1 juv. A (22 mm), 499 (23-31 mm), 6 juv. 9$ (14-21 mm), Mt Anne Plateau,

Ahyong: Endemic Tasmanian Mountain Shrimps

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1000 m asl, coll. S. Eberhard, 12 Jan 1987; TMAG G6496, 13 (25 mm), 1 $ (29 mm),
Deep Thought Cave (MA10-1), Mt Anne, 42°56.0'S 146°26.5'E, at 180 m in, 1000 m
asl, coll. S. Eberhard, 13 Jan 1987; TMAG G6416, 1 juv. £ (9 mm), Search Camp, Mt
Anne, 42°56.8'S 146°25.4'E, 1240 m asl, coll. J. Bludhorn, 4 Mar 1972; TMAGG6430,
1 juv. S (21 mm), Search Camp, Mt Anne, 42°56.8’S 146°25.4’E, 1240 m asl, coll. J.
Bludhorn, 4 Mar 1972; QVM 10:49052, 1 £ (21 mm), Croaking Lake, Remote Peak,
42°58.07'S 146°03.05’E, benthos, 820 m asl, coll. S. Chilcott, 13 Nov 1986; TMAG
G6358, \S (25 mm), 1 juv. <3 (16 mm), 1$ (32 mm), 6 juv. $$ (9-15 mm), small
creek draining into Lake Edgar, 43°02.9'S 146°20.7’E, 300 m asl, coll. R. Swain etal.,
3 Jun 1970; TMAG G6453, 233 (24-25 mm), 4 juv. 33 (16-20 mm), 2$$ (24-28
mm), 3 juv. $$ (12-16 mm), Lake Fortuna, Western Arthurs, 43°07.6FS 146°13.68’E,
1240 m asl, coll. Project Raleigh, Western Arthurs Team, 15 Jan 1987; TMAG G6354,
29$ (21 mm), 1 juv. £ (17 mm). Lake Cygnus, Western Arthurs Range, 43°07.80'S
146°14.16'E, coll. P. Hamr & L. Cook, 7 Nov 1990; TMAGG6451, 1 juv. 3 (19 mm),
8$$ (20-29 mm), Lake Cygnus, Western Arthurs Range, 43°07.80'S 146°14.16'E,
coll. Project Raleigh, Western Arthurs Team, 16 Jan 1987; AMP993U, 1 3 (damaged),
1 juv. $ (damaged), Square Lake, Western Arthurs, 43°09.08'S 146°16.09'E, amongst
cobbles near shore, 860 m asl, coll. S. Jarman; TMAG G6322,1$ (20 mm), 3 juv. $ $
(12-19 mm), Square Lake, 43°08.59'S 146°15.58'E, 860 m asl, coll. Western Arthurs
Team, Project Raleigh, 17 Jan 1987; TMAG G6437, 10 33 (18-24 mm), 23 juv. 33
(9-17 mm), 27$$ (19-37 mm), 33 juv. $$ (8-18mm), Square Lake, Western Arthurs,
43°08.6'S 146°15.6'E, 860 m asl, coll. R. Swain, 4 Feb 1971; TMAG G6447,2 juv. 33
(15-16 mm), 4 juv. $ $ (8-16 mm). Square Lake, WestemArthurs, 43°08.6'S 146° 15.6'E,
860 m asl, coll. M. Fenton, Jan 1971; NMV J40508, 4$$ (24-43 mm), Lake Oberon,
WestemArthur Ranges, 43°09.1'S 146°16.0'E, 840 m asl, coll. G. Poore, 28 Feb 1990;
AM P99310, 13 (damaged), 1$ (damaged). Lake Oberon, 43°08.92'S 146°16.10'E,
amongst cobbles near shore, coll. S. Jarnian; AM P99309,2$ $ (damaged). Haven Lake,
WestemArthurs, 43°10.35’S 146°19.92'E, coll. S. Jarman; TMAGG6436, 333 (15-18
mm), 1 juv. S (10 mm), 6$$ (18-20 mm), 1 juv. $ (12 111m), Haven Lake, Western
Arthurs, 43°10.3'S 146°20.0'E, 900 m asl, coll. M. Fenton, Jan 1971; TMAG G6417,
1 $ (20 mm), Prom Lake, 43°10.0'S 146°21.6'E, 820 m asl, coll. D. Gotts, 27 Jan 1970;
TMAG G6452,5 $ $ (19-26 mm). Lake Ceres, WestemArthurs, 43°08.53'S 146° 15.07'E,
780 m asl, coll. Project Raleigh, WestemArthurs Team, 17 Jan 1987; TMAGG6425,9
juv. 33 (15-17 mm), 5$$ (21-26 mm),3 juv. $$ (15-17 mm), Lake Sirona, Western
Arthurs, 43°10.0'S 146°21.6'E, 1020 m asl, coll. M. Fenton, Jan 1971; AM P99308, \$
(damaged), Lake Picton, 43°09.56'S 146°38.23'E, coll. S. Jarman; TMAG G6317, 10
juv. 33 (15-28 mm), 20 $$ (24-33 mm), 11 juv. 99 (14-21), Lake Picton, 43°09.5'S
146°38.3'E, 900 m asl, 23 Jan 1969; TMAG G6424,1 3 (23 mm), 2 juv. 33 (26-28 mm),
25$$ (23-36 mm). Lake Picton, 43°09.5'S 146°38.2'E, 900 m asl, P. Tyler, Jan 1969;
NMV J42445, 1 juv. 3 (21 mm), 5$$ (25-27 mm), Hanging Lake, near Federation
Peak, 43°16.7'S 146°27.8'E, 1140 m asl, coll. I. Stuart, 17 Jan 1974.

Mt Field, Junee-Florentine Karst. TMAG G6435, 3$$ (18-20 mm), 1 juv. 3 (12
mm), 1 $ (19 mm), 1 juv. $ (12 mm), Robert Tarn, Tam Shelf, Mt Field, 42°40.7'S
146°34.2'E, sample 1, 1200 m asl, coll. I. Wilson & J. Ong, 25 Jan 1970; TMAG
G8172, 233 (22-30 mm), 2 juv. 33 (14-16 mm), Robert Tam, Tarn Shelf, Mt Field,
42°40.7'S 146°34.2'E, 1200 m asl, coll. T. Walker, Mar 1972; QVM:2016:10:0003,
\$ (23 mm), Mawson Plateau, Mt Field, 42°41.4'S 146 0 35.1'E, 1270 m asl, coll. C.
Reid, 1975; QVM 10:12456, 1$ (28 mm), 2 juv. $$ (10-15 mm), Khazad Dum
(JF4-17), Junee-Florentine karst, streamways, 42°42.6'S 146°33.6'E, 700 m asl, coll.
S.M. Eberhard et al, 27 Jun 1989; AM P99305, 1$ (shrivelled, poor condition, c.
30 mm), Risby Basin Cave (Ray Benders Cave) (RB-X4), Risby Basin Karst, SW
of Maydena, 42°46.6'S 146°36.9'E, underside of large boulder in stream, dark zone,
998-11, coll. A. Clarke, 13 Sep 1998.

Arve Valley- Hart:\ TMAG G2214, 1$ (25 mm), Arve Loop Road, Arve Valley,
43°07.7'S 146°44.9'E, from creek, 390 m asl, coll. R. Shoobridge, 14 Feb 1980;
TMAG G6382, 24 juv. 33 (13-28 mm), 8$$ (26-32 mm), 39 juv. $$ (8-24 mm),
1 indet juv. (7 mm), Hartz Mountains [almost certainly from northern Hartz], coll. R.
Swain & G. Bert, Feb 1970.

Wellington Range: TMAG G6402, 533 (22-31 mm), 10 $$ (27-34 mm), Myrtle
Forest Creek, 42°51.6'S 147°10.3'E, 440 m asl, coll. R. Swain, Jun 1969; TMAG
G6403, 533 (28-33 mm), 14 juv. 33 (17-26 mm), 17$$ (24-38 mm), 3 juv. $$
(18-21 mm), Myrtle Forest Creek, 42°51.6'S 147°10.3'E, 440 m asl, coll. R. Swain,
Jun 1969; TMAGG983, 1933 (18-25 mm), 12$ $ (14-26 mm), Sorell Creek, Myrtle
Gully [Myrtle Forest], Collinsvale, 42°53.8'S 147°15.4'E, 400 m asl, coll, museum
staff, 9 Dec 1964; TMAG G6315, 4 33 (24-28 mm), 6$$ (25-32 mm), Myrtle
Forest, Collinsvale, 42°51.6'S 147°10.3'E, 440 m asl, coll. R. Swain, Sep 1969; AM
P4143, \S (32 mm), 1$ (34 mm), Mt Wellington, 42°53.8'S 147°14.5'E, coll. E.A.
Briggs, pre 1918; AM P14772, 233 (24—26 mm), 1 juv. 3 (19 mm), 1$ (21 mm),
Mt Wellington, 42°54'S 147°14'E, from University of Sydney Biology Dept in 1964;
AM P14773, \S (36 mm), Mt Wellington, from University of Sydney Biology Dept
in 1964, possible Haswell label; AM G1779, 1 3 (24 mm), 3$$ (24-27 mm), 5 juv.
$$ (14-20 mm), summit of Mt Wellington, 42°54'S 147°14'E, pres. C. Hedley, pre
1898; AM P2266, 13 (34 mm), Mt Wellington, 42°54'S 147°14'E, pres. E.G. Goddard;
AM P2551, hS (32 mm), 4 juv. 33 (19-20, 22 mm), 3$$ (23-24 mm), 4 juv. $$
(12-20 mm), Mt Wellington, 42°54'S 147°14'E, coll. T.T. Flynn; USNM 59126 (ex
AM P2551), 233 (24-25 mm), Mt Wellington, coll. T.T. Flynn; USNM 78433, \$
(29 mm), Mt Wellington, snow pools in swamp, from Mel Ward; OM Iv. 1395, 2>33
(15—25 mm), 15$$ (8-28 mm), no data; USNM 25030, 1 3 (27 mm), 1$ (25 mm),
“Lakes (4000 ft) Tasmania”, G.M. Thomson; AM P9217, 11 juv. 33 (18-29 mm),
12 juv. $$ (19-27 mm), Wishing Well, Mt Wellington, 42°55.67'S 147°14.76'E,
1450 feet asl [442 m], coll. C. Anderson, A. Musgrave, G.P Whitley, 23 Jan 1928;
AM PI0724, 233 (19-22 mm), 6 juv. 33 (16-18 mm), 2$$ (19-24 mm), i3 juv.
$$ (8-17mm), Mt Wellington, 42°54'S 147°14'E, coll. F.D. Manning, Jan 1935; AM
P56375, (32 mm), 3 juv. 33 (14-18 mm), 1 $ (20 mm), 6 juv. $ $ (12-16 mm), Mt
Wellington, small pools (running water) 2500 ft [750 m], coll. J.W. Evans, Dec 1938;
AMP82859,1^(24 mm), 2 juv. 33 (9-11 mm), 1$ (23 mm), 15 juv. $$ (8-12 mm),
Mt Wellington, 12Mar 1997; TMAGG794,1 3 (23 mm), back of Mt Wellington, coll.
G.E. Nicholls, 16 Dec 1933; WAM C58162, 2$$ (29-34 mm), Mt Wellington, coll.
J. Searle; WAM C367 (ex No. 6613), 3 juv. $$ (16-17 mm), Mt Wellington, coll. J.
Searle, 29 Jan 1913; SAMAC473,2$$ (shrivelled, previously dried; c. 25-26 mm),
2 juv. $$ (9-13 mm), 2 indet juv. (6 mm), North West Bay River, Mt Wellington,
42°55.3'S 147°11.2'E, 2700 ft [810 m], coll. Prof. Osborn; AM P9218,2 33 (30-32
mm), 3 juv. 33 (20-22 mm), 5$ $ (28-34 mm), 7 juv. $ $ (15-23 mm), Fern Tree
Glen, Mt Wellington, 42°55.5'S 147°15.7'E, coll. C. Anderson, A. Musgrave, G.P.
Whitley, 23 Jan 1928; TMAG G6450,3c S3 (30-33 mm), 3 juv. 33 (16-22 mm), 6$$
(26-40 mm), 4 juv. $$ (18-23 mm), St. Crispins Well, Mt Wellington, 42°55.76'S
147°12.57'E, 640 m asl, coll. R. Swain, 14 Feb 1971; SAMAC8445, 13 (21 mm),
2$$ (22-28 mm), Huonville, creeks, coll. R.T.T.

Orford : TMAGG120,1 juv. 3 (c. 17 mm), 1 juv. $ (16mm),“?Orford, east coast, 1926”.
No data: TMAG, 1 juv. 3 (14 mm), 2 juv. $ $ (16-19 mm), label faded, from R. Swain.

Description. Eyes with well-developed cornea, pigmented,
wider than and longer than half length of stalk (epigean
specimens) to slightly reduced, slightly narrower than stalk,
half length of stalk (in some subterranean forms); stalk with
subparallel margins.

Rostrum narrow in adults, apex blunt.

Pleonites with pleura sparsely setose, rounded; pleura
1-2 unarmed; pleuron 3 usually unarmed, at most with
small serration; pleura 4-5 unarmed or with 0-3 and 0-6
small spines, respectively, usually unarmed or with 1 spine

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Figure 30. Anaspides swaini Ahyong, 2015, male holotype, 27 mm, Weld River, AM P73042. (A) cephalothorax, dorsal view; (B) pleonite
6, telson and left uropod; (C) pleonites 4-6 pleura, right lateral view; (D) pleonites 4-5 pleura, left lateral view; (E) right antennule; (F)
right antenna; (G) labrum, anterior view; (H) right mandible; (I) right mandible incisor process; (J) paragnaths, anterior view; (K) right
maxillule; (L) right maxilla. Scale: A-F = 1.0 mm; G-L = 0.5 mm.

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Figure 31. Anaspides swaini Ahyong, 2015. (A-P) male holotype, 27 mm, Weld River, AM P73042; (Q-R) female paratype, 28 mm, Weld
River, AM P73043; (A) right thoracopod 1 (maxilliped); (B-H) right thoracopods 2-8; (I-J) right pleopod 1 endopod, lateral and ventral
views; (K) right pleopod 2 endopod, lateral view; (L) right pleopod 4, anterior view; (M) right pleopod 5, anterior view; (N-P) pleonites
3-5 median sternal processes; (Q-R) female gonopore, right lateral and ventral views. Scale A-H, L, M = 2.0 mm; N-R = 1.4 mm.

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Figure 32. Anaspides swaini Ahyong, 2015, selected features. ( A-D) male, 19 mm, Robert Tarn, Mt Field, TMAGG6435. (E-F) paratype
female, 22 mm, AM P73043; (G-J) female, Arve Valley, TMAG G2214; (K-N) female, 25 mm, Vale of Rasselas, TMAG G6400; (0-0)
male, c. 23 mm, Square Lake, AM P99311; (R-U) female, 18 mm, Lake Pedder, TMAG G136; (V-Y) male, 27 mm, Lake Tahune,
Frenchmans Cap, AM P99165. A, Q, S, anterior cephalothorax. B, G, K, V, telson. C, E, J, M, U, X, pleonal pleura. D, F, I, N, O, T, W,
scaphocerite. H, L, Y, rostrum. R, telson and right uropod. Scale = 1.0 mm.

on pleuron 5. Pleonite 5 posterior tergal margin (usually)
unarmed or with 3-7 small spines either side of midline,
setose. Pleonite 6 posterior margin weakly to fully spinose,
setose; posterolateral margin setose, rounded, with or without
minute denticle. Pleonal stemites 3-4 with distinctly bilobed
median processes between pleopod bases, widest on sternite

3; sternite 5 with narrow, weakly emarginate lobe.

Telson length and width subequal or longer than wide,
pentagonal, widest proximally; lateral margins sinuous in
dorsal outline, distally subparallel to convergent; transition
from lateral to posterior margin obtusely angular; posterior
margin angular to slightly rounded, blunt medially; posterior

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357

Figure 33. Anaspides swaini Ahyong, 2015, selected features. (A-D) male, 33 mm, St. Crispins Well, North West Bay River, Mt Wellington,
TMAG G6450; (E-F) juv. male (16 mm), juv. female (23 mm), St. Crispins Well, Mt Wellington, TMAG G6450; (G-J) male, 22 mm,
Sorell Creek, Myrtle Forest, TMAG G983; (K-N) male, 24 mm, Butlers Gorge, YPM 9195; (O-P) indet juv, 8 mm, Mt Rufus, AM P72842;
(Q-T) female, 30 mm, Mt Rufus, AM P72842. A, K, O, anterior cephalothorax. B, E-G, L, P, Q, telson. C, I, M, S, pleonal pleura. D, J,
N, T, scaphocerite. H, R, rostrum. Scale: A-N, Q-T =1.0 mm; O-P = 0.5 mm.

spine row with 19-54 slender, evenly graded, closely spaced
spines, longest medially.

Antennule inner flagellum about 0.2 x body length (19-20
articles in holotype); article 7 inner margin obtusely angled in
adult males, with 2 long, slender clasping spines proximally;
outer flagellum 0.4-0.6 x body length (78-80 articles in
holotype) in epigean specimens, 0.5-0.7 x in subterranean
specimens. Antennal flagellum 0.3-0.4 x body length (57-58
articles in holotype) in epigean and subterranean specimens;

scaphocerite elongate, ovate, lateral spine slightly distal
to midlength; apex reaching as far as midlength of distal
peduncular article.

Pleopods 1-4 or 5 with endopod in adults. Adult male
pleopod 1 distally widened, scoop-like, lateral margins
expanded, obscuring retinacular lobe in lateral view.

Uropodal protopod dorsally unarmed or with 1 or 2 small
spines; exopod with 2-4 movable spines on outer margin near
position of partial diaeresis; exopod length about 2.5-3 times

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Figure 34. Anaspides swaini Ahyong, 2015, selected features. (A-D) male, 30 mm, Deep Thought Cave, Mt Anne, TMAG G6496; (E-H)
female, 28 mm, Khazad Dum Cave, Mt Field, QVM 10:12456; (I-L) female, 24 mm, Bill Nielson Cave, QVM 10:12144; (M-O) male, 21
mm, Kutikina Cave, TMAG G6499; (P-R) juv. female, 8 mm, Kutikina Cave, TMAG G6497; (S-U) indet juv, 7 mm, Capricorn Cave, Mt
Ronald Cross, TMAG G6498. A-D, E-H, I-L, anterior cephalothorax, telson, pleonal pleura and scaphocerite. M-O, P-R, S-U, anterior
cephalothorax, telson and scaphocerite. Scale: A-0 = 1.0 mm; P-U = 0.5 mm.

width, slightly wider than endopod, apex rounded, narrow
to relatively broad.

Measurements. Male (n = 433) 6-34 mm, female (n = 794)
6-40 mm, indet (n = 13) 5-7 mm.

Remarks. Anaspides swaini is distinguished from other

congeners by the combination of the angular posterior
margin of the telson, blunt or minutely spinose posterolateral
angles of pleonite 6, and the presence of two clasping spines
in adult males (Fig. 30B,C,E). Anaspides swaini ranges
widely in southern Tasmania, from the Wellington range in
the southeast, southwest to the Snowy and Arthur Ranges,

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359

flanking lakes Pedder and Gordon, and north to the vicinity
of Lake St Clair (Fig. 36).

Three subtly different morphological forms of A. swaini
are recognized here. Form 1 (Figs 29-31, 32A-U, 34A-H),
corresponds to A. swaini sensu stricto and has short spines
along the upper posterior margins of pleonite 6 (and often
pleonite 5 in specimens from the vicinity of Weld River, Vale
of Rasselas and caves in the vicinity of Mt Field), 1-3 small
spines on the pleura of pleonites 4-5, no endopod on pleopod
5 (except in specimens from the Vale of Rasselas, and some
specimens from Mt Mueller and Federation Peak), and a
usually trifid proximal tooth on the right mandibular incisor
process. Form 1 has a southern range, essentially around
the periphery of Lake Gordon and Lake Pedder, from Lake
Rhona to Mt Field (where it may overlap with A. richardsoni )
and Mt Mueller to the Snowy Mountains, Federation Peak,
the Arthur Ranges and at least as far north as Coronation
Peak on the southwestern side of Lake Pedder. Note that
the distributional overlap between A. richardsoni and A.
swaini at Mt Field is largely epigean versus subterranean,
respectively, though the two species are occasionally
sympatric in surface waters of Mt Field. Specimens labelled
as possibly from “?Orford” are juvenile A. swaini , with
antennular modifications as yet incomplete in the male. As
argued by O’Brien (1990), the Orford locality is almost
certainly erroneous, being well outside of the known range
of A. swaini (Fig. 36). Moreover, the specimens correspond
most closely to specimens from the Vale of Rasselas, having
a spinose pleonite 6 and pleura 4-5 (larger juvenile), and an
endopod on pleopod 5.

Form 2 (Figs 32V-Y, 33K-T, 34I-U) has a northern
range largely beyond Lakes Gordon and Pedder. On the
surface, Form 2 ranges from the western vicinity of Lake
St Clair including the Cuvier Valley and Mt Rufus south to
Butlers Gorge and Wentworth Hills and Frenchmans Cap; it
continues further south in caves in the Nicholls Range karst
(Bill Nielson) and Franklin River karst (Kutikina), where it
is apparently isolated from surface forms (Eberhard et al .,
1991). Specimens of Form 3 typically have a few spines on
pleonite 6, unarmed pleura 4-5 (occasionally 1 or 2 small
spines on pleuron 5), presence of the pleopod 5 endopod in
adults (variable in specimens from Frenchmans Cap) and a
bifid right proximal mandibular incisor tooth.

Form 3 (Fig. 33A-J), has a southeasterly range, stretching
from the western and northern Wellington Range, including
the North West Bay River catchment of Mt Wellington, to at
least the Huonville area; it may overlap with A tasmaniae at
Mt Wellington (see Remarks under account of A. tasmaniae).
Form 3 has similarly minimal pleonal spination as Form 2,
but usually lacks the pleopod 5 endopod and usually has a
trifid right proximal mandibular incisor tooth as in Form 1.
Form 3 also frequently has a less angular posterior margin
of the telson than Forms 1 and 2.

The three forms, however, are not strictly discrete
morphologically, with some intergrading in parts of their
ranges. For instance, at Lake Tahune and Frenchmans Cap, at
the southern end of the epigean range of Form 2 (northern),
the condition of the pleopod 5 endopod is variable, being
present or absent on one or both sides. Similarly, specimens
from the Vale of Rasselas in the northern range of Form 1
(southern) share features of Forms 1 and 2 in the pleonal
spination of the former and presence of the pleopod 5
endopod of the latter. In addition, the presence of the
pleopod 5 endopod in some specimens from Federation Peak

in the southern part of the range of Form 1 is anomalous.
Morphological and distributional continuity between Form
1 and Form 2 is consistent with the likely persistence of A.
swaini in periglacial lakes formed to the west of Lakes St
Clair and King William during the Pleistocene glaciations
that dominated most of the Central Plateau and adjacent
areas (Kiernan, 1990). These lakes and associated glaciers
fed the Franklin and Gordon Rivers in which A. swaini is
widespread. Forms 1 and 2 both occur to the west of the
biogeographic discontinuity known as Tyler’s Line (Shiel
et al., 1989; Mesibov, 1994; Andrew, 2005), and Form 3
to the east.

Overall, A. swaini from the southwest generally have
a more spinose pleon and usually lack the pleopod 5
endopod whereas northern and southeastern specimens are
minimally spinose, and generally with (northern) or without
(southeastern) the pleopod 5 endopod. Each of these forms
might represent separate species or subspecies, but until more
detailed population data are available, they are considered
to represent a single wide ranging species.

Within the three broad forms of A. swaini identified
here, specimens are rather consistent morphologically. The
largest Butlers Gorge specimen (female, 36 mm) is aberrant,
however, in having a posteriorly rounded rather than angular
telson, and the 34 mm female has an abnormal pleopod 5
exopod that is basally trifurcate. A male from Lake Tahune
(29 mm, AM P99165) has the right pleopod 3 endopod
developed like the modified pleopod 2 endopod. Two lots
of A. swaini labelled as from “Lake St Clair?” and “Lake St
Clair” collected in 1937 and 1941, respectively, are probably
from creeks in the vicinity of the lake rather than the lake
itself as argued by Nicholls (1947), Williams (1965a) and
O’Brien (1990). All specimens from Lake St Clair proper,
including juveniles, exhibit the characteristic pleon and
telson spination of A. spinulae.

Like other epigean species of Anaspides, A. swaini also
occurs in caves (Fig. 34): Mt Anne (Deep Thought Cave),
Mt Field (Junee-Florentine and Risby’s Basin systems),
Nicholls Range (Bill Nielson Cave), Franklin River karst
(Kutikina) and Mt Ronald Cross karst (Capricorn Cave).
These subterranean specimens resemble epigean forms, and
at most show more elongate antennular flagella, slightly
reduced corneal size (Fig. 34A), and sometimes reduced
pigmentation, unlike more strongly cave adapted populations
of A. richardsoni and A. jarmani , or obligate troglobites
such as A. clarkei and A. eberhardi, with noticeably reduced
corneas and pigmentation. Specimens, from Khazad Dum
(Fig. 34E-H) and Risbys Basin caves, presently known only
from have a more rounded posterior telson margin,
armed pleonite 5-6 terga and usually armed pleura as
well; they are strongly pigmented as in epigean specimens.
Eberhard etal. (1991) delineated three morphological types
of telson in Anaspides : a “normal” form as exhibited by
epigean populations; a “cave” type, in which the telson spines
are few in number, stout and widely spaced (as in A. clarkei
and A. eberhardi ); and an “intermediate” form, recorded
from Capricorn Cave (Fig. 34S-U) and Deep Thought
(Fig. 34A-D). Re-examination of the “intermediate” form
specimens showed the Mt Anne telson to be of the “normal”
type, and the Capricorn Cave specimen to be a very early
stage juvenile (with normal spination for its stage) in which
the telson ornamentation is yet to be fully developed. No
specimens have so far been observed with a telson that
could be considered as intermediate between the “normal”

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Records of the Australian Museum (2016) Vol. 68

Figure 35. (A) Anaspides tasmaniae (Thomson, 1893), female, Mt Wellington, specimen not preserved; (B) A. c/arkei Ahyong, 2015, female,
24 mm, Eastern Passage streamway of Little Grunt Cave and Exit Cave, QVM 10:49168; (C) A. clarkei Ahyong, 2015, male, Lake Pluto,
Wolf Hole, specimen not examined; (D) A. richardsoni sp. nov., female, Tarn Shelf, Mt Field, specimen not preserved; (E) A. richardsoni
sp. nov., female, 32 mm, Kellys Pot, Mole Creek karst, TMAG G6463; (F) A. richardsoni sp. nov., female, 35 mm, Marakoopa Cave, Mole
Creek karst, specimen not preserved. Photo credits: A, Kristi Ellingsen, used under a Creative Commons Attribution-NonCommercial
license (http://creativecommons.Org/licenses/by-nc/2.0/); B, E, ©Stefan Eberhard; C, F, ©Arthur Clarke; D, ©Simon Grove, TMAG.

and “cave type”, with the possible exception of an aberrant
specimen of A. clarkei with asymmetrically developed
spination (Fig. 8W-Y).

Sexual maturity (indicated by development of secondary
sexual features) is usually reached by 18-23 mm body length
in both sexes, typically 20-21 mm. Unusually, however,
at Lake Picton, parts of the Snowy Range including Lake
Skinner, Hartz area, Lake Rhona, and two Mt Wellington
localities (St Crispins Well, Wishing Well), secondary sexual

characteristics are not expressed until very late, at sizes
well above that which individuals are otherwise sexually
mature (24-33 mm, usually 28 mm body length or above).
Additionally, development of secondary characteristics in
these immature males seems to be particularly attenuated,
with an incomplete complement of antennular clasping
spines. Because of their relatively large size, these
immature male A. swaini with as yet incomplete antennular
modification could be overlooked as A. richardsoni , which

Ahyong: Endemic Tasmanian Mountain Shrimps

361

(N

■nT

no

'=t

145°E 146°E 147°E 148°E

Figure 36. Distribution of Anaspides. Dubious Orford record of A. swaini indicated by

has only one antennular clasping spine in adult males. The
causes of the late onset of sexual maturity are not known.

Distribution. Southern Tasmania from the Weld River,
Snowy Mountains region, Mt Field and Mt Wellington
(North West Bay River catchment) to the Western Arthurs,
throughout the Franklin-Gordon drainages, north to Lake
Rhona and Frenchmans Cap, Mt Rufus and the vicinity of
Lake St Clair; 300-1440 m asl (epigean), 30-1000 m asl
(subterranean).

Discussion

Anaspides is demonstrably more diverse than previously
supposed, with at least seven species now recognised.
Secondary sexual characters of the male antennules and
pleopodal endopods proved instrumental to taxonomic
delineations. Anaspides is now known as far north as
Great Western Cave, Gunns Plain (41°18’S) (Fig. 36). The
southernmost records are from the far south at Precipitous

Bluff (43°29'S), westernmost records are from the Tyndall
Range (145°35'E), and easternmost records from the Oatlands
area (c. 146°22'E) and Mount Wellington (147°17'E). The
previous doubtful east coast record from Orford is almost
certainly erroneous (see account of A. swaini).

Throughout most of the range of the genus, Anaspides
has entered subterranean habitats; some normally epigean
species also occur in caves but two wholly troglobitic
species are also recognized here, A. clarkei and A.
eberhardi. Williams (1965b) emphasized the role of caves as
oligothermal refuges, especially during the warmer climate
of the Tertiary, and perhaps during glacial maxima too, so
it is significant that the specimens from the lowest altitudes
at present are also from caves (e.g., A. clarkei , 70 m, Exit
cave; A. swaini , 30 m asl, Bill Nielsen Cave) including the
northernmost record from Great Western Cave, Gunns Plain
(A. richardsoni , 109 m asl). The distributions of species
are largely discrete, with minor overlap of A. richardsoni
and A. swaini in the Mt Field area, and A. swaini and A.
tasmaniae at Mt Wellington, and with the two obligate cave

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species (A. clarkei and A. eberhardi ) in close geographic
proximity to respective, closely related epigean species
(A. jarmani and A. richardsoni ). Glacial expansion and
contraction during the Pleistocene has almost certainly
influenced the distributions of epigean and subterranean
species of Anaspides. Today Anaspides distributions broadly
correspond to the biogeographical discontinuity known as
Tyler’s Line, which runs approximately diagonally between
the 146th and 147th parallel, marking a significant geological
discontinuity and dividing the drier eastern from the wetter
western parts of Tasmania (Shiel etal., 1989; Mesibov, 1994;
Rees & Cwynar, 2010). Anaspides jarmani and A. clarkei ,
A. spinulae , and two forms of A. swaini occur to the west of
Tyler’s line. Anaspides tasmaniae, A. eberhardi and one form
of A swaini occur east of Tyler’s Line. Anaspides richardsoni
also occurs largely to the east of Tyler’s Line, crosses over
in the northern part of its distribution, extending westwards
to the West Coast Range.

Based on the extensive series of Anaspides examined
here, the largest so far assembled, the taxonomy and
distributions of the species are significantly clarified.
Nevertheless, important questions remain. The several
morphological forms identified within each of the most
widely ranging species (A. richardsoni , A. swaini and A.
jarmani) might represent simple phenotypic variatnts or
indicate more significant population differentiation. This is
particularly apropos to A. richardsoni , which, in addition
to non-spinose epigean forms, includes three cave forms
at Mole Creek and markedly spinose fonns similar to A.
spinulae on the western Central Plateau. The relationship
between Central Plateau A richardsoni and A spinulae itself
requires further investigation. In this connection, additional
sampling is required to fill distributional “gaps”, as in the
Cradle Mountain area between the Walls of Jerusalem and
the West Coast Range, for which few specimens, especially
adult males are known. Similarly, sampling is sparse in the
north-western part of the range of A. swaini including a
number of cave sites, for which only juveniles are available,
and between Mt Field and the Western Lakes on the Central
Plateau, for which no records of A. richardsoni are presently
available. More detailed delimitation of the distributions
of Anaspides tasmaniae and A. swaini on the Wellington
Range is required.

The most recent conservation assessments of Anaspides
assumed A. tasmaniae to be widespread throughout much of
Tasmania (Wells etal., 1983;Horwitz, 1990; O’Brien, 1990).
The revised taxonomy of the genus, however, indicates that
the conservation status of all species of Anaspides requires
review. Ironically, A. tasmaniae may have the narrowest
range of all epigean species, so far limited to the eastern
drainages of Mt Wellington. In this regard, the possible
translocation of A. swaini to the lower Browns River on the
southeastern face of Mt Wellington is of potential concern
(see Remarks under account of A. tasmaniae). Other narrow
range species are A. spinulae from Lake St Clair, the
subterranean A. clarkei from the Hastings-Ida Bay karsts
and A. eberhardi from the Junee-Florentine karst system.
Anaspides jarmani , A. richardsoni and A. swaini have the
widest ranges, particularly the latter two. Because much of
the range of Anaspides is encompassed by the Tasmanian
World Heritage Area, a degree of protection from agriculture
and forestry already exists (Horwitz, 1990). Other major
biotic factors, however, such fish predation, warrant further

scrutiny, particularly on the Central Plateau, where trout
predation may have already had significant impact on
Anaspides populations (Lake & Knott, 1973; Williams,
1974; O’Brien, 1990).

The distributions of species determined herein are based
on all available specimens, some from localities at which
Anaspides might no longer occur. Therefore, given the
current rate of environmental change, whether from land-use
practices, climate warming or trout predation, further surveys
are required to determine the current distributions of species
of Anaspides in order to best inform management decisions.

Acknowledgments. Thanks are due to Andrew Hosie (Western
Australian Museum), Thierry Laperousaz and Rachael King
(SAMA), Genefor Walker-Smith (formerly TMAG), Kirrily
Moore (TMAG), Judy Rainbird (QVMAG), Eric Lazo-Wasem
(YPM), and Simon Wylie and Anusha Beer (OM) for the loan
of speciimens. Karen Reed and Rafael Lemaitre are thanked for
their hospitality at the USNM in 2016. Jessica O’Donnell (AM) is
gratefully acknowledged for preparing Fig. 36 as are Arthur Clarke,
Kristi Ellingsen, Simon Grove and Stefan Eberhard for use of their
images in Fig. 35. Jane Andrew, Arthur Clarke, Stefan Eberhard,
Simon Jarman, Alastair Richardson, Stefan Richter and Roy Swain
are thanked for collecting many of the specimens used in this study
and for useful insights and discussion. Brian Timms and Rachael
Peart are gratefully acknowledged for their constructive comments
on the manuscript, and Shane McEvey for his expeditious treatment
of the manuscript. This study was partially funded by a grant from
the Australian Biological Resources Study. This is a contribution
from the Australian Museum Research Institute.

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Manuscript submitted 13 October 2016, revised 24 October 2016, and
accepted 24 October 2016.

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certify that the research described has adhered to the Australian
Museum’s Guidelines for Research Practice —or those of their
home institution providing they cover the same issues, especially
with respect to authorship and acknowledgment. While under
consideration, a manuscript may not be submitted elsewhere.

More information and examples are freely available at our
website:

http://dx.doi.org/10.3853/issn.2201-4349
Editor, Records of the Australian Museum
Australian Museum Research Institute
1 William Street, Sydney NSW 2010, Australia
editor@austmus. gov. au

7 December 2016

Stock no. 016R68G

Australian Museum Research Institute
1 William Street, Sydney NSW 2010
scientific publications freely accessible at
http://dx.doi.org/10.3853/issn.2201-4349
ISSN 0067-1975 (print) 2201-4349 (online)

AUSTRALIAN MUSEUM