EIGHT
Phylum
ARTHROPODA
SUBPHYLUM CRUSTACEA
shrimps, crabs, lobsters, barnacles, slaters, and kin
W. RICHARD WEBBER, GRAHAM D. FENWICK, JANET M.
BRADFORD-GRIEVE, STEPHEN H. EAGAR, JOHN S. BUCKERIDGE,
GARY C. B. POORE, ELLIOT W. DAWSON, LES WATLING, J. BRIAN
JONES, JOHN B. J. WELLS, NIEL L. BRUCE, SHANE T. AHYONG,
KIM LARSEN, M. ANNE CHAPMAN, JØRGEN OLESEN, JU-SHEY
HO, JOHN D. GREEN, RUSSELL J. SHIEL, CARLOS E. F. ROCHA,
ANNE-NINA LÖRZ, GRAHAM J. BIRD, W. A. CHARLESTON
‘N
Scyphax ornatus,
an endemic coastal slater.
Shane Ahyong
98
o group of plants or animals on the planet exhibits the range of
morphological diversity seen among the extant Crustacea.’ This
provocative quote from Martin and Davis (2001) highlights at least
one attribute of the group. Nevertheless, the body plan of the Crustacea has a
number of unifying characteristics, including a five-segmented head with two
pairs of antennae and an elongate body that may be divided into two more-orless distinct sections – generally the thorax or ‘body’ and the pleon or ‘abdomen’.
Each of these sections bears multisegmented appendages (mostly limbs) that
are primitively biramous (forked) but some are uniramous in many groups.
Brusca and Brusca (2002) gave a succinct summary of the characteristics of the
subphylum. In addition to enormous diversity of form, crustaceans exhibit a
great range of sizes (exceeded only by molluscs, which can claim the largest
individual invertebrate in the form of the colossal squid), from minute interstitial
and parasitic forms (e.g. Tantulocarida) measuring as little as a tenth of a millimetre
to giant crabs, lobsters, and isopods with a body size of up to half a metre in length
or breadth and weighing up to 20 kilograms. By virtue of their edibility, many
crustaceans are prized items on restaurant menus around the world.
They are an ancient group, dating from at least the Early Cambrian (Chen et
al. 2001), and have diversified abundantly since then. Calculations of the number
of named living species of Crustacea range from approximately 50,000 to 67,000.
Estimates of the potential number of species range from 10 to 100 times that
number. The smaller species, such as those of the Peracarida and Copepoda may
eventually be found in numbers comparable to those of the insects on land. By
way of an example, the Isopoda currently number approximately 11,000 species,
but estimates suggest that as many as 50,000 species of Isopoda could exist on
coral-reef habitats alone (Kensley 1988), a figure close to the current total for all
Crustacea, while Wilson (2003) estimated a total of 400,000 deep-sea species!
Clearly, with thorough documentation, crustacean diversity will be found to be
huge.
Five (Brusca & Brusca 2002) or six (Martin & Davis 2001) classes of Crustacea are recognised. Whichever classification is used, only the cave-dwelling
PHYLUM ARTHROPODA
CRUSTACEA
Remipedia have not yet been found in New Zealand waters. As one moves
down the taxonomic hierarchy from class to species, the level of endemism
increases. The New Zealand fauna currently stands at 2974 known species,
of which at least 485 have not yet been named or described. This number is
very conservative, and more than a thousand additional species will surely be
discovered. Most major groups of Crustacea (orders) are to be found in New
Zealand waters, though many families and genera will be found to be absent,
particularly among those groups with strong warm-water representation, such
as the commercially and gastronomically desirable ‘prawns’. Prawns of the
family Penaeidae (notably Penaeus and Metapenaeus) and portunid crabs of the
genera Portunus and Scylla are rare or absent.
Class Branchiopoda: Fairy shrimps, water fleas, and kin
The approximately 1000 species of branchiopods (‘gill feet’) mostly inhabit
fresh water (Dumont & Negrea 2002). They cover a wide range of body form
from many-segmented, ancient-looking taxa – generally the larger-bodied
forms such as Anostraca (fairy shrimps), Notostraca (tadpole shrimps), and
‘Conchostraca’ (clam shrimps) – to more-modified short-bodied taxa like the
Cladocera (water fleas). The larger Branchiopoda do not collectively form a
natural, evolutionary group but have a general similarity (many segments and
same structure of trunk limbs) and are almost all adapted to a short life-span
in temporary pools.
There are more than 250 species of Anostraca (fairy shrimps) worldwide
(Dumont & Negrea 2002), none of which is naturally represented in New
Zealand (Chapman & Lewis 1976) although the brine shrimp Artemia
franciscana has apparently been introduced into saline Lake Grassmere near
Blenheim. They are all relatively slow and graceful forms that swim with the
back facing the bottom (opposite to most other Crustacea) while they use their
11 pairs of trunk limbs, beating in metachronal (wave-like) fashion, for both
swimming and filtration.
The Notostraca (tadpole shrimps) comprises about 10 species worldwide,
one of which (Lepidurus apus viridis) is found in New Zealand. One of the
most striking features of notostracans is the large, flattened dorsal carapace
that originates immediately behind the head and overhangs a part of the
body. Behind the carapace is a relatively long (sometimes very long), flexible
and limbless abdomen that ends in a pair of superficially segmented tail-like
processes. At the front end, the carapace has a conspicuous so-called ‘dorsal
organ’ (used for osmoregulation). The first and second antennae – which often
have sensory functions in the Crustacea – are much reduced in size in the adult,
and the sensory function has been taken over by the very long endites (innermost
branches) of the first pair of biramous trunk limbs. All notostracans have basically
the same lifestyle. In contrast to most other branchiopods, notostracans are not
filter-feeders, but remain near the bottom, where they use the heavily chitinised
parts of the anterior trunk limbs to handle detritus and small organisms (Fryer
1988).
It has recently been shown that the former order ‘Conchostraca’ is most
likely to be paraphyletic, having given rise to descendant evolutionary lineages
(Braband et al. 2002; Olesen 1998, 2000; Spears & Abele 2000; Richter et al. 2007).
The taxonomic rearrangement of Martin and Davis (2001) recognises the order
Diplostraca, with four suborders – Laevicaudata, Spinicaudata, Cyclestherida,
and Cladocera – of which only the Cladocera and Spinicaudata are represented
in New Zealand, the latter by a species of Eulimnadia. All diplostracans have
the body and legs enclosed between a large, sometimes bivalved carapace. The
biramous second antennae are used for swimming, while the phyllopodous
(leaf-like), often serially similar, trunk limbs are used for filtration. The most
speciose group in New Zealand is the Cladocera, discussed below.
Tadpole shrimp
Lepidurus apus viridis (Notostraca).
Stephen Moore
Eulimnadia marplesi (Diplostraca).
After Timms & McLay 2005
99
NEW ZEALAND INVENTORY OF BIODIVERSITY
Summary of New Zealand crustacean diversity
A query (?) following an entry in the column for alien species indicates that alien status is suspected for some but not confirmed.
Taxon
Described
living
species +
subspecies
Known
undescribed/
undetermined
species
Estimated
unknown
species
Adventive
species
named +
unnamed
Endemic
species
Endemic
genera
Branchiopoda
Anostraca
Notostraca
Diplostraca
Cephalocarida
Maxillopoda
Ascothoracida
Acrothoracica
Rhizocephala
Thoracica
Tantulocarida
Branchiura
Pentastomida
Copepoda
Calanoida
Cyclopoida
Mormonilloida
Harpacticoida
Siphonostomatoida
Monstrilloida
Ostracoda
Palaeocopida
Podocopida
Myodocopida
Malacostraca
Leptostraca
Stomatopoda
Anaspidacea
Bathynellacea
Lophogastrida
Mysida
Thermosbaenacea
Amphipoda
Isopoda
Tanaidacea
Cumacea
Euphausiacea
Decapoda
44
1
1
42
1
661+2
2
1
8
77
3
1
1
568
252+1
100
1
130
85+1
0
356
3
275
78
1,425+1
3
8
2
5
5
17
0
439
358
40
51
19+1
480
5
0
0
5
0
139
1
0
3
6
0
0
0
129
9
4
0
99
16
1
86
0
82
4
255
2
0
4
3
1
1
0
64
67
77
24
0
12
7
0
0
7
1
2,067
7
2
30
20
8
0
0
2,000
290
500
3
850
330
27
320
0
200
120
2,665
2
20
5
5
3
50
5
800
1,000
300
110
15
150
3?
1?
0
2?
0
16?
0
0
0
3
0
1
1
11?
6?
5?
0
0
0
0
3
0
3
0
23
0
1
0
0
0
0
0
11
7
0
1?
0
4
5
0
0
5
1
153
1
1
4
34
2
0
0
111
10
8
0
63
30
0
89
3
61
24
850
0
2
5
8
0
11
0
268
331
12
66
0
147
0
0
0
0
1
5
0
0
0
2*
0
0
0
3
0
0
0
3***
0
0
7
0
6
1
85+10
0
0
1
0
0
0
0
48+10
19**
0
7*
0
10
Totals
2,488+3
485
~5,060
46?
1,097
98+10
*
including one new undescribed genus
** including two new undescribed genera
*** including three new undescribed genera
100
PHYLUM ARTHROPODA
CRUSTACEA
Order Diplostraca: Suborder Cladocera – water fleas
The Cladocera is generally believed to be a monophyletic group within the
Branchiopoda (Martin & Cash-Clark 1995; Olesen 1998; Taylor et al. 1999; Spears
& Abele 2000; Martin & Davis 2001), a notion that was called into question
by Fryer (1987) when providing detailed diagnoses for all branchiopod ‘orders’
(the rank was changed by Martin & Davis 2001). The Cladocera is by far the
most diverse and speciose group within the Branchiopoda, with approximately
640 species worldwide (Korovchinsky 2000), which is more than half of all
branchiopod species described.
Historically, Sars (1865) had recognised four tribes within the Cladocera
– the Haplopoda, Ctenopoda, Anomopoda, and Onychopoda – which are
basically still accepted as monophyletic groups; these groups are now treated
as infraorders (Martin & Davis 2001). The Anomopoda is the most speciesrich, with at least five families (the number varies depending on the author), 75
genera (Dumont & Negrea 2002), and approximately 560 species (Korovchinsky
2000); the Ctenopoda has eight genera and 47 species (Korovchinsky 2000),
the Onychopoda 10 genera with 34 species (Rivier 1998), and the Haplopoda
is monotypic with only one species (Leptodora kindtii – not represented in New
Zealand).
The four infraorders are rather different in their general morphology, which
means that cladocerans are difficult to characterise overall. They are in general
small, free-living crustaceans ranging from about 0.2–5.0 millimetres in length
(with the exception of Leptodora kindtii, which is a giant at one centimetre
long). Most are somewhat compact in appearance (except for L. kindtii and
some Cercopagididae, an onychopod family not represented in New Zealand).
They have a bivalved carapace (sometimes modified) with one compound eye,
small tubular unsegmented antennules (Ilyocryptus excepted), large branching
antennae, and a distinctive pair of so-called ‘postabdominal setae’ (similar setae
are seen in other branchiopods). They swim using their antennae. The Ctenopoda
and Anomopoda are somewhat alike and both have a bivalved carapace that
covers the body (but not the head), a pair of curved caudal claws, and five to
six (Anomopoda) or always six (Ctenopoda) flattened leaf-like trunk limbs that
are used to filter food particles from the water. In the Ctenopoda the six trunk
limbs show serial similarity (as in the ‘large’ branchiopods), while the trunk limbs
of the Anomopoda have undergone remarkable evolutionary modifications in
relation to food selection, with each limb in many cases being different from
its neighbour limb (Fryer 1963, 1968, 1974, 1991). The remaining two groups,
the Haplopoda and Onychopoda, are also somewhat alike, having, in contrast
to all other branchiopods, narrow-footed segmented trunk limbs – four pairs in
the Onychopoda and six pairs in the Haplopoda, used for predation or at least
for selective feeding. Olesen et al. (2001) have shown how the segmented trunk
limbs of the Haplopoda (Leptodora kindtii) have been derived secondarily from
the typical phyllopodous limbs of other branchiopods. Both the Haplopoda and
the Onychopoda have a relatively small carapace that does not cover the trunk
limbs.
In New Zealand, as elsewhere, freshwater cladocerans (water fleas) can often
be found in great abundance in open water or at the weedy edges and bottom
deposits of lakes, ponds, and stream backwaters (Chapman & Lewis 1976). A
child with a scoop-net can easily capture a good supply for a home aquarium.
A few species are known from brackish and nearshore ocean environments
(Rivier 1998). Among the freshwater species, some are strictly planktonic, others
are bottom-dwelling, and Scapholeberis (Daphniidae) lives against the surface
film. Simocephalus (Daphniidae) has the distinctive habit of interrupting its
swimming and hanging down from algal filaments by a hooked bristle on one of
the swimming antennae (e.g. Fryer 1991). Daphniids are specialist filter-feeders,
while chydorids and many macrothricids feed by scraping particles off substrata
Water flea Daphnia dentifera (Cladocera).
Barry O’Brien
101
NEW ZEALAND INVENTORY OF BIODIVERSITY
Summary of New Zealand crustacean diversity by environment
Taxon
Branchiopoda
Anostraca
Notostraca
Diplostraca
Cephalocarida
Maxillopoda
Ascothoracida
Acrothoracica
Rhizocephala
Thoracica
Tantulocarida
Branchiura
Pentastomida
Copepoda
Calanoida
Cyclopoida
Mormonilloida
Harpacticoida
Siphonostomatoida
Monstrilloida
Ostracoda
Palaeocopida
Podocopida
Myodocopida
Malacostraca
Leptostraca
Stomatopoda
Anaspidacea
Bathynellacea
Lophogastrida
Mysida
Amphipoda
Isopoda
Tanaidacea
Cumacea
Euphausiacea
Decapoda
Totals
Terrestrial
species
Fully
freshwater
species
Marine/
estuarine
species
0
0
0
0
0
2
0
0
0
0
0
0
1*
1
0
0
0
1**
0
0
1
0
1**
0
120
0
0
0
0
0
0
47***
72
0
0
0
1
41
0
1
40
0
68
0
0
0
0
0
1
0
67
11
21
0
35
0
0
37
0
37
0
90
0
0
6
8
0
0
54
17
1
0
0
4
8
1
0
7
1
730
3
1
11
83
3
0
0
629
250
83
1
193
101
1
404
3
319
82
1,470
5
8
0
0
6
18
402
336
116
75
19
487
123
236
2,614
*
internal parasite of mammal
** damp forest litter
*** including 11 supralittoral species
Water flea
Ilyocryptus sordidus (Cladocera).
From Chapman & Lewis 1976
102
using their trunk limbs. Genera in the infraorders Onychopoda and Haplopoda
are predaceous or at least raptorial feeders (Rivier 1998).
Cladocerans are able to produce non-fertilised (parthenogenetic) eggs that
develop in a brood-pouch under the carapace and hatch as miniature adults.
Females may continue to moult and grow after reaching sexual maturity, unlike
copepods and ostracods. Cladocerans reproduce sexually as well as asexually
and produce resting eggs after males have appeared in the population; these
eggs undergo a period of dormancy before development begins. In the case of
the Anomopoda, resting eggs are protected by a part of the mother’s carapace,
which is shed together with the eggs as an ephippium. The appearance of males
is probably triggered by environmental conditions.
PHYLUM ARTHROPODA
CRUSTACEA
Summary of New Zealand fossil crustacean diversity
Taxon
Maxillopoda
Acrothoracica
Rhizocephala
Thoracica
Ostracoda*
Archaeocopida
Palaeocopida
Podocopida
Myodocopida
Malacostraca
Phyllocarida
Eumalacostraca
Isopoda
Decapoda
Described
fossil
species +
subspecies
Known
Endemic
undescribed/ species
undetermined
species
Endemic
genera
61
0
0
61+3
284
0
1
283
0
67
7+1
60
4
56
19
4
1
14
127
2
0
124
1
44
1
43
0
43
60
1
0
59
22
0
1
21
0
61
7
54
4
50
2
0
0
2**
5
0
0
5
0
8
0
8
1
7
412
190
143
15
Totals
* Several species range to the present day; these are also in the Recent checklist.
** undescribed new genera
The end-chapter list of New Zealand Cladocera is based on the work of
Chapman and Lewis (1976) for freshwater species and the records of Krämer
(1895) and Jillett (1971) for marine species. The marine forms particularly need
revising, as most of Krämer’s species are not well known. The zoogeography
of freshwater zooplankton in Australasia (Bayly 1995 and references therein)
suggests that the New Zealand cladoceran fauna reflects the fact that New
Zealand split from Antarctica during the Late Cretaceous. New Zealand,
Australia, and South America completely lack the predaceous-raptorial
families Polyphemidae and Cercopagididae (Onychopoda), the Leptodoridae
(Haplopoda), and the Holopedidae (Ctenopoda). It seems likely that these
families evolved in Laurasia after splitting from Pangaea (Bayly 1995). On
the other hand, the Anomopoda, well-represented in New Zealand, are a
very ancient group (from at least 130 million years ago) that was probably
distributed over Pangaea.
Class Cephalocarida
The Cephalocarida was introduced as a new crustacean subclass by Sanders
(1955) for a tiny, primitive-looking species taken off the Atlantic coast of North
America. Since then, very few additional species have been discovered, and
the most recent treatments recognise only one family with five genera and 10
species worldwide (Hessler & Wakabara 2000; Martin & Davis 2001). All are very
small, measuring only 2–4 millimetres in length. The swimming limbs barely
differ from one another, with the endemic New Zealand genus Chiltoniella being
the least modified. The class is generally regarded as one of the more primitive
of the living Crustacea.
Most species have been recorded from silty seafloors. In general, their biology
is poorly known. New Zealand’s sole species, endemic Chiltoniella elongata, is
known from the Hawke’s Bay region (Knox & Fenwick 1977).
Chiltoniella elongata (Cephalocarida).
From Knox & Fenwick 1977
103
NEW ZEALAND INVENTORY OF BIODIVERSITY
Class Maxillopoda
Barnacles, seed shrimps, oar-footed bugs (copepods), and related parasitic
groups – these are all examples of maxillopod crustaceans. They are a disparate
lot, and carcinologists (crustacean specialists) are still arguing over whether or
not they are a single evolutionary lineage (monophyletic). Apart from some
barnacles, most species are small or minute. Most feed by means of mouthparts
called maxillae (instead of using trunk limbs as filtration devices), barnacles
again being a notable exception. Other characteristics of maxillopods include
a basic body plan of five head and 10 trunk segments followed by a terminal
telson. Abdominal segments usually lack appendages; elsewhere on the
body, appendages are usually branched (biramous). As a group, maxillopod
crustaceans are very important – economically, as in the case of many marinefouling barnacle species, and more especially ecologically because of their shear
abundance. Copepods, for example, are the most numerous crustaceans in
open-ocean waters.
Subclass Thecostraca
Cutaway view of Calantica spinilatera showing
the long bristly feeding limbs (cirri) with smaller
mouthparts to the lower left of the cirri.
From Foster 1979
104
This subclass comprises representatives of two infraclasses in New Zealand –
the Ascothoracica and Cirripedia (‘curly footed’). The latter includes barnacles,
sessile crustaceans that use their trunk limbs to catch food particles. Most New
Zealanders will be familiar with the acorn barnacles that carpet the upper zones
of rocky seashores or, annoyingly, boat hulls, and perhaps the stalked goose
barnacles that attach to floats and other buoyant objects, but few will know of
the tiny burrowing and parasitic thecostracans.
Minute borings in mollusc shells, attributed to barnacles, have been well
documented since Darwin (1854a) collected and described specimens during
his voyage on HMS Beagle. Originally a number of parasitic organisms were
included within this group of ‘burrowing barnacles’, e.g. the Ascothoracica
and Rhizocephala (Newman et al. 1969), but these latter two taxa have been
subsequently shown to possess spermatozoa, nauplius larvae, and newly
settled cypris stages that are very different from barnacles. Following the reevaluation of the Cirripedia by Newman (1987, 1996), the Ascothoracica and
Rhizocephala are no longer considered as barnacles by some specialists; on
the other hand, Martin and Davis (2001), Buckeridge and Newman (2006),
and Lützen et al. (2009) treat the Rhizocephala as a superorder of Cirripedia.
Ascothoracicans are represented in New Zealand by two species of starfish
parasites (Palmer 1997); living rhizocephalans, virtually unknown in New
Zealand until very recently, comprise 11 species (Brockerhoff et al. 2006; Lörz
et al. 2008; Lützen et al. 2009).
The burrowing acrothoracicans possess a soft carapace, with calcareous plates
reduced or absent. There are about 40 known species worldwide, including one
endemic New Zealand species. All live buried in calcareous shells of a wide range
of marine invertebrates, including molluscs, echinoderms, corals, bryozoans, and
other barnacles. The group has a fossil record extending back to the Devonian
(Tomlinson 1987), although no pre-Mesozoic taxa are known from New
Zealand. As the fossil record of acrothoracicans is based solely upon burrows,
two distinct acrothoracican nomenclatures have developed, one ichnomorphic,
the other biological. This may lead to some confusion, as trace-fossil names such
as Zapfella have equivalents such as Australophialus. Both systems are used in
this review of the New Zealand fauna because the relationship between fossils
and living species is unclear.
The familiar thoracican barnacles are classified into four orders with 81 living
species in New Zealand – the stalked (pedunculate) Ibliformes, Lepadiformes,
and Scalpelliformes, and the generally squat, nonstalked Sessilia, comprising
the acorn (balanomorph) barnacles, wart (verrucomorph) barnacles, and the
PHYLUM ARTHROPODA
Brachylepadomorpha (confined to deep-ocean hydrothermal vents and not yet
known from New Zealand).
Most barnacles are hermaphrodites, although in some species the
‘typical’ hermaphrodite form may also carry minute or dwarf males within
the capitulum (see below). These dwarf males possess either reduced or no
appendages and capitular plates, being essentially packages of male gonads.
Sexual differentiation does occur in some species, e.g. endemic Idioibla idiotica,
(Ibliformes).
The pedunculate forms are the most ancient of the barnacles. They are
characterised by a stalk (peduncle), by which they attach themselves to the
substratum. A series of calcareous plates, together forming a capitulum, are found
on top of the peduncle of most species, enclosing most of the soft tissue of the
animal. A careful examination of this area verifies the evolutionary placement
of the barnacles within the crustaceans, as the animal is effectively arranged
head down, with its jointed limbs (cirri) extending out through a slit (orifice) in
the capitulum wall. When the barnacle is submerged, the cirri extend into the
surrounding water, netting planktonic food.
As the number and arrangement of capitular plates varies considerably
between taxa, they are of considerable value in classification. In the goose
barnacle Lepas (Lepadiformes) there are five plates: paired terga and scuta with
a single carina, arranged in a single whorl. However, in species like Calantica
spinosa (Scalpelliformes) the number of capitular plates varies from 11 to more
than 50, and these are arranged in two or more whorls. In taxa like Calantica
and Anguloscalpellum, the peduncle is armoured with small overlapping plates
or scales. In contrast, there are no plates or overlapping scales on the peduncle
in Lepadiformes. The most primitive order of living thoracicans is the Ibliformes,
with predominantly chitinous rather than calcareous plates. Of the five living
genera, three of them are found in New Zealand, including the endemic genus
Chitinolepas from Spirits Bay (Buckeridge & Newman 2006).
The Verrucomorpha are a group of barnacles that, because of their asymmetry, have intrigued cirripede workers since Darwin (1854b). Although they are
amongst the most primitive Sessilia that are likely to be encountered as fossils,
they are as yet unconfirmed from the New Zealand Mesozoic. They are, however,
known from the Cretaceous of Australia (Buckeridge 1983). The Verrucidae are
represented in New Zealand waters by species of Altiverruca and Metaverruca,
both of which possess six calcareous plates. The lid (operculum) comprises just
two articulating plates, the shell wall being made up of the remaining four: a
fixed tergum and fixed scutum, plus rostrum and carina. Unlike other Sessilia,
each wall plate in verrucids joins with its adjacent plate by interlocking ribs. The
distribution of verrucid genera tends to conform to depth, with Verruca species
characteristic of shallow coastal waters, Metaverruca to midshelf environments,
and Altiverruca to the continental slope and deeper. Some verrucid species also
have symbiotic or commensal relationships with other invertebrates, and these
may be host-specific, e.g. Brochiverruca on cnidarians and Rostratoverruca on
cidaroid urchins (Buckeridge 1997). This appears to be the situation with an asyet-undescribed verrucid from northern New Zealand waters that inhabits the
coral Ellanopsammia rostrata.
When one considers balanomorph or acorn barnacles, the image many
people have is of a limpet-like creature commonly attached to vessel hulls.
Although barnacle fouling on ships is well known, it represents only a small
proportion of their distribution. They are best seen as ubiquitous opportunists
of the marine environment attached to a great variety of living and inanimate
objects. Barnacles include species specialised for attachment to whales, sea
snakes, turtles, corals, sponges, and other crustaceans.
Many shallow-water acorn barnacles are known to have variable tolerances
to both high temperatures and desiccation. Because of this, species in the
intertidal zone may be found distributed in distinctive bands, e.g. on exposed
CRUSTACEA
Idioibla idiotica.
John Buckeridge
Chitinolepas spiritsensis.
From Buckeridge & Newman 2006
105
NEW ZEALAND INVENTORY OF BIODIVERSITY
Coronula diadema, a barnacle that grows
on whales.
John Buckeridge
rocky shores, where Chamaesipho brunnea forms bands in the uppermost
intertidal and Epopella plicata at mid- to low tide.
The balanomorph shell is made up of two parts: a rigid calcareous wall
comprising four or more parietal plates, and an operculum or lid generally made
up of paired scuta and terga. The opercular plates articulate to permit extension
of the cirri between them during feeding. They also enable the animal to seal
itself off from the environment in times of stress (e.g. predation, desiccation). As
with the stalked barnacles, the plates are very important in identifying species.
Parietal plates may be solidly calcified (e.g. Austrominius), calcareous with internal
chitinous laminae (e.g. Epopella), calcareous with one row of vertical tubes (e.g.
Balanus), or calcareous with chitin, arranged in multiple rows of tubes as in
Tetraclitella (Buckeridge 2008). The number of parietal plates is also significant,
with four in Austrominius, Epopella, and Tetraclitella and six in Austromegabalanus,
Balanus, Chamaesipho, Coronula, Megabalanus, and Notobalanus.
The elements of barnacle anatomy and morphology, forming the basis of
our modern classification and understanding, were elucidated by none other
than Charles Darwin. His outstanding work on these creatures had a very strong
influence on the ideas that eventually led to his revolutionary book On the Origin of
Species. Indeed, Darwin was so amazed by the profusion and ubiquity of barnacles
in the Cenozoic that he described Tertiary seas as‘abounding with species of Balanus
to an extent now quite unparalleled in any quarter of the world’. (In Darwin’s time,
although most sessile cirripedes were ascribed to the genus Balanus, he was able to
demonstrate groupings of similar taxa through the use of ‘varieties’.)
That Darwin was infatuated with barnacles is clear, and he put much else
aside to work on them: ‘I have for the present given up Geology, and am hard
at work at pure Zoology and am dissecting various genera of Cirripedia, and
am extremely interested in the subject.’ [Letter to Dieffenbach, February 1847].
But it was not always an agreeable infatuation: ‘I have now for a long time
been at work on the fossil cirripedes, which take up more time than the recent:
confound and exterminate the whole tribe; I can see no end to my work.’ [Letter
to Hooker, 1850]. Darwin did persist, both with his monographs on fossil and
living cirripedes (Darwin 1851a,b, 1854a,b) and his Origin of Species. Darwin’s
second cirripede volume was dated 1851 but came out quite late in 1852. His
works endure as a monument to scholarship, and remarkably, one and a half
centuries later, still provide the intellectual platform from which we are able to
develop our present-day understanding of Earth’s biodiversity.
Infraclass Ascothoracica
Adult female of
Dendrogaster otagoensis.
From Palmer 1997
106
These curious creatures are primitive among thecostracans, ectoparasitic on
feather stars and sea urchins, and endoparasitic within some corals and sea stars.
Females have a much-reduced thorax and abdomen and a simplification or loss
of limbs. The carapace is enlarged and grossly distorted, being much-branched
and unrecognisable as belonging to a crustacean. Males are tiny and more
recognisably crustacean in form, resembling larvae. They have a well-segmented
body enclosed in a carapace and greatly elongated testes and and are found
within the mantle cavity of females.
Ascothoracicans were unknown in New Zealand until Palmer (1997) found
two species inhabiting sea stars off the Otago coast. Dendrogaster otagoensis was
described as a new species, infesting Asterodon miliaris. Of a collection of 159 sea
stars taken from the coast over an 11-month period, 124 (78%) were infested
with the parasite. Found inside the arms and disc of the sea star, there can be as
many as 15 female parasites, with their convoluted carapaces over 20 millimetres
across, causing some atrophy of the sea-star’s digestive caecae and gonads. Up to
19 creamy-white males 2.9–3.5 millimetres long occur inside the female parasite.
A second species, Dendrogaster argentinensis, was also found off Otago,
infesting 96% of 152 specimens of the sea star Allostichaster insignis quite severely.
PHYLUM ARTHROPODA
CRUSTACEA
This particular parasite, previously known from southern South America and the
Falkland Islands, can fill much of the sea-star’s body cavity, comprising up to
28% of the wet weight. Gonads in such specimens are absent, and digestive
caecae are severely atrophied. Curiously, specimens of A. insignis in other parts of
its range (Cook Strait to the Auckland Islands) have never been noted as having
such parasites, so it would be interesting to know what conditions promote such
infestations in Otago waters.
Dendrogaster belongs to one of three families in the ascothoracican order
Dendrogastrida. Palmer (1997) also mentioned an unpublished Te Papa (Museum
of New Zealand) record of an undescribed member of the Synagogidae, one of
three families in the only other ascothoracican order, Laurida.
Infraclass Cirripedia: Barnacles
Superorder Acrothoracica
Apart from the study by Batham and Tomlinson (1965) on Australophialus
melampygos, there has been little work done on New Zealand acrothoracicans.
They are a very difficult group to work with, particularly as most occurrences
are known only by their tiny borings. Australophialus melampygos is often found
infesting paua (Haliotis iris) and mussel (Perna canaliculus) shells, commonly in
very large numbers (up to 3350 borings noted in a single paua shell. The family
Cryptophialidae was revised by Tomlinson (1969), who introduced Australophialus
to incorporate the austral members (including A. melampygos) of Cryptophialus
that possessed four rather than three pairs of terminal cirri (feeding appendages).
Existing literature infers that acrothoracicans have very low diversity in
the New Zealand region. Further, they appear to be somewhat host-specific,
and whilst this is not generally a problem where a host is a common marine
invertebrate, there is cause for concern if the host is over-fished. Both Haliotis
iris (paua) and Perna canaliculus (green-lipped mussel) are extensively harvested
as a food source, and although they are now widely cultured in marine farms,
the new aquacultural environment does not appear to provide the habitat so
favoured by Australophialus melampygos in nature. The likelihood that the shellinfesting population represents more than one species should not be overlooked,
especially in light of acrothoracicans’ poorly mobile larval phase (which may
account for its absence from the Chatham Islands). The distribution of these
molluscs extends from Northland to Stewart Island; although both species range
well into the subtidal, A. melampygos is not known much below low tide, its
preferred habitat.
Australophialus melampygos falls within a group of southern acrothoracicans
including A. tomlinsoni from the Antarctic and A. turbonis from South Africa.
Newman and Ross (1971) considered the cirral arrangement of these taxa to
be more generalised (and therefore phylogenetically older) than other Cryptophialidae, inferring a Southern Hemisphere origin for the family. However,
rather than a South African centre of cryptophialid diversification, abundant
cryptophialids in some turritellid gastropods within the Pakaurangi Formation
(Early Miocene), Kaipara Harbour, should not rule out the New Zealand region
as a potential centre of dispersal.
Superorder Rhizocephala
Rhizocephalans are wholly parasitic. They have little similarity with other
cirripedes, or indeed other crustacean adults, as there are neither appendages
nor segmentation (e.g. Høeg & Lützen 1995, 1996). A rhizocephalan consists of
a sac-shaped body, the externa, which is mainly involved in reproduction and
is attached to the outside of the host’s abdomen. The host is always another
crustacean, in most instances an anomuran or brachyuran crab. A mouth
and a digestive tract are absent and nutrients are taken up from the host’s
interior by an internal trophic root system (or interna) which is distributed
Australophialus melampygos removed from
its excavation in a shell; five dwarf males
attached middle right.
Modified from Batham & Tomlinson 1965
Briarosaccus callosus, a saccular rhizocephalan
parasite under the abdomen of the king crab
Paralomis hirtella.
Dianne Tracey
107
NEW ZEALAND INVENTORY OF BIODIVERSITY
Sacculina sp., a saccular rhizocephalan parasite
under the abdomen (folded back) of the crab
Metacarcinus novaezelandiae.
Annette Brockerhoff
within the haemolymph of the host (Høeg & Lützen 1995). The externae are
most often attached singly or a few together to the host’s abdomen, but some
rhizocephalans are colonial and in such species many small externae may attach
to the abdomen, appendages, or other parts of the host body (Høeg & Lützen
1993, 1996). Despite their bizarre appearance, rhizocephalans are related to the
non-parasitic barnacles, which they resemble in reproducing via short-lived
planktonic nauplii and/or cypris larvae (Høeg & Lützen 1993).
Apart from sparse records in the literature, rhizocephalans were almost
unknown in New Zealand until the 2000s; there are now at least 10 genera
and 11 species (Brockerhoff et al. 2006; Lörz et al. 2008; Lützen et al. 2009).
Decapod host species belong to the families Paguridae, Lithodidae, Galatheidae,
Chirostylidae, and Callianassidae. Parthenopea vulcanophila (Lützen et al. 2009),
is the first rhizocephalan recorded from the vicinity of active cold seeps.
The recently discovered New Zealand rhizocephalans are registered in the
invertebrate collections of the National Institute of Water and Atmospheric
Research (NIWA) and the National Museum of New Zealand Te Papa Tongarewa,
Wellington (NMNZ). Some of the specimens could not be identified because they
were in turn infected by species of Cryptoniscinae, a subfamily of hyperparasitic
isopods. In the final stage of this relationship of a parasite on a parasite the
rhizocephalan host is no longer recognisable (Øksnebjerg 2000).
Recent gene-sequencing studies on the Rhizocephala have indicated that
the conventional grouping of its members is in need of rearrangement (Glenner
et al. 2003; Glenner & Hebsgaard 2006). Since these findings have not yet
resulted in a taxonomic revision, the traditional division of the Rhizocephala
into the orders Kentrogonida and Akentrogonida is followed in the end-chapter
checklist; as a consequence of the study by Glenner and Hebsgaard (2006),
however, Parthenopea is included in the Akentrogonida.
Superorder Thoracica
On 3 October 1769, in calm seas some 300 kilometres off what is now known
as Mahia Peninsula, HM Bark Endeavour, under the command of James Cook,
retrieved ‘one peice of wood coverd with Striated Barnacles Lepas Anserina?’
(Banks 1962). This was not only the first record of barnacles from New Zealand
seas, but also one of the first records of marine life from the region. In an editorial
footnote to Banks’s journal, J. C. Beaglehole stated that Daniel Solander (the
naturalist who accompanied Banks) considered the species to be Lepas anserifera.
The next major scientific expedition to New Zealand was in 1827, when the
Astrolabe collected extensive natural history material, including barnacles.
The barnacles were subsequently described by Quoy and Gaimard (1834) as
Anatifera spinosa, Anatifera elongata, and Anatifera tubulosa (now respectively
known as Calantica spinosa (Quoy & Gaimard), Lepas testudinata Aurivillius, and
Heteralepas quadrata (Aurivillius)). The first endemic New Zealand barnacle to be
described was, therefore, C. spinosa.
In 1839 the New Zealand Company appointed Ernst Dieffenbach as surgeon
and naturalist on the Tory. Dieffenbach made extensive biological collections
during his time in New Zealand, and included in these were barnacles. These
were later compiled by J. E. Gray into a Fauna of New Zealand and listed as an
appendix to Dieffenbach’s Travels in New Zealand (Gray 1843). Gray recorded
nine thoracicans, now known as C. spinosa, L. testudinata, H. quadrata, Coronula
diadema, Epopella plicata, Tetraclitella depressa, Tubinicella major, and two unidentified species of Balanus.
Shortly after this, Darwin’s four comprehensive monographs on living and
fossil cirripedes were published. Darwin had collected New Zealand barnacles
from the Bay of Islands during the voyage of HMS Beagle, which, along with
British institutional material, resulted in 14 species being listed from the New
Zealand region. Ten were new to science, of which Austrominius modestus,
108
PHYLUM ARTHROPODA
Notobalanus vestitus, and Notomegabalanus decorus are endemic to New Zealand.
Darwin included a complete description of the endemic species Chamaesipho
columna, which had previously been described from material supposedly
collected from Tahiti (Spengler 1790). Spengler’s original description was,
however, incomplete, as the shells he possessed were without opercula or soft
tissue. In Foster and Anderson (1986), the status of C. columna was reviewed
and it was concluded that Spengler’s material came from New Zealand, where
it is endemic. (They renamed the Australian species previously attributed to C.
columna as Chamaesipho tasmanica.)
The last major systematic work of the 19th century that dealt with New
Zealand barnacles was based upon specimens obtained during the 1873–76
HMS Challenger expedition. In an expedition report, Hoek (1883) described
five new species, now known as Amigdoscalpellum costellatum, Anguloscalpellum
pedunculatum, Gymnoscalpellum intermedium, Smilium acutum, and Verum
novaezelandiae. During the early to mid-20th century, numerous descriptions
of new records for the region, generally for single species, were published
and a full list of these was given by Foster (1979). The latter work is the most
comprehensive study ever written on living New Zealand Thoracica. In it, Foster
listed a fauna of 61 species, nine (including a new subspecies) of which were
new, one was a new name, and 15 species were recorded for the first time from
New Zealand waters. Foster also made valuable observations on the geographic
distribution, zonation, and ecology of barnacle species. In the 14 years following
his 1979 monograph, Foster described a further two new species and add records
of eight taxa not previously known from New Zealand waters (Foster & Willan
1979; Foster 1980, 1981; Foster & Anderson 1986). Brian Foster died suddenly in
1992, tragically cutting short what was, up to that time, a prolific and invaluable
career in barnacle systematics and biology. Since then, J. S. Buckeridge, a
student of Foster, has continued study of the New Zealand fauna, frequently
in collaboration with W. Newman. The systematics of barnacles was reviewed
by Buckeridge and Newman (2006), in which the Iblidae was identified as
the most ancient family of Thoracica. Significantly, it was the discovery of
an extraordinary but minute new species from New Zealand, Chitinolepas
spiritsensis, that provided the impetus for this work, which demonstrated that
the New Zealand region not only has a diverse living thoracican fauna but also
one of the most primitive.
Although not specifically focussing on the New Zealand fauna, Newman’s
(1979) publication is an inspired revision of the phylogenetic and biogeographic
relationships between barnacles of the Southern Ocean. His work led to a
reappraisal of the entire fauna, with many of the proposed taxonomic concepts
incorporated in Buckeridge (1983). The evolving nature of systematic biology
results from an ongoing reappraisal of relationships between taxa. As our
understanding of barnacle phylogeny becomes more sophisticated, this often
creates the need to provide new names for species. The overview herein is based
upon the comprehensive review of Cirripedia by Newman (1996), in which
subgenera are elevated to full generic status. Consequently, species like Elminius
modestus and Austromegabalanus decorus are now listed as Austrominius modestus
and Notomegabalanus decorus respectively. A recent publication reviews the status
of the Elminiinae and identifies Austrominius as a tetraclitoid, returning it closer
to Epopella, where Darwin (1854) had originally perceived it to be (Buckeridge &
Newman 2010).
There are 81 species of Recent thoracican cirripedes known from the New
Zealand EEZ. Of these, six are currently undescribed. Four are stalked barnacles,
comprising two species of Scillaelepas (Calanticidae) one of which conforms
to a southern group of primitive scalpellids, and two species of Scalpellidae;
an unusual undescribed verrucid is likely to represent a new genus; and a
possible new species of Acasta (Archaeobalanidae) remains to be determined (J.
Buckeridge is currently reviewing this genus of sponge-inhabiting barnacles). All
CRUSTACEA
Chamaesipho columna.
Dennis Gordon
Smilium zancleanum, with plates on the righthand side removed to show the cirri.
John Buckeridge
109
NEW ZEALAND INVENTORY OF BIODIVERSITY
Metaverruca recta.
John Buckeridge
Ashinkailepas kermadecensis.
From Buckeridge 2009
110
species referred to as new in the end-chapter checklist are held in the collections
of the NIWA Invertebrate Collection, Wellington.
The vertical zonation of thoracican barnacles on New Zealand surf shores
has been well documented (e.g. Morton & Miller 1968). The zonation is not
always consistent, however, with ranges expanding/contracting in the absence/
presence of other taxa (Foster 1979). Nevertheless, there are generalisations
that can be made, and these provide useful ecological benchmarks: chthamalids
are found higher on the shore than all other thoracicans; below them, and
overlapping somewhat, are the tetraclitids; further down the shore the lower
range of the tetraclitids overlaps the balanids. This chthamalid-tetraclitid-balanid
arrangement appears to be fairly uniform on both temperate and tropical shores
(Foster 1974, 1979). Cantellius septimus, a widespread Indo-Pacific species, has
been found in Montipora coral off Raoul Island (Kermadec Ridge), representing
the most southerly record of a coral-inhabiting barnacle (Achituv 2004).
Some species are epizoic on cetaceans. Conchoderma auritum, C. virgatum,
and Coronula species attach to whales and three species of the latter genus are
found in the New Zealand fossil record.
The isolation of New Zealand since the late Mesozoic has led to high regional
endemism in taxa that evolved during the Late Cretaceous–Early Cenozoic.
This is no more evident than in the thoracican barnacles (Buckeridge 1996a,b,
1999a). Although 40% of the Recent species listed are endemic, the figure is a
little misleading, as the current distribution of New Zealand species such as
Austrominius modestus to include Australia and Europe has almost certainly been
achieved via shipping. What is particularly significant about the New Zealand
region is the high proportion of endemics that are phylogenetically primitive.
The percentage of balanomorph and verrucid taxa that have their earliest (fossil)
records in New Zealand is impressive, with 73% of all primitive sessilians with a
generic age earlier than the Miocene being first recorded here (Buckeridge 1996a).
There are several species of thoracican barnacles that may be termed ‘living
fossils’, i.e. they have fossil records extending back at least to the Early Miocene.
Two of these, Chionelasmus darwini and Notobalanus vestitus extend back to
the Eocene and Oligocene, respectively; two others, Metaverruca recta and
Chamaesipho brunnea, to the earliest Miocene. The order Ibliformes extends back
to the Permian and the Neolepadinae to the Jurassic.
Sampling of deep-sea cirripedes from the New Zealand EEZ is far from
comprehensive, but 13 species are known from depths greater than 1500 m, the
deepest of which are Gymnoscalpellum intermedium (to 2505 m) Amygdoscalpellum
costellatum (to 3120 m), and Verum raccidium (to 4405 m) according to NIWA
database records. Specimens have often been made available as bycatch from
the fishing industry or from research cruises. Recent discoveries include the
neolepadine Vulcanolepas osheai from ca. 1500 metres depth in the volcanically
active Brothers Caldera (in the Havre Trough northeast of the Bay of Plenty)
and a related taxon, Ashinkailepas kermadecensis (Buckeridge 2009), from a coldwater seep at 1165 m on the western flank of the Kermadec Ridge. Both of these
taxa have specialisations, like long filamentous cirri, that permit them to feed
on bacteria, the most abundant food source in the area, living on the barnacle
exteriors and around the vents and seeps (Suzuki et al. 2009). Bathylasmatids
such as Tetrachaelasma tasmanicum, although not yet formally recorded from
within the New Zealand EEZ, almost certainly occur here. This taxon was recently
described from 3600 metres on the southeastern Tasman Rise (Buckeridge
1999b) where it is widely distributed as disassociated shells that are very similar
to isolated plates collected from New Zealand waters; in the absence of living
tissue the latter material has not been placed to species.
Although the total number of thoracican barnacle species from New Zealand
is not high compared with the numbers of species of taxa such as the Bryozoa
and Mollusca, it is high compared with cirripede faunas from other regions. In
particular there is a broader representation of known cirripede taxa (especially
PHYLUM ARTHROPODA
CRUSTACEA
phylogenetically primitive taxa) than in any region of comparable size, and there
is a disproportionately large number of species, both living and fossil, that have
their earliest records in New Zealand (Buckeridge 1996a).
Palaeontology and paleoecology
Acrothoracica
Acrothoracican burrows are known to occur in thick-shelled bivalves (e.g.
trigoniids) of Late Triassic age from Nelson and Southland (H. J. Campbell pers.
comm.) and belemnite guards (e.g. Belemnopsis alfurica) of Late Jurassic age from
Kawhia. These can be attributed to the ichnogenus Zapfella, to which the burrow
shapes generally conform; however, their true biological relationships remain
unclear and, as such, no move is made to classify them at ordinal level or below.
The Triassic record extends the range of Zapfella from that provided in Häntzschel
(1975) of ‘Jurassic to Tertiary’. Burrows are also known in Early Miocene deposits
from the Auckland region, e.g. Waiheke Island (J. A. Grant-Mackie pers. comm.),
and in turritellid gastropods from the Pakaurangi Formation, Kaipara Harbour.
The later burrows appear indistinguishable from modern Australophialus borings,
to which genus they are tentatively assigned.
Rhizocephala
Perhaps surprisingly, given their parasitic lifestyle, rhizocephalans are detectable
in the fossil record and are known from the New Zealand Miocene. Feldmann
(1998) studied a large number of beautifully preserved specimens of the large
xanthoid crab Tumidocarcinus giganteus. Several males had abnormally broad
abdomens, which is normally attributable to the parasitic castration induced by
the parasite.
Thoracica
Thoracican barnacles have a fossil record extending back to the Paleozoic, but
not in New Zealand. The pedunculate order Cyprilepadiformes is known from
the Silurian, attached to a eurypterid, and other thoracicans are known from the
Early Devonian and the Pennsylvanian (upper Carboniferous) (Newman et al.
1969; Buckeridge 1983; Foster & Buckeridge 1987; Newman 1996; Buckeridge
& Newman 2006). There is no record of Paleozoic cirripedes from the entire
New Zealand–Australian–Antarctic region, the first such record being Eolepas?
novaezelandiae from Middle Triassic strata of Southland (Buckeridge 1983).
Although there are rare scalpellomorphs of Jurassic age, it is not until
the Cretaceous that significant records are known – locally abundant, as-yetundescribed remains of Cretiscalpellum? are known from Middle Cretaceous
rocks in the Coverham area. These scalpellomorphs are preserved in association
with species of the large bivalve Inoceramus, upon which they appear to have
been growing. Hence, apart from a new verrucid from the Cretaceous of the
Waipara River in central Canterbury, the only barnacles known from the
New Zealand Mesozoic are stalked ones. Surprisingly, even though there are
barnacle-rich horizons in the Paleocene of the Chatham Islands, there are no
barnacles of Mesozoic age known from there. This is not likely to have resulted
from a paucity of appropriate facies, as there are some excellent Late Cretaceous
fossiliferous horizons present on Pitt Island that could have been expected to
have provided an appropriate environment for scalpellomorphs. At present,
it must be concluded that the absence of a Cretaceous barnacle fauna reflects
incomplete paleontological knowledge, and this provides an impetus for further
fieldwork on the islands.
Reconstruction of the fossil barnacle
Anguloscalpellum euglyphum (Oligocene).
John Buckeridge
Cenozoic barnacles
The New Zealand Cenozoic barnacle fauna is dominated by balanomorphs.
The first fossil cirripede to be described from New Zealand strata was the giant
111
NEW ZEALAND INVENTORY OF BIODIVERSITY
balanomorph Bathylasma aucklandicum, from Early Miocene strata near Auckland.
The locally abundant, but generally disarticulated plates of this sessile barnacle
were however, initially described as a pedunculate (Hector 1888). A quarter of
a century was to pass before the true nature of the remains was established,
in a paper wherein the author also described two new endemic species now
known as Anguloscalpellum ungulatum and Smilium subplanum (Withers 1913)
(see Jones 1992). In the early 1920s, Withers, working from the British Museum,
was commissioned by the then Geological Survey of New Zealand to produce a
monograph of the fossil cirripedes of New Zealand (Withers 1924). This listed 18
species, of which only 15 were truly fossil, and seven of these were both new and
endemic to New Zealand. In 1953, he published his last major work that dealt
specifically with cirripedes from New Zealand (Withers 1953). This included a
revised list of the New Zealand fossil fauna, arranged according to stratigraphic
horizons. He listed 15 species, none of which was new. Interestingly, he omitted
the record for ‘Balanus amphitrite’ that he included in his 1924 monograph, but
added the record for what is now Pristinolepas harringtoni. No reason is given
for his omission of ‘Balanus amphitrite’, which is now recognised in the New
Zealand fossil record as Amphibalanus variegatus. In all, Withers described nine
fossil cirripedes from the region, all of which are endemic.
Many limestones are so enriched with balanomorph remains that they may
justifiably be termed ‘barnacle coquinas’. The first horizons with locally abundant
balanomorphs are of late Paleocene age, occurring as lenses in the Red Bluff
Tuff of the Chatham Islands. In some of these lenses, the barnacle Pachylasma
veteranum is also the dominant macrofossil, with the other macrofauna primarily
being teeth of the elasmobranch fish Isurus sp. plus brachiopod and bivalve
shells. Although barnacle-rich horizons are also recorded in the Early Oligocene
(Cobden Limestone, West Coast), and Early Miocene (basal Cape Rodney
Formation, Auckland), it is the Pliocene coquina limestones of the North Island
East Coast that are singularly spectacular, e.g. the Pukenui and Castlepoint
Limestones, which contain extensive horizons dominated by Fosterella tubulatus
and Notobalanus vestitus. These coquinas outcrop at Rangitumau and Castlepoint respectively (both in the Wairarapa), and have extensive beds in which
F. tubulatus comprises more than 50% of the total mass. There are no modern
equivalents of these deposits, although lesser shell banks of N. vestitus and
Notomegabalanus decorus are today accumulating in the outer Hauraki Gulf near
the Mokohinau Islands. It is inferred by Beu et al. (1980) that these deposits
originated in subtidal settings dominated by strong currents, in a Pliocene sea
occupying the East Coast Inland Depression. These Pliocene ‘barnacle coquinas’
are not only impressive from a cirripedological perspective, they are also the
greatest accumulation of fossil crustaceans known!
Because barnacle species tend to be distributed along clearly delineated
depth, salinity, and temperature zones, their presence as fossils can be most
useful in paleoecological reconstruction. There are, however, some trends in
the ‘preferred’ environments of some taxa over time, e.g. species of the genus
Pachylasma are currently restricted to deep water, with the shallowest living
species of the group not known from less than 55 metres. In the Paleocene,
however, Pachylasma veteranum is known to have lived in very shallow water,
along with a diverse fauna of bryozoans, molluscs, and cnidarians, well within
the photic zone (Buckeridge 1983, 1999a). A similar pattern can be observed
with species of Bathylasma, which also occupied upper subtidal environments
in the Paleogene, but are now exclusively mid- to outer-shelf species. Indeed,
this change, which was interpreted by Buckeridge (1983) as ‘migratory’, is now
viewed more as a result of having been excluded (or outcompeted) from the
shallower-water environments by ‘modern’ balanomorphs. Modern taxa such as
Austrominius modestus have a higher metabolism and an earlier onset of sexual
maturity, which has permitted the species to aggressively exploit desirable
shallow-water niches. This has left refugial chthamalids (such as Chamaesipho
112
PHYLUM ARTHROPODA
columna and Chamaesipho brunnea) occupying upper littoral niches, and
pachylasmatines (such as Pachylasma scutistriata and Bathylasma alearum) midto outer-shelf environments (Buckeridge 1999a).
By the Late Miocene, it appears that thoracican barnacles occupied much
the same habitats as their modern counterparts (including as epibionts on
other crustaceans – Glaessner 1960, 1969). As a consequence, the zonation of
modern balanomorphs is useful in the reconstruction of the fossil depositional
environments that existed in the Late Cenozoic, e.g. in the barnacle-rich Titiokura Limestone of the eastern North Island Te Aute Limestone Complex. The
Titiokura Limestone (Beu 1995), outcropping in the northwest of Hawke’s Bay,
is characterised by a mixed assemblage of barnacles, including Pachylasma sp.,
Notomegabalanus miodecorus, and the inferred intertidal taxon Epopella cf. plicata.
The depositional environment at that time is, however, considered to have been
at more than 100 metres depth. The geological processes operating at the time
resulted in the build-up of shallow-water sediments on the upper shelf to a point
at which the accumulation became unstable. Sediments and faunas were then
mobilised, to be transported and deposited alongside deeper-water elements as a
mixed thanatocoenosis (death assemblage).
The sessile Balanomorpha are not known from strata older than the
Paleocene, with the first of these, Bathylasma rangatira and Pachylasma veteranum,
being recorded from the Chatham Islands (Buckeridge 1983). There has been
considerable conjecture concerning the origins of the balanomorphs, which
diversified and spread very rapidly in the Early Cenozoic. Buckeridge (1996a,
1999a) proposed that the Chatham Islands was a centre of sessilian diversification during the Paleogene, with taxa evolving in the warm shallow seas that
characterised the environmental conditions for strata like the Red Bluff Tuff. New
Zealand has a remarkable fossil cirripede fauna, with the phylogenetically early
taxa Eolasma, Chionelasmus, Waikalasma, Pachylasma, Bathylasma, Tetraclitella,
Palaeobalanus, Notobalanus, Chamaesipho, and Notomegabalanus having their
earliest records here.
As with the Recent fauna, there are a number of publications describing
single new species of New Zealand fossil Thoracica. These are listed in the
historical review provided in Buckeridge (1983), which also revised and
improved current knowledge of the New Zealand and Australian fossil
cirripede faunas. Buckeridge listed 69 fossil taxa from New Zealand, of which
36 were new. Of these, 94% (i.e. all but two) are endemic to New Zealand.
Since 1983, Buckeridge has described a further six species of fossil cirripedes
(Buckeridge 1984a,b, 1991, 1999a, 2008), and in addition has a further four new
taxa awaiting formal description.
CRUSTACEA
Waikalasma juneae (Miocene).
From Buckeridge 1983
Economic aspects of barnacles
Marine fouling
The first ‘close encounter’ some New Zealanders may have with barnacles is
when they need to remove fouling organisms from the hulls of their recreational
or fishing vessels. Barnacles are opportunistic organisms that colonise almost any
available surface in the marine environment. Boats and ships provide excellent
surfaces for suspension-feeders – a platform within the upper subtidal zone that
generally coincides with oxygenated, predator-poor, plankton-rich waters. In
addition, the mobile substratum facilitates dispersal.
Exotic fouling species in the New Zealand environment are generally
introduced through commercial shipping. It is in this way that the widespread
species Amphibalanus amphitrite, A. variegatus, and Lepas anatifera were introduced many decades ago. Lepas anserifera, Fistulobalanus albicostatus, Amphibalanus reticulatus, Megabalanus rosa, M. volcano, and Tetraclita squamosa japonica
were introduced on oil-drilling platforms (Foster & Willan 1979) but none appears
to have become naturalised in New Zealand waters. Hosie and Ahyong (2008)
113
NEW ZEALAND INVENTORY OF BIODIVERSITY
reported the establishment of the Australian species Austromegabalanus nigrescens and its South American congener A. psittacus at Taharoa and Wellington
respectively.
Research into the development of antifouling systems has intensified as a
result of a greater understanding of the deleterious ecological impact of traditional
antifouling paints such as tributyltin (Buckeridge 1998). Preliminary results
indicate that low-level ultrasonic transmitters have the potential to restrict organic
accumulation on certain hulls.
Barnacles as a food source
Although balanomorph barnacles such as the very large South American
Austromegabalanus psittacus are considered a delicacy, they do not occupy a
similar place in modern New Zealand cuisine. There is evidence, however,
that barnacles were once eaten by Maori, as they are often found in middens
(Foster 1986). In most cases, it appears that this was not through deliberate
harvesting; rather it was incidental to the harvesting of other seafood such as
Perna canaliculus (green-lipped mussel). This is no doubt a reflection of the
small size of most shallow-water New Zealand barnacles – many hundreds of
Austrominius modestus would need to be collected to make even a small meal.
Nevertheless, somewhat larger species such as Notomegabalanus decorus and
Epopella plicata may occasionally have been deliberately collected as a dietary
supplement (Foster 1986).
Environmental monitoring
Thoracican barnacles have a number of properties that may prove to be invaluable to humans. One that is currently under development is their use as
environmental indicators. Common shallow-water fouling species such as
Austrominius modestus and Epopella plicata are invaluable in monitoring environmental changes to marine systems during urbanisation (e.g. at Auckland’s
Long Bay–Okura Marine Reserve). A high metabolic rate, rapid onset of maturity, and frequent spawning make Austrominius modestus an excellent species
for gauging the impact of human activities.
Biotechnology
Another feature of thoracican barnacles that has intrigued scientists is the
means by which they attach themselves to surfaces. Barnacles are known to
grow on a very wide range of materials, both natural and synthetic. Their ability
to successfully adhere to flexible and elastic materials like plastic sheeting and
fibreglass is of specific interest, for if the nature of this ‘organic adhesive’ is
determined and commercially manufactured, it will have obvious use in fields
such as dentistry.
Barnacles that are commensal or symbiotic with other marine organisms
may need to produce chemicals to prevent the host overgrowing them. This is
particularly the case with sponge-inhabiting taxa like Acasta and coral-inhabiting
taxa like Brochiverruca. Isolation of chemical deterrents may be invaluable in
the design of new drugs for restricting or reducing cell growth in other species,
including humans.
Subclass Tantulocarida: Tantulocarids
Tantulus larva of Deoterthron dentatum
attached to an antenna seta of its ostracod host.
From Huys 1990
114
Nearly 30 years ago, a new maxillopodan subclass was created by Boxshall and
Lincoln (1983) to accommodate, amongst others, three tiny parasitic crustaceans
discovered in the New Zealand region (Bradford & Hewitt 1980; Boxshall &
Lincoln 1983; Lincoln & Boxshall 1983). They infect benthic and hyperbenthic
crustaceans such as amphipods. Tantulocarids are minute ectoparasites, not
exceeding half a millimetre (0.04–0.40 millimetre) in length, with a unique dual
life cycle that is completed, without moulting, on a crustacean host (Huys et al.
PHYLUM ARTHROPODA
CRUSTACEA
1993). There are now five recognised families with more than 20 genera and
about 30 species worldwide (Ohtsuka & Boxshall 1998), notably with several
taxa being recently documented from Japan (Huys et al. 1992; Huys et al. 1994;
Ohtsuka & Boxshall 1998).
While there have been no further records of tantulocarids from New Zealand,
it is very likely that more species of this subclass will be discovered as the benthic
and benthopelagic fauna of the New Zealand region becomes better studied.
Subclass Branchiura: Branchiurans
Branchiurans are parasitic on marine and freshwater fishes. They resemble
copepods in many respects but differ in some important features. Unlike
copepods, they have compound eyes and lateral head lobes, the opening of
the genital ducts lies between the fourth pair of thoracic limbs, and they have
a proximal extension to some of the exopodites (outer branch) of the thoracic
limbs. They are good swimmers and females deposit their eggs on stones and
other objects. The larvae differ little from the adult. Argulus has a pair of suckers
on the maxillae and a poison spine in front of the proboscis. One introduced
species has been recorded from goldfish in New Zealand (Hine et al. 2000). It is
likely that more species will be discovered.
Argulus japonicus.
Note the paired suckers.
Kenneth M. Bart
Subclass Pentastomida: Tongue worms
Tongue worms are obligatory parasites of reptiles, mammals, and birds, inhabiting
their respiratory tracts (nasal passages and lungs). Particularly prevalent in the
tropics, there are no native species in New Zealand, but one introduced species
has been reported (Tenquist & Charleston 2001). This is Linguatula serrata, whose
most regular host is the dog. It is rare in New Zealand, but developmental stages
have also been reported from the brown hare, European rabbit, house cat, and
sheep (Thomson 1922; Gurr 1953; Sweatman 1962).
Globally, there are about 130 species, ranging in length from about 3 to
150 millimetres or more and generally transparent or yellow to red-coloured.
Like most parasites, their body form is simple and wormlike. Blood is their only
food. The jawless mouth (sometimes protruding) and two pairs of lobe-like
appendages with claws give the appearance of five orifices, hence, penta- (five)
stomida (mouths). Long treated as a separate phylum of invertebrates, tongue
worms are now regarded as highly modified crustaceans, based on sperm
and larval morphology, the nervous system, and DNA studies. Some very
convincing fossils of apparent larval pentastomids from the Late Cambrian give
no evidence of a crustacean relationship, leading Maas and Waloszek (2001)
to question it. On the other hand, recent mitochondrial DNA sequencing
supports the evidence from sperm that pentastomids are most closely related
to the Branchiura (Lavrov et al. 2004).
Tongue worm Linguatula serrata.
Composite from various sources
Subclass Copepoda: Copepods
Copepoda (oar-footed bugs) are small crustaceans that are common in aquatic
and semi-aquatic environments, both marine and freshwater. Zoogeographical
data indicate that copepods are ancient arthropods (Dussart & Defaye 1995) and
fossils are known from the lower Cretaceous (Huys & Boxshall 1991). They have
undergone extensive adaptive radiation and include a wide variety of openwater, bottom-dwelling, herbivorous, predatory, and parasitic forms. Copepods
can often be extremely abundant and have been estimated to be among the most
numerous animals on earth, mostly because of their dominance in the plankton
of oceans and lakes. There are a number of excellent accounts that give general
information on copepods. The comprehensive monograph by Huys and Boxshall
(1991) deals especially with morphology and evolution, while Williamson
115
NEW ZEALAND INVENTORY OF BIODIVERSITY
Calocalanus pavo.
After Giesbrecht 1893
116
(1991) and Dussart and Defaye (1995) concentrate on the structure, function,
and taxonomy of freshwater species. Coull and Hicks (1983) and Mauchline
(1998) provide detailed information on the biology of harpacticoid and calanoid
copepods, respectively, especially the marine species. These references are the
main sources of the following notes.
The name ‘Copepoda’ is derived from two Greek words (kope, oar, and podos,
foot), hence oar-footed. Copepods are typically small, mostly in the range 0.5–
5.0 millimetres. Free-swimming forms may achieve a mimimum size of only 0.2
millimetres (some Oncaea) or a remarkable 18 millimetres (a Valdiviella species),
but some parasites are even larger. The body is usually approximately cylindrical
and segmented, and divided into three parts—cephalosome, metasome, and
urosome (equivalent to head, trunk, and abdomen). There are 10 pairs of
appendages on both the cephalosome and metasome, used for both feeding and
locomotion (some of these appendages also have a sensory function), and the
urosome ends in two bristle-bearing caudal rami. Uniquely among crustaceans,
copepods have a flat plate that connects the basal segments of each pair of
swimming legs. This plate is probably why copepods can have a rapid jumping
mode of movement. In all copepods the first thoracic segment (bearing the
maxillipeds) is incorporated in the cephalosome, unlike other maxillopodans.
The presence of a uniramous (unbranched) antennule is also a fairly reliable
copepod characteristic. In male copepods the first antennae can be typically
geniculate (with a prominent elbow), and are used to grasp the female during
mating. The antennae, mandibles, maxillules, maxillae and maxillipeds are used
in feeding. A wide variety of food types are utilised, including detritus, bacteria,
algae, rotifers, nematodes, naidid oligochaete worms, crustaceans, and larval
fish, and the structure of the feeding appendages varies in association with
diet. The mechanics of feeding are complex, although copepods are probably
fundamentally raptorial and use their mouthparts to grasp food particles.
Many species, however, especially calanoids, are suspension-feeders and use
the mouthparts to create water currents that bring food particles towards the
copepod. Smaller particles are then captured passively and directed towards
the mouth by bristles on the maxillipeds, maxillae, and maxillules, while larger
particles are individually grasped by ‘fling and clap’ movements of the maxillae
that grasp both the particle and a packet of water surrounding it and remove the
water by an inward squeeze.
Reproduction is usually sexual, and sperm are transferred from male
to female in a sac-like spermatophore (a few harpacticoids can reproduce
parthenogenetically). Egg sacs are probably not an ancestral condition of
Copepoda as many groups lack true egg sacs. Nevertheless, in many copepods the
eggs are carried in one or two egg masses, sacs, or strings until hatching. Under
favourable conditions, multiple clutches of eggs can be produced, at intervals of
a few days or weeks, so that each female may produce tens to hundreds of eggs
in a lifetime. The egg hatches into a nauplius larva and the life-cycle typically
includes six naupliar stages and six copepodite stages, the last of which is the
adult stage. There is a marked metamorphosis between the last nauplius and the
first copepodite stage. Development may sometimes be abbreviated, especially in
parasites. Copepods are relatively long-lived compared to other microcrustaceans.
Development times from egg to adult are typically in the order of 1–6 weeks, but
may take several months, and the lifespan of adults may be from one to several
months. Developmental times are markedly affected by temperature and food
levels. Some copepods have resting stages that enable avoidance of detrimental
environmental conditions and dispersal. Calanoids and harpacticoids produce
resting eggs that have a thick shell and which can survive extended periods of
dormancy and dryness. In cyclopoids and some harpacticoids, copepodites may
enter diapause and encyst in bottom sediments.
There are 11 orders, approximately 213 families, 1763 genera, and 11,956
species worldwide (Humes 1994; Ho 2003). The Harpacticoida alone comprises
PHYLUM ARTHROPODA
54 families, about 599 genera, and about 4400 species (J. Wells, unpublished data
updating Wells 2007). The Calanoida has 42 families with about 2000 species
(Boltovskoy et al. 1999); in the Poecilostomatoida there are 55 families, 359
genera, and about 1770 species (Ho 2003); and in the Siphonostomatoida there
are 45 families, 377 genera, and about 1840 species (Ho 2003). The known New
Zealand copepod fauna comprises 698 species, of which the Calanoida is the best
known with 261 species, nine of which are undescribed. There are only 230 species
of Harpacticoida, with about 99 of them undescribed; the remaining orders are
also very poorly known.
Copepods live in a remarkable number of environments. These include not
only marine and freshwater planktonic realms but in or on aquatic sediments,
in association with plants, forest litter, and damp moss, in subterranean habitats
or anchialine (isolated-marine) caves, and deep-sea hydrothermal-vent settings,
but also in association with other animals as commensals or parasites.
In the marine plankton, calanoid copepods (‘insects’ of the sea) are
extremely abundant. Some typical New Zealand examples are Acartia ensifera,
Calanus australis, Centropages aucklandicus, and Paracalanus indicus. They are
adapted to swimming in the water column and are fine-particle feeders in nearsurface waters, eating mainly phytoplankton and protozoans. Carnivorous or
detritivorous forms occupy deeper water-layers down to the deepest trenches.
In the water column we also find forms that are not strictly free-living but live
associated in some way with surfaces – the sea floor, the underside of sea ice, or
on other planktonic animals.
The freshwater plankton in New Zealand is dominated by calanoid copepods
of the family Centropagidae, which are widespread and very abundant in lakes,
ponds, and the lower reaches of larger rivers. Many of the species also occur in
Australia, although there are at least three endemic species. Calamoecia lucasi
and Boeckella dilatata are typical lake dwellers while B. triarticulata is found in
ponds. As in marine habitats, the freshwater calanoids are suspension-feeders
on algae and protozoans, although at least some of the boeckellids are also
predatory on small zooplankters such as rotifers and nauplii. A few cyclopoid
copepods also live in fresh water, although they are usually sparser than the
calanoids. They are probably mostly omnivores, consuming both animals and
algae. Some are found mainly in the bottom waters and are probably strays from
the benthic and littoral areas.
In aquatic sediments, copepods (mainly harpacticoids) live either permanently within the sediment or alternate between the sediment and its surface,
browsing on the microflora associated with the sediment particles or with the
accompanying detritus. In well-oxygenated coarse-grained sediments such as
beach sand, specialised copepods (again, mainly harpacticoids) are part of the
‘interstitial fauna’ that lives within the interstices of this habitat. This habitat is
commoner in marine sediments than in freshwater sediments, although it does
exist in river systems and their ground waters where a strong intra-sediment
water flow occurs. Most families of Harpacticoida have representatives in all of
the above habitats, with specialisations for the interstitial habitat having evolved
many times in different lineages. These trends exist among the New Zealand
fauna to the same extent as they do elsewhere and are represented by numerous
endemic and non-endemic species. An extremely important characteristic of this
fauna is that, with very few exceptions, the entire life-cycle is benthic and the
larvae are not dispersed large distances by water movements. This not only must
affect their ecology but must also impact on population genetics and eventually
on phylogeny. As a result we should expect a high level of endemism.
Many copepods are associates of plants. In the marine intertidal zone
many harpacticoids live in association with seaweeds and sea grasses and
are highly specialised for life on the surface of the fronds. Members of the
Porcellidiidae, Peltidiidae, and Tegastidae, for example, are especially adapted
to this environment; each family is well represented in New Zealand. In the
CRUSTACEA
Centropages aucklandicus – female at top
(left profile and dorsal views), male below,
with modified antenna for copulation.
From Bradford-Grieve 1999
117
NEW ZEALAND INVENTORY OF BIODIVERSITY
Paramphiascopsis waihonu.
From Hicks 1986
118
littoral areas of freshwater lakes, ponds, and running waters, cyclopoids and
harpacticoids are abundant on and amongst macrophytes. Damp terrestrial
situations are exploited by cyclopoid and harpacticoid copepods. These include
damp soil, forest litter, sphagnum bogs, liverwort and moss clumps, and the
pools between the leaves of bromeliads. Only the harpacticoids from this cryptic
fauna have been extensively studied in New Zealand, and in these the same
trends exist as elsewhere in the world; most species belong to cosmopolitan
genera in the predominantly freshwater family Canthocamptidae, and most are
endemic.
Copepods live in groundwater and can be caught in springs, wells, and pools
in caves. In New Zealand these habitats have not been extensively surveyed
(Chapman & Lewis 1976) and nothing is known about the copepods except that
parastenocaridids have not been found, despite extensive searching (Schminke
1981a). Overseas, the Parastenocarididae (Harpacticoida) is a large family of ca.
270 species (190 of them currently placed in the genus Parastenocaris) that mostly
inhabit the interstices of groundwater. These habitats range from the water
table beneath beaches and sand banks, including a few fully marine beaches,
to brackish systems such as the Baltic Sea, and riverine and lacustrine inland
systems, above and below ground.
Recently the study of deep-sea hydrothermal vents and marine caves
has revealed many interesting copepods of great importance to the study of
evolutionary relationships between the various groups of copepods, as they are
amongst the most primitive forms. Because isolated marine caves are not yet
known in New Zealand and the microscopic fauna of New Zealand hydrothermal
vents has not yet been studied, these types of copepods have not been recorded
here.
In thermal waters of the central North Island only one copepod, the endemic
cyclopoid Paracyclops waiariki, is known. It is restricted to Lake Rotowhero,
which has seasonal temperatures varying between 29.5° and 37.5° C and an
average pH of 3.1.
Nearly half of all known copepod species live in symbiotic relationships with
other organisms. It is evident that commensalism and parasitism have evolved
independently several times in the class, even within an order. Copepods parasitise
virtually every phylum of animals from sponges and cnidarians to vertebrates
including mammals. They also have a range of associations from external and
internal parasitism to varied forms of commensalism. For example, two species
of endemic New Zealand harpacticoids are associated with macroinvertebrates
– Porcellidium tapui on hermit crabs and Alteuthoides kootare on sponges. It
is interesting to note that these genera are highly adapted for clinging to a
substratum and are genuinely ‘phytal’ in this respect. This particular association
with macroinvertebrates is almost certainly of the same type as with marine plants,
i.e. using them as a substratum on which bacteria, fungi, and microalgae grow
abundantly. Similarly, Paramphiascopsis waihonu is known only from a sample
of spent elasmobranch embryo cases (taken at 1116 m), where many specimens
occurred along with a gastropod mollusc; an association with the gastropod
is unlikely and it is most probable that both are feeding on detritus and decay
products within the case. Paramphiascopsis comprises several other species that
have been taken in association with ascidians, polychaetes, gorgonians, and
decapod crustaceans but many species are also known from algae and sediments.
Harpacticoids are also found in burrows in wood inhabited by the gribble
(Limnoria spp.), where the nature of the association is unclear (Hicks 1988a),
with some authors arguing for an obligate commensal relationship and others
believing the attraction for the copepod is the microhabitat created by the
gribble. Evidence for the latter is the presence of copepods in decaying wood no
longer occupied by Limnoria, but the fact remains that the copepod species have
never been found in habitats that have not been associated with the gribble. Five
species, of which four are endemic, occupy this habitat in New Zealand waters.
PHYLUM ARTHROPODA
CRUSTACEA
Importance of copepods
In both marine and fresh waters worldwide, abundant copepods form a vital
link in the food web that leads from minute algal cells or phytoplankton and
small protozoans (e.g. Chapman & Green 1987; Bradford-Grieve et al. 1998)
to the largest fishes, and some whales in the oceans. Many commercial and
non-commercial marine fish (and some crustaceans) are utterly dependent on
copepods as a food source during a portion of their larval life. For example, in New
Zealand it has been shown that the larvae of hoki (Macruronus novaezelandiae),
which forms the basis of the largest New Zealand fishery, feed on copepod adults
(e.g. Calocalanus) and copepodites almost exclusively (Murdoch 1990). With their
large mouth size, hoki larvae actively select copepods such as Calocalanus and
Paracalanus (Murdoch & Quigley 1994). For inshore benthos and for migratory
fish, estuaries and lagoons are typically the critical location for this life-history
phase. In a New Zealand estuary, Parastenhelia megarostrum is a principal prey
item for young post-metamorphic flatfish during the first six months of their
lives (Hicks 1984). The very smallest fish feed on the naupliar stages while larger
specimens have an increasing proportion of older copepods in their guts. In
lakes, copepods are an important part of the diet of smelt (e.g. Stephens 1984,
Chapman & Green 1987), which in turn form a major part of the diet of rainbow
trout. Copepods can be so abundant that their faecal pellets, produced at a rate
of several per hour, are an important source of food for detritus feeders. Copepod
grazing can significantly reduce the densities of at least some algal species (e.g.
Edgar & Green 1994) and it has been suggested that they may have potential
in the biomanipulation of the effects of eutrophication in lakes (Edgar 1993).
Copepods are increasingly being used as test organisms in ecotoxicological testing.
In New Zealand, the freshwater species Calamoecia lucasi, Boeckella delicata, and
Mesocyclops sp. have been shown to be very sensitive to pentachlorophenol (Willis
1998) and the latter two species have been recommended as suitable candidates
for the development of routine testing protocols involving acute and chronic
endpoints (Willis 1999).
Copepods can be important economic pests when they parasitise commercial
species. This is especially the case overseas, where ectoparasitic copepods of the
families Ergasilidae and Caligidae (‘sealice’) infect salmonids reared in sea cages,
causing damage and sometimes death of valuable aquacultured product reared
in marine areas (Johnson et al. 1997). In New Zealand, copepod ‘sealice’ are not
yet a problem in salmon culture (Hine & Jones 1994) but the causative copepod
genera are present in the farms (Jones 1988a). Copepods of the family Sphyriidae
are also of economic importance in that the anterior portion of the copepod is
buried in the musculature of the host fish, while the posterior portion bearing
egg strings trails from a hole in the skin. Skinning machines do not remove the
‘head’ from the fillet causing wastage and customer complaints.
In freshwaters, the ergasilid Abergasilus amplexus infests a wide variety of
fish including longfinned and shortfinned eels, smelt, inanga, goldfish, and
perch (e.g. Jones 1981). Two other parasitic copepods, Thersitina inopinata and
Paeonodes nemaformis, are rather enigmatic (McDowall 1990). Thersitina inopinata
is known only from its free-swimming males, while P. nemaformis, although
endemic, is known to parasitise only introduced brown trout and salmon. The
exotic copepod Lernaea cyprinacea has been recorded from introduced goldfish.
Free-living copepods are also known to be intermediate hosts in the life-cycles
of tapeworms of freshwater fish. The initial stages of Amurotaenia decidua,
which parasitises bullies, occur in Macrocyclops albidus (Weekes 1986) and
planktonic copepods are secondary hosts in the life-cycle of Ligula intestinalis,
the pleurocercoid of which infests both rainbow trout and bullies (Weekes &
Penlington 1986).
Copepods can be disease vectors for human parasites in tropical climates.
But conversely they can also carry the fungi or sporozoans that parasitise
Abergasilus amplexus.
From Jones 1981.
119
NEW ZEALAND INVENTORY OF BIODIVERSITY
malarial mosquitoes. Copepods have been implicated in the spread of viruses
through fish populations (Mulcahy et al. 1990). Freshwater copepods of the
genera Mesocyclops and Macrocyclops have been used for control of the containerbreeding mosquito species of Aedes, Anopheles, and Culex. So far, no examples of
these kinds of relationships have been noted in New Zealand.
Zoogeography of the New Zealand copepod fauna
Calanus australis (female)
From Bradford-Grieve 1994
120
Marine plankton
Very few marine planktonic copepods are endemic to the New Zealand region.
The distribution of pelagic Copepoda (Bradford & Jillett 1980; Bradford et al.
1983; Bradford-Grieve 1994, 1999a) in the region appears to be maintained by
a combination of factors probably related to their occurrence in water masses
in some way or other. The physiological requirements of a species (temperature
tolerances, ability to breed in differing temperature regimes, nutritional
requirements for growth and breeding) and their behaviour (vertical migration
in relation to particular water masses or physical-oceanographic phenomena)
all contribute to the patterns we observe. An additional factor (plate tectonics)
was probably important in the occurrence of some neritic plankton species in the
New Zealand region.
Some species have a clearly coastal distribution. Among the New Zealand
epipelagic calanoids, only species of Acartiidae, Calanidae, Centropagidae,
Clausocalanidae, Paracalanidae, Pontellidae, and Temoridae contain coastal forms
that are rarely encountered in oceanic waters. Endemic coastal species such as
the calanoids Acartia ensifera, A. jilletti, A. simplex, and Centropages aucklandicus
and the poecilostomatoid Corycaeus aucklandicus are confined to New Zealand
waters, whereas Gladioferens pectinatus, Labodocera cervi, and Sulcanus conflictus
are confined to Australia and New Zealand. Calanus australis is found in at least
New Zealand and southeastern Australian coastal waters, where it is essentially
restricted to the mid-shelf (Bradford 1985). It seems possible that many of these
species had common ancestors with close relatives in other temperate neritic
parts of the world as far back as the Oligocene, when equatorial sea temperatures
were low (Bradford 1979). Paracalanus indicus is restricted to coastal waters, with
maximum concentrations occurring close to shore (Bradford 1985), although
this species possibly has a broad tropical/subtropical distribution. Clausocalanus
jobei and Temora turbinata also have a tropical/subtropical distribution whereas
Drepanopus pectinatus has a coastal distribution around subantarctic islands.
Relationships to water masses are most clearly seen among oceanic epipelagic
species. Nevertheless, in the New Zealand region some oceanic species are capable
of responding rapidly to the heightened productivity of coastal waters and may
attain maximum numbers close to the coast, obscuring their oceanic affinities.
Examples of this type of distribution are seen in the calanoids Nannocalanus
minor and Clausocalanus ingens and the cyclopoid Oithona similis.
Warm-water (tropical) oceanic epipelagic species usually have a cosmopolitan distribution if they are able to breed at a range of latitudes extending to 40°
S, whereas those with breeding ranges restricted to lower latitudes (e.g. Euchaeta
rimana) are not circumglobal in their distribution because of the geographical
barriers (South America and Africa) presented to their distribution. In tropical
or subtropical waters, epipelagic calanoid species with distributions extending
to 40° S and sometimes as far as the Subtropical Front are Aetideus giesbrechti,
many Calocalanus species, Clausocalanus arcuicornis, C. lividus, C. parapergens, C.
paululus, C. pergens, Eucalanus hyalinus, Mecynocera clausi, Nannocalanus minor,
Neocalanus gracilis, Pareucalanus sewelli, Pareuchaeta acuta, P. media, Rhincalanus
nasutus, and Subeucalanus crassus. Species with a warm-temperature (transition zone) Southern Hemisphere distribution include Aetideus pseudarmatus,
Clausocalanus ingens, Pareucalanus langae, and possibly Neocalanus tonsus and
Calanoides macrocarinatus. Species with subantarctic distributions include Cala-
PHYLUM ARTHROPODA
CRUSTACEA
nus simillimus, Clausocalanus brevipes, Neocalanus tonsus, and Subeucalanus longiceps. Species with Antarctic–subantarctic distributions include Aetideus australis,
Clausocalanus laticeps, and Rhincalanus gigas.
Marine sediments
Throughout the world the copepod fauna of marine sediments (predominantly
harpacticoids) is well known only for the intertidal and shallow sea areas.
Detailed data are available for only a few sites of more than a few metres
in depth, mostly in Europe, although scattered information is known for all
depths down to almost the bottom of the deepest trenches. Even for intertidal
and sublittoral areas, most of the world outside Atlantic Europe, the western
Mediterranean, and a few locations on the eastern coast of the Americas
is poorly known or even totally unknown. A reasonably comprehensive
survey of the North and South Islands of New Zealand has been carried
out, but the results have yet to be fully published and many species remain
unnamed. Furthermore, assessment of the zoogeographic relationships of the
New Zealand fauna is made impossible by the almost complete absence of
information from Australia and New Caledonia. All that can be said at this
time is that it seems unlikely that New Zealand will harbour many endemic
genera (though that will depend on the attitude of future taxonomists towards
taxon definitions).
Freshwater plankton
In New Zealand, most freshwater calanoids (eight species of Boeckella and one
of Calamoecia) belong to the family Centropagidae, the non-marine members
of which are mainly confined to Australasia, the subantarctic, the Antarctic
Peninsula, and parts of South America (Bayly 1992). Only three of these species
are found only in New Zealand (Jamieson 1998); the others also occur in
Australia. A further four species are considered to be resident natives (Boeckella
dilatata, B. propinqua, B. triarticulata, and Calamoecia lucasi) whereas B. minuta
and B. symmetrica may have invaded New Zealand since European colonisation
(Banks & Duggan 2009). Recently, the diaptomid cross-hemisphere invaders
Skistodiaptomus pallidus and Sinodiaptomus valkanovi have been recorded in
constructed water bodies (Duggan et al. 2006; Banks & Duggan 2009; Makino
et al. 2009).
Bayly (1995 and references therein) concluded that the present-day
distribution of freshwater and brackish Centropagidae can be interpreted as being
a result of the colonisation of southern-hemisphere inland waters from marine
and then brackish-water ancestors at a time when Australia, New Zealand, and
South America were still linked to Antarctica, and Africa, Madagascar, and India
had already drifted northwards. The absence of the Diaptomidae from New
Zealand, most of Australia, and all of Antarctica also appears to be related to
the timing of the separation of these landmasses from Pangaea in relation to the
evolution of this family.
The distribution of calanoids in the major lakes is probably well known
(Chapman & Green 1987; Jamieson 1988, 1998; Bayly 1992; Banks & Duggan
2009) but has yet to be fully examined in smaller habitats, especially ephemeral
pools and the less-accessible high-country tarns. Most species show relatively
clear habitat segregation. Calamoecia lucasi is widespread in northern, central,
and western parts of the North Island, where it is found in streams, ponds, and
large rivers. It also lives in a few small lakes in Northern Nelson. Calamoecia
ampulla, a widespread species in Australia, is known only from one unverified
South Island record (Bayly pers. comm.). Of the Boeckella species, B. minuta, B.
symmetrica, and B. tanea have restricted distributions in the North Island. Boeckella
tanea is found only in Northland, B. symmetrica in a pond near Auckland, and B.
minuta in the Waikato River hydroelectric reservoirs and water-supply reservoirs
in Wellington. It has been suggested that B. symmetrica and B. minuta may be
121
NEW ZEALAND INVENTORY OF BIODIVERSITY
Abdiacyclops cirratus, an endemic
cyclopoid genus andspecies from a
subterranean well in Canterbury.
From Karanovic 2005
122
recent immigrants from Australia (Chapman & Green 1987) and this may apply
to C. ampulla too. Boeckella propinqua occurs mainly in central and northern areas
of the North Island but, like C. lucasi, its distribution also extends to the tip of the
South Island. Boeckella hamata occurs throughout the southeastern part of the
North Island, the eastern part of the South Island, and southern Westland, mainly
in reservoirs and coastal lakes. Boeckella triarticulata has a similar distribution but
apparently does not co-occur with B. hamata. It is found mainly in ponds and
reservoirs in eastern parts of the South Island from Canterbury to Otago, with
one record from Hawke’s Bay in the North Island. Boeckella delicata has a disjunct
distribution, occurring in Northland and the Waikato region of the North Island
and also on the west coast of the South Island. Boeckella dilatata occurs only in
the South Island, mainly in glacial lakes and in associated reservoirs. It also has
a disjunct distribution and is found only in northern and southern areas of this
island. Unlike the usual situation elsewhere in the world, co-occurrences of two
or more species of calanoids in one lake are rare, and most lakes have only one
calanoid. In the North Island, there are a few co-occurrences of C. lucasi and B.
delicata, C. lucasi and B. propinqua, and C. lucasi and B. minuta, and in the South
Island B. triarticulata and B. dilatata, B. triarticulata and B. hamata, and C. lucasi
and B. propinqua in a few habitats (Chapman & Green 1987; Jamieson 1998;
Banks & Duggan 2009).
Various attempts have been made to explain the distributional patterns of the
New Zealand freshwater calanoids (summarised by Jamieson 1998) and, until
recently, most of these used dispersalist biogeographical ideas. Banks and Duggan
(2009) have highlighted the role of constructed lakes and ponds in facilitating
inter-and intracontinental invasions of calanoid species. Maly (1984) suggested
that distributions resulted from probabilities of immigration and extinction that
were assessed from clutch sizes and the likelihood of predation by fish. Maly
(1991) modified these ideas to include the number of existing populations and
concluded that dispersal was probably not important over long distances but
may be important at local scales. Jamieson (1988) explained the distribution of
Boeckella dilatata, B. hamata, and B. triarticulata by relating differences in their
ecological requirements and dispersal abilities to vicariant events. More recently,
Jamieson (1998) has provided a convincing explanation for the distribution of
these three species and B. delicata based on panbiogeographic methods. She
showed that their distributions are correlated with the three principal pre-Late
Cretaceous technostratigraphic terranes that, over the last 150–200 million years,
have come together to make up New Zealand. Boeckella dilatata and B. delicata
occur in lakes and ponds on the Tuhua and Caples Terranes and B. hamata and B.
triarticulata on the Torlesse Terrane. The species overlap at the terrane margins.
The present-day disjunct distributions of B. dilatata and B. delicata are thus
thought to result from tracks arcing out to sea.
The species pairs on the different terrane groups are thought to differ in
ecology; in particular B. delicata and B. hamata are suggested to have a higher
salt tolerance than either B. dilatata or B. triarticulata, thus enabling sympatry.
Localised dispersal presumably explains the overlap of species at the terrane
margins. Jamieson’s panbiogeographic approach would seem to have considerable potential for explaining distributions of the remaining calanoids. It is
clear, however, that ecological information remains important for explaining
distributions of sympatric species. Ecological studies of life-histories and food
requirements have been made of some species (e.g. Green 1975; Forsyth &
James 1984; Jamieson 1986; Chapman & Green 1987; Burns 1988; Jamieson &
Burns 1988; Xu & Burns 1991; Burns & Xu 1990; Twombly et al. 1998; Couch
et al. 1999), but much more remains to be done. The effects of post-European
colonisation, with altered fish communities and changing trophic status of lakes,
on distributional patterns are not known.
The cyclopoid copepod fauna is very poorly known taxonomically and
ecologically. A few cyclopoids are found in the lake plankton, but their
PHYLUM ARTHROPODA
CRUSTACEA
populations are usually either sparse or seasonal and little is known about them.
There are no equivalents of the large-bodied Cyclops (in the strict sense) of many
Northern Hemisphere lakes.
Mesocyclops leuckarti has been recorded from various North Island lakes
(Green 1974, 1976; Jamieson 1977; Chapman & Green 1987; Greenwood et al.
1999), but it is likely that these records were not of the nominate species as M.
leuckarti does not occur in the Southern Hemisphere (Kiefer 1981). Bayly (1995)
has suggested that its correct identity is possibly M. australiensis. Macrocyclops
albidus occurs in low numbers in the Rotorua and Taupo lakes (e.g. Chapman
1973; Forsyth & McCallum 1980), in the lakes of the Waitaki River system, and in
other South Island lakes (Stout 1978; Burns & Mitchell 1980). Eucyclops serrulatus
is found in the plankton of Lakes Hayes and Johnson (Burns & Mitchell 1980) and
Acanthocyclops robustus in the plankton of Lake Mahinerangi (Mitchell 1975). It
still can be concluded that, until a revision is made of the freshwater cyclopoids,
no valid assessments of biogeographical relationships can be made. Nevertheless,
Karanovic (2005) held it to be highly likely that the cosmopolitan cyclopoids
Acanthocyclops robustus, Diacyclops bisetosus, Eucyclops serrulatus, and Paracyclops
fimbriatus were accidentally introduced to New Zealand by early European settlers
in barrels of fresh water. Jamieson (1980a, b) conducted experimental studies of
predatory feeding and development rates of Mesocyclops sp.
Plant associates
In marine systems the term ‘plant associates’ means the fauna associated with
macroalgae and sea grasses and is usually called the phytal habitat. In addition,
a few species have been found associated only with decaying wood (from wharf
piles to driftwood dredged from depths of 1100 metres). These perhaps should
be included in the phytal fauna as it is most probable that the role of the living
or dead plant is primarily as a substratum for the copepods’ food supply, namely
bacteria, fungi, and microalgae attached to the plant. However, in this regard the
phytal fauna is little different from the true benthos, which relies on these food
sources attached to particles of the sediment.
Most of the species do not show obvious morphological adaptations
to the phytal habitat. In those that do, the adaptations are usually to enable
the animal to attach itself more effectively to the plant. Very few species seem
actually to damage the plant or to be directly feeding on its tissues. Many genera
that contain species found among algae have other species living on or in the
adjacent benthic sediment. Many species are found equally often among algae
and in sediments without associated plant growth. Also, it is known that many
of the species washed from samples of macroalgae and sea grasses are actually
associated with the sediment and detritus that becomes trapped in the interstices
of the plant and thus are really part of the sediment fauna. Even many of the
truly phytal species that do show adaptations to that environment have been
shown to leave the plant for mating; this may partially explain the relative rarity
of males in collections of these species.
In the marine system, about 45% of the described phytal species are endemic.
Only a few undescribed species currently exist in collections, which may partly
be a consequence of inadequate collecting and cataloguing. Notwithstanding,
the phytal fauna is quite well known ecologically (e.g. Hicks 1977, 1988b) and,
while it is very probable that many species remain to be discovered, the main
outlines of the fauna are well known. Unfortunately, the phytal fauna of adjacent
marine regions is as poorly known as their sediment fauna and similar remarks
about understanding zoogeographical relationships apply. The comments below
on endemism in the sediment fauna apply equally to the phytal but the lack of
regional collecting makes it futile to try to estimate the true level of endemism.
The situation in freshwater and terrestrial systems is much the same.
Some copepods (cyclopoids and harpacticoids) probably use plants mainly
as the substratum on which their food grows, but much less is known about
Goniocyclops silvestris (female).
From Karanovic 2005
123
NEW ZEALAND INVENTORY OF BIODIVERSITY
their ecology. Certain copepods are found associated with aquatic vegetation
in lakes and ponds, and with mosses (Harding 1958; Chapman & Lewis 1976).
In semiterrestrial situations such as mossy banks and the edges of waterfalls or
in damp forest litter and decaying wood, some copepods (such as Goniocyclops
silvestris and a variety of harpacticoids) are found; most are apparently endemic
but this fauna has still to be properly examined (Chapman & Lewis 1976).
The fish parasite Caligus pelamydis,
from barracouta.
From Hewitt 1963
Animal associates
It is difficult to make any definitive statement about the zoogeography of animal
associates because the commensal and parasitic copepod fauna of marine
invertebrates in New Zealand and neighbouring seas is very poorly known.
For example, known New Zealand siphonostomatoid species diversity is only
29% of that in European seas, and even less for cyclopoids and harpacticoids,
whereas, based on what is known for well-studied high-level Animalia taxa
in both regions, New Zealand species diversity matches or exceeds that in
European waters (Gordon et al. in press). The end-chapter checklist of New
Zealand species in these copepod orders is annotated to indicate the type of
relationship and host.
Species identifications of parasitic copepods from fishes of neighbouring
seas are, in many cases, awaiting critical review. For example, Trifur lotellae in
New Zealand would appear to be identical to Trifur physiculi from Australia.
There are many other such examples. Also, the parasitic copepod fauna of marine
invertebrates in New Zealand and neighbouring seas is almost totally unknown.
Nevertheless, Jones (1988b) examined the then known parasitic copepod fauna
and concluded that endemism on teleosts at the generic level was very low (2%)
and there were no endemic genera on elasmobranchs (sharks).
The freshwater parasitic copepod fauna consists of only three species –
Abergasilus amplexus and two very rare or extinct species, Thersitina inopinata and
Paeonodes nemaformis. Abergasilus is an endemic estuarine genus common in, and
known only from, Lake Ellesmere and the Chatham Islands lagoon. It has close
affinities with South American genera. Thersitina has been found only once, in
a plankton sample from Lake Poerua (Percival 1937). Paeonodes nemaformis has
been found only twice, both times in South Westland on introduced salmonids
(Hewitt 1969). The genus has also been found in Africa and is apparently closely
related to Mugilicola, found in South Africa, India, and Australia (Boxshall 1986).
The native hosts of Thersitina and Paeonodes are unknown, despite extensive
searching. It is concluded that the parasitic copepod fauna of marine vertebrates
is derived from the wandering of host fishes and reflects the strong links with
Australia and the island chains to the north (Jones 1988a,b).
Endemism
One key element in the occurrence of endemism in New Zealand is the
paleogeography of the region. The freshwater, brackish, and inshore copepod
faunas illustrate the key elements of such reconstructions (Lewis 1984; Bayly
1995). The absence of the calanoid family Diaptomidae and presence of
freshwater species of Centropagidae in Australia, New Zealand, South America,
and Antarctica indicates that the period when these land masses were still
linked but already separated from Africa, Madagascar, and India (120–80 million
years ago) is crucial in reconstructing the evolution of Boeckella, Calamoecia, and
Gladioferens in New Zealand and other southern hemisphere regions. These
events, and the subsequent submergence of New Zealand in the Oligocene (35
million years ago) were probably responsible for speciation and the currently
observed endemism (Bayly 1995).
The connection between New Zealand and Antarctica was broken during
the Late Cretaceous. Three of eight New Zealand species of Boeckella are endemic
to New Zealand (Maly & Bayly 1991) and it is likely that this genus inhabited
the fresh waters of the ancestral landmass when it separated from Antarctica.
124
PHYLUM ARTHROPODA
CRUSTACEA
By the Late Oligocene, nearly all of the New Zealand landmass (possibly all of
it according to Landis et al. 2008) was submerged. Significant extinctions will
have occurred at this time, accounting for the relatively impoverished fauna of
New Zealand compared with that of Tasmania. On the other hand, the multiple
vicariant events associated with the production of a diminishing New Zealand
archipelago in the Oligocene might have been expected to result in some
speciation and the currently observed endemism if not all of the landmass was
in fact submerged.
We predict that a higher degree of endemism than is currently recorded will
be discovered amongst freshwater and benthic copepods when the less wellknown groups are revised. But we need to introduce here a note of caution
in this discussion of endemism. While the number of endemic species indeed
reflects the evolutionary history of a particular fauna, in practice the number of
such species recognised by past and present taxonomists depends on the interpretation of morphological variability within a species, especially where there is
discontinuous distribution and not enough morphomolecular information for
phylogenetic analysis.
Marine plankton
Very few marine planktonic species are endemic to New Zealand. The main
reason for this is that most species are oceanic and are relatively widespread
in a global sense, ranging from circumglobal subantarctic and Indo-Pacific to
distributions encompassing all the world’s oceans. Only a few coastal calanoid or
cyclopoid species are endemic to New Zealand waters (Acartia ensifera, A. jilletti,
A. simplex, Centropages aucklandicus, and Corycaeus aucklandicus). The cyclopoid
Corycaeus aucklandicus is endemic to coastal waters of northern New Zealand.
Freshwater plankton and benthos
Only three freshwater calanoid species are endemic – Boeckella dilatata, B.
hamata, and B. tanea; the other seven species also occur in Australia. Only two
(Metacyclops monacanthus, Paracyclops waiariki) of the 19 cyclopoid species are
known to be endemic to New Zealand. All others are supposedly cosmopolitan
or Australasian. Notably, several genera recorded from Australia, some with
multiple species (Apocyclops, Australocyclops, Ectocyclops, Mixocyclops, Neocyclops,
Thermocyclops), have not yet been recorded from New Zealand. Some studies
(see Bayly 1995) have shown much greater degrees of differentiation and
endemicity than previously recognised in microcrustaceans, and it is evident
that more stringent resolution of morphotypic variation of the New Zealand
freshwater cyclopoids is required before their status can be assessed. Presumed
‘cosmopolitan’ species may be so only because of widespread and indiscriminate
misuse of authoritative (?northern hemisphere) taxonomic references. As
noted earlier for Mesocyclops leuckarti (discovered to be a species complex by
Kiefer (1981) and not represented by the nominate species in the Southern
Hemisphere), comparable species groups may be found in other ‘cosmopolitan’
species. An on-going global revision of the Cyclopoida (e.g. Dussart & Defaye
1995; Einsle 1996) will help resolve some of the problems. This series should be
consulted as a guide to the global literature on cyclopoid genera and families,
and in particular for the accepted modern level of taxonomic discrimination.
Acartia ensifera.
After Bradford-Grieve 1994
Marine sediments
Approximately 50% of the described harpacticoid species are endemic, but at
least three times as many species remain undescribed in collections, and it is
reasonable to estimate that at least 75% of these will prove to be endemic new
species. It would seem, therefore, that the rate of endemism in New Zealand
is high compared, for example, to the British Isles (as an example of another
island group of comparable size), where probably it is less than 10%. But this
comparison is meaningless. The British fauna has been investigated for much
125
NEW ZEALAND INVENTORY OF BIODIVERSITY
longer and at much greater intensity. As a result, it is known to contain at least
four times as many species. Further, and very importantly, the British Isles are
close to the shores of northwestern Europe, where the fauna is also very well
known and shares many species with Britain. New Zealand is distant from its
nearest neighbours. This, and its geological history since separation from the rest
of Gondwana, may well have increased the level of endemism, but the lack of
data from Australia (where the fauna is very poorly known) undoubtedly inflates
the current estimates.
The limited amount that is known about the benthopelagic calanoid fauna
indicates that there may be some degree of endemism (e.g. Bradford 1969;
Bradford-Grieve 1999b) in the New Zealand region. Nevertheless, in the deep
sea the perception of endemicity may reflect the paucity of sampling of nearbottom faunas worldwide.
Cryptic habitats
Freshwater harpacticoids in New Zealand have been collected mainly from
clumps of moss or liverworts or similar vegetation in streams, the littoral of
ponds and lakes, or from wet banks close to water bodies and in damp forest
in leaf litter. Of the 19 named species in the end-chapter checklist, 17 are
endemic, but relatively little collecting has been carried out and large areas of
the country remain unexplored. The total fauna is likely to be many times the
recorded number of species, but it is probable that a very high level of endemism,
and of localised distribution of species, may be found. It will be interesting to
see if their distribution supports the panbiogeographic explanation for the
distribution of freshwater planktonic Calanoida (Jamieson 1998). The presence
of small cyclopoid species has also been noted, but only one has been identified
to species and the true extent of this fauna cannot be estimated at this time
(Chapman & Lewis 1976).
Gaps in taxonomic knowledge of copepods and scope for
future research
Platycopioida
This order is not known in the New Zealand region. It is possible that
platycopioids will be found when the benthopelagic realm is properly sampled,
because they have been found in other temperate, shallow-water, near-bottom
habitats. Other genera have been found in marine caves in Bermuda so their
relatives might not be expected to occur in New Zealand.
Metridia lucens (Calanoida).
From Bradford-Grieve 1999
126
Calanoida
The marine pelagic calanoid copepod fauna of New Zealand is fairly well known,
mainly from the work of Janet Bradford-Grieve. The end-chapter crustacean
species list incorporates results from Bradford and Jillett (1980), Bradford et
al. (1983), and Bradford-Grieve (1994, 1999a,b). Their data are augmented
by information in the revisions of the Aetideidae (Markhaseva 1996) and
Euchaetidae (Park 1995). All these works incorporate other records of 19th- and
20th-century workers.
A number of calanoid families have not been recorded in the New Zealand
region. This may partly reflect lack of extensive sampling. For example, the poor
sampling of benthopelagic habitats at all depths is probably responsible for
the absence of the Diaixidae, Discoidae, Hyperbionychidae, Mesaiokeratidae,
Parkiidae, Pseudocyclopiidae, Ridgewayiidae, and Ryocalanidae, although it is
likely that the New Zealand fauna does include some species from a number
of these families. The apparent absence of isolated marine (anchialine) caves in
New Zealand probably explains the absence of the Boholinidae, Epacteriscidae,
and Fosshageniidae.
Species of Parapontellidae have been recorded only from the North Atlantic
PHYLUM ARTHROPODA
CRUSTACEA
Ocean and from deep waters of the Malay Archipelago, so this rare family may
not occur in the New Zealand region.
Other families are absent from the New Zealand fauna for paleogeographic
reasons. The Diaptomidae are known from fresh waters in most of the world
apart from New Zealand, most of Australia, and all of Antarctica (Bayly 1995).
Pseudodiaptomids are brackish to marine species, widespread in other parts of
the world but present in the Australasian region only in northern Australia.
The taxonomy of the freshwater planktonic calanoids is reasonably well
known (Chapman & Green 1987), although genetic studies using modern
techniques are required to assess whether there has been cryptic speciation in
any of the geographically widespread and disjunct species and in those shared
with Australia (cf. Boileau 1991). Ecological studies are still in their infancy, and
for all species much more needs to be known about autecology (e.g. growth
and reproduction, feeding rates, behaviour, life-history strategies, population
dynamics, etc.), and contributions to community and ecosystem dynamics (e.g.
competitive interactions, predation effects, production rates, contribution to
food chains, nutrient cycling, etc.).
Misophrioida
Members of this order have not been recorded from New Zealand. It is possible
that they might be found when marine benthopelagic habitats are more
extensively sampled.
Cyclopoida
This order now includes the Poecilostomatoida (Boxshall & Halsey 2004).
Cyclopoids have been relatively little studied in New Zealand – knowledge of
the marine, freshwater, and brackish non-parasitic Cyclopoida is very scattered
and inadequate.
Early records of freshwater Cyclopoida were summarised by Hutton (1904)
and amplified by Chapman and Lewis (1976). The synonymies and taxonomic
arrangement given by Dussart and Defaye (1985) in their checklist of the world
free-living Cyclopoida were taken into account in compiling the New Zealand
list. In addition, the revision of the Paracyclops fimbriatus complex (Karaytug &
Boxshall 1998) and the records of Roper et al. (1983) were noted. The commoner
New Zealand taxa in ponds and lakes are known but both their generic and
species status need re-examination in view of the recent taxonomic revisions
of supposedly cosmopolitan genera (Morton 1985; Dussart & Defaye 1995).
The underground and cryptic fauna is unknown taxonomically apart from
Goniocyclops silvestris in forest litter (Harding 1958), and genera and species
described by Karanovic (2005), but other undescribed species are known. Entries
in the end-chapter checklist accompanied by a question mark are doubtful old
records that need further investigation.
Checklists entries of the free-living marine planktonic families Oithonidae,
Corycaeidae, and Sapphirinidae of the New Zealand region are based on the
unpublished records of Janet Bradford-Grieve; the identities of the species need
more detailed study. The species of Oncaeidae are known from the work of
Heron and Bradford-Grieve (1995).
Another group of families comprises mainly marine parasites or associates
of other animals. For example, Hemicyclops (a near relative has been discovered
in New Zealand but is undescribed) has a typical cyclopoid body form and lives
in loose associations with other marine organisms (e.g. polychaetes), sharing
their burrows. There has been some work on fish parasites in New Zealand but
the fauna is essentially unknown or undescribed – an extensive collection of
Sarcotaces spp., made by Jones in the 1980s and 1990s from around New Zealand,
remains in the Auckland Museum collection awaiting description.
The parasitic families Archinotodelphyidae, Chordeumiidae, Cucumaricolidae, Mantridae, Ozmanidae, and Thespesiopsyllidae and the marine benthic
Oncaea media (Cyclopoida).
From Heron & Bradford-Grieve 1995
127
NEW ZEALAND INVENTORY OF BIODIVERSITY
family Cyclopinidae are not known from New Zealand. The freshwater parasitic
family Lernaeidae is represented by only Lernaea cyprinacea, which was introduced with ornamental fish (Boustead 1982). The commensal Ascidicolidae and
Notodelphyidae, living in association with tunicates, are known from only two
collections (Schellenberg 1922a, b; Jones 1974, 1979). It is certain that many more
cyclopoid associates of marine invertebrates remain to be found and described.
Data on the occurrence of commensal and parasitic forms have been collated
here using the works of Thomson, Hewitt, Jones, Pilgrim, and Ho as described
above. In general, we can say that the symbiotic copepods of New Zealand
are very poorly known, particularly those occurring in association with marine
invertebrates. Certainly, those parasitic on marine fishes are better known than
those parasitic or commensal on/in other hosts, but we still cannot say that fish
copepods are well known in New Zealand. There is currently nobody working
on symbiotic copepods in New Zealand.
Gelyelloida
The two known species of this order are found in subterranean waters of France
and the order is unlikely to be found in New Zealand.
Mormonilloida
This order contains only two species that are usually found at mesopelagic depths.
Mormonilla phasma has been recorded off the east coast of northern New Zealand.
Mormonilla phasma (Mormonilloida).
After Giesbrecht 1893
Artotrogus gordoni (Siphonostomatoida).
From Kim 2009
128
Harpacticoida
Early contributions to knowledge of New Zealand’s fauna were made by
Thomson (1878a,b, 1882), Brady (1899), Sars (1905), Brehm (1928, 1929), Farran
(1929), Lang (1934), and Harding (1958). More recent additions to the fauna
have been made by Barclay (1969), Hicks (1971, 1976, 1986, 1988a,c), Lewis
(1972a,b; 1984), Wells et al. (1982), Hicks and Webber (1983), and a number of
other authors. Hicks has also contributed a body of ecological and biological
information on the phytal harpacticoid fauna. Included herein are unpublished
records of freshwater species from Dr Maureen Lewis, and marine species
from Drs John Wells and Geoff Hicks. When the presently undescribed species
in existing collections are worked up, our knowledge of the sediment-dwelling
harpacticoids of seashores will be reasonably good, but much work still needs
to be done on the marine phytal fauna (mainly nationwide collecting to
establish distributional patterns). As is common worldwide, there is very little
knowledge of the sediment or phytal faunas of the sublittoral and deeper.
Lack of extensive exploration may be responsible for the absence of some
families. It is highly probable that Argestidae, Cerviniinae (Aegisthidae),
Cletopsyllidae, and Nannopodidae will be found in shelf and deep-water
sediments and Longipediidae and Metidae associated with seashore plants
and algae. On the other hand, the absence of the Parastenocarididae may be for
geological reasons.
Only a fraction of New Zealand’s freshwater and damp terrestrial locations
has been surveyed. It is to be expected that the number of species in the fauna
will be at least tripled, and New Zealand’s geological history makes it likely that
a number of intriguing questions of zoogeography and phylogeny will arise as
a result. The harpacticoid fauna of New Zealand’s ground waters is completely
unknown, yet cave systems exist that are comparable to the species-rich karst
formations of Europe.
Of particular note is the paucity of information on the fauna of the far
offshore islands from the Kermadecs to the Chathams and subantarctic islands.
Siphonostomatoida
All Siphonostomatoida are parasites or associates of other animals and the order
is mainly marine. Most work has been done in New Zealand on the parasites
PHYLUM ARTHROPODA
of fish, but this work is nowhere near complete. Almost nothing is known
of the vast proportion of this order likely to live in association with marine
invertebrates. We estimate that there are many species waiting to be discovered
in the New Zealand siphonostomatoid fauna. There is currently nobody working
on symbiotic copepods in New Zealand.
Commensal and parasitic forms have been collated here using the works of
Thomson, whose major work was published in 1890 and whose collection is still
housed in the Otago Museum (Thomson 1890). Gordon Hewitt also published
extensively in the 1960s (Hewitt 1963, 1967, 1968, 1969) and, later, one of his
students, Brian Jones, continued (1979, 1981, 1985, 1988b, 1991); his collection,
including many undescribed species, is now in the Auckland Museum. A large
collection was amassed at Kaikoura by students of the University of Canterbury
under Bob Pilgrim (Pilgrim 1985) and some of that material was worked up by
Ju-Shey Ho (Ho 1975, 1991; Ho & Dojiri 1987). The compilation given in the
end-chapter crustacean species is based on the parasite list of Hewitt and Hine
(1972), Pilgrim (1985), and the unpublished collection records of Jones.
Monstrilloida
All Monstrilloida have internal parasitic naupliar and early postnaupliar stages
and free-swimming, non-feeding adults. The known hosts are polychaete worms
and prosobranch molluscs. Members of this order have been noted in the New
Zealand fauna although there are no published records and descriptions.
CRUSTACEA
An unidentified species of Monstrilloida.
Geoff Read
Conclusions
There are few copepod taxonomists in New Zealand and none is able to
work full-time on the subject. The greatest gaps in our knowledge copepod
diversity are in the orders Cyclopoida, Harpacticoida, Siphonostomatoida, and
Poecilostomatoida, especially concerning copepods as symbionts and parasites.
These can be filled only by sampling little-studied environments, namely phytal,
freshwater, deep-water, damp-terrestrial groundwater, and offshore islands.
Sampling of benthopelagic and deep-sea habitats will yield records of hitherto
undiscovered families and orders.
Because copepods are ecologically and economically so important, there is
tremendous scope to understand the roles they play in the different ecosystems
that they occupy, and to understand their impact on the other organisms with
which they live in association, some of which are directly exploited by humans.
Class Ostracoda: Seed shrimps, mussel shrimps
Ostracods are tiny bivalved crustaceans that are widely distributed in the oceans, in
fresh waters, and, rarely, in terrestrial situations. Food-mediated seasonal blooms
in some freshwater habitats can result temporarily in vast numbers. Their shape
confers on them the common name seed shrimps or mussel shrimps. Species subclass Podocopa range from 0.2 to 1.5 millimetres in length, while modocopids are
often much longer, reaching an extreme of 30mm in Gigantocypris. Their shells,
strengthened by deposition of calcium carbonate amongst the layers of cuticle, also
fossilise well; in fact, ostracods are the most abundant arthropods in the fossil record,
with a body plan that has been conserved at least since the Silurian. The shells
can be brightly coloured and highly sculptured, making them attractive creatures
to study, especially with a scanning electron microscope. They have an indistinctly
segmented body like most arthropods, with paired appendages that are adapted for
a variety of functions. Their identification is normally a specialist occupation.
They are very useful organisms, as knowledge of their taxonomy and
distribution can be applied to studies of ecology and to environmental monitoring
in relation to water quality, water depth, salinity levels, and temperature, as well
as in stratigraphy. The number of specialists studying this group of animals is
declining even though there is great potential for their usefulness. There are
Hemicytherura pentagona (Pleistocene).
Stephen Eagar
129
NEW ZEALAND INVENTORY OF BIODIVERSITY
approximately 22,000 living and fossil species in the Catalog of Ostracoda
published by the American Museum of Natural History and estimates of likely
global diversity suggest more than 62,000 species in total. Of the described living
species, 7000 belong to subclass Podocopa and 600 to subclass Myodocopa
(Cohen 1998). There are many more species yet to be found in New Zealand,
both living and fossil, in all environments.
Ostracods live in most aquatic environments and even, in the case of one
New Zealand species – the bright yellow Scottia audax – in the damp leaf litter
of the forest (Chapman 1961). Freshwater species live for between one season
(as ponds dry) and three years. Marine species similarly live for one season
to two years. Many marine planktonic ostracods constitute food for fish and
species of one family (Entocytheridae, represented in New Zealand by a single
species) are commensal on fish and other arthropods. Some myodocopids are
bioluminescent but none have yet been found in New Zealand.
The first description of an ostracod, by Carl Linnaeus (1746), was very
generalised. A figure was published in 1753, but the ‘father’ of the study of
ostracods is regarded as O. F. Müller who, in a 1785 monograph on Entomostraca
from Denmark and Norway, produced good descriptions and figures of
freshwater ostracods.
Cymbicopa hanseni.
From Brady 1898
130
History of study in New Zealand
Currently, the New Zealand living ostracod fauna stands at 442 species (including
86 undetermined), mostly marine but also comprising 37 freshwater and one
terrestrial species. This tally is the product of many zoological studies since 1843;
actual descriptive taxonomy has proceeded in pulses. The first species to be studied,
by William Baird, was a relatively large (1.94 millimetres body length) freshwater
species (Candonocypris novaezelandiae), often found in ponds and drinking
troughs for farm animals (Baird in White & Doubleday 1843). It was collected
by naturalist-explorer Ernst Dieffenbach. Baird (1850) was also responsible for
describing the large (6.5 millimetres) marine species Leuroleberis zealandica sent
to him by Rev. Richard Taylor of Waimate, one of the early settlers. George M.
Thomson, teacher, Member of Parliament, and an amateur naturalist, produced
the first locally published paper on ostracods from the Dunedin district in 1879.
The first global oceanographic voyage of HMS Challenger (1873–1876) brought
the ship into New Zealand waters and into Wellington Harbour for sampling.
The results were published by Brady (1880). With the general establishment of
the New Zealand colony, there was by the end of the 19th century an exchange of
information between naturalists in New Zealand and Europe who were keen to
document the fauna. So material was sent away for identification. Norwegian G.
O. Sars (1894) published on freshwater species contained in dried mud and Brady
(1898), living in Newcastle, England, received some marine specimens from New
Zealand. Owing to the paucity of New Zealand ostracod taxonomists, this practice
continued well into the 20th century with Brehm (1929) in Austria, Kornicker
(1975) in the USA, and Hartmann (1982) in Germany providing identifications.
One consequence is that many of the type specimens of New Zealand species
reside in overseas institutions.
The freshwater ostracod fauna was reviewed by Chapman (1963) and
Chapman and Lewis (1976), and Scarsbrook et al. (2003) briefly summarised the
ecology of New Zealand groundwaters in which ostracods occur but which are
poorly known.
The podocopids and platycopids from the shallow intertidal to outer shelf
have been the most intensively studied ostracods because they are also the most
accessible (e.g. Morley & Hayward 2007). As mentioned above, ostracods are
useful for environmental monitoring. They are sensitive to small changes in
salinity and water quality and respond negatively to pollution. One study of a
New Zealand waste outfall has shown the effects of sewage on a coastal ostracod
fauna (Eagar 1999).
PHYLUM ARTHROPODA
CRUSTACEA
The planktonic myodocopids, which require specialist zoological knowledge,
has been treated in monographs by Poulsen (1962, 1965) and Kornicker (1975,
1979) and in research studies by Deevey (1982). The first halocyprids were not
recorded until Barney (1929). This group, together with the deep-sea podocopids,
had received the least attention, but the recent study by Jellinek and Swanson
(2003) has significantly increased knowledge of the latter.
Fossil species have followed a similar pattern of study. The earliest paper was
by Jones (1860) on some tertiary species from Orakei. A bulletin by Chapman
(1926) was issued by the New Zealand Geological Survey for Cretaceous and
Tertiary species, but he used European names. His records are therefore not
explicitly included in the following checklist, but the species are probably still
represented there as synonyms of other workers’ identifications. Benson (1956)
recorded the occurrence of ostracods in late Middle Cambrian rocks from New
Zealand, based on F. H. T. Rhodes’s identification of their remains in a limestone.
The preservation did not permit accurate identification. Simes (1977) recorded
a phosphatic or phosphatised specimen from the limestone of the Upper
Cambrian Anatoki Formation, and silicified ostracods were recorded by Marden
et al. (1987) from the Triassic (Norian age). No other records whatsoever are
available for any specimens from the Ordovician to the Jurassic.
Good fossil faunas are now known from sediments of Cretaceous age at
several localities and these have been published recently (Dingle 2009). There
have been a large number of papers on the systematics and paleoecology of
New Zealand region Tertiary Ostracoda from the mid-1950s onwards (Swanson,
1969; Ayress 1990, 1991, 1993a,b,c, 1995, 1996; Ayress & Warne 1993; Ayress et
al. 1994, 1995, 1997, 1999; Ayress & Drapala 1996). These faunas are rich, easily
obtained, and interesting as they can be tied into other paleontological work.
Most of the ostracod species in the end-chapter fossil checklist are therefore
Tertiary species. The first publications to illustrate New Zealand ostracods using
scanning electron microscopy came later (Swanson 1979a,b, 1980). The endchapter checklist following builds on the one published by Eagar (1971).
Features of the New Zealand ostracod fauna
Many Cenozoic marine species are endemic, long-ranging, and even still
living. Presuming that they have not evolved a tolerance to changed ecological
conditions, it can be assumed that the paleoeviromental conditions in which
they lived were the same as now. Of particular interest are species of the
endemic living-fossil genera Manawa and Puncia (Punciidae). Similar in shape
and ornamentation to some Paleozoic genera, they are found living in shallow
water off the north and east coasts of New Zealand. They provide insight into
the soft-part anatomy of a group of ostracods (order Palaeocopida) that has
otherwise been extinct for a long time (Hornibrook 1963; Swanson 1990; Horne
et al. 2005).
Freshwater species are rare as fossils. Many species are swamp- or ponddwellers and are not found on lake margins; inasmuch as ostracod shells
are very soluble in the acid conditions of swamp deposits, their chances of
preservation there are small. Further, most of New Zealand was submerged by
the Late Oligocene and there were relatively few lakes, along with limited means
of dispersal, available in the geological past (Hornibrook 1955; Eagar 1995a).
Once colonisation from Europe was established, trout, salmon, and carp were
introduced from Europe via Australia and it is likely that ostracod eggs travelled
as hitchhikers to New Zealand on the damp media used to transport the fish
(Eagar 1994). There is one non-marine saline species – Diacypris thomsoni (see
Bayly & Williams 1973) – from Sutton, Otago, in salinity conditions of up to
15 parts per thousand. Guise (2001) discovered in the Avon-Heathcote Estuary,
Christchurch, a new endemic genus of brackish-water ostracod (Swansonella)
that tolerates higher salinities.
Lateral view of valve of Puncia sp. (upper)
and ventral view of Manawa staceyi,
both from Cavalli Islands.
Kerry Swanson
131
NEW ZEALAND INVENTORY OF BIODIVERSITY
There are now more opportunities for introducing ostracods into New
Zealand. Resting eggs that can withstand desiccation may even be transported
by aircraft on footwear and camping gear. In addition to European freshwater
species, several other species have an Australasian distribution. One marine
species discovered close to shipping ports in the North and South Islands may
have been brought in ballast water (Eagar 1999).
Few studies have been made of the anatomy of New Zealand ostracods.
These were mostly on myodocopids (Poulsen 1962, 1965, Kornicker 1975,
1979) and to a lesser extent to the freshwater species (Podocopida: Cyprididae)
(Chapman 1963; Eagar 1995b; Rosetti et al. 1998), with a few ventures into the
marine podocopids (e.g. Brady 1902; Swanson & Ayress 1999).
Class Malacostraca
This class contains more than half of all known species of crustaceans, including
the aristocrats – the giant spider crabs of Japan with their 3-metre leg span
(vying with fossil eurypterids as the largest of all arthropods) and the New
Zealand packhorse rock-lobster (Sagmariasus verreauxi) at 20 kilograms – and
krill, one of the most ecologically critical malacostracans in marine food webs,
slaters, and tiny sand-hoppers. Malacostracans are very unevenly divided into
three subclasses – Phyllocarida, Hoplocarida, and Eumalacostraca.
Subclass Phyllocarida: Phyllocarids
Order Leptostraca
Levinebalia fortunata.
From Wakabara 1976
132
The Leptostraca is the sole living order of the Phyllocarida, a group of Crustacea
with a long geological history (Rolfe 1969), possibly extending back as far as the
Cambrian, some 600 million years ago (Briggs 1992). Despite new conclusions
from DNA analyses as to their place in crustacean evolution (Spears & Abele
1999), the Leptostraca may still be regarded as ‘living fossils’ indicative of the
times and conditions in which the so-called primitive arthropods lived (Hessler
& Schram 1984; Dawson 2003b). They are known from the New Zealand
Ordovician (Chapman 1934), and the presence of several living species of
Leptostraca in the region is of considerable interest. Using the small-subunit 18S
ribosomal-DNA gene of 10 representative foliaceous-limbed Crustacea, Spears
and Abele (1999) concluded that the Phyllocarida are true malacostracans,
which diverged fairly early from the main lineage. This result is consistent with
the pioneer work of Claus (1888) and Calman (1909) and with Manton’s (1934)
study of embryology, and also corroborates the views of Dahl (1987, 1991) of the
Leptostraca as an early offshoot.
The late British zoologist Sir Alistair Hardy (1956) vividly recalled the
excitement of his first encounter with one of the little crustaceans, Nebaliopsis
typica, found in great depths but rarely collected, and then usually dead and
very damaged. It had only ever been seen alive on one occasion – on the
Swedish Antarctic Expedition in 1904 – until a second specimen was collected
from the Discovery II fifty years later. The Leptostraca, wherever they have
been found subsequently, have continued to excite and interest zoologists and
paleontologists alike.
A paleontological summary of the Phyllocarida was made by Rolfe (1969).
Monographs on the Leptostraca as a whole have been made by Claus (1888) and
Cannon (1960), and these still have their usefulness, but a new and compact text
has been produced (Dahl & Wägele 1996). More recently, the relationships of
the leptostracan genera were examined by Olesen (1999) and by Walker-Smith
and Poore (2001), who revised the families and genera. The latter authors also
provided a complete listing of all species of Leptostraca together with keys to
the families and genera. Some 42 species of living Leptostraca are recognised
PHYLUM ARTHROPODA
at present, divided into three families – Nebaliopsidae (genera Nebaliopsis,
Pseudonebaliopsis ), Paranebaliidae (named only in 2001, containing Paranebalia,
Levinebalia, and Saronebalia), and Nebaliidae (with five other genera). Many
species of Nebalia and Paranebalia remain undescribed as yet (Dahl & Wägele
1996).
Leptostracans are small, usually 4–12 millimetres in length although one
species, Nebaliopsis typica, can exceed 35 millimetres. They are characterised by
the possession of a relatively large, bivalved carapace, hinged on the midline and
held together by an adductor muscle. The carapace loosely covers the abdomen
and part of the thorax, and is attached by a hinged rostral plate covering the head
and closing the anterior gap of the carapace itself. Long anteriorly projecting
antennae are used for swimming, the antennal flagellum in males being as long
as the body. There are eight pairs of foliaceous, leaf-like thoracic limbs that also
provide a feeding mechanism and may be modified in the female in the form of a
fan of plumose setae forming a basket-like chamber for brooding eggs between
the ventral regions of the valves of the carapace. The first four pairs of pleopods
are well developed and biramous whereas the 5th and 6th pairs are small and
uniramous. The abdomen ends in two characteristic long and articulated tail
spines or furci. In contrast with all the six abdominal segments possessed by
all other Malacostraca, the Leptostraca have a 7th segment and this lacks any
appendages. The telson may be considered an 8th segment.
Relatively little is known of the life-history, growth rates, or physiology of
most leptostracans. Useful observations have been made by Cannon (1927),
Rowett (1943, 1946), Martin et al. (1996), Vetter (1996a), and Wägele (1983).
Manton (1934) worked on the embryology of Nebalia bipes, helping to elucidate
phylogenetic relationships of the Phyllocarida (Dahl 1987; Spears & Abele 1999).
Linder (1943) described some larval stages, which could be useful for recognition
in sorting plankton samples. Leptostracans play a significant role in benthic
production (Rainer & Unsworth 1991; Vetter 1996a,b; MacLeod et al. 2007).
The unusual marine rotifer Seison is often found epizoic on leptostracans. None
has yet been discovered in New Zealand but it would be worth checking local
Nebalia to ascertain their presence or absence.
Leptostracans are widely distributed as a group. Individual species may be
limited or widespread in depth range and geographically, but taxonomic caution
needs to be observed in the case of the purportedly wide-ranging species. Dahl’s
(1990) analysis of the Nebalia longicornis complex showed that it comprised at
least 10 different species. Walker-Smith (1998) reviewed the genus Nebaliella,
describing the first known Australian species. In her unpublished Honours
thesis, she recognised six new species and a new genus of Leptostraca from
Australia (Walker-Smith pers. comm. 2000).
Present-day leptostracans live in a variety of habitats, including under
intertidal stones, with decaying seaweed or dead shell, in crab pots, on mangrove
shores and coral reefs, and in subtidal sandy plains or muddy sand. A non-New
Zealand species, Speonebalia cannoni, is the only leptostracan to be recorded from
a groundwater habitat. Nebalia hessleri lives in enriched sediments and detrital
mats with low oxygen levels in submarine canyons off southern California.
Here they form the highest density ever reported for a macrofaunal assemblage,
namely 1.5 million per square metre. In northwestern Spain, Moreira et al.
(2009) reported six species of leptostacans in subtidal sediments, the largest
number of species recorded in a single area. Dahlella caldariensis occurs among
mussels and vestimentiferan worm tubes, swimming above clumps of animals
at hydrothermal vents.
CRUSTACEA
Nebalia longicornis.
From Thomson 1879
The New Zealand leptostracan fauna
The New Zealand fauna currently consists of five species in four of the 10 known
genera. Unfortunately, little is known of the true numbers of taxa represented in
133
NEW ZEALAND INVENTORY OF BIODIVERSITY
any one geographic area, but the indications are that New Zealand could well be
shown to have a higher diversity.
The first to be recorded and named in New Zealand was Nebalia longicornis,
based on a single mature male collected in Otago Harbour (Thomson 1879a).
It was subsequently described in more detail, based on records from 8–10
metres depth in Dunedin Harbour and 20 metres at Stewart Island (Thomson
1881). This later paper by Thomson (with its slightly different figure) appears
to have been overlooked by all subsequent authors. Nebalia longicornis was
inadequately described and illustrated according to Dahl (1990), and great
taxonomic confusion subsequently resulted from attempts to apply this name
to later records of Nebalia from other parts of the world. Since Thomson’s type
specimen could no longer be found, Dahl redescribed the species based on a
female collected from Otago Harbour in 1965, thereby fixing Nebalia longicornis
Thomson, 1879a as a member of the New Zealand fauna. Thomson (1913) noted
his Nebalia longicornis as found in Otago Harbour and frequently taken outside
the Otago Heads in trawl-nets.
Thiele (1904) reported a specimen of what he considered to be Nebalia
longicornis from Akaroa Harbour. Dahl (1990) examined this specimen and
found it to be a species of Nebalia (then in his genus Sarsinebalia) but in too
damaged a condition to be able to describe further. Thiele had also recorded
juvenile Nebaliella antarctica from Akaroa Harbour but apparently this specimen
has not been re-examined.
In 1907, W. Benham collected a juvenile Nebalia from Musgrave Harbour
on the Auckland Islands that Chilton (1909) attributed to N. longicornis as then
understood. Another specimen was taken at Port Ross, Auckland Island, in 1914
during the Mortensen Expedition (Stephensen 1927). Calman (1917) reported
two immature specimens of Leptostraca collected in 1911 at Terra Nova Stations
130 and 135 off Three Kings Islands and in Spirits Bay [given incorrectly by Dahl
(1990) as Stns 10 and 15]. Dahl (1990) has since examined these specimens,
concluding that one is a Nebalia and the other a Sarsinebalia.
Morton and Miller (1968) described a Nebalia as a member of the protected
sandy-beach fauna, one of the small filter-feeding Crustacea that live in the fine
sands of the lower beach. They also illustrated it as the prey of the small shallowwater cephalopod Sepioloidea pacifica.
The only other work on New Zealand leptostracans has been the description
of Levinebalia fortunata (Wakabara 1976, as Paranebalia) based on 16 females
collected by trawl nets at 420–660 metres depth in canyons off Otago Peninsula,
representing a marked extension to the known bathymetric range of the genus.
Apart from Prof. John Jillett at Otago (see Dahl 1990) no-one has conscientiously
searched New Zealand habitats for leptostracans. It is likely that deliberately
intensive collecting will reveal not only great extensions of the range of the
already listed forms but undescribed species as well. Morton (2004) suggested
searching for leptostracans in black anaerobic sediments with decaying algae
and carrion-baited traps may also be useful (Lee & Morton 2005), especially for
assessing population densities.
Chapman (1934) described several species from Ordovician rocks in
Fiordland, based on numerous specimens. They have never been studied since
and are listed in the end-chapter checklist of fossil New Zealand Crustacea
under the generic names recommended by Rolfe (1969).
Subclass Hoplocarida
Mantis shrimp Heterosquilla tricarinata.
Shane Ahyong
134
Order Stomatopoda: Mantis shrimps
Mantis shrimps are among the most aggressive and behaviourally complex
crustaceans. All are active predators and mark one of the very few radiations
of obligate carnivores within the Crustacea. The general morphology of mantis
PHYLUM ARTHROPODA
shrimps has been described by Holthuis and Manning (1969), and characteristic
features are the triflagellate antennules, well-developed stalked eyes, and the
greatly enlarged, raptorial second maxillipeds. The name mantis shrimp stems
from these large and powerful raptorial claws. Prey is captured by ‘spearing’ or
‘smashing’, depending on whether the dactyl of the raptorial claw is extended
or kept folded during the strike. (Think of the dactyl as a finger, opposing the
thicker ‘thumb’ of the claw.) Hence the two modes of prey-capture define the
‘smashers’ and the ‘spearers’ among mantis shrimps (Caldwell & Dingle 1976).
The strike of the raptorial claw is among the fastest known of animal movements,
being completed in 3–5 milliseconds, and the strike of large species of ‘smashers’
may break aquarium glass.
Vision in mantis shrimps is strongly developed. In most species, the cornea
is divided into two halves by a midband of ommatidia, enabling binocular vision
with each eye. Additionally, the midband ommatidia in many families enable
colour vision and detection of polarised light (Marshall 1988).
Most stomatopods live in temperate or tropical shallow marine habitats, but
several species also range into subantarctic waters, and a few tropical species may
occur in brackish water. Seven superfamilies are recognised: Bathysquilloidea,
Erythrosquilloidea, Eurysquilloidea, Gonodactyloidea, Parasquilloidea, Lysiosquilloidea, and Squilloidea. Most members of the Gonodactyloidea occur on
coral reefs where they shelter in or under boulders and coral. The bathysquilloids
are known only from deep outer-shelf waters. Members of other superfamilies
generally burrow in flat sandy and muddy harbour bottoms and sea-floors.
The Stomatopoda comprises the only living order of Hoplocarida, two other
orders (Aeschronectida and Palaeostomatopoda) being known only as fossils.
Compared with other major crustacean groups such as the Decapoda, the fossil
record of the Hoplocarida is relatively poor but it appears that the hoplocarids
originated in the Devonian and the Stomatopoda proper first appeared during
the Carboniferous. Recognisably modern stomatopods, with well-developed
raptorial claws, did not appear until the Mesozoic (Holthuis & Manning 1969;
Hof 1998; Hof & Schram 1998).
Over the past three decades, the taxonomy of the Stomatopoda has been
extensively revised, largely through the work of the late R. B. Manning, who
recognised five living superfamilies (Manning 1995). Ahyong and Harling
(2000) provided the most recent phylogenetic study. At present, more than
450 species in more than 100 genera, 19 families, and 7 superfamilies are
recognised.
The stomatopods of the Atlantic have been monographed and are well
known (Manning 1969, 1977), while those of the eastern Pacific were treated
relatively comprehensively by Schmitt (1940) and Hendrickx and SalgadoBarragán (1991). Stomatopod diversity in the Indo-West Pacific region, however,
is more poorly known. The most important major works for this region are those
of Kemp (1913) on the Indian fauna, Manning (1995) on the Vietnamese fauna,
and Ahyong (2001) on the Australian fauna. The Indo-West Pacific fauna has
been extensively studied in the past decade (e.g. Ahyong 2002a,b,c; Ahyong &
Naiyanetr 2002; Ahyong et al. 2008).
CRUSTACEA
Pterygosquilla schizodontia.
Shane Ahyong
The New Zealand fauna
New Zealand’s mantis shrimps are known from only a few studies, the most
important of which are those of Miers (1876), Chilton (1891, 1911a) and
Manning (1966). Manning (1966) recognised three species from New Zealand
and its offshore islands: Pterygosquilla schizodontia, Heterosquilla tricarinata, and
Acaenosquilla brazieri (as Heterosquilla brazieri). He also remarked that Squilla
tridentata Thomson, 1882, synonymised with H. tricarinata by Chilton (1891),
was probably a distinct species. Ahyong (2001) recognised Thomson’s species
as distinct under the combination Heterosquilla tridentata. Other additions to the
135
NEW ZEALAND INVENTORY OF BIODIVERSITY
New Zealand stomatopod fauna are Hemisquilla australiensis (Stephenson 1967),
Odontodactylus brevirostris (Manning 1991), and the striking 30-centimetre-long,
scarlet deep-sea species Bathysquilla microps (O’Shea et al. 2000). Therefore,
seven species are presently recorded from New Zealand.
The commonest species are Heterosquilla tricarinata (known around both
main islands and Chatham, Stewart, Campbell, and Auckland Islands, generally
in intertidal sand or mudflat burrows) and Pterygosquilla schizodontia (central New
Zealand to the Auckland Islands, burrowing in subtidal sand and mud). Their
biology has received little scientific study. Larval development of Pterygosquilla
schizodontia was studied by Pyne (1972). Several studies have been conducted on
H. tricarinata including those of Fussell (1979), Greenwood and Williams (1984),
and Williams et al. (1985).
The New Zealand stomatopod fauna is relatively small, and this is consistent
with the primarily tropical distribution of most species. Neverthless, low diversity
may also reflect low collecting effort. Study of collections from northern island
groups in New Zealand territorial waters should reveal numerous additional
faunal records. The Japanese mantis shrimp Oratosquilla oratoria has become
established in some North Island estuaries and is the first exotic species of
Stomatopoda to be detected in New Zealand waters. New species and numerous
additional distribution records will be reported in a forthcoming revision of the
New Zealand Stomatopoda by Shane Ahyong.
Subclass Eumalacostraca
Superorder Syncarida
Orders Anaspidacea, Bathynellacea
The Syncarida constitutes a group of tiny crustaceans that may be regarded as
living fossils, with a geological history extending as far back as the Carboniferous
(Dover 1953; Drummond 1959; Brooks 1969; Schram & Hessler 1984; Uhl 1999,
2002; Jarman & Elliott 2000; Dawson 2003a). They are little known to most
biologists, the exception being the large-sized Anaspides, found in Tasmania,
which has attracted much interest and attention largely because of its accessibility
in open waters rather than the subterranean habitat in which most syncarids live.
The Syncarida were first made known to science by the report of a fossil
species, Uronectes fimbriatus, in Europe. Their relationships and place in the
crustacean hierarchy remained a matter of contention until Packard (1885, 1886)
gave them separate status as the Syncarida. Much later, Brooks (1962, 1969) finally
settled the status of the fossil as one of three orders constituting the superorder
Syncarida, and Schminke (1975) related them to the living orders. Schram (1984)
subsequently reviewed and revised the fossil species, which range in time from
the Early Carboniferous (Uhl 2002) to the Early Permian in Europe and North
America, the Late Permian of Brazil, and the Triassic of Australia, corresponding
to the former landmass of Laurentia prior to the formation of Pangaea.
New Zealander George Malcolm Thomson, a noted amateur scientist,
teacher, and politician, is generally credited with the discovery and description
of the first living syncarid – Anaspides tasmaniae, which he discovered when
visiting Tasmania in January 1892. He was of the opinion that his discovery was
a schizopod shrimp (Thomson 1894). However, Calman (1896) said this new
crustacean was no schizopod and supplemented Thomson’s description in some
detail, comparing Anaspides with fossils from Illinois and Germany that Packard
(1885) had already placed in his new group, Syncarida. Calman concluded that
Anaspides was, in fact, a living representative of primitive malacostracans that
had flourished widely in Paleozoic times
Ironically, however, living syncarids had in fact been discovered some years
previously when Vejdovský (1882, 1889) published a description of the tiny
Bathynella that he had found two years earlier in a well in Prague. Calman (1899)
136
PHYLUM ARTHROPODA
subsequently recognised Bathynella as a syncarid, but little more was known
until 1913 when Chappuis (1915) found more specimens in a well near Basle.
He placed them in a new taxon, Bathynellacea. Syncarids were soon found to
occur in many places throughout Europe, in wells, springs, or streams in caves
(Chappuis 1939) as well as in Australia, New Zealand, Japan, North and South
America, and elsewhere.
Although Thomson turned out not to be the first discoverer of a living
syncarid, the finding of such an ancient form of crustacean living in Tasmania did
excite many subsequent workers (up to the present day), resulting in a substantial
number of publications on aspects of their morphology, development, ecology,
and relationships – and even a poem in the style of Longfellow dedicated to
Anaspides (Mesibov 2000). In essence, there have been two approaches to the
study of the Syncarida, one concentrating on the relatively tiny subterranean
and interstitial forms (basically the order Bathynellacea), and the larger, openwater taxa of Australia (order Anaspidacea, which also includes the subterranean
Stygocarididae). General accounts of the Syncarida can be found in Siewing
(1959), Noodt (1964), McLaughlin (1980), Schminke (1982), Schram (1986), and
Coineau (1996, 1998).
Within the Eumalacostraca, the Syncarida are distinguished by the absence
of a carapace, an elongate body form (more or less cylindrical in the subterranean
forms), with a thorax consisting of seven or eight segments, the first segment
being fused to the head in some groups. The abdomen consists of six segments
and a telson, or five segments followed by a pleotelson formed from the fusion
of the 6th segment with the telson.
The order Anaspidacea contains four families: Anaspididae, Kooningidae,
Psammaspididae, and Stygocarididae. Only the last of these has been found
in New Zealand. They include the largest of the syncarids, with a body length
ranging from about 1 to 50 millimetres. The Bathynellacea contains two families,
the Bathynellidae and the Parabathynellidae, which are both represented in the
New Zealand fauna as it is presently known. They are very much smaller in size
than the anaspidaceans, ranging from about 0.4 to 3.5 millimetres.
The body form of syncarids is reflected in the habitats in which they are
found: the tiny forms, with slender, cylindrical bodies, devoid of pigment and
eyes, are found in caves and underground waters, whereas the much larger
forms, such as Anaspides, found in surface waters are shrimp-like.
Living syncarids comprise more than 200 species worldwide (Camacho
& Valdecasas 2008), although fresh explorations and more refined collecting
techniques are already increasing this number. There are many species of syncarids
collected from eastern Australian caves and karst areas awaiting identification and
description (Thurgate et al. 2001) and such may be the case for New Zealand.
Syncarids have the reputation of being rare animals, although the pioneer
investigations by Chappuis (1943) on Bathynella in Hungary showed that
numerically rich collections could be made at individual sites. Much of the alleged
rarity is a consequence of their small size (which is why early investigators in
New Zealand such as Chilton did not find them) and their largely subterranean
habits. Schminke (1986) has said that those who know how to sample their
habitats ‘today have lost the impression of dealing with rare animals.’ Syncarids
are globally widespread; Schminke (1986) listed all the species then known, with
their locations. New taxa continue to be described Camacho 2005a,b; Cho 2005;
Cho et al. 2005, 2006; Camacho et al. 2006; Cho & Schminke 2006).
While some Syncarida inhabit open- and surface-water habitats (Camacho
& Valdecasas 2008), it is acceptable to say that syncarids are characteristic of
subterranean habits throughout the world, whether groundwater (as revealed
by sampling wells, springs, and gravel river margins), or caves with streams and
sandbanks providing living space in the interstitial spaces between sediment
grains.
Syncarids have been recorded from springs in Australia (Knott & Lake
CRUSTACEA
Stygocaris townsendi.
From Scarsbrook et al. 2003
137
NEW ZEALAND INVENTORY OF BIODIVERSITY
Notobathynella longipes.
From Schminke 1978
138
(1980), and in New Zealand they occur in similar situations as well as from
groundwater in wells (Scarsbrook et al. 2003), just as did the first-discovered
European living syncarids. Many syncarids have been collected from caves,
although in New Zealand only Stygocaris townsendi has been described from
such a habitat (Morimoto 1977). Karst landscapes throughout the world provide
habitats for syncarids.
Information on the development, life-history, and habits of syncarids is still
quite limited. So far as the Anaspidacea are concerned, most of the developmental
studies have been done on Anaspides tasmaniae, by Hickman (1937), with other
aspects covered in other studies, for instance Dohle (2000). The biology of
bathynellaceans is less well known, but what is known has been summarised
by Coineau (1996). In feeding, Anaspides has a filtering mechanism, used in
conjunction with collecting particles by scraping detritus with its limbs. Smith
(1908) noted that Anaspides was an omnivorous feeder, eating dead insects as
well as each other, but mainly feeding on algal slime and submerged mosses
and liverworts. The habitat of Tasmanian anaspidaceans, notably Allanaspides
hickmani and A. helonomus, is under continuing threat (Driessen et al. 2006).
Compared to the amount of information regarding the general biology
and ecology of the anaspidacean syncarids, there is virtually nothing recorded
about the lifestyle and habits of the Bathynellacea. What is known has been
summarised by Coineau (1996), and Camacho (1992) has outlined the abiotic
characters of the subterranean environment in which most of bathynellaceans
live.
Camacho (2006) noted 256 species and subspecies of extant Syncarida, 95%
of which are subterranean in habitat. In addition to the two living orders is the
order Palaecaridacea, which is entirely fossil.
The order Anaspidacea comprises five families, of which three are confined
to Australia. These include: Anaspididae, with five genera – Allanaspides, Anaspides, Paranaspides, Anaspidites (Triassic, Australia), Koonaspides (Lower Cretaceous,
Australia); Koonungidae, with two genera – Koonunga, Micraspides; Psammaspididae, with two genera – Eucrenonaspides, Psammaspides; and Stygocarididae,
with four genera – Oncostygocaris (Chile), Parastygocaris (Argentina), Stygocarella
(New Zealand), and Stygocaris (Australia, New Zealand, Chile). The 21 living
species of Anaspidacea are confined to the Southern Hemisphere. Anaspides tasmaniae is of particular interest in the context of mitochondrial DNA studies, in
which it has been demonstrated that there may be at least three cryptic species
(Jarman & Elliott 2000).
The order Bathynellacea comprises two families, both distributed widely
throughout the world, totaling 66 genera an 219 species: Bathynellidae, with
more than 20 genera (including Bathynella, of which there are New Zealand
representatives) and more than 80 described species; and Parabathynellidae,
with about 32 genera and more than 90 species (also recorded from New Zealand in the genera Atopobathynella, Hexabathynella, and Notobathynella). As discussed by Camacho et al. (2002), there have been two contrasting views as to
the systematic position of the bathynellids as being either within the superorder
Syncarida or as a separate suborder Podophallocarida in infraclass Eonomostraca. These Spanish researchers’ molecular studies in Spain on a cave-dwelling
bathynellid, Iberobathynella (Espanobathynella) magna, have now provided a
nucleotide sequence that supports a basal position for the Bathynellacea with a
clear distinction from the Syncarida, placing them in the Podophallocarida but
retained in the Eumalacostraca.
Schminke (1986) postulated that the Syncarida originated in the marine
environment from whence they invaded freshwater by two independent lines,
living first in surface waters and then invading the groundwater habitat. He
developed the ‘zoea’ theory (Schminke 1981b) in which it was suggested that
the Syncarida originally passed through a series of larval stages and through
neoteny reached sexual maturity at a stage corresponding to the zoea larva of
PHYLUM ARTHROPODA
CRUSTACEA
the penaeid prawns (Decapoda). Schminke (1972) had previously demonstrated,
by a study of all the then-known species of Hexabathynella (but which did
not include the subsequently discovered H. aotearoae of New Zealand), all of
which were known to occur close to the sea, that syncarids did not invade the
freshwater interstitial habitat from sandy marine beaches. Presumably, some of
the more recently discovered occurrences of Hexabathynella aotearoae indicate
secondarily derived habitats. This species is closest evolutionarily to Australian
H. halophila (Camacho 2003).
Biogeographically, the breakup of the ancient supercontinent Gondwana has
been invoked to explain some of the distributions between northern and southern
hemispheres and within the austral landmasses (Schminke 1973, 1974, 1975, 1980,
1981a; Williams 1986). Subsequent information about the distribution and phylogeny of the various syncarid groups can be found in Coineau (1996), Camacho
and Coineau (1989), Camacho et al. (2000), and Guil and Comacho (2001).
The New Zealand fauna
In 1967 and 1968, visiting German scientist Kurt Schminke searched for
syncarids quite widely throughout New Zealand, taking almost 200 samples
from interstitial freshwaters at 11 different localities (Schminke & Noodt 1968;
Schminke 1973). Of these, 36 yielded syncarids in the families Bathynellidae,
Parabathynellidae, and Stygocarididae. In his unpublished thesis, Schminke
(1971) included two new forms of Bathynella, a species and its subspecies (as yet
not formally described), collected from the Tauherenikau River in the Wairarapa
and from the Orari River in South Canterbury. In his major work on the evolution,
taxonomy and biogeography of the world fauna of the Parabathynellidae,
Schminke (1973) listed his collecting locations in New Zealand with descriptions
and distribution maps of four new species from New Zealand: Atopobathynella
compagana, Hexabathynella aotearoae, Notobathynella chiltoni, and N. hineoneae.
Schminke (1978) subsequently reported on a collection, made by by G. Kuschel
of the former DSIR Entomology Division, which included a female bathynellid
from a bore in Nelson, and two females of Notobathynella. He also noted two
more specimens of Atopobathynella compagana and described Notobathynella
longipes from wells at Motueka. In the Anaspidacea, Schminke (1973) mentioned
at least three unidentified New Zealand species of Stygocarididae in one new
genus, later describing Stygocarella pleotelson (Schminke 1980) and noting 16
localities from which other unidentified specimens had been collected. During
a brief trip to New Zealand in 1975, Morimoto (1977) collected syncarids at four
South Island locations, finding three species of Stygocaris, of which S. townsendi
was described as new. More recently, in a NIWA study of the New Zealand
groundwater fauna (Scarsbrook et al. 2000), syncarids appeared to be widespread
in interstitial habitats in alluvial groundwaters in Hawkes Bay and Canterbury,
both within the margins of gravel riverbeds and in the deeper (10–20 metres)
ground water.
Thus, the New Zealand syncarid fauna, as presently known from limited
sampling, consists of at least four species of Anaspidacea – Stygocaris, and one
or possibly more species of Stygocarella. The Bathynellacea is represented by
what appear to be quite abundant and widespread species of Bathynellidae
(Bathynella), none formally described, and three genera of the Parabathynellidae
– Atopobathynella and Hexabathynella (each with one described species), and
Notobathynella (at least four species, three of them named). It is highly likely
that the New Zealand syncarid fauna will be found to be much more extensive,
if only in terms of the distribution of the already described species, all of which
are endemic.
Stygocarella pleotelson.
From Schminke 1980
Gaps in knowledge of Syncarida
Not only is taxonomic knowledge of the New Zealand Syncarida incomplete;
even less is known about their ecology and special adaptations to the several
139
NEW ZEALAND INVENTORY OF BIODIVERSITY
kinds of habitats in which they occur. It is apparent that a geographically widely
distributed syncarid fauna exists in New Zealand’s ground waters. The brief
venture into cave collecting by Morimoto (1977), taken with what is known of
the distribution of syncarids in Europe and Australia, suggests the prospect of
further exciting discoveries locally in this particular habitat. Cave systems and
karst-type landscapes with sink holes and sunken streams are common in
many parts of New Zealand (Crossley et al. 1981), and there is a very strong
fraternity of recreational cavers, some of whom have already contributed to
scientific knowledge of cave life. There is a real challenge to use the technical
expertise of such people to look for these fascinating ‘living fossils’; a preliminary
guide to promote such work was issued by the New Zealand Department of
Conservation (Hunt & Millar 2001). The results of a 15-year study of Spanish
cave fauna by Camacho (2000) shows what could be achieved by a systematic
approach towards elucidating New Zealand’s subterranean syncarid fauna.
There is a growing appreciation worldwide of the importance of groundwater
organisms as environmental indicators of water quality, not to mention the
scientific interest of these organisms in their own right (Danielopol 1992;
Marmonier et al. 1993; Danielopol et al. 2000; Gibert et al. 1994; Jones &
Mulholland 2000; Scarsbrook et al. 2000, 2003) and the need to understand karst
landscapes and their fauna from a conservation perspective (William & Wilde
1985) and cave life in general (Vandel 1964; Ford & Cullingford 1976; Sasowsky
et al. 1997; Culver 1982; Camacho 1992; Juberthie & Decu 1994–2001). ‘Living
fossils’ carry appealing overtones in the public imagination (Dawson 2003a),
so the demonstration of the existence even of such tiny forms as the syncarids
could be another highlight to make known.
Orders Lophogastrida and Mysida (‘Mysidacea’): Opossum
shrimps
An estuarine species of Tenagomysis.
Stephen Moore
The Mysidacea are shrimp-like but have a number of characters, including
a ‘brood pouch’, that distinguish them from other crustaceans of similar
appearance. They are mainly marine, living in all oceans from great depths to
brackish coastal waters, and there is a small number of freshwater species. They
are of limited commercial importance and therefore not as familiar as the decapod
shrimps and prawns. Mysidacea may, however, be very common, particularly in
estuaries and coastal waters, where they often congregate in large swarms, and
are of considerable importance as primary consumers and as food of fishes.
Historically, the Mysidacea comprised a single order with two suborders
– Lophogastrida and Mysida. The two groups differ in important ways and
there is now debate over whether they are mono- or polyphyletic (having one,
or more than one, ancestor). Some workers question whether the Mysida,
which contains the great majority of Mysidacea, even belongs in the large
malacostracan superorder Peracarida, with the Lophogastrida; see Martin and
Davis (2001) for a discussion of mysid classification. These authors discarded the
Mysidacea, raising the two suborders to ordinal status, a decision followed here.
Even so, the two groups have many characters in common and, since relatively
few species (24) have been recorded from New Zealand waters, are discussed
here collectively.
Historical studies
Mysidacea have been recognised since the late 18th century. The taxonomic
literature is scattered and deals mostly with northern hemisphere faunas and
least with that of the Indian Ocean and Australasia. Major contributors include
Tattersall and Tattersall (1951), Gordan (1957), Mauchline and Murano (1977),
Mauchline (1980), and Müller (1993).
The history of New Zealand mysidacean taxonomy is brief. The first
published record is that of Thomson (1880), who described Siriella denticulata.
140
PHYLUM ARTHROPODA
CRUSTACEA
Kirk (1881) described Mysis meinertzhagenii, but the type and further evidence of
its existence have not been found since. Thomson (1900) described Tenagomysis
novaezealandiae from brackish water near Dunedin, and Calman (1908) attributed
an immature mysidacean specimen to the genus Pseudomma, apparently not
identified since. Tenagomysis tenuipes Tattersall, 1918, from Carnley Harbour,
Auckland Islands, brought the early list of mysidaceans known with certainty
to occur in New Zealand to three. Next, Walter Tattersall’s (1923) report on the
Mysidacea of the 1910 Terra Nova Expedition to Antarctica and the Southern
Ocean added 12 species. Eight were new, seven of which belonged to Tenagomysis,
and the remaining species was named Gastrosaccus australis, the first and so
far only named species of the genus from New Zealand. New records for New
Zealand of previously described species of Mysida included Euchaetomera
oculata, E. typica, and Siriella thompsoni. Chilton (1926) presented an overview
of New Zealand Mysidacea to that date. Later, Olive Tattersall (1955) identified
Boreomysis rostrata and Euchaetomera zurstrasseni from New Zealand waters and
Hodge (1964) redescribed Tenagomysis chiltoni. The most recent addition to the
fauna was that of Tenagomysis longosquama (Fukuoka & Bruce 2005).
Walter Tattersall (1923) appears to be the first to have recorded a species
of Lophogastrida, Paralophogaster glaber, in New Zealand. Apart from a record
of Lophogaster sp. from Te Papa (Museum of New Zealand) collections, the
remaining records are from Fage (1941) reporting on mysidaceans caught by the
1928–30 Dana Expedition, all in the family Gnathophausiidae: Gnathophausia
elegans, G. zoea, Neognathophausia ingens, and N. gigas.
Clearly there are more mysidacean species to be described from New
Zealand. Small numbers of specimens have been collected, with most material
in New Zealand held at the University of Otago and Auckland University of
Technology (Jocqué & Blom 2009).
Morphology, species, and endemism
The carapace is well developed in Mysidacea and covers the thorax, as it does in
euphausiids and decapod shrimps, but is fused with the anterior three or four
thoracic segments only; the back of the carapace can simply be lifted to expose
the posterior four or five thoracic segments. They have a shrimp-like abdomen
with fully developed or reduced pleopods, and the telson and paired uropods
form a tail fan. Mysidacean eyes are compound and stalked although in a few
deep-water species they are reduced to immovable plates. The antennules are
always biramous and most have an antennal scale. Of the eight pairs of thoracic
appendages, the first one or two are modified as maxillipeds. The remaining
six or seven pairs form legs and generally bear swimming exopods. A feature
of female mysidaceans is their large leaf-like oostegites, on the inner side of
some or all of the legs, which overlap to form a brood chamber or marsupium
(recalling the name opossum shrimp) beneath the thorax, in which eggs are laid
and the young develop. Both orders have all these characters in common.
In the Lophogastrida, however, gills are present on some or all of the legs,
pleura (‘side plates’) are present on the abdominal segments, and the pleopods
are well developed and usually unmodified in both sexes. Lophogastrids also
have seven pairs of oostegites but lack statocysts in the endopods of the uropods.
All species of Lophogastrida live offshore in meso- and bathypelagic habitats,
with many being hyperbenthic (living close to the bottom) in deep water. The
largest mysidaceans belong to this order and most occur throughout the world’s
oceans but are less often seen than species of Mysida, because of their oceanic
existence.
A characteristic of the Mysida (excluding all 33 species of the Petalophthalmidae) is the presence of a pair of balancing organs or statocysts, in the
telson. Situated near the base of each uropodal endopod, statocysts are an obvious
feature, distinguishing mysids from similar animals such as euphausiids (krill).
Mysids also lack gills and the pleopods of females are reduced or rudimentary;
Female Neognathophausia ingens
(Lophogastrida).
From Sars 1885
141
NEW ZEALAND INVENTORY OF BIODIVERSITY
those of males are variously modified. Like the lophogastrids, many mysids have
seven pairs of oostegites, but there are fewer pairs in some subfamilies of the
Mysidae, including the Gastrosaccinae, Mysinae, and Siriellinae, which between
them contain 16 of the 18 species of Mysida recorded from New Zealand. Mysids
occur throughout the marine environment to deep oceanic trenches but are
particularly concentrated in coastal regions, and 24 species have colonised fresh
waters around the world.
Adult Mysidacea range considerably in size from 2–3 to 350 millimetres
long. The largest are in the Lophogastrida but most species belong to the Mysida
and are appreciably less than 100 millimetres long. The few species recorded
from New Zealand almost cover this range, with mature females of Tenagomysis
macropsis as small as 3.2 millimetres long (Greenwood et al. 1985) and the largest
of all mysidaceans, Neognathophausia gigas, also being recorded in New Zealand
waters (Fage 1941).
Around 1000 species of Mysidacea have been described worldwide, the great
majority in the order Mysida, with some 51 in the Lophogastrida. Twenty-four
species have been recorded in New Zealand waters, representing both orders
(see end-chapter checklist). Of the three lophogastrid families, the Eucopiidae
are not yet known here. Of the four families of Mysida, two are found in New
Zealand – the Petalophthalmidae (one unnamed species) and Mysidae (all other
species). Globally, this is a very large family, with ca. 870 species. Four of the six
subfamilies occur in New Zealand.
As might be expected in a worldwide group inhabiting a wide diversity
of habitats, endemism reflects distribution; no species of the oceanic order
Lophogastrida is confined to New Zealand whereas endemism is high in species
occupying coastal and littoral waters. Twelve of the 18 species (~67%) of New
Zealand Mysida are endemic, including all 10 species of Tenagomysis (Müller
1993), but although the genus was first described from Otago (Thomson 1900)
it is no longer restricted to New Zealand; five species are known from either
Australian or African shores. While Siriella denticulata is endemic, S. thompsoni
is cosmopolitan in its distribution, as one of a minority of epipelagic Mysidacea.
The five non-endemic New Zealand Mysida are offshore species, the shallowest
among them being Euchaetomera typica, another pelagic species, found between
the surface and 380 metres. The distributions of the two unnamed species of
Mysidacea are not known. Neither Petalophthalmus sp. from deep offshore water
nor Lophogaster sp. in a typically oceanic genus is likely to be endemic.
Male Neognathophausia gigas (Lophogastrida).
From Sars 1885
142
Ecology and distribution
Distributional records of named New Zealand Mysidacea are, for the most
part, far from comprehensive, although there are probably records of littoral
species in unpublished environmental reports from various parts of the country.
Apparently the only records of Paralophogaster glaber are those of Tattersall
(1923) offshore of Cape Maria van Diemen and the Three Kings Islands in the
far north. Te Papa collections indicate that Neognathophausia ingens is common
around central New Zealand at least as far south as Banks Peninsula, N. gigas
is present off the east coast of the North Island, and Gnathophausia zoea in the
Bay of Plenty and on the outer Challenger Plateau west of Cook Strait. The
deepest record of any of the mysidacean species found in New Zealand waters
is that of G. zoea, at 6050 metres (Müller 1993) at a non-New Zealand locality.
The majority of mysidacean species are found on the inner shelf and in coastal
and littoral areas and form an abundant component of estuary zooplankton.
Many have very localised distributions and can form dense concentrations
among rocks and algal beds. Ingles (1973) encountered Tenagomysis macropsis
in high numbers in association with red algae in Pauatahanui Inlet. All 10
Tenagomysis species in New Zealand are coastal pelagic or littoral, and in some
cases freshwater dwellers. Tenagomysis macropsis is widespread around New
PHYLUM ARTHROPODA
CRUSTACEA
Zealand, from Spirits Bay eastwards almost to the Chatham Islands (Tattersall
1923) and south to Foveaux Strait although the maximum recorded depth of
the species is only 24 metres (Bary 1956). New Zealand’s southernmost species,
T. tenuipes, is so far known only from Foveaux Strait and east of Stewart Island
(Bary 1956), and from Carnley Harbour, Auckland Islands.
New Zealand has no strictly freshwater species but Tenagomysis chiltoni
passes through its life-cycle in at least one completely freshwater locality – Lake
Oturi, near Waverley, southwestern North Island (Hodge 1964). Thomson (1900)
had originally collected T. chiltoni from fresh and saline waters in Otago. Jones
et al. (1989) confirmed that this species also frequents saline waters in the AvonHeathcote Estuary (Christchurch) but is an upper estuarine species and was
seldom found in salinities greater than 20 parts per million (ppm). Chapman and
Lewis (1976) reported T. chiltoni and T. novaezealandiae as living in brackish water
below the Paratya curvirostris (Decapoda) zone in streams. Jones et al. (1989)
indicated a salinity-correlated ecological separation between T. chiltoni and T.
novaezealandiae in the Avon-Heathcote Estuary with the former in upper reaches
and the latter mid- to upper estuarine. In this study and that of Greenwood
et al. (1985), T. macropsis was found throughout the estuary and had no linear
correlation with salinity range along a transect from 4.1 to 30.1 ppm. In his work
in Pauatahanui Inlet, Ingles (1973) found distinct differences in distributions
between three species in the Horokiwi Stream – T. macropsis occurred in the
estuary proper, entering only the mouth of the stream, T. novaezealandiae centred
around the mouth and lower part of the stream, while Gastrosaccus australis was
highest upstream, not moving as far as the mouth.
Tenagomysis macropsis, the most abundant species in the Avon-Heathcote
Estuary, occurs in greatest numbers at salinities between 16.9 and 19.2 ppm, but
it is clearly euryhaline as Bary (1956) found it in great numbers in Foveaux Strait
(ca. 60,000 individuals in one plankton tow). The results from overnight surface
samples in a tideway, taken during his survey of mysidaceans and euphausiids
east and south of the South Island, indicated daily vertical migrations by T.
macropsis and T. tenuipes. The numbers of both species at the surface (including
juvenile T. macropsis), peaked around 2 a.m. Bary’s is the only study published to
date on vertical distributions of New Zealand Mysidacea.
Swarming is characteristic of mysidaceans (though not as densely as
euphausiids) and more complex than it may appear. Mauchline (1980) discussed
possible reasons for this behaviour. Concentrations probably result when
physical and chemical factors in the water make some areas more habitable than
others. Salinity, food availability, light or dark, and age class are all components
of swarming behaviour. Conditions change regularly in estuaries and dispersed
populations can be forced to aggregate in restricted areas at low tide. Ingles’s
(1973) work on T. macropsis in Pauatahanui Inlet suggested a relationship between
shoaling and the tidal cycle. Breeding aggregations also take place, probably
more so in deep-water species because littoral mysidaceans regularly aggregate
for other reasons but breed at the same time. Data gathered by Greenwood
et al. (1985) suggested that T. macropsis may undertake seasonal migrations,
in common with littoral mysids in other parts of the world (Mauchline 1980).
Mature T. macropsis females move up-estuary with the rise of temperature in
spring whereas Roper et al. (1983) found them closer to the estuary mouth
in winter. Aggregation of females over the summer breeding season suggests
that this is for breeding purposes. Swarming in currents can also lead to the
segregation of age classes, which have differential swimming rates. Swarms are
of all shapes from globular to elongated and can be very extensive horizontally
in the water but only a few centimetres thick (Mauchline 1980).
Reproduction and development
Mysidacea do not have planktonic larvae as most euphausiids and decapods do.
Instead, development of embryos and larvae takes place in the marsupium, from
143
NEW ZEALAND INVENTORY OF BIODIVERSITY
which they emerge as juveniles. Mating usually, if not always, involves the male
using its fourth pleopods to deposit sperm in the female marsupium (Mauchline
1980) and eggs are fertilised as they are laid in the marsupium. The number and
size of resulting embryos depends upon the size of the eggs and the female and
on water temperature and season. Meso- and bathypelagic species tend to have
larger eggs and produce somewhat fewer young than epipelagic and coastal
species, relative to body size.
The embryo (developmental stage 1) grows to some extent in the egg
membrane, moults into a stage 2 (eyeless) larva, and passes through a third (eyed)
stage to moult into a juvenile ready to emerge from the marsupium. Juveniles
grow to become adults without passing through further stages, although the
abdomen increases in proportion to the cephalothorax, and the appendages
and telson undergo gradual changes as well. Jones et al. (1989) found the sizes
of embryos in Avon-Heathcote Estuary species to be in accord with the range
generally found for coastal forms. A range of embryo numbers was also carried
in the marsupia of the Tenagomysis species: 4–25 in T. macropsis, 6–19 in T.
novaezealandiae, and 22–39 in T. chiltoni.
In T. tenuipes from Foveaux Strait, Bary (1956) found that females (up to
19.9 millimetres long) shed juveniles of 4.2 millimetres length. Those of T.
macropsis from the same area were about 2.5 millimetres long, mature females of
T. macropsis being less than half the length of T. tenuipes. Greenwood et al. (1985)
found emerging larvae of T. macropsis to average only 1.47 millimetres in length
in the Avon-Heathcote Estuary.
Tenagomysis chiltoni.
After Chapman & Lewis 1976
144
Food and predators
Dietary studies of Mysidacea are limited and none has been carried out on any
New Zealand species, although Chapman and Lewis (1976) considered that
Tenagomysis chiltoni and T. novaezealandiae might be detritus feeders. Chapman
and Thomas (1998) subsequently reported predatory feeding in T. chiltoni.
Mouthparts and thoracic appendages are variously modified in relation to diet.
Some species are strict filter feeders, some specialise in grazing phytoplankton,
and some are carnivores concentrating on certain substrata such as algae, but most
are more opportunistic and eat a considerable range of the organic material in
their environments. Mauchline (1980) tabulated the diets of 25 species of mainly
northern hemisphere mysids. Though by no means comprehensive, his table
showed the major importance of organic detritus, significantly supplemented
by diatoms, other algae, copepods, and other crustaceans. Probably most New
Zealand shallow-water species have similarly generalised diets, but a few
specialised feeders are indicated. One of the most extreme modifications of feeding
appendages is of the mandibular palp in Petalophthalmus species. It is greatly
elongated, projecting well beyond the antennae. Carnivorous Petalophthalmus
armiger pierces its prey and sucks out the internal contents (Mauchline 1980).
Lophogaster typicus is incapable of filter-feeding, having mouthparts modified to
feed on large lumps of food material on the surface of sediments, suggesting that
New Zealand Lophogaster sp. could have a similar diet.
Filter-feeding is common in Mysidacea and is accomplished using setose
mouthparts and thoracic appendages. The animals ‘stand on their heads’ above
soft substrata, creating a current using the thoracic exopods and filtering
particles from the stirred-up sediment. In common with euphausiids (see
section on Euphausiacea), some Mysidacea employ a ‘food basket’, formed by
the mouthparts and anterior thoracic appendages, in which food items collected
using the mandibular palps are retained until they are chewed and swallowed.
Some species follow diel feeding rhythms, with certain species feeding
by day, others only at night. Gastrosaccus australis individuals caught by Jones
et al. (1989) were virtually all taken at night in the Avon-Heathcote Estuary,
suggesting that they feed nocturnally instead of competing with the three
Tenagomysis species by day.
PHYLUM ARTHROPODA
CRUSTACEA
Mysidacea are important links in the food web between primary producers
(e.g. bacteria and microalgae) and secondary consumers, especially in coastal
waters. They therefore play a critical role in the cycling of energy through
the detrital pathway (Jones et al. 1989). Mysids especially are eaten by a very
wide variety of fish and also by decapod crustaceans, seabirds, cetaceans, and
other predators. Data on predation of lophogastrids is limited because they
live offshore but the size and appearance of the largest species facilitates their
recognition in stomach contents. Albacore tuna eat Neognathophausia ingens.
Deep-sea hyperbenthic rattails eat mysidaceans including N. gigas, a species
also reported in fin whale stomachs (Nemoto 1959 in Mauchline 1980) and N.
ingens has been found in the stomachs of pigmy sperm whales stranded in New
Zealand (Te Papa data). Weddell seals and gentoo penguins are known to eat the
Antarctic mysid Antarctomysis maxima and, intriguingly, yellow-nosed albatross
near Tristan da Cunha have been found with N. ingens and N. zoea in their guts
(Mauchline 1980). It seems these otherwise extremely deep-living lophogastrids
may undertake diel migrations near enough to the surface to be captured
by albatrosses. Deep benthic and midwater prawns including Aristaeopsis
edwardsiana, Pasiphaea tarda, and Aristaeomorpha foliacea, found in New Zealand
waters, have also been found to eat mysidaceans.
Mauchline (1980) cited many studies of the diets of coastal fish that indicated
the major significance of mysids as food items. He also noted that mysids tended
to be underestimated as prey items because their remains were often mistaken
for euphausiids. Little information on mysidaceans in the diets of New Zealand
fish is apparent, although Griffiths (1976) reported that introduced European
perch in the Selwyn River (Canterbury) eat high numbers of T. novaezealandiae.
Estuaries such as Avon-Heathcote and Pauatahanui are important as fish
nurseries and there is little doubt that the mysids that concentrate there are
an important part of their diets. In lakes of the Waikato district, mysids are an
important part of the diet of smelt (Northcote & Chapman 1999). Along the
coast, seahorses (Hippocampus abdominalis) ingest Tenagomysis similis along with
amphipods and the shrimp Hippolyte bifidirostris (Woods 2002), all found in the
subtidal kelp beds in which seahorses live.
Mysidacea employ defensive strategies to avoid being eaten, including, as in
shrimps and lobsters, tail flexing. While transparent and virtually invisible when
swimming, mysids have chromatophores – pigment cells – that enable them to
adopt camouflage colours and blend with algae, rocks, or sand. Lophogastrids are
uniformly bright red, so can avoid detection by exploiting the lack of penetration
of red light in sea water, as do many meso- and bathypelagic decapods. Swarming
may also confer some protection on mysidaceans by reducing the number of
targets apparent to their attackers.
A wide range of ecto- and endoparasites have been reported in Mysidacea.
Very common endoparasites are ellobiopsid protozoans (phylum Myzozoa) such
as Thalassomyces fasciatus, found in N. gigas, N. ingens, and G. zoea. Choniostomatid
copepods parasitise mysidaceans, and epicaridean isopods, particularly of the
family Dajidae, are common ectoparasites. Juvenile and small male dajids live in
the host’s marsupium among the developing larvae.
Economic aspects
Mauchline (1980) reported that thousands of tons of Neomysis intermedia, N.
japonica, and Acanthomysis mitsukurii are harvested each year in Japan; N. intermedia from brackish lakes is the most important of these and is cooked, dried and
eaten. There do not appear to be any other major fisheries for Mysidacea but several species are or have been fished in South-east Asia, China, and Korea by local
fishers, who net them when they swarm. Some species have been reared successfully in laboratories, and freshwater species have been successfully transferred to
other rivers or lakes as food for fish. It is also possible that some Mysidacea have
colonised other habitats by transferring there on ships’ hulls or in ballast water.
145
NEW ZEALAND INVENTORY OF BIODIVERSITY
Future work
There is clearly a need for further taxonomy followed by biological research on
the Mysidacea of New Zealand before we can gain a reasonable appreciation
of their diversity. Historically, New Zealand has never had the services of a
mysidacean specialist but the need for such work is surely increasing given the
importance of mysids in the marine economy, particularly as a major food of fish.
Once Mysidacea currently held in collections are analysed, further assessment
of their diversity, numbers, and roles in the region will require sampling gear
and strategies appropriate to the collecting of these generally small and easily
damaged animals.
Order Amphipoda: Beach fleas, sand hoppers, and kin
The freshwater amphipod Paracalliope fluviatilis.
Stephen Moore
Paradexamine houtete.
From Barnard 1972a
146
Amphipods are the among the most ubiquitous crustaceans, inhabiting diverse
environments from the depths of the oceans’ trenches to high altitudes on
mountains, living in situations as varied as plankton in the open seas, burrowers
in surf beaches, litter-dwellers on forest floors, epizoites on the skin of whales
and dolphins, and cryptic inhabitants of subterranean aquifers more than 20
metres below ground level. Amphipods are likely to be found in almost all
aquatic habitats, as well as on land wherever water is freely available or humidity
is high. In many of these situations, species are numerous and numbers high,
frequently overwhelmingly so. It is surprising, therefore, that they have received
relatively little scientific attention.
The name of order is derived from two Greek words – amphi, both or of two
kinds, alluding to the forward orientation of the anterior legs and the backward
and/or lateral orientation of the posterior legs (Stebbing 1888), and podos, foot.
The relative neglect of amphipods as subjects for scientific study in New
Zealand may be because of two related attributes – their biodiversity is bewildering
and different species are often not easily distinguished by the untrained eye.
The trained worker, on the other hand, finds the myriad variations on the basic
morphology fascinating, continually generating questions about relationships
between taxa and the selective value of the differences in morphological
structures.
The basic amphipod body plan is difficult to define because of the group’s
diversity. Amphipods are distinguished from other peracarids (malacostracan
crustaceans that brood their eggs and young) by the following combination
of characteristics: body generally laterally compressed, carapace absent, eyes
sessile and usually lacking cuticular facets, pereon (thorax) with seven pairs of
unbranched limbs, pereopods (legs) 1–4 orientated anteriorly, pereopods 5–7
directed posteriorly, pereopods 1–2 usually modified as subchelate (grasping)
gnathopods, coxal gills present on some pereopods, pleon (abdomen) segments
1–3 with multi-articulate swimming appendages (pleopods), usually biramous,
pleon segments 4–6 (urosomites) with stouter, biramous uropods, the final
urosomite with a distinct telson.
Some 6000 species in about 120 families are known worldwide (Barnard &
Karaman 1991). Estimates suggest that several thousand species await discovery
and scientific description, despite more than 100 new species being described
annually, on average, during the mid-1980s. The order is divided into three
suborders – Ingolfiellidea, Gammaridea, and Hyperiidea; caprellids (formerly
Caprellidea) are now regarded as specialised gammarideans.
Historical overview
Knowledge of the New Zealand amphipod fauna began with Dana’s (1852,
1853–55) descriptions of a few species, but accelerated with G. M. Thomson’s
and Charles Chilton’s work. Thomson’s (1879b) first paper was followed by
14 more over the next 34 years; that of Chilton (1882a) was succeeded by 52
papers by 1926, although not all dealt with New Zealand species. Thomson and
PHYLUM ARTHROPODA
Chilton’s (1886) ‘Critical list of the Crustacea’ contained 71 amphipod species
names: 63 gammarideans, four hyperiids, three caprellids, and one cyamid.
Chilton was the strongest influence on early New Zealand amphipod
systematics. He himself was influenced by Della Valle’s (1893) attempt to combine
many of the world’s Gammaridea into fewer species and he treated many New
Zealand species as variants of extrinsic taxa (Barnard 1972a). This tendency was
exacerbated in his later career by his acquaintance with research on phenotypic
variation of amphipods at Plymouth (England). This led him to regard many New
Zealand species as phenotypes of sub-cosmopolitan species (Barnard 1972a) or
as variants of local species (Fenwick 2001a). Significant contributions were also
made by Stebbing (1888, 1910) through his work on local collections made by
the Challenger and Thetis Expeditions. Also notable are Walker’s (1908) work on
subantarctic material, K. H. Barnard’s (1930) studies of Terra Nova Expedition
collections from the far north of New Zealand, and Stephensen’s (1927) and
Nicholls’s (1938) studies of subantarctic amphipods.
A new phase of New Zealand amphipod systematics began in the 1950s
with D. E. Hurley’s detailed papers (1954–75) on gammarideans, hyperiids, and
cyamids. Several problems were resolved, new species described, and many
previously described species clarified. Extensive collections from New Zealand’s
deep waters were made during the Danish Deep-Sea Expedition, 1950–52, on the
Galathea. Dahl’s (1959) and Barnard’s (1961) reports on these collections added
considerably to knowledge of our fauna. In none of the preceding investigations,
however, was there any attempt to collect amphipods widely in New Zealand
waters in order to gain understanding of species’ distributions. This, however,
was the approach followed by J. L. Barnard during his 1967–68 visit. The resulting
monograph (Barnard 1972a) made a preliminary assessment of the biogeography
of the New Zealand gammaridean fauna, described numerous new taxa, and
provided the most comprehensive guide to date of the fauna (although its focus
was algae-living amphipods). Barnard’s visit and monograph stimulated much
subsequent local interest in the gammaridean fauna (Cooper 1974; Cooper and
Fincham 1974; Hurley and Cooper 1974; Fincham 1974, 1977; Lowry 1979, 1981;
Fenwick 1976, 1977, 1983; Myers 1981; Lowry and Fenwick 1982, 1983; Moore
1983a,b, 1985; Lowry and Stoddart 1983a,b, 1984).
New Zealand freshwater amphipods were studied by Hurley (1954a,
c, f) over this period, as were terrestrial amphipods (Hurley 1955a, 1957a, c).
Bousfield (1964) and Duncan (1968) also investigated the terrestrial amphipods.
Subsequently, Duncan (1994) substantially reviewed this group, recognising
several new genera and species.
Elements of the New Zealand hyperiid fauna were reported by Stebbing
(1888) and K. H. Barnard (1930). After about 1950, hyperiids and caprellids were
usually investigated and reported separately from gammarideans, with Fage
(1960), Shih (1969), and Hurley (1955b) exploring the fauna more fully. Much
of this information is brought together in Vinogradov’s (Vinogradov et al. 1996)
substantial review of the world hyperiids, with Zeidler (2003a, b, 2004a, b, 2006,
2009) refining the group’s systematics and adding further new records. The New
Zealand caprellids were reviewed by McCain (1969) and he described one new
species subsequently (McCain 1979).
Amphipod diversity in New Zealand currently stands at 500 species, of
which 64 are undetermined or undescribed.
Amphipods in the ecology of New Zealand
The general abundance of amphipods means that, despite their small individual
size, collectively they are important in the ecology of many ecosystems, especially
as food for larger animals. Huge densities of amphipods are found among New
Zealand seaweeds, in which they often dominate the associated fauna (Fenwick
1976; Taylor 1998).
Several studies have demonstrated the importance of gammaridean and
CRUSTACEA
Cyamis boopis.
From Hurley 1952
Parawaldeckia angusta.
From Lowry & Stoddart 1983a
147
NEW ZEALAND INVENTORY OF BIODIVERSITY
Waitomo manene.
From Barnard 1972a
Themisto gaudichaudi.
From Stebbing 1888.
148
hyperiid amphipods as food for fish and birds in New Zealand. Amphipods were
the most frequently utilised food item among 26 species of common northern
New Zealand reef fishes (Russell 1983). Indeed, Jones (1988, p. 454) considered
‘the importance of gammaridean amphipods as a food source … startling’ for
juvenile fish. They were the principal food item for adults of several species and
formed important secondary foods for the others (Russell 1983). These amphipods
were mostly gammarideans and caprellids but some planktivorous fishes ate a
few hyperiids. Small fish species were most dependent upon amphipods for food.
Amphipods were eaten by 75–90% of specimens and comprised 40–60% of diet by
volume in the various triplefin species (Russell 1983). A few large species also fed
extensively on amphipods. Over half of all red moki, blue moki, trevally, goatfish,
and juvenile snapper ate gammaridean amphipods, which made up 40%, 38%,
51%, 55%, and 62%, respectively, of their food by volume (Choat & Kingett 1982;
Russell 1983). A similar study at Kaikoura (Duffy 1989) confirmed the importance
of amphipods as food for fishes and showed their increased consumption by fishes
inhabiting brown seaweeds of semi-sheltered, southern shores.
Amphipods are important food for some fishes inhabiting soft bottoms and
estuaries also. Adults of nine species of fish in the Avon-Heathcote Estuary all ate
amphipods, although they were a common (> 10%) food item for three species
only – common sole (13%), cockabully (68%), and common bully (74%) (Webb
1973). Although amphipods were scarce in the diets of yellow-bellied and sand
flounders in the estuary (Webb 1973), their juveniles fed almost exclusively
(92–96% of food items) on the small tube-dwelling amphipod Paracorophium
excavatum (Nairn 1998). Offshore, however, larger amphipods were common
items (33%) in adult yellow-bellied flounders’ diets (Knox & Fenwick 1978).
Fish also eat pelagic hyperiid amphipods. Warehou, banded rattails, javelin
fish, black oreos, southern blue whiting, carinate rattails, small-scaled brown
slickhead, and small-scaled nototheneids all include amphipods as substantial
components of their diets. Many of these fishes fed extensively on amphipods
when smaller (up to 37% of food weight and eaten by up to 75% of small fish),
with individual fishes taking larger prey as their sizes increased (Gavrilov &
Markina 1981; Clark 1985; Rosecchi et al. 1988; Clark et al. 1989). Amphipods
were a minor element of the diets of several other deeper-water fishes, notably
hoki, smooth oreos, smooth rattails, and orange roughy. Pelagic fishes are the
usual predators of these amphipods, but benthic fishes may feed extensively
on hyperiids when swarms are carried into shallow water. At The Snares,
the demersal telescope fish, as well as spotties, banded wrasse, and benthic
nototheneid cod, fed intensively on hyperiids (Themisto gaudichaudi, Hyperietta
luzoni) and krill swarming close to the surface (Fenwick 1978).
The importance of amphipods in freshwater fishes’ diets varies with species,
amphipod abundance, abundance of other prey items, and fish size. Longfinned and short-finned eels, whitebait (Galaxias maculatus), mudfish, common
smelt, and brown trout all eat small numbers of the common stream amphipod,
Paracalliope fluviatilis (McDowall 1968; Eldon 1979; Ryan 1986; Jellyman 1989;
Sagar & Glova 1995, 1998; Hicks 1997). Typically, amphipods comprise less than
5% of whitebait food, but more are eaten with increasing fish size (McDowall
1968). Amphipods are commoner in the diets of whitebait closer to estuaries
than those further upstream and, in some rivers, amphipods comprise up to 45%
of the diet (McDowall 1968). Similar variation in the consumption of amphipods
occurs in eels. Amphipods (Paracalliope fluviatilis and the brackish Paracorophium
excavatum) may be a major (70%) or minor (< 0.01%) food for short-finned eels,
depending upon the specific habitat, season, and eel size, with amphipods being
most important for small eels 100–190 millimetres long. Similarly, juvenile brown
trout feed preferentially on amphipods, which make up 80% of food items of
trout inhabiting tree-lined sections of some rivers.
Birds also feed on marine and estuarine amphipods. A number of oceanic
birds typically feed extensively on hyperiid amphipods. Red-billed gulls, cape
PHYLUM ARTHROPODA
CRUSTACEA
pigeons, Buller’s mollymawks, and sooty shearwaters fed on hyperiid swarms at
The Snares, with the latter two diving below the surface to catch them at times
(Fenwick 1978). Fairy prion chicks are fed a diet comprising 14% amphipods by
weight, diving petrels consume 17% by weight of amphipods, and grey-faced
storm petrels at the Chatham Islands include four species of amphipods in their
diet (Prince & Morgan 1987).
Numerous other New Zealand birds eat amphipods as larger or smaller
components of their diets. For example, most penguins are believed to include
these crustaceans in their diets (Croxall & Lishman 1987). In North American
estuaries, some migratory waders consume 10,000–22,000 corophiid amphipods
per day (Wilson 1989). Related species (plovers, dotterels, and wrybills) in New
Zealand probably eat appreciable quantities of amphipods. Ground-foraging,
insectivorous birds (e.g. robins, fernbirds, tits, and wekas), as well as blackbirds
and song thrushes, are almost certain to include land hoppers from among plant
litter in their diets. In addition, gulls and other birds probably capture beachfleas from amongst wrack at times.
Diversity of New Zealand amphipods
Ingolfiellidea
Ingolfiellids are highly specialised, mostly small (< 3 but up to 14 millimetres
long), worm-like animals adapted to living interstitially in marine and freshwater
sediments, as well as in groundwaters. Marine species occur from the intertidal
to the deep sea. Widely regarded as very primitive amphipods, over 30 species
are known from two families. They are reported from most continents, including
Australia, and two species from New Zealand interstitial marine habitats
(Schminke & Noodt 1968) remain undescribed.
Caprellidea
In a detailed cladistic analysis, Myers and Lowry (2003) demonstrated that
caprellids and cyamids are specialised corophiidean amphipods. They are
discussed separately here but the end-chapter checklist follows Myers and
Lowry. The Caprellidea includes two distinct families, both found worldwide –
the skeleton shrimps (Caprellidae) and whale lice (Cyamidae). Whale lice live
ectoparasitically on whales and dolphins, whereas caprellids are benthic and
often extremely abundant among algal fronds and on bryozoans, hydroids, and
sea stars intertidally and on shallow marine bottoms. Each group’s body form
is very different, although both possess rudimentary abdomens and vestigial
abdominal limbs. Whale lice have short, flattened bodies with powerful limbs
adapted to grasp their hosts’ skin firmly. Caprellids have long slender bodies and
their last three pairs of legs, grouped posteriorly, are modified for grasping the
substratum, leaving their anterior legs and antennae free for feeding.
Caprellids are quite diverse, with about 85 genera worldwide (McCain
& Steinberg 1970; Laubitz 1993). The New Zealand skeleton-shrimp fauna
comprises just eight species in six genera, belonging to two subfamilies (McCain
1969, 1979; Guerra-García 2003). Half (four) of these species are endemic. Eight
species of whale lice in four genera are known from New Zealand (Hurley 1952;
Lincoln & Hurley 1980), whereas the worldwide cyamid fauna comprises 27
described species in six genera (Martin & Heyning 1999). If, however, cyamids
known to occur on whale and dolphin species reported from New Zealand
waters are considered, the total cyamid fauna may number some 19 species in
all six known genera.
Hyperiidea
Hyperiid amphipods are purely pelagic, living freely in the ocean or associated
with other pelagic invertebrates, from the surface to abyssopelagic depths (>
7000 metres) (Vinogradov et al. 1996). Species living near the surface typically
Whale louse Scutocyamus antipodensis.
From Lincoln & Hurley 1980
Phronima sedentaria.
From Hurley 1955b
149
NEW ZEALAND INVENTORY OF BIODIVERSITY
Iphinotus typicus.
From Barnard 1972a
Raukumara rongo.
From Barnard 1972a
150
make diurnal vertical migrations from below 200 metres depth to spend the
hours of darkness within the surface 50 metres.
A great variety of body shapes occurs within the hyperiids, making them
extremely difficult to characterise. Large eyes and/or an inflated head and
variously reduced first thoracic segments or pleon and urosome are common
(e.g. Hyperiidae), although the opposite is true in others (e.g. Scinidae). The
very compact forms of many surface dwellers (e.g. Platyscelidae) contrast with
the needle-like shapes of Rhabdosoma species. Lengths also vary widely from
2.5 millimetres (e.g. Hyperietta luzoni) to over 150 millimetres for the extremely
elongate Rhabdosoma armatum.
Some hyperiids live on and within one or a few species of jellyfish,
siphonophores, and ctenophores. The relationship between host and amphipod
seems uncertain, but the consistent pairings of some species (e.g. Hyperia
macrocephala is found only on the jellyfish Desmonema gaudichaudi) indicate
commensalism. Host tissues and other prey items in the guts of these amphipods
suggest that the amphipods behave opportunistically, with no obvious advantage
to the host. Species of the family Phronimidae apparently eat the viscera of
pelagic tunicates, siphonophores, and heteropods and use the prey’s transparent
covers as a refuge against predators and for rearing their eggs.
Over 240 species of hyperiid in more than 72 genera and 23 families are
known from the world’s oceans. It is difficult to characterise the New Zealand
fauna because of the hyperiid pelagic habitat. Many hyperiids have very wide
distributions (Vinogradov et al. 1996), so it seems inevitable that most widely
distributed species will be found in local waters eventually (Zeidler 1992),
depending upon movements of the specific water masses with which they
tend to be associated (Young & Anderson 1987). Thus, New Zealand’s hyperiid
fauna probably exceeds the reported 94 species in 49 genera reported from our
surrounding seas (Hurley 1955b; Kane 1962; Vinogradov et al. 1996; Zeidler
(2003a, b, 2004a, b, 2006, 2009) and a total fauna in excess of 100 species seems
probable.
Gammaridea
The Gammaridea is the most abundant, ubiquitous, and diverse of the amphipod
suborders. More than 5800 species in about 1100 genera are known, some from
hadal depths exceeding 10,000 metres (Dahl 1959) and others higher than
4000 m above sea level (Stebbing 1888). Gammarideans range in length from
about 2–3 millimetres to a whopping 340 millimetres for the abyssal Alicella
gigantea (Barnard & Ingram 1986). Large size appears to be associated with
higher dissolved-oxygen concentrations in cold-water habitats, and warmwater faunas are dominated by very small species. These amphipods also seem
most abundant and diverse in temperate to cool climates, with tropical faunas
being relatively inconspicuous, although surprisingly diverse (Thomas 1993).
Gammarideans are often referred to as the laterally compressed amphipods.
Land-hoppers, beach-fleas, and many aquatic amphipods certainly have
the typical shape. However, several tube-dwelling and nestling genera have
elongated, more vermiform, shapes. Burrowers in surf beaches (urothoids
and some phoxocephalids) are wide-bodied, presumably for stability in highenergy habitats. Iphinotus typicus is even more flattened. Its limpet-like shape
adapts it for life on the fronds of smooth brown seaweeds on New Zealand’s
turbulent rocky shores.
Marine and freshwater gammarideans are predominantly free-living and
benthic. A few are planktonic and others form close associations with algae,
hydroids, bryozoans, and a variety of other invertebrates. Members of some
families build tubes, nests, or columns from strands of material secreted from
glands in their anterior legs, variously incorporating mud, sand, shell, bryozoan
fragments, and other particles from their habitats. Species of yet other families
PHYLUM ARTHROPODA
characteristically burrow in soft sediments, at times burrowing to more than 200
millimetres beneath the sediment surface. Scavenging, detritivory, and omnivory
are the predominant feeding habits, but predation, ectoparasitism on fish, and
herbivory also are known (Bousfield 1987; Enequist 1949; Lowry & Stoddart
1983b; Sainte-Marie & Lamarche 1985; Haggitt 1999).
The New Zealand gammaridean fauna (including caprellids and cyamids)
comprises 401 species (62 undescribed) in 192 genera (10 unnamed), belonging
to 55 families. [Figures below indicate that New Zealand’s total gammaridean
amphipod diversity is probably 3–4 times geater than the total reported here.]
This equates to about 5.6% of the world’s described species and 15.8% of world
genera, representing over a third of all families. Some 74% of the species (296)
are endemic, as are ~29% (55) of the genera. The fauna inhabiting each of three
major habitats in New Zealand is discussed separately below.
CRUSTACEA
Paracentromedon? whero.
From Fenwick 1983
Terrestrial amphipods
All terrestrial species belong to the Talitridae, the only amphipod family to have
successfully occupied terrestrial habitats worldwide. These amphipods inhabit
gardens, forest floors, and grasslands, where they live in litter, under trees and
rocks, or in burrows that they construct themselves. Some 36 species in 10 genera
occur in New Zealand (Duncan 1994; Fenwick & Webber 2008). Beach fleas
are usually considered with terrestrial species, and 11 species in three genera
are known from shore environments, although their revision seems overdue.
Most New Zealand talitrids are endemics, but there are at least three aliens.
New Zealand species range in length from c. 5–6 to > 50 millimetres for the
giant subantarctic Notorchestia aucklandiae. Land hoppers and beach fleas occur
throughout New Zealand, including the subantarctic islands, from sea level to
over 2000 metres.
Freshwater amphipods
Some 53 species (~30 undescribed) in nine named (and 10 additional unnamed
new) genera belonging to eight families are reported from freshwater habitats
in New Zealand (Fenwick 2001a,b). Several undescribed species from hypogean
water (saturated sediments beneath or beside streams and rivers (hyporheic)
and groundwater) are currently under investigation and others from epigean
(surface) waters await description (Fenwick 2000). Within these habitats,
amphipods are often surprisingly abundant, but have received little attention.
This relative neglect probably reflects their small adult sizes (3–6 millimetres body
length), although two hypogean species (Phreatogammarus fragilis and Ringanui
toonuiiti) grow to over 20 millimetres long. All New Zealand freshwater species,
five named genera, ca. 10 unnamed genera, and three families are endemic.
Marine and estuarine amphipods
The New Zealand marine and estuarine amphipod fauna comprises some 365
species. Amphipods inhabit every conceivable habitat in the sea, although few
species live in estuaries. They are predominantly benthic, living in and on mud
and sand and rocky bottoms, as well as among other invertebrates and algae.
The total diversity of the New Zealand marine amphipod fauna is difficult to
estimate, but is likely to comprise at least three times the presently known
species. Of the known marine fauna, 194 species (~53%) and 35 genera (19%)
are endemic.
Special features of the New Zealand gammaridean fauna
Biodiversity and abundance
Amphipods are frequently a major component of marine benthos, especially in
cool-temperate to cold-water environments. New Zealand is no exception in
this respect. A study of animals inhabiting the green alga Caulerpa brownii at
Patuki roperi.
From Fenwick 1983
151
NEW ZEALAND INVENTORY OF BIODIVERSITY
Ringaringa littoralis.
From Cooper & Fincham 1974
Paracrangonyx compactus.
From Fenwick 2001
152
Kaikoura on the South Island east coast revealed a fauna dominated by huge
numbers of amphipods – up to 12,000 per 200 grams wet weight (handful) of
alga (Fenwick 1976). Some 61 species occurred in this specific habitat. Amphipod
abundance increased dramatically with increased exposure to wave action, but
fewer species predominated. Thus, the fauna at more sheltered sites comprised
lower densities, with more species having more equal abundances.
Shallow sand bottoms at Kaikoura illustrate amphipod abundance in
another near-shore habitat. Four species of amphipods and a large myodocopid
ostracod comprise most of the fauna in this habitat. Amphipod densities average
about 6000 per square metre, fluctuating from a winter low of 4000 to a summer
high of more than 12,000 per square metre (Fenwick 1985). Crowding of these
crustaceans is reduced by each species occupying a different depth in the
sediment (Fenwick 1984) – cryptically coloured, surface-skipping Patuki roperi
lives in the top 25 millimetres of sand, smaller white Ringaringa littoralis dwells
at about 40 millimetres depth, bright red Paracentromedon? whero inhabits middepths (50–80 millimetres), and large Protophoxus australis overlaps at mostly
65–85 millimetres. Leuroleberis zealandica, a very large ostracod, is most abundant
at 75–100 millimetres depth. Species’ mean depths in the sediment change
slightly between sand ripples (150–200 millimetres high) and troughs, as well
as with season.
Amphipods are a significant component of surf-zone faunas on New
Zealand’s exposed beaches, such as in Pegasus Bay (Fenwick 1999). These small,
frail-appearing crustaceans not only survive in these highly turbulent situations,
but some species are found nowhere else. Amphipod densities peak just outside
the zone of wave break, at about six metres depth in Pegasus Bay. Biodiversity of
the amphipod fauna changes markedly with depth and, hence, changes in waveinduced turbulence, with most species abundant in only one depth zone. All but
one of the abundant inshore (3–10 metres depth) species are free-living active
burrowers of the family Phoxocephalidae.
These three studies demonstrate some key aspects of marine amphipod
biodiversity. Perhaps most significantly, amphipods are a very important
component of faunas inhabiting many of the shallow marine habitats around
New Zealand. Not only are amphipods abundant in many of these habitats, but
also their biodiversity is high. Individual species of amphipods are very sensitive
to small changes or variations in their environments, resulting in marked
changes in faunas within and between habitats. Species within some families
exhibit very different tolerances of environmental factors, indicating that species
or genus may be more useful levels of taxonomic resolution for amphipods in
ecological investigations.
New Zealand Phoxocephalidae
Phoxocephalids are the typical amphipods of the surf beaches and sandy shores
that make up so much of New Zealand’s coastline. Fifteen (88%) of the 17
phoxocephalid species known from New Zealand are endemic. Eight (53%)
of the 15 genera to which these species belong are endemic and monospecific.
This generic diversity and endemism is remarkably high. Museum collections
indicate that the fauna includes 15 or more undescribed species, indicating over
30 species of phoxocephalids in New Zealand.
The Australian shallow-water phoxocephalid fauna consists of 89 species in
26 genera (comprising 40% of the known phoxocephalid species worldwide),
with 23 of these genera endemic (Barnard & Drummond 1978; Barnard &
Karaman 1983). Despite the high biodiversities of both the Australian and the
New Zealand phoxocephalid faunas, there is little overlap between the two. Only
one shallow-water genus (Booranus?) seems to be shared between New Zealand
and Australia, although three deep-water genera (Cephaloxoides, Harpiniopsis,
Protophoxus) and two of their species are found on both sides of the Tasman Sea.
Australia is regarded as the epicentre of phoxocephalid evolution because
PHYLUM ARTHROPODA
of high diversity of species and genera and high generic endemism (Barnard &
Karaman 1983). The subantarctic islands of South America are the only other
centre of phoxocephalid radiation, with distinctive attributes present among
its species and genera. New Zealand’s location between Australia and South
America indicates that the New Zealand phoxocephalid fauna is likely to be both
diverse and of special biogeographic interest.
Groundwater amphipods
Late in the 19th century the biological world was intrigued by Chilton’s (1882a,b,
1884, 1894) reports of crustaceans living within aquifers of the Canterbury
Plains. Following this initial work, the groundwater received scant attention.
Subsequent workers, including Chilton himself (e.g. 1912, 1924), apparently
assumed no additional species, assigning specimens to known taxa without
critical examination.
During the 1970s Guillermo Kuschel of the former DSIR surveyed
groundwater faunas by pumping wells throughout the country. Ten new
gastropod mollusc, 71 mite, and two water-beetle species were described from
these collections (Scarsbrook et al. 2003). The several amphipods from Kuschel’s
collections await full investigation, but preliminary work (Fenwick 2000) revealed
several new taxa. Current collecting effort indicates the existence of a further
20–30 species of groundwater amphipods.
The described hypogean (groundwater) amphipod fauna of New Zealand
comprises four species in three endemic genera (two of which have epigean
representatives) each belonging to quite different families. Two of the hypogean
families are endemic. Given the number of species, this fauna seems remarkably
diverse at generic and familial levels. Preliminary work indicates that the New
Zealand hypogean amphipod fauna appears dominated by paraleptamphopids
and is very different to that of Australia, where hadzioids and crangonyctioids
predominate (Bradbury & Williams 1997). Taxonomic work on these collections
is required to determine the true diversity, to determine relationships with the
Australian freshwater amphipod fauna, and to make the fauna accessible to
ecologists.
Should we be surprised by a high diversity of groundwater amphipods in
New Zealand? Groundwater volumes in New Zealand are huge and probably
several times greater than volumes within surface waters (lakes and rivers). For
example, the groundwater of the Golden Bay region is estimated to approximate
the volume of water in Lake Taupo. There are extensive aquifers beneath most of
the Canterbury Plains to depths of 350–550 metres. This is not simply all water,
but variably porous gravels with water moving through interstices. Obviously,
there is a huge volume of water beneath the plains. Other parts of the country
also comprise large plains of porous alluvial gravels (e.g. Waimea Plains around
Nelson, the Heretaunga Plains of Hawke’s Bay) containing extensive aquifer
systems. Given the very large habitable volumes available and the apparent
barriers to dispersion between each groundwater system, a high amphipod
biodiversity should not be unexpected.
Investigations at one site in Canterbury indicate that groundwater amphipods
help to maintain the quality of Canterbury’s groundwater (Fenwick et al. 2004).
The three known amphipod species, as well as a large subterranean isopod
(Phreatoicus typicus), congregate at sites of organic enrichment from sewageoxidation-pond effluent. A series of field and laboratory experiments showed
that these animals browse on non-living organic slime layers from sediment
and stone surfaces (Fenwick 1987). Extrapolation of experimental results using
conservative estimates of crustacean densities indicates that the two dominant
amphipods remove large amounts of organic carbon annually in the vicinity of
the disposal area.
Further understanding of the biology of these valuable groundwater
systems depends on documenting and monitoring their biodiversity to facilitate
CRUSTACEA
Paracrangonyx winterbourni.
From Fenwick 2001
Ringanui toonuiiti.
From Fenwick 2006
153
NEW ZEALAND INVENTORY OF BIODIVERSITY
ecological studies for improved policy formulation and management decisionmaking. Fundamental to this is better taxonomic knowledge of the fauna.
Polycheria obtusa.
From Barnard 1972a
Rakiroa rima.
From Lowry & Fenwick 1982
154
Biogeography of the freshwater fauna
Some New Zealand freshwater amphipods have attracted considerable interest
from workers seeking to untangle phylogenies and relationships between
faunules of Gondwana and other landmasses. Two endemic genera are of special
interest. Phreatogammarus was seen as ‘an amazing antiboreal morphological
counterpart of the Holarctic crangonyctids’ (Barnard & Barnard 1983, p. 51), a
group now largely confined to North America. This genus was considered to
be ‘perhaps the most primitive [living] gammarid’ (ibid., p. 420) that is ‘now a
perfect relict’ (Barnard & Barnard 1982, p. 264). The absence of any significant
amphipod fossils increases the significance of Phreatogammarus to evolutionary
biologists. The morphologies of both Phreatogammarus and Paraleptamphopus, a
modern derivative from a Phreatogammarus-like ancestor (Barnard & Barnard
1983), are incompletely known. Thus it is difficult to establish the relationships
of these two genera with other genera.
Other New Zealand freshwater amphipod genera are also distinctive
and have intriguing faunal relationships. Paracalliope, a genus with three
New Zealand species and Australian, Philippine, New Caledonian, and Fijian
representatives, is calliopiid-like, but sufficiently distinctive to justify placement
in a separate family, the Paracalliopiidae, which has one other genus (Barnard
& Karaman 1982, 1991). The endemic genus Chiltonia, together with the
closely related Afrochiltonia, Austrochiltonia, and Phreatochiltonia, comprise the
subfamily Chiltoniinae from New Zealand, Australia, and South Africa (Barnard
1972b). Yet another endemic genus poses biogeographic and phylogenetic
problems. Bousfield (1977) moved the genus Paracrangonyx into his superfamily
Bogidielloidea, re-assigned it to the Crangonyctoidea (Bousfield 1978), thence
(Bousfield 1982, 1983), along with three other disparate genera, to the family
Paracrangonyctidae within his superfamily Liljeborgioidea. Barnard & Barnard
(1983, p. 52) placed Paracrangonyx among the bogidiellid gammaroids ‘for the
moment’. Following careful analysis, Koenemann and Holsinger (1999) found the
genus to be most closely related to three genera from each of Western Australia,
Madeira, and East Africa. After reviewing these placements and rediagnosing
the genus, Fenwick (2001b) concluded that the relationship of Paracrangonyx
to other genera remains uncertain, but that it belongs within the crangonycoid
cluster and is close to the Paramelitidae, as well as showing relationships to
other genera of Australian hypogean amphipods.
Many of these taxa have not been re-examined since their first collection.
The original specimens of some species are in very poor condition and the
illustrations and descriptions of some are inadequate. Consequently, many older
taxa must be revised before descriptions of new taxa can take place.
Special associations
The ecology of New Zealand amphipods is generally poorly known and few
associations with other invertebrates are reported. Gammaridean amphipod
associations with other crustaceans, ascidians, sponges, hydroids, echinoids,
molluscs, and other organisms elsewhere are well documented (e.g. Vader
1978, 1984, 1996) and some New Zealand amphipods probably live in similar
associations.
The corophioid amphipod Pagurisaea schembrii occurs only on the hermit
crab Paguristes pilosus, where up to 50 at a time live among the dense setae on
the host’s chelipeds, walking legs, and carapace (Moore 1983a). The amphipods
apparently do not steal their host’s food but use their specially modified antennae
to capture food particles from the host’s respiratory current whilst sheltering
within the host’s setae and shell.
Some amphipods are found almost exclusively on algae, but the nature of
PHYLUM ARTHROPODA
the relationships between amphipods and the algae is uncertain. Many species
are found on more than one species of alga, as well as on foliose invertebrates
(hydroids, bryozoans). This suggests that many amphipods use their hosts more
as a substratum than as a partner in some interdependent association. Species
of the tube-building genus Notopoma found at Kaikoura illustrate this apparently
non-obligate relationship. Notopoma fallohidea lives only on the green alga
Caulerpa brownii at relatively sheltered sites (Lowry 1981). One of its congeners,
N. harfoota, is extremely abundant on the same alga in more severe wave action,
but lives on other algae also. A third Kaikoura species, N. stoora, is most abundant
on the foliose bryozoan Costaticella solida, although a few occur on Caulerpa.
Another New Zealand amphipod, Orchomenella aahu, bores into stipes of
the kelp Ecklonia radiata to eat up to 22 milligrams per day of the more palatable
(low phenolic content) internal tissues (Haggitt 1999). These amphipods remain
within the stipe, reproducing several times. Whole families of as many as 300
individuals, comprising several generations, live within most infected plants.
This association seems opportunistic because O. aahu is also an active scavenger
of animal tissue (Lowry & Stoddart 1983b).
The large subantarctic amphipod Rakiroa rima appears to live only within
empty sponge-covered shells of a large barnacle (Megabalanus campbelli)
(Lowry & Fenwick 1982). Similarly, some cryptic species such as Acontiostoma
tuberculata, Ocosingo fenwicki, and Stomacontion spp. are known only from among
collections of subtidal encrusting sponges (Lowry & Stoddart 1983b). It is
uncertain whether these are commensal associations or whether the conditions
sought by the amphipods are found coincidentally in close proximity to these
other organisms. Some have, however, evolved specialised morphological and
reproductive adaptations to their inquilinous life-styles. For example, species
of Ocosingo and Stomacontion have specialised piercing mouthparts (Lowry &
Stoddart 1984). Acontiostoma and some Stomacontion species undergo a sex
change to ease the problems of finding a mate; small sexually mature males
change into reproductive females as they grow larger (Lowry & Stoddart 1983b,
1984, 1986).
The place of some amphipods in various food-webs makes them ideal
intermediate hosts for parasites. The common freshwater amphipod Paracalliope
fluviatilis is the intermediate host for a parasitic nematode (Hedruris spinigera)
commonly found in long-finned and short-finned eels, smelt, brown mudfish
(Hine 1978, 1980; Jellyman 1989), and whitebait (McDowall 1968). Infection rates
of the nematode in these fishes (up to 38% for short-finned and 70% for longfinned eels) are often directly related to abundances of the amphipod and the
incidence of Paracalliope fluviatilis or smelt in the fishes’ diets (McDowall 1968;
Hine 1978). This amphipod is also the intermediate host for three additional
parasites of freshwater fishes – Acanthocephalus galaxii, Coitocaecum anaspides, and
at least one species of hymenolepid cystocercoid (Hine 1978). Similar amphipod–
parasite relationships are almost certain to occur among marine species.
These observations show some of the diverse relationships between amphipods and other organisms. Other relationships, notably those between widely distributed hyperiid amphipods and various other planktonic invertebrates
(salps, tunicates, medusae), plus those between cyamids and their cetacean
hosts, are not considered. Numerous other relationships between New Zealand
caprellid and gammaridean amphipods and various parasites, other invertebrates, and algae are likely to be described in the future.
Alien species
Relatively few invasive amphipods (11 species) have been reported in New
Zealand. Among the hyperiids, the potential for a species to invade seems
extremely low; ships’ ballast water seems the only feasible vector, but the
likelihood of hyperiids surviving within ballast water for any appreciable time
seems remote. Certainly, exotic species may arrive fortuitously as ephemeral
CRUSTACEA
Ocosingo fenwicki (anterior at left,
head hidden by large lateral coxae).
from Lowry & Stoddart 1984
Caprella equilibra.
From McCain 1968
155
NEW ZEALAND INVENTORY OF BIODIVERSITY
Ericthonius pugnax (antennae broken).
From Just 2009
Gammaropsis typica.
From Barnard 1972a
156
transients within water masses not normally entering our region. Such arrivals
seem destined to disappear when their water masses are displaced by the more
usual regime.
One New Zealand caprellid, Caprella mutica, is a very recent invader (Willis
et al. 2009), another species (Caprella equilibra) is cosmopolitan, and a third
(Caprellina longicollis) is widespread in southern waters (McCain 1969, 1979).
Caprellids’ usual association with sessile fouling invertebrates at sites of high
water movement suggests that the latter two caprellids could arrive on the fouled
hulls of ships and, thus, may be invaders. Equally, several additional cyamids may
be found in New Zealand in the future. Whale hosts of several more species are
known from New Zealand waters, but these small, apparently rare, amphipods
are collected infrequently.
One land hopper, Arcitalitrus sylvaticus, has been imported from Australia.
It is established in urban and disturbed habitats of northern New Zealand,
displacing native land hoppers to become the principal land hopper in domestic
gardens in Wellington and Auckland (Duncan 1994). The species has failed to
become established in Christchurch, despite at least two separate introductions
via potted plants.
There is no evidence of any exotic amphipods invading New Zealand’s fresh
waters. A few gammarideans have been introduced to harbours, however, via
ships. Two corophioids, Monocorophium acherusicum and Apocorophium acutum,
are cosmopolitan and ‘trace out some of the major shipping routes, particularly
that from England through the Mediterranean and Suez Canal to South Africa’
(Hurley 1954f), indicating that both are invaders. Ericthonius pugnax, another
tube-building corophioid, is probably another invader because, although its
distribution is less readily explained (New Zealand and Indonesia), the species
was not discovered in New Zealand until 1923, some 70 years after its original
description.
Two additional corophioids have been reported as invaders in New Zealand.
Paracorophium brisbanensis and an unidentified species of Corophium were found
in brackish waters of the upper reaches of Tauranga Harbour. Both were regarded
as adventives because neither was reported from New Zealand previously, they
were not found at any of 92 similar sites surveyed around the country, both
Tauranga populations had ‘remarkably limited genetic variability’, and juveniles
dominated their population structures (Stevens et al. 2002).
Another notable alien amphipod, distributed nearly globally, is the woodboring Chelura terebrans. First found in New Zealand in Auckland Harbour
(Chilton 1919), this small amphipod bores into most human-made wooden
structures around the world (Barnard 1955). Chelura, along with Limnoria isopods
and boring molluscs (Teredo species), wreaks havoc on wharf piles, rapidly boring
into the timber and weakening any wooden structures. Apart from Chilton’s
(1919) original records, there appear to be no other reports of this species from
New Zealand, although it is certain to be more widespread.
Three additional aliens were found in the sea chest (a large recess in a ship’s
hull for seawater intake pipes) of a vessel from the tropical Pacific that was slipped
at Nelson in September 1999. These were Stenothoe gallensis and Elasmopus rapax,
two known tropicopolitan species, and an unidentified species belonging to the
cosmopolitan genus Podocerus. The first two species were abundant and included
mature males, gravid females, and juveniles. There is no information on whether
any of these species has become established in Nelson or elsewhere in New
Zealand, despite repeat surveys.
In general, it seems extremely difficult to determine whether marine species
with wide distributions are invaders (become established on new shores through
dispersal by human activities) or simply arrived by natural dispersal. Several other
New Zealand species have variably wide extrinsic distributions, but the ecologies
of only a few seem likely to equip them for dispersal on the hulls of ships. Tubebuilders and nestlers, especially corophioids, are the most likely candidates. For
PHYLUM ARTHROPODA
example, Gammaropsis crassipes was described from shallow bays and harbours
in eastern Australia in 1881 but not reported from New Zealand until 1920,
suggesting possible introduction. Recent invasions by algae, as well as longterm climatic changes, suggest that the potential for permanent establishment
by amphipod invaders will increase in the future.
Monocorophium sextonae was considered to be a successful New Zealand
invader of European shores (Hurley 1954f), although this has recently been
questioned (Costello 1993; Bousfield & Hoover 1997). First described from
Plymouth and Wembury in 1937, this amphipod was present, albeit unrecognised,
in Chilton’s (1921) material (Hurley 1954f). Crawford (1937) remarked that the
‘abundance of this species is the more surprising since it is not present in the rich
collections of Corophium made from the same dredging grounds in 1895–1911. It
seems possible, therefore, that it is not indigenous at Plymouth … I cannot guess
its original locality’. In revising these species of the family Corophiidae, Bousfield
and Hoover (1997) considered that M. sextonae ‘is almost certainly endemic to
the eastern North Atlantic and Mediterranean regions, from whence it has been
spread by commerce to world-wide temperate marine waters’.
Amphipods in environmental investigations
Diverse approaches are used to assess and manage human impacts on the
aesthetic and life-sustaining qualities of natural environments. Use of plants and
animals as bioindicators is increasingly common because of the sensitivities and
broad-spectrum responses of some species. Amphipods are ideal bioindicators
for shallow marine environments (Conradi et al. 1997) because they are
ecologically (trophically) important, tend to be numerically dominant within
many habitats, have specific niche requirements, have generally low mobility
and dispersive capabilities, and are known to be sensitive to several pollutants
and toxicants. Indeed, Thomas (1993) reported that ‘[a]mphipods are so useful
as bioindicators that U.S. Government agencies now require their identification
to species in permitting operations such as oil leases and outfalls.’ In addition,
individual species of amphipods may serve as very useful assays for pollutants
(Lamberson et al. 1992). Several US agencies employ amphipods in bioassays to
test toxicities and specific contaminant levels independent of chemical analyses
and environmental surveys, particularly for marine environments.
Many of New Zealand’s estuarine and marine amphipods fulfil all of
Thomas’s (1993) criteria for effective biomonitors (e.g. Fenwick 1976, 1985;
Hickey & Martin 1995; Nipper & Roper 1995; Nipper et al. 1998). This is also
true for some terrestrial (e.g. Rainbow et al. 1993) and freshwater species (Hunt
1974). Environmental survey research in New Zealand, however, continues to
look at the total fauna and these investigations follow a trend of identifying and
enumerating taxa to family level only (Somerfield & Clarke 1995) in attempts
to reduce costs by minimising the taxonomic expertise and time required for
identifications. Some workers (Thomas 1993; Conradi et al. 1997) advocated
focusing on the amphipods alone in surveys of shallow marine environments
and, certainly, their identification to species seems worthwhile in such surveys.
There has been no specific examination of the merits of using amphipods alone
for such surveys in New Zealand, and identification tools and knowledge of the
group are inadequately developed for this to become a viable, standard approach
in the short term.
New Zealand estuarine amphipods (Paracorophium excavatum, P. lucasi) have
been used in bioassays of sediment contamination and toxicity (Nipper & Roper
1995). Additional studies (Nipper et al. 1998; De Witt et al. 1999) revealed the
robustness of this assay approach, which is now used extensively. Only recently,
however, has the taxonomy of these two species been resolved (Chapman et al.
2002), illustrating that taxonomic knowledge of New Zealand’s amphipod fauna
is often inadequate for reliable ecological applications.
CRUSTACEA
Puhuruhuru aotearoa.
From Fenwick & Webber 2008
Paracorophium excavatum.
From Barnard 1969
157
NEW ZEALAND INVENTORY OF BIODIVERSITY
Gaps in knowledge and future research
New Zealand’s amphipod fauna is important ecologically on land, in fresh
waters (especially groundwaters), and in marine habitats where species fill vital
roles in food-webs and often provide appreciable direct or indirect economic
benefits. Amphipods also offer considerable potential as bioindicators of
environmental quality in many habitats. Obviously, the potential for ecological
and environmental research using amphipods is huge, even when only the
more urgent or applied issues are considered. Equally, the scope for academic
investigation of amphipods is enormous.
Despite all this, their systematics is very incomplete, hindering attempts
to work with the group. Certainly, the land-hoppers appear well known as a
result of Duncan’s (1994) work, but the beach fleas require equivalent treatment.
Freshwater amphipods require urgent attention in view of our scant knowledge
of this group and the huge environmental pressures on fresh waters. Known
species require extensive redescription and revision to facilitate work on the
>50 new taxa in collections. Several other new species exist in other freshwater
habitats that await collecting.
The marine gammaridean amphipods of shallow and continental-shelf
waters comprise another substantial gap. Collecting has been sparse and the
fauna at no one location is well known. Even the distribution of the algaldwelling species along New Zealand is poorly known, despite Barnard’s (1972a)
work. Amphipod faunas of shallow soft seafloors are very poorly known. A study
in Pegasus Bay showed that 28% of species in the 4–10-metre depth zone are
undescribed (Fenwick 1999). Similarly, less than 30% of the 98 species in a series
of collections off Kaikoura are known and the unknown ones include several
new genera. Also, just 24% (10 of 42 species identified by a leading taxonomist)
of amphipods in another study of New Zealand shelf benthos were known to
science (Probert & Grove 1998).
Amphipod research in New Zealand thus offers considerable scope for
both economically important issues and questions of more academic interest.
However, the present status of the group’s taxonomy hinders the successful
development of this work, as well as discouraging many ecologists from using
amphipods as ideal subjects for environmental and ecological investigations. The
future, therefore, requires not just more taxonomy, but also the development of
interactive guides and keys to overcome these barriers and make the local fauna
accessible to non-specialists. This is particularly true for hypogean and other
freshwater amphipods, given their role in maintaining the quality of groundwaters
and the urgent need for effective management of this economically important
resource in the face of increasing demands and human-induced threats.
Order Isopoda: Slaters, fish lice, and kin
Fish micropredator
Aega monophthalma (Cymothoida).
From Bruce 2009
158
The most diverse range of body plans of all the nine peracaridan orders, if not of
all crustacean orders, is shown by the Isopoda, named, however, for the relative
sameness of limbs (Greek isos, equal, like; podos, foot).
Only one of the isopod suborders, Oniscidea, is familiar to most people.
The oniscideans are commonly called woodlice, slaters, pillbugs, or roly-polies.
However, the order is predominantly marine, being less well-represented in
estuarine and fresh waters. There are fewer common names for the marine
groups but sea-lice, fish doctors, tongue-biters, and sea-centipedes are applied
to some families. No common name, except isopod, applies to all members of
the order.
Life-styles vary. Free-living predators, marine filter-feeders, scavengers in
forest leaf-litter and on the sea floor, and various parasitic forms are represented
in the order. The isopods have succeeded in two unusual habitats besides the
shallow marine environments where most crustaceans are typically found. One
is the land, where woodlice, slaters, and phreatoicideans are most often the sole
PHYLUM ARTHROPODA
CRUSTACEA
crustacean representatives in some habitats, and the other is the deep sea, where
the suborder Asellota has radiated into a variety of bizarre forms.
Although they are often said to be ‘dorsoventrally flattened’ while their
close relatives the amphipods are ‘laterally flattened,’ there are many exceptions;
some are cylindrical, others laterally compressed, and others extraordinarily
ornamented. The smallest isopod adults are c. 1 millimetre long, many are in the
range 4–12 millimetres, and the largest are deep-sea scavengers of the genus
Bathynomus, growing to an astonishing 400 millimetres!
The only sure way to tell an isopod from an amphipod is that isopods lack
strongly chelate first legs and have only one pair of uropods (tail appendages)
and a free second thoracic segment. Character interpretation can be difficult,
however, because uropods vary considerably. They may be flat limbs that lie in
the same plane as the pleotelson, or enclose the pleopodal gills, or have any
of several other forms. Technically, Isopoda are defined as follows: eyes sessile
(not stalked); carapace absent; one pair of maxillipeds; seven pairs of pereopods
(legs), without exopods (an outer branch); abdomen clearly differentiated from
thorax and divided into a pleon of five segments (sometimes some fused) and
pleotelson (fused pleonite six and telson); pleopods 1–5 similar or anterior pair
operculiform, forked; one pair of uropods.
Isopods are of interest to marine biologists because of the important roles
they play in ecosystems, especially on the sea floor. Here, species of many families
are important scavengers of decaying material. Isopods of the family Cirolanidae
are critical in cleaning up decaying dead fish (Bruce 1986a; Brusca et al., 1995;
Keable 1995). Fish-lice of the family Cymothoidae are flesh- and blood-feeders
that attach to the skin of living fishes. Aegids and juvenile gnathiids are bloodsucking micropredators of fishes, and in the tropics gnathiids can be so abundant
that fishes attend cleaning stations where wrasses remove and eat them. Seacentipedes (Idoteidae) feed on algae. The diverse Sphaeromatidae feed on living
and dead material of all sorts. Many isopods are ideal food for many bottomliving fishes such as flounders and skates.
One family of economic significance is the Limnoriidae (gribble). These
are wood-borers, formerly of ships but now only of wooden piles and wharves.
Like timber borers on land, gribble make galleries throughout the timber and
weaken it considerably (Menzies 1957; Cookson 1991). Species of Sphaeroma
(Sphaeromatidae) behave similarly. Another important group, at least to
gardeners, is the terrestrial slaters or woodlice. While most feed innocuously on
decaying leaves and wood they can become so abundant as to attack vegetables
and other garden plants.
Diversity of New Zealand Isopoda
The world’s isopod fauna exceeds 10,000 described species but the actual number
of species is certainly several times this. There are big gaps in knowledge of the
deep sea, the tropics, and some families with small individuals. The New Zealand
fauna totals only 426 living species (and four fossil species) but it appears that
few shallow-water isopod groups are well covered taxonomically. It would not
be surprising if many species of Sphaeromatidae, Cirolanidae, Gnathiidae,
anthuroids, Asellota, and Valvifera remain to be discovered, especially from
shelf depths. Even so, the number of already described species (353) somewhat
exceeds that of South Africa (cf. 275 species in Kensley 1978) but, not surprisingly,
is far fewer than in Australia (1,118 species; Poore 2002, 2005). South African
and Australian isopods have attracted greater taxonomic attention than those
in New Zealand. As is the case for many marine and terrestrial animals, New
Zealand isopods are largely endemic.
The only habitat that is relatively well known is intertidal and subtidal
rocky shores, but even here the Asellota have been largely ignored. Museum
collections from The Snares (partly described by Poore 1981) contain several
undescribed species of small asellotes and more such species could be expected
Brucerolis hurleyi (Sphaeromatidea).
From Storey & Poore 2009
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NEW ZEALAND INVENTORY OF BIODIVERSITY
throughout New Zealand. While the benthos of the New Zealand continental
shelf has been thoroughly sampled, the gear used has not deliberately targeted
small invertebrates, and collections available for study seem not particularly
diverse for isopods. NIWA collections appear from superficial examination to
be far less rich than, for example, those from comparable habitats in Bass Strait
at similar latitudes in Australia. Museum Victoria, Melbourne, houses a benthic
collection that includes c. 250 species of isopods from sediments (Poore unpubl.).
There are even fewer species described from the continental slope. Poore et al.
(1994) recognised 359 species, mostly undescribed, from this habitat off the
southeastern coast of Australia and a similar number could be expected for the
New Zealand slope. Several species from bathyal depths north of New Zealand
were described from collections of the Galathea Expedition but the rest of the
EEZ is virtually unsampled. Another habitat as yet largely unexplored is fresh
water in limestone caves; sphaeromatids are known from this environment near
Nelson, South Island (Sket & Bruce 2004).
Three species of isopod fossils have been recorded from New Zealand
(Grant-Mackie et al. 1996; Hiller 1999; Feldmann & Rust 2006).
Numerous families, 120 at last count in the world fauna, are arranged in a
complex hierarchy within suborders (Martin & Davis 2001). Most of the widely
used suborders are monophyletic groups, but the one that has traditionally
included the most familiar marine species, Flabellifera, is not (Wägele 1989;
Brusca & Wilson 1991; Brandt & Poore 2003). Here, Brandt and Poore’s (2003)
classification is followed and the suborder Flabellifera is superseded by the
three suborders Cymothoida, Limnoriidea, and Sphaeromatidea. Three other
previously recognised suborders are subsumed within Cymothoida – Epicaridea
as superfamilies Bopyroidea and Cryptoniscoidea, Anthuridea as superfamily
Anthuroidea, and Gnathiidea as family Gnathiidae. Hurley and Jansen (1977)
provided an effective key to identify some families but their classification is now
out of date. Modern faunal treatments, also using the older classification, can be
found in Kensley (1978) or Kensley and Schotte (1989). Only 49 families have so
far been recorded from New Zealand.
Joeropsis sp. (Asellota).
From Hurley & Jansen 1977
Suborder Asellota
Some 93 New Zealand species are known, of which 36 remain unnamed or not
fully determined. They have diverse shapes. Diagnostic characters include: coxal
plates usually minute; one (rarely two or three) pleonites free, others fused;
uropods attached posteriorly. Asellotes are common but small, difficult to find, and
even harder to identify. A microscopic examination of tufts of algae from sheltered
marine environments will often reveal several species of asellotes, rarely more
than two millimetres long. Others live in freshwater streams. Globally, almost
30 diverse families exhibit an exceptional range of form on the floor of the deep
sea. Some species are quite bizarre, with extraordinary ornamentation. Several
species from the deep sea near New Zealand were described from collections of
the Danish research ship Galathea (Wolff 1956a, 1962) but only one family from
this environment in New Zealand has been treated in detail (Lincoln 1985). The
identity of many of the species recorded from subantarctic New Zealand may be in
doubt until specimens are compared with those from other islands or continents.
Globally, Wilson and Wägele (1994) listed all known asellote species and provided
a key to the genera of Janiridae, an important shallow-water family, and Cohen
(1998) did the same for Dendrotiidae. The diverse Munnopsididae has been
treated in part by G. D. F. Wilson (1989), the Stenetriidae by Serov and Wilson
(1995), Pseudojaniridae by Serov and Wilson (1999), Joeropsididae by Just (2001),
and Paramunnidae by Just and Wilson (2004, 2006).
Suborder Phreatoicidea
Nine New Zealand species are known, all endemic, and in endemic genera. They
160
PHYLUM ARTHROPODA
CRUSTACEA
are laterally flattened. Other diagnostic characters include: coxal plates extending
ventrally; five pleonites free; uropods rod-like and attached posteriorly. Peculiar
to southern continents and islands, phreatoicids comprise an unusual group of
freshwater and terrestrial species. They superficially resemble amphipods but
differ in having only one pair of uropods as well as other isopod features. Most
of the New Zealand fauna was dealt with by Nicholls (1944), with one species
described in detail by Wilson and Fenwick (1999). The suborder was reviewed by
Wilson and Keable (2001).
Suborder Cymothoida
Comprising sea-lice, fish-lice and other mobile scavengers, predators, and
microparasites, 116 described and 16 undetermined New Zealand species are
known. Diagnostic characters: usually dorsoventrally flattened but otherwise
diverse; mandibular molar blade-like or reduced; coxal plates expanded and
free or reduced; five pleonites free or variously fused; uropods usually forming
tail fan with pleotelson, rotating in horizontal plane and in broad contact with
pleopods. All are marine, but habits and shapes vary. Numerous authors have
contributed to knowledge of cymothoidan families in New Zealand, notably
the Cirolanidae (Jansen 1978; Bruce 1986a, 2003, 2004a; Svavarsson & Bruce
2000; Keable 2006), Cymothoidae (Bruce 1986b), Gnathiidae (Cohen & Poore
1994; Svarvasson 2006), Tridentellidae (Bruce 1988, 2002), and Aegidae (Bruce
1983, 2004b, 2009a). The suborder contains four superfamilies – Anthuroidea,
Bopyroidea, Cryptoniscoidea, and Cymothooidea.
Some 21 described New Zealand species of Anthuroidea are known (in
the families Anthuridae, Expanathuridae, Hyssuridae, Leptanthuridae, and
Paranthuridae). Diagnostic characters include: shape elongate and cylindrical;
coxal plates indistinguishable from pereon wall; pleonites fused or free; uropodal
exopod attached proximally on peduncle and dorsally arched over pleotelson.
Anthuroids live in sediment and on macroalgae, although the New Zealand
species Cruregens fontanus is unusual in living in artesian and river waters (Wägele
1982). Very few species had been described until the work of Wägele (1985). The
family arrangement follows Poore (2001a), who synthesised many papers and
whose earlier work, principally on the Australian fauna, is relevant.
The superfamily Bopyroidea comprises parasitic isopods of crustaceans,
with 13 described New Zealand species in the family Bopyridae. Diagnostic
characters include: individuals sexually dimorphic, females usually
asymmetrical, males minute; mouthparts reduced; branchial parasites of
crabs, shrimps etc., but also of other crustaceans and some hyperparasites of
other bopyroideans. Page (1985) studied New Zealand species. Few modern
taxonomists have tackled this confusing group, but Markham (1985) and other
papers by this author are a good introduction.
The largest superfamily in New Zealand is Cymothooidea, with 93 species
(15 unnamed or not fully determined) in the families Aegidae, Anuropidae,
Cirolanidae (with endemic genus Pseudaega), Cymothoidae, Gnathiidae, and
Tridentellidae. The largest of these, with 37 species, is the recently monographed
Aegidae (Bruce 2009a), a family of micropredators mostly associated with
fishes. The Cryptoniscoidea has just five species in New Zealand, in the families
Crinoniscidae and Hemioniscidae (Hosie 2008).
Suborder Limnoriidea
These are wood-boring isopods, sometimes called gribble, with nine New
Zealand species all in a single family, Limnoriidae, reviewed by Cookson (1991).
Mandibles are specially modified, the body is cylindrical, and pleonites are free.
Wood is not their only target in New Zealand. Limnoria limnorum caused the
1916 failure of the Cook Strait submarine cable when some individuals bored
through the gutta-percha that was around the inner cable core.
Neophreatoicus assimilis (Phreatoicidea)
From Hurley & Jansen 1977
Cruregens fontanus (Cymothoida).
From Hurley & Jansen 1977
Dorsal (upper) and ventral (lower) views
of Athelges lacertosi (Cymothoida), a parasite
of the hermit crab Lophopagurus lacertosus.
From Pike 1961
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NEW ZEALAND INVENTORY OF BIODIVERSITY
Suborder Sphaeromatidea
These comprise marine pillbugs in general, with 81 described New Zealand
species known, including 61 species of Sphaeromatidae. Diagnostic characters:
usually dorsoventrally vaulted, occasionally flattened, sometimes able to
enroll; coxal plates well developed; pleonites variously fused; uropods usually
forming tail fan with pleotelson, rotating in vertical plane and excluded from
branchial cavity. All are marine, but habits and shapes vary. Notable taxonomic
contributions include those on the Sphaeromatidae (Hurley & Jansen 1977)
and the enigmatic, sometime sphaeromatid, genus Paravireia, herein placed as
incertae sedis (Jansen 1973; Brökeland et al. 2001). A sphaeromatid species is host
to a fecampiid flatworm, Kronborgia isopodicola, described from Kaikoura, the
adults of which live in the body cavity of Exosphaeroma obtusum (Blair & Williams
1987; Williams 1988).
Suborder Valvifera
These include the so-called sea-centipedes and other bizarre forms, comprising
25 described New Zealand species. The form of the uropods, as long plates
attached to the side of the abdomen and tightly enclosing all the pleopods,
defines the valviferans. Most are marine, but the three species of Austridotea are
among the few freshwater members of the suborder (Chadderton et al. 2003).
Some forms are ornately decorated. The only family-level reviews are by Poore
and Lew Ton (1990, 1993) and Poore and Bardsley (1992). The family arrangement
follows Poore (2001b).
Pseudarcturella chiltoni (Valvifera)
From Hurley & Jansen 1977
Suborder Oniscidea
These are the land-dwelling woodlice, slaters, and pillbugs, with 72 described
New Zealand species known. Four species are naturally occurring non-endemics
and six others are introduced. Diagnostic characters: usually flattened but
sometimes able to roll up; five pleonites usually free; pleopods highly modified
for air-breathing. Oniscideans are exclusively terrestrial and are the only
crustacean group to compete successfully with other arthropods on land. Seven
pairs of legs immediately reveal that they are not insects or millipedes. There
are examples high up on the seashore but none is truly marine. Although damp
places, and under leaves and decaying logs, are favoured habitats, some overseas
species are known from deserts. Like all isopods, oniscideans rely for respiration
on their pleopods, which are kept damp with a variety of water-conservation
measures. Most species are scavengers on dead plant litter but some can be pests
in gardens. There are numerous families including five genera and many species
endemic to New Zealand. But the most commonly seen species are introduced
from Europe. The New Zealand fauna was reviewed by Hurley (1950) and one
family revised by Green (1971). Some of the names listed by Hurley are now out
of date and the present review follows the taxonomy of Green et al. (2002).
Historical overview of isopod studies
The first scientific collection of isopods in New Zealand was made by the
French biologists J. R. C. Quoy and J. P. Gaimard when the l’Astrolabe, captained
by Dumont d’Urville, visited in 1826. They discovered two shallow-water
sphaeromatids from algae, described 13 years later as Isocladus armatus and
Cassidina typa in a significant publication on isopods by H. Milne Edwards
(1840). Earlier publication dates appear in the New Zealand checklist but these
are of species either introduced to the country or of species described from
elsewhere. Later, the United States Exploring Expedition visited New Zealand
on its 1838–42 round-the-world voyage, and numerous species of marine
animals were described by its chief scientist, James D. Dana. Among these
are 19 species of isopods (Dana 1852b, 1853–55). The first review of the New
Zealand crustacean fauna (Miers 1876) listed 28 isopod species in 16 genera.
When a second review was completed 10 years later by Thomson and Chilton
162
PHYLUM ARTHROPODA
CRUSTACEA
(1886), 60 species of isopods had by then been described, many by these two
authors. A third checklist and key (Hurley 1961) listed 168 species; the increase
in the intervening years was contributed largely by results from foreign deepsea expeditions like the British HMS Challenger (1873–76) and Danish Galathea
(1952). By 2009 the number had grown again, largely as a result of the work
of New Zealand-based taxonomists Desmond Hurley and Peter Jansen in the
1970s and Niel Bruce in the 2000s, as well as overseas workers with an interest
in specific families (J. Just, R. Lincoln, G. C. B. Poore, and J.-W. Wägele).
Special features of the New Zealand isopod fauna
Some 38 isopod families have marine representatives in the New Zealand
fauna. Gaps can be explained by inadequate collecting. For example, it is safe
to say that most deep-water asellote families will be recorded once appropriate
sampling is done. It is possible that the Ancinidae and Corallanidae might one
day be found in New Zealand. Four small families from the southwestern Pacific
(Bathynataliidae, Hadromastaciidae, Keuphyliidae, and Phoratopodidae) are
so far not recorded from New Zealand. The Serolidae, rich in species in shelf
environments in Australia (Harrison & Poore 1984; Poore 1985, 1987), the
southwestern Pacific (Bruce 2009b), and Antarctica (Brandt 1988; Wägele 1994),
is represented in New Zealand by only a relatively small number of deep-water
species, several of which have been described (Bruce 2008; Storey & Poore 2009).
The Gondwanan affinities of the fauna are evident in the largest families,
Sphaeromatidae and Cirolanidae, where genera found in other Gondwanan
continents dominate. This is clear too in Plakarthriidae, a family known only from
three species, one each in South America, New Zealand, and southern Australia
(Poore & Brandt 2001). The same is true for the terrestrial families, with many
Palaearctic oniscidean families absent and strong radiation of southern ones. The
Phreatoicidea is a typical high-level Gondwanan taxon, being confined to New
Zealand, Australia, and India.
New Zealand isopods are largely endemic – 100% of freshwater species,
86% of terrestrial species, and almost 77% of marine species. The endemicity of
some taxa reflects the long isolation of the fauna from Australia, the continent
from which it last separated 85 million years ago. Close relatives (perhaps
sister species) of New Zealand species are found in Australia within several
families, e.g. Austrarcturellidae, Idoteidae, Leptanthuridae, Phreatoicoidae,
Plakarthriidae, and Sphaeromatidae. Much less is known about relationships
among other apparent endemics. Many species from the shelf and deep sea are
known only from type specimens from a single sample, so their true distribution
is unknown. But even here evidence is emerging that endemism is truly high.
For example, none of the anthurideans or haploniscid and dendrotiid asellotes
described from New Zealand occurs in Australia (Cohen 1998; unpublished
material and catalogues).
Non-endemic species fall into two groups – those apparently naturally
widespread, and those thought to be introduced. The idoteids Batedotea elongata
and Paridotea ungulata have been identified from algal communities in New
Zealand and Tasmania and another, Idotea metallica, is cosmopolitan on oceanic
algal wrack (Poore & Lew Ton 1993). Several other species may occur naturally
in New Zealand and Australia and sometimes also elsewhere, e.g. Natatolana
pellucida (Cirolanidae), Limnoria rugosissima, L. tripunctata (Limnoriidae), and
Cymodoce convexa (Sphaeromatidae). Several species of aegid micropredators of
fishes and at least three species of cymothoid fish ectoparasites seem widespread
in the Tasman Sea (and sometimes beyond), as are their host species. A deep-sea
gnathiid, Bathygnathia vollenhovia, which occurs on both sides of the Tasman Sea
(Cohen & Poore 1994), is certainly naturally distributed. For other seemingly
widespread species, identifications are suspect until type material has been
compared. Specimens of the New Zealand sphaeromatid Pseudosphaeroma
campbellense identified from Australia (Harrison 1984) may be specifically
Paridotea ungulata (Valvifera).
From Hurley & Jansen 1977
163
NEW ZEALAND INVENTORY OF BIODIVERSITY
different (Poore 1994; Bruce & Wetzer 2008). This suspicion is especially valid
for some species recorded from the New Zealand subantarctic but whose type
locality is elsewhere, e.g. the sphaeromatids Exosphaeroma gigas and Cymodocella
tubicauda (Hurley & Jansen 1977; Brandt & Wägele 1989).
The most familiar isopods of gardens and farmland, the woodlice and
pillbugs, are definite imports from Britain or continental Europe, namely
Armadillidium vulgare, Porcellionides pruinosus, and Porcellio scaber. They arrived
with garden plants or simply as stowaways with the first Europeans. An export
of a slater has occurred, too – the styloniscid Styloniscus otakensis to Australia’s
Macquarie Island (van Klinken & Green 1992).
Alien marine isopods
For marine isopods the presence in New Zealand of exotics is ambivalent,
although the ability to be transported to and from New Zealand with fouling
on ships is certain. Cranfield et al. (1998) recorded three isopods as potentially
introduced to New Zealand. The first, Australian species Cymodoce tuberculata
(Sphaeromatidae), recorded by Chilton (1911b) from a plank of the ship Terra
Nova in Lyttelton, seems not to have become established in New Zealand. The
second, a species of wood-boring gribble, Limnoria tripunctata (Limnoriidae),
has potentially been distributed by shipping between widespread localities
around the world but its origin is unknown (Cookson 1991). The third, Limnoria
rugosissima, is a borer of algal holdfasts, not of timber, so is more likely to be
distributed between southern Australia and New Zealand by drifting kelp. On
the other hand, Limnoria quadripunctata (not listed by Cranfield et al. 1998)
was first described from Europe and now globally recognised; its origin is more
probably Southern than Northern Hemisphere (Cookson 1989; Poore & Storey
1999). Likewise, Sphaeroma quoianum (Sphaeromatidae), another wood-borer
and its commensal, Iais californica (Janiridae), could have been distributed
similarly. Eurylana arcuata (Cirolanidae) is possibly a New Zealand species
introduced to Australia (or vice versa) and to North America (Bowman et al.
1981).
The affinities of the New Zealand fauna can only be understood if the
taxonomy is accurate. Two species of Phalloniscus (Oniscidae) erroneously
recorded from Australia, P. kenepurensis and P. punctatus, were excluded by
Bowley (1935) and Green (1961). Deto marina (Scyphacidae), recorded from New
Zealand by Schultz (1972), is endemic to Australia.
Order Tanaidacea: Tanaids
Apseudes larseni.
From Knight & Heard 2006
164
Tanaids (there is no common name) are very small, shrimp-like creatures.
They are mostly in the 2–5 millimetre range but adults of a few species can
be as small as half a millimetre or as long as 75 millimetres (Gamo 1984).
There are three living orders, the members of which exhibit characteristic
morphologies and, to some extent, lifestyles. Species of Neotanaidomorpha
are free-living surface dwellers, while those of Tanaidomorpha are largely tube
dwellers and the Apseudomorpha are mostly burrowers or crawlers. The first
two segments of the thorax are covered by a carapace forming, with the head, a
cephalothorax. The first thoracic segment supports a small pair of maxillipeds,
the second a distinctive pair of chelipeds, and each of the third to seventh
segments bears a pair of pereopods. The first pereopod may be adapted for
burrowing in the suborder Apseudomorpha, equipped with spinning glands
for tube construction in the suborder Tanaidomorpha, or may be a simple
‘walking leg’ in the suborder Neotanaidomorpha. Sexual dimorphism is often
evidenced in the chelipeds and the claw of the left cheliped can be greatly
enlarged in the males of some species of Apseudomorpha. Each of the first
five abdominal segments normally carries pleopods but these may be absent
in many deep-sea species. The final pleonal segment is fused with the telson
PHYLUM ARTHROPODA
(forming a pleotelson) and carries a pair of uropods. Respiration takes place
over the inner surface of the carapace.
As with other peracarid crustaceans such as isopods, amphipods, and
cumaceans, tanaids carry their fertilised eggs and mancae (post-larval juveniles)
within a ventral marsupium. In most groups this is formed out of four pairs of
oostegites, attached to the first four pairs of pereopods. This is not the case in
the Tanaidae, examples of which that are common in intertidal habitats; in this
family the marsupium is seen as a ventral pair of elongate sacs (or sometimes
just one sac). Similarly, species of Pseudotanaidae, common in the deep-sea,
have only a single pair of oostegites arising from the fourth pair of pereopods.
There is also some evidence to show that in some burrowing-tubicolous groups
(such as the Typhlotanaidae) the female constructs a mucous brood pouch in
which she and her young live (G. Bird unpubl.).
Tanaids are usually detritivores or grazers but some taxa are filter-feeders
and opportunistic predation on smaller invertebrates (such as foraminiferans or
juvenile echinoderms) may be common. Only a few species are considered to
be parasitic but none are obligate parasites. Tanaids are preyed upon by a large
number of other organisms including polychaetes, other crustaceans, migratory
birds, and a large number of juvenile and adult fish such various rat-tails and
grenadiers in the deep sea (Bird unpubl.)
Identification of tanaids is notoriously difficult, complicated by their small size
and sexual and developmental variation (Larsen 2005) along with widespread
and intense convergent evolution. So far, 25 families, more than 200 genera, and
more than 1000 species have been described, but it is estimated that the order
contains several thousand undescribed species, most of which are suspected to
live in the deep sea. Tanaids live almost exclusively in marine or brackish habitats,
with just a few species in fresh water. They occupy a wide range of depths. Marine
species can be found intertidally among coralline algae, crevices, holdfasts, and
in rock-pools. Shallow-water and shelf forms can be found in sand and mud,
although tanaid sand-faunas are typically sparse. Tanaids are very common and
species-rich in deep-sea oozes and some live in deep-ocean trenches to hadal
depths exceeding 9000 metres (Kudinova-Pasternak 1972).
Apart from those species that are attached to floating objects, all tanaids are
benthic, but some have short-lived males that can be found swimming above
the seafloor in their search for females. Tanaids are free-living, tube-dwelling,
burrowing, or live in association with other organisms in a variety of relationships.
Some live as epifauna on solitary corals (Sieg & Zibrowius 1988), colonial corals
and hydroids (Bacescu 1981), live scallops (Brown & Beckman 1992), oysters
(Bamber 1990), barnacles (Reimer 1975), and even sea turtles (Caine 1986). Some
species are true symbionts, living together with gastropods (Howard 1952),
tube-dwelling sea cucumbers (Larsen 2005), in the canals of sponges (Hassack
& Holdich 1987), and as cleaning commensals on mobile bryozoan colonies
(Thurston et al. 1987). Tanaids may also have their own epifaunal associates such
as stalked protozoans (Gardiner 1975) or bivalves (Warén & Carrozza 1994) and
deep-sea species can carry foraminiferans embedded in the cuticle. They may be
parasitised internally by nematodes and externally by copepod-like tantulocarids
(Larsen 2005).
The New Zealand fauna is so poorly known that even an approximate
assessment is difficult but, if comparison is made with a similar area and range
of habitats, based on the Rockall-Biscay region of the Northeast Atlantic (G. Bird
unpubl.), then 250–300 species are possible. The cryptic habits of the group and
the small number of active specialists globally and in New Zealand suggest that
this state of affairs may continue for some time although progress is now being
made. Knowledge of the New Zealand fauna is still largely based on the older
published records of Chilton (1882c, 1883), Thomson (1880, 1913), Stephensen
(1927), Wolff (1956b), and Lang (1968). As a consequence, there are only about
20 authoritative records among the species in the end-chapter checklist. The
CRUSTACEA
Sinelobus stanfordi.
From Chapman & Lewis 1976
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NEW ZEALAND INVENTORY OF BIODIVERSITY
remainder are unpublished records or undescribed species based on studies by
Graham Bird, Elizabeth Hassack and the late Jürgen Sieg. Amongst these records
are a number of undescribed species (indicated in the end-chapter checklist by
bracketed numbers) and several new genera, the family affiliation of which is not
currently available. This list is a snap-shot view and highly provisional. A few old
records have been reappraised in the light of current tanaid taxonomy (Larsen
& Wilson 1998, 2002; Knight & Heard 2006; Bird 2008). The New Zealand fauna
also contains one of the few known freshwater tanaids – Sinelobus stanfordi from
lakes in the Rotorua district.
Order Cumacea: Comma shrimps
The common name for cumaceans alludes to one of their distinctive features,
i.e. resemblance to a comma when preserved. That is, they have an enlarged
front section (head and part of the thorax) followed by a rather narrow posterior
section (remainder of thorax and abdomen).
Comma shrimps live on the seafloor with their bodies generally slightly
submerged in the sediment. They feed on diatoms, pieces of seaweed,
foraminiferans, and detritus, which they collect from the sediment surface. For
the most part, they will stay hidden in the sediment during the day, and some
will make extended trips into the overlying water after sunset. The reasons for
these excursions are not precisely known, but include moulting and searching for
mates. In fact, in some cumacean families, the body morphology of the mature
male is completely modified for swimming, suggesting that at that stage the
animal rarely visits the sediment. Swimming cumaceans are vulnerable to fish
predation, and mature males are commonly found in fish stomachs.
The cumacean body is one of the more modified of the higher crustaceans.
Anteriorly, the head and three segments of the thorax are covered with a carapace.
As a result, the normal feeding appendages of the head are augmented by three
thoracic appendages (known as maxillipeds) that are also used for feeding. The
first of these is also highly modified for respiration. That is, the epipod, which
is not present in amphipods and is reduced in isopods, is greatly enlarged in
cumaceans as a branchial lobe. Respiration occurs as the branchial lobe is moved
back and forth underneath the sides of the carapace.
The remaining thoracic segments bear appendages that function as walking
legs. In some cases, especially in mature males, these legs will also have an
outer branch, the exopod, that is used to aid in swimming. The abdomen is
generally long and thin. Abdominal appendages are either pleopods, if they
occur on one or more of the first five segments, and uropods when present on
the last segment. Pleopods are not present in the females of species that occur
in New Zealand, and may or may not be present on some or all segments in
the males. A final, post-abdominal segment, the telson, may be present as a
separate structure, or it may be fused to the last abdominal segment.
Cumaceans are rare in the fossil record. There are two species known from
the Jurassic, but they are more or less similar to a modern cumacean family,
suggesting that the group as a whole is quite old. On the other hand, cumaceans
are among the last of their line to have evolved, so it possible that all peracarids
were present by the end of the Paleozoic.
As with other members of the superorder Peracarida, cumaceans carry their
young in a brood pouch, with the young hatchling looking like a miniature
version of the adult minus the last pair of thoracic legs. Because of this direct
development, cumacean species are generally not very widespread, and some
genera are restricted to individual continents or ocean basins. Some families,
such as the Bodotriidae and Nannastacidae, are primarily warm-temperate to
tropical, while others such as the Lampropidae and Diastylidae are most diverse
in colder oceans. All families are represented in the deep sea, but lampropids
show the greatest diversity in that environment.
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PHYLUM ARTHROPODA
CRUSTACEA
New Zealand Cumacea
The first cumaceans known from New Zealand were described by George
Thomson (1892), who had spent a couple of days dredging in the Bay of Islands
in 1883. Not being able to sort the material for some time, his two species went
undiscovered for several years. It would be another decade before Zimmer (1902)
would describe an additional two species, collected by Prof. Dr Thilenius from
the Bay of Plenty and deposited in the Berlin Museum. The biggest contribution,
to this day, of our knowledge of New Zealand cumaceans was made by W. T.
Calman, who, over a 10-year period (Calman 1907, 1908, 1911, 1917), described
17 species from material sent to him by G. M. Thomson and Henry Suter. Norman
Jones, a prolific cumacean worker, described a new species and added a new
record from the Chatham Islands area (Jones 1960). He added five new species
and two new records to the New Zealand fauna in his now classic monograph
covering material in the collections of the former New Zealand Oceanographic
Institute (now part of NIWA), the Zoology Departments of Auckland and
Canterbury Universities, and the then Dominion Museum, Wellington (Jones
1963). A further eight deep-water species were described by Jones (1969) from
material collected in the Tasman Sea by the Galathea Expedition.
Over the intervening 31 years, many samples containing cumaceans have been
taken in the waters of New Zealand’s EEZ and stored in the NIWA Invertebrate
Collection, Wellington. Until this present review, no one had taken the challenge
of working up this material. Most of the new material examined was collected in
the deep waters of the New Zealand microcontinent and contains much that is
new, both at species and genus levels. From these collections, four new species of
Gynodiastylidae were found and described in a recent monograph of the family
by Gerken (2001). Several other new taxa have been sorted from the collections
and will be described in future papers.
Of the eight currently recognised cumacean families, only six are represented
in New Zealand waters. (The Ceratocumatidae is known only from abyssal
depths in the Atlantic and Indian Oceans and the Pseudocumatidae are so
far exclusively Eurasian–Atlantic in distribution.) The Gynodiastylidae is the
smallest of the families represented in New Zealand, with only seven species,
and the Diastylidae the largest, with 19 species formally known (and at least
another six species remain to be characterised). Some remarks are now offered
for each family, based on historical records as well as new findings from NIWA
material.
Family Bodotriidae: Subfamily Bodotriinae. Members of this subfamily occur
in all oceans, primarily in shallow water, but also in the deep sea. New Zealand
is quite unusual in having only one (Cyclaspis) of the 13 genera represented in
its fauna. This is most likely because the other genera are primarily warm-water
and have invaded temperate waters only at the edges of their distributions.
Because of the long isolation of the New Zealand microcontinent, temperatewater invasion would have been difficult. On the other hand, Cyclaspis is
found in tropical to cool-temperate shelf waters as well as the cold waters of
the deep sea, so its radiation in New Zealand waters might be expected. The
level of endemicity is high in absolute numbers, but species in this genus are
usually found in one, maybe two, zoogeographic provinces. Few new species are
likely to be found in shelf waters, with most additions to the fauna coming from
bathyal depths. If another genus is to be added, it will most likely be something
completely new.
Family Bodotriidae: Subfamily Vaunthompsoniinae. This subfamily is largely
austral in its distribution and is found from tropical-shelf habitats to cold bathyal
waters. Only one New Zealand shelf species is known, and it is not endemic.
One of the two bathyal species is endemic, as are both abyssal species. It is
unlikely that more than one or two additional shelf species will be found, but
the deep-water fauna could continue to contribute new genera and species.
Family Diastylidae. Of the seven genera represented, one (Colurostylis) is
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NEW ZEALAND INVENTORY OF BIODIVERSITY
Some New Zealand representatives of cumacean families.
Bodotriidae: A (female), B (male), Cyclaspis elegans; C (female), D (male), Cyclaspis thompsoni.
Diastylidae: E (female), Diastylis acuminata (Diastylidae); F (female), G (male), Colurostylis pseudocuma.
Gynodiastylidae: H (female), Gynodiastylis milleri. Lampropidae: I (female), Hemilamprops pellucida.
Leuconidae: J (female), K (male), Paraleucon suteri. Nannastacidae: L (female), Campylaspis rex; M (male), Nannastacus pilgrimi.
A–K, M, from Jones 1960; L, from Gerken & Ryder 2002
168
PHYLUM ARTHROPODA
CRUSTACEA
endemic. The others are broadly distributed in the colder waters of the world
ocean. The genera Makrokylindrus and Vemakylindrus are exclusively bathyal or
deeper. Specific endemicity is very high (18 of 19 known species) for this family
considering the widespread nature of the genera. In addition, diastylids are
very abundant and at least one or two individuals can be found at any benthic
sampling station.
Family Gynodiastylidae. This is a predominantly southern hemisphere family
(but ranges as far west as the Persian Gulf and east to Japan) and exhibits its
greatest radiation in southern Australia. There are seven endemic species in New
Zealand shallow waters, of which three are in the widespread genus Gynodiastylis.
One of the new species, in the genus Allodiastylis, was found at bathyal depths.
Family Lampropidae. The lampropids are a worldwide, cold-water, primarily
deep-sea group. The taxonomy of the family is in need of serious revision, so some
of the species found in the current study may be assigned to new endemic genera
when revision is completed. Prior to this study only one lampropid, Hemilamprops
pellucidus, was known from New Zealand. It is a widely distributed southern
hemisphere species. Bathyal waters, however, have so far produced eight new
species and one new genus (Gerken 2010), suggesting that the Chatham Rise
and Campbell Plateau have much higher-than-average lampropid diversity.
Family Leuconidae. This family has very high generic endemicity (three of six
genera) in New Zealand, especially in shelf waters. Further, the endemic genera
are morphologically advanced within the family, anchoring a group (clade)
where the male second antenna becomes reduced in length and modified so
it can be used to grasp the female during mating. This trend continues in other
eastern Pacific genera, with the second antenna possessing a more complete
grasping structure in one Japanese genus and finally culminating in a western
North American slope-dwelling genus where the grasping structure is all that
is left of the appendage. All species of leuconids are endemic, with the single
exception of Eudorella truncatula, which is surely an introduced species, broadly
distributed in the North Atlantic and North Pacific. This family does not seem to
be well represented in New Zealand bathyal samples, in contrast to what is seen
in northern hemisphere waters.
Family Nannastacidae. There are two groups of genera in this family in New
Zealand – deposit-feeding Cumella and its relatives and carnivorous Campylaspis
and its relatives. Of the deposit-feeders, only one genus, Scherocumella, has been
found in shallow waters, and two genera were found in the bathyal samples. This
group seems to be under-represented in New Zealand. In contrast, there are at
least six species of the carnivorous genus Campylaspis and two of Procampylaspis.
The radiation within these genera is typical of that seen in other shelf and
slope cold-water environments in both northern and southern hemispheres.
All species in this family are endemic. The finding of a species of Styloptocuma
extends the range of this genus into the Pacific.
In summary, there are two groups of cumaceans in the New Zealand fauna –
the highly endemic species and genera of shallow water and the continental shelf,
and the bathyal and abyssal species that belong to genera and families that are
widespread throughout the cold deep waters of the world. Notably, within one
family, the Leuconidae, there has developed a specialised morphology among the
males that seems to have spread northwards in the eastern Pacific, culminating
in advanced forms in Japan. Finally, New Zealand lacks representatives of many
warm-temperate genera, even though it has a warm-temperate zoogeographic
province and the Kermadec Islands within its EEZ. This may be a consequence
of the geological history of the microcontinent, which, after it became isolated,
went through a cooling period, thus eliminating resident warm-water species.
Gaps in knowledge of New Zealand Cumacea
The cumacean fauna of New Zealand’s EEZ currently comprises 31 genera (two
not yet named) and 74 species, not all formally named. Of these, about half, i.e.
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NEW ZEALAND INVENTORY OF BIODIVERSITY
15 genera and 37 species, are from shelf waters. In 1999, a brief collection by Les
Watling in a few areas of the North and South Islands produced one new species
of Colurostylis. Additional collecting is probably not likely to result in the addition
of more than 10 new species from shelf depths, with the possible exception of
Stewart Island and the subantarctic islands, which so far remain unexplored with
respect to cumaceans. The relatively few samples (ca. 15) obtained by Watling
have so far yielded 31 new species and two new genera, with the Diastylidae
still to be studied in detail. None of the species in the new NIWA and Watling
samples can be matched to the eight species Jones (1969) described from the
Tasman Sea, suggesting either that there is a high level of endemicity between
the east and west deep waters of New Zealand or that the deep-water fauna is
very diverse. Neither of these hypotheses is unlikely. Because they brood their
young, cumacean species are highly restricted to zoogeographic provinces in
shallow water, and may well be restricted to individual tectonic plates in deep
water. Since cumacean diversity is generally highest in the Southwestern Pacific,
one might expect the overall diversity of bathyal waters to be much higher, at
least by a factor of two, than that which has been observed to date. In addition,
the lack of correspondence between the shallow New Zealand and southern
Australian faunas lends credence to the fact that there is little natural waterborne transport of cumaceans. Most likely the shelf-dwelling cumaceans of
New Zealand evolved in situ from whatever stock was present after Zealandia
(the New Zealand continental mass) separated from Antarctica about 56 million
years ago.
Order Euphausiacea: Krill
Stylocheiron abbreviatum.
After Sars 1885
170
We’ve all heard of ‘krill’, shrimp-like crustaceans congregating in vast swarms
in cooler latitudes of both hemispheres, and famous as whale food. The term
krill was originally used by Norwegian whalers for the northern hemisphere
cold-water euphausiids Meganyctiphanes norvegica and Thysanoessa inermis
(Mauchline & Fisher 1969) but is now applied to all species of the order
Euphausiacea. ‘Euphausiids’ is itself an unusual word because the ending ‘-ids’
is commonly reserved for family names, not orders. But all except one species of
Euphausiacea belong in just one family, the Euphausiidae and, based on longterm use, ‘euphausiids’ is here to stay. The Euphausiidae contains 85 species and
the Bentheuphausiidae one species.
The Euphausiacea is notable among the crustacean orders because all the
species have conceivably been described. One or two new species may yet be
discovered, but only eight have been added in the last 50 years, two in the last 30,
with the very deep-water Thysanopoda minyops Brinton, 1987, the most recent.
However, in some species, particularly in the genus Stylocheiron, up to six distinct
‘forms’ are recognised (Brinton et al. 1999). A few species such as Euphausia
similis and E. similis var. armata are also extremely similar. In some cases these
forms and species are geographically separate and in others overlapping. It is
unclear what the taxonomic significance of the forms is, but new taxonomic
techniques such as gene-sequence analysis may resolve this problem. If so, it
seems likely that any future changes in the number of euphausiid species are
more likely to result from redefinition of current taxa than from new discoveries.
There is a further, informal subdivision of the family Euphausiidae, with Brinton
et al. (1999) listing several ‘species groups’ within five of the larger genera based
on morphological similarity. The 19 species found in New Zealand waters are
named in one or another of these groups.
Krill are of great importance in the marine economy because of their vast
numbers. They constitute a major proportion of oceanic biomass, are major
grazers of phytoplankton and consumers of small zooplankton, and are
themselves essential in the diets of whales, fish, seals, seabirds, and even people.
PHYLUM ARTHROPODA
Morphology and distinguishing characters of krill
Krill are rather uniform in appearance and easily distinguished from other
crustaceans. Their morphology is well illustrated and described in several
publications, including Baker et al. (1990), who gave a particularly clear overview
of their structure, and Brinton et al. (1999). Only the more distinctive characters
are described here. Typical of shrimp-like crustaceans, krill are adapted to a
natant (swimming) life-style, having an elongate body with the cephalothorax
covered by a carapace, a six-segmented abdomen, and a telson with uropods
that form a tail fan. They also have moveable eyes, biramous first and uniramous
second antennae, and, behind the mandibles, two pairs of maxillae. There are
eight pairs of thoracic limbs. Each has a two-segmented outer exopod and a fivesegmented inner ‘leg’ but the posteriormost pair of limbs (eighth pair) is reduced
to lobes in all but Bentheuphausia amblyops. The form of the seventh pair of limbs
also varies between genera. While the first pair of limbs is used in the manner
of maxillipeds they are similar in form to those behind. Abdominal segments
1–5 bear a pair of pleopods, the first pair in males being modified to form a
handlike copulatory organ (petasma). This is used to transfer sperm packages
to a midventral female structure (thelycum). The petasma and thelycum are
diagnostic of species although they can be difficult to examine and other, more
accessible, structures are generally used for this purpose if they are present and
undamaged. Of particular use in this respect are the proximal three segments
of the antennule (the antennular peduncle), which may bear a lappet having
a characteristic shape or number of spines. The peduncle is usually present in
collected specimens and used in combination with other characters.
Krill are easily distinguished from other shrimp-like crustaceans in having
the gills exposed below the edges of the carapace, rather than covered by it.
Euphausiid gills stem laterally from the first (coxal) segment of the thoracic limbs
and become larger, more branched, and more obvious posteriorly.
A second distinctive character is the presence of movable light organs called
photophores (the name Euphausiidae indicates they emit ‘true light’), which
are distributed in the same pattern throughout the order. Only the two very
deep-water species Bentheuphausia amblyops and Thysanopoda minyops lack
photophores; all others have a photophore on the carapace beneath each eyestalk
and two pairs ventrally on the thorax, adjacent to the second and seventh limbs.
Most also have four single photophores ventrally on abdominal segments 1–4,
but in species of Stylocheiron only one abdominal photophore is present, on the
first segment.
The cuticle is thin, flexible, and mostly smooth, with a small spine behind the
eye and one or two pairs of denticles (tiny spines) on the sides of the carapace
in some species. The front is rounded or produced into a simple sharp rostrum
that is small in comparison to many other shrimp-like crustaceans. Some species
have a keel behind the rostrum, there may be low-profile dorsal spines and
keels on the third to sixth abdominal segments, and, in a few species, some
characteristic sculpturing of the abdominal pleura (side-plates). Krill otherwise
lack the variety of rostra, spines, and keels found in many decapod shrimps and
mysidaceans but they still have rather unusual, distinguishing characters.
Two groups of the Euphausiidae can be distinguished by the shape of the
eyes, which are round or almost so in one group and divided by a constriction
into upper and lower lobes in the other (Baker et al. 1990). The genera Euphausia,
Meganyctiphanes, Nyctiphanes, Pseudeuphausia, and Thysanopoda have round eyes,
while Nematobrachion, Nematosceles, and Stylocheiron have bilobed eyes. One
genus, Thysanoessa, has a mixture of both eye types. There is also a consistent
relationship between eye shape and the form of the thoracic limbs (Baker et al.
1990) – species with bilobed eyes have one or two pairs of thoracic limbs greatly
elongated while round-eyed species do not. Stylocheiron eyes are the oddest
of all – four New Zealand species have eyes with enlarged crystalline cones
making them tube- or pear-shaped. While lacking obvious cones, the eyes of S.
CRUSTACEA
Thysanopoda acutifrons.
From Holt & Tatersall 1906
171
NEW ZEALAND INVENTORY OF BIODIVERSITY
abbreviatum are also pear-shaped and those of S. maximum dumbbell-shaped.
Fully grown krill range in length from < 10 millimetres (e.g. Stylocheiron
affine) to the largest, Thysanopoda spinicaudata, which reaches 150 millimetres
(Brinton et al. 1999). In New Zealand, the smallest is probably S. suhmi at 6–7
millimetres; the largest so far recorded is Thysanopoda cornuta, which can reach
120 millimetres.
Classification
Martin and Davis (2001) placed the order Euphausiacea, with the Decapoda and
Amphionidacea, in the superorder Eucarida, well separated from the Mysidacea
and other orders of the Peracarida. Brinton et al. (1999) recorded earlier
recognition of the similarities between krill and the pelagic decapod shrimps of
the Sergestidae (suborder Dendrobranchiata). Krill and sergestid shrimps have
free-swimming nauplius larvae, metamorphose to the post-naupliar larval stage,
have reduced posterior thoracic limbs, and have a petasma in the male and
thelycum in the female. However, Brinton (1966) had suggested these similarities
might reflect parallel evolution rather than a close relationship. A recent analysis
of ribosomal DNA sequences in krill (Jarman et al. 2000) indicates that they may
be more closely related to the Mysida than to the Sergestidae, which accords
with Brinton’s suggestion.
Thysanoessa gregaria.
After Sars 1885
172
Discovery and diversity of New Zealand krill
Most krill are oceanic in distribution, with consequent low endemicity, and no
species is confined to the New Zealand region, so the history of studies of species
recorded in the region is mostly international. The first species recorded from the
New Zealand region were those collected by the 1873–76 Challenger Expedition
(Sars 1883, 1885). Sars’s reports included 12 of the 21 species now known from
New Zealand waters (see end-chapter checklist). H. J. Hansen (1905a,b–1911)
described many species in several papers published in the early 20th century,
including five species that occur in the New Zealand EEZ.
Tattersall (1924) provided the first list of seven New Zealand krill species
gleaned from the reports of Sars (1883), Thomson (1900), and Hansen (1911)
and added six more collected by the Terra Nova Expedition of 1910. Soon after,
Chilton (1926) listed them again but included two species that Tattersall had
reported, although rather unclearly, as occurring only in Australian waters
(Pseudeuphausia latifrons and Euphausia tenera). Neither has been recorded from
New Zealand since, meaning Chilton’s (1926) list more accurately gives 13 New
Zealand species. The remaining 12 recorded species have resulted from surveys
of pelagic faunas and plankton off New Zealand’s coasts (Roberts 1972; Bradford
1972; Bartle 1976; Robertson et al. 1978). The work of Bartle (1976) focused on
krill in Cook Strait and is the most extensive study of the New Zealand fauna to
date. Four new records are included in the current checklist from collections held
at the Museum of New Zealand.
The only identification guide to krill that includes the New Zealand
region was produced by Kirkwood (1982), but, apart from the early works
listing New Zealand species and referred to above, no taxonomic works on
krill in New Zealand waters have appeared. Sheard (1953) reported in detail
on the taxonomy, distribution, and development of the Euphausiacea with
particular emphasis on the Australasian species Nyctiphanes australis. A
number of recent papers have reported on aspects of the biology of N. australis
in southern New Zealand waters and/or included useful distributional and
biological observations (e.g. Bary 1956, 1959; Jillett 1971; Bradford 1972; Dalley
& McClatchie 1989; McClatchie et al. 1989, 1990, 1991a,b; Murdoch 1989;
O’Driscoll 1998a,b; O’Driscoll & McClatchie 1998).
Research on krill biology continues around the world, especially on species
of economic importance such as Euphausia superba, but the review of Mauchline
and Fisher (1969) remains the major source of information. These authors brought
PHYLUM ARTHROPODA
CRUSTACEA
together a large and disparate literature on all aspects of euphausiid biology, and
Mauchline (1980) updated this. Baker et al.’s (1990) guide to the world’s krill species
is indispensable. It includes a good brief description of euphausiid anatomy and
well-illustrated keys to the species. A paper on krill fisheries of the world (Nicol
& Endo 1997) was recently published by FAO, and an easy-to-use CD by Brinton
et al. (1999), giving illustrated identification of species, synonymies, references
and distribution maps, was published by UNESCO.
The genus best represented in the New Zealand region is Stylocheiron. Half
of the 12 species known globally occur in New Zealand waters, whereas only two
(20%) of 10 Thysanoessa species have been recorded here. Nyctiphanes australis
is one of four and Nematobrachion flexipes one of three species in their genera.
Two of seven species of Nematosceles (29%) and five of 14 Thysanopoda species
(36%) are present. Euphausia, the largest euphausiid genus with 31 species,
is represented in New Zealand waters by just six species and one subspecies
(22%). Records from New Zealand include three ‘round-eyed’ genera (Euphausia,
Nyctiphanes, Thysanopoda) and three genera with bilobed eyes and elongated
legs (Nematobrachion, Nematosceles, Stylocheiron). Both Thysanoessa species found
in New Zealand waters also have bilobed eyes.
Species recorded in the literature as present, and species believed to be
correctly identified, are listed in the end-chapter checklist, but this probably
does not give the full picture. Other species are very likely to occur in New
Zealand waters. Brinton (1962a) and Brinton et al. (1999) have given Pacificwide and worldwide distributions of krill. Because they are typically offshore
and pelagic in habit, mostly with wide geographic distributions, these
distributional data and maps are, of necessity, generalised. Records from
outside New Zealand’s EEZ suggest that some species may range within the
EEZ boundary, and shading on some maps in both works (Brinton 1962a;
Brinton et al. 1999) indicates that they do. It is possible, though unlikely, that
one or two species have been recorded from New Zealand in food studies of
their many predators (fish, birds and whales), not reviewed here. Unrecorded
krill species likely to be present include some medium-to-large sized species
that may escape capture; not all krill swarm, and swarming species are easier
to catch. Some species also live at depths where fine mesh nets are seldom
deployed. The deep-living species Nematosceles tenella and oceanic N. atlantica fit
these criteria and have yet to be found in New Zealand waters.
More species of Thysanopoda are also likely to be present in New Zealand
waters. Mesopelagic T. astylata, T. cristata, T. orientalis, and T. pectinata occur
widely in the Pacific to about 35° S and a few Thysanopoda species are mesoor bathypelagic and seldom sampled, e.g. T. spinicaudata, found at 2000–3000
metres. Species such as T. cristata are sparsely distributed and not caught
regularly. Distributional records in Brinton et al. (1999) suggest at least some
of these species may occur in the deep offshore waters of New Zealand but
have yet to be collected, which is also the case for Bentheuphausia amblyops
(Bentheuphausiidae) found throughout the Pacific to 54° S.
The species considered above live either in tropical or subtropical waters
or are bathypelagic. Several species present in colder, Antarctic circumpolar
water lying south of the Subantarctic Convergence (Euphausia superba, E.
frigida, E. triacantha, Thysanoessa macrura, and T. vicina) must also come close
to encroaching on the southern areas of New Zealand’s EEZ. However, Morris
et al. (2001) have shown that the Subantarctic Front (Convergence) forms a
boundary between the colder, fresher Antarctic water to the south and warmer
saltier subantarctic water to the north of the front. This abrupt, hydrographic
and biological barrier extends deeply into the water column and is apparently
a permanent phenomenon. The front also skirts the southern edge of the
Campbell Plateau, 200 kilometres south of Campbell Island. This suggests
that these circumpolar species are unlikely to be found within the EEZ, except
perhaps as stragglers.
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NEW ZEALAND INVENTORY OF BIODIVERSITY
Nyctiphanes australis, a small species with adults 10–17 mm long and first
recorded in New Zealand more than a century ago (Thomson 1900), is probably
the best-known euphausiid of New Zealand waters, being abundant around the
main islands and south to The Snares. It has also been studied more than any
other species occurring here or in Australian waters, where it is also plentiful
from New South Wales to South Australia including Tasmania.
The New Zealand species of Euphausia are all small to medium-sized; as
adults, E. recurva is smallest at 10–14 millimetres long; E. longirostris, the largest,
can reach 34 millimetres. Euphausia similis and E. similis armata, both 22–26
millimetres long as adults, are difficult to distinguish but the latter is more often
caught and is one of the commonest krill species encountered in New Zealand.
Three of the five species of Thysanopoda found in New Zealand waters are new
records (T. cornuta, T. egregia, T. monacantha). The largest of these is T. cornuta at
50–120 millimetres adult length; purple-red T. egregia reaches 50–62 millimetres,
and T. obtusifrons is the smallest at 18–23 millimetres (Brinton et al. 1999).
The identity of Stylocheiron longicorne is complicated by the existence of
three ‘forms’ – a North Indian Ocean form, a short form, and a long form. The
latter is present in New Zealand waters and throughout all three main oceans,
while the short form is almost as widespread and may occur in northern New
Zealand. Stylocheiron longicorne is also one of three species of the ‘S. longicorne
species group’ (Brinton et al. 1999) in New Zealand waters, the other two being
S. elongatum and S. suhmi.
Nyctiphanes australis.
From Sars 1885
174
Ecology and distribution of New Zealand krill species
Most krill live in the upper layers of the oceans or in coastal areas. Because they
are pelagic at all stages in their life cycles and strongly influenced by currents
and environmental factors (light intensity, oxygen saturation, temperature,
salinity, and food availability), they tend to be confined to certain water-masses.
The majority of species undertake daily migrations, swimming upwards into
shallower strata of the water column by night and back down before daylight.
Most species are omnivores and feed day and night. Upward migration at night
into shallower waters may enable consumption of phytoplankton, while retreat
to deeper layers during daylight probably helps to avoid pelagic predators.
Krill are well known for swarming, which they do at regular seasonal
intervals or irregularly (Mauchline 1984). Aggregations form at or below the
surface for feeding or reproduction and swarming by Nyctiphanes australis
during the breeding season is well developed. Swarms of N. australis have been
found in harbour and coastal waters of Otago in summer and autumn and a
very dense swarm of about four cubic metres was photographed off The Snares
by Fenwick (1978). Such swarms tend to be patchy and ephemeral (O’Driscoll &
McClatchie 1998) but can be huge and occasionally wash ashore. The largest of
a series of strandings of N. australis on Otago Harbour beaches in January 1990
was estimated to be ca. 100 tonnes (McClatchie et al. 1991b). Euphausia similis
armata also intermittently strands in large numbers. In March 1985 and February
2002, millions of live individuals were washed ashore at Waikanae Beach north
of Wellington. Drifts were hundreds of metres long and ‘ankle deep’, as reported
by locals, who also observed gulls gorging themselves on the windfall. The krill
had apparently been brought ashore by unusual wind and current patterns in
the Cook Strait area.
Although krill actively swim, they are classified as plankton because they
are moved about by currents, but the larger-sized species may behave more as
nekton. Nyctiphanes australis lives mainly over the continental shelf and further
inshore than other species recorded in the New Zealand region (Bary 1956;
Blackburn 1980; Brinton et al. 1999). Offshore transport of N. australis is limited
by coastal currents running parallel to the coast and by behaviour generated
by environmental factors, possibly including vertical movements that place the
krill in currents that retain them near the coast (Bradford 1979). Murdoch (1989)
PHYLUM ARTHROPODA
and O’Driscoll and McClatchie (1998) found that N. australis off Otago became
entrained in an anticlockwise gyre off Blueskin Bay and are most numerous in
low-salinity coastal waters resulting from river runoff. Bary (1956) observed that
the species tolerates a wide salinity range and also penetrates semi-enclosed
waters such as Otago and Wellington Harbours and the Marlborough Sounds.
Nyctiphanes australis undertakes diel vertical migrations from below 150 metres
into the top 40 metres of the water column (Bartle 1976) and Bradford (1979)
observed that N. australis off Kaikoura was able to exist in water temperatures
from 8–10° to 23°C.
All species of Euphausia recorded in New Zealand waters are oceanic with
a circumglobal distribution in the Southern Hemisphere. Only one subspecies,
Euphausia similis similis, occurs in both hemispheres; the remaining New
Zealand representatives of the genus are confined to the Southern Hemisphere,
with each distributed in a circumglobal band. South of the Equator E. similis
similis ranges from 25°S to 55°S (Brinton et al. 1999), which coincides with the
northern and southern extremities of the EEZ and encompasses the distribution
of its co-subspecies E. similis armata. Both subspecies inhabit depths of 0–300
metres but it is not clear if either migrates vertically. Baker (1965) observed what
seems to be an inverse relationship between the numbers of the two subspecies
and Bartle (1976) suggested this may reflect a difference in depth as he found
E. similis similis mostly in the upper 100 metres of Cook Strait while E. similis
armata was mainly deeper.
Euphausia longirostris, E. lucens, and E. spinifera also occur north and south
of the Subtropical Convergence in New Zealand waters (Bary 1956; Bartle 1967;
Robertson et al. 1978; James 1989). Euphausia recurva is a more tropical species
found as far south as Cook Strait (Bartle 1976) and is bi-antitropical in the major
oceans, meaning it is distributed both north and south of the Equator but not
across it, although it can be found at lower latitudes than 20° S and 20° N. On
the other hand, E. vallentini is a colder-water species, recorded by Brinton et al.
(1999) from 50°–60° south of mainland New Zealand, but also found within or
just to the north of the Subtropical Convergence Zone off Kaikoura (Bradford
1972).
Recognition of Nematobrachion boopis in New Zealand waters was only a
matter of time since it is very widespread in the three main oceans from 42°N to
50°S. It is the deepest-living species in its genus, the adults being mesopelagic at
300 metres or more, but it also performs daily migrations. Nematobrachion flexipes
is a deeper mesopelagic species (100–600 metres). It is very widespread though
more patchily distributed than N. boopis (Brinton et al. 1999).
Two species of Nematosceles are found in New Zealand – N. megalops and N.
microps. The former is a warm-temperate species found in all main ocean basins
in the Southern Hemisphere and in the North Atlantic. Nematosceles microps is
widespread in warm-temperate seas in all three main oceans between 40° N and
35° S (Brinton et al. 1999) but has been recorded only once off northern New
Zealand (Tattersall 1924).
Stylocheiron elongatum is widespread in all oceans from 40° N to 35° S
(Brinton et al. 1999) although Bartle (1976) collected two juvenile specimens
from Cook Strait. He did not consider this unusual since waters of subtropical
origin are known to penetrate southwards along the Hikurangi Trench into
Cook Strait at 300–500 m, the appropriate depth for S. elongatum. Stylocheiron
carinatum, S. suhmi, and S. abbreviatum have been recorded only in northern
New Zealand waters (Tattersall 1924) but S. maximum is very widespread in the
three main oceans. Its distribution encompasses New Zealand to 63° S in the
Pacific Ocean (Brinton et al. 1999) although Robertson et al. (1978) found it only
north of the Subtropical Convergence east of central New Zealand. Stylocheiron
maximum is mesopelagic, being mostly caught at depths exceeding 400 metres,
while S. carinatum occupies near-surface waters above 140 metres both day and
night (Brinton et al. 1999).
CRUSTACEA
Nematoscelis megalops.
After Sars 1885
175
NEW ZEALAND INVENTORY OF BIODIVERSITY
Thysanoessa gregaria is biantitropical in all three oceans, is found throughout
New Zealand waters, and has been caught regularly in eastern and southern
areas (Bartle 1976; Bary 1959; Bradford 1972; Murdoch 1989). While it is
usually found above 150 metres depth, Bartle (1976) noted that it is deeper in
subtropical than subantarctic waters and suggested it also undergoes extensive
vertical migrations. Brinton et al. (1999) indicated that it occupies thermocline
waters, rising and falling with them day and night, and that it has been found
as deep as 1200 metres. Roberts (1972) identified Thysanoessa macrura at the
Auckland Islands but Brinton et al. (1999) placed this species in circumpolar
Antarctic waters south of 55° S. It seems likely that Roberts was dealing with
T. vicina rather than T. macrura since the two species are difficult to distinguish
and, according to Brinton et al. (1999), T. vicina overlaps and occurs north of T.
macrura to 50° S.
Thysanopoda cornuta has been found at scattered locations in the three main
oceans at 1200–2500 metres depth, while larvae and juveniles are present at
700 metres or deeper. Thysanopoda egregia occurs at 800–2000 metres, while T.
monacantha is mesopelagic at 300–400 metres, rising into the upper layers at
night. Like several other widespread krill found at these depths, T. monacantha
requires water fully saturated with oxygen and is absent from oxygen-deficient
areas of the northern Indian and eastern central Pacific Oceans (Brinton 1962b).
Thysanopoda obtusifrons inhabits the low-nutrient central water masses of the
main oceans and is found up to 140 metres deep at night, migrating below 300
metres during the day.
Stylocheiron longicorne.
After Sars 1885
176
Breeding and development of krill
Krill sexes are separate. During mating, a sperm package is transferred to the female
and sperm are stored in a reservoir until eggs are laid and fertilised externally.
In the species of Nematobrachion, Nematosceles, Nyctiphanes, Pseudeuphausia,
Stylocheiron, and Tessarabrachion, eggs are attached to the posterior three pairs
of thoracic limbs until they hatch at the second nauplius (metanauplius) larval
stage. As in other Nyctiphanes species, N. australis females not only retain their
eggs until this stage, but also secrete a paired, membranous ‘egg sac’ to hold the
eggs (Brinton et al. 1999). Nematosceles megalops lays 220–250 small eggs per
brood and Stylocheiron species 2–50 larger eggs (Mauchline & Fisher 1969), both
taxa being represented in New Zealand. In the remaining genera (58 species),
the first nauplius hatches from eggs that are shed directly into the water. Thus
krill have two nauplius stages, but in those with attached eggs the first stage is
passed through in the egg.
Nauplius larvae swim using their antennae, and all subsequent developmental
stages through to the adult are pelagic. The nauplius metamorphoses to the first
of three calyptopus stages in which the abdomen develops its full complement
of six segments, a telson and uropods. Throughout the calyptopus phase the
eyes remain beneath the carapace, and locomotion continues to be provided by
the antennae. The final calyptopus moults to the first of several furcilia stages
in which the eyes become stalked and free of the carapace, the antennae are
no longer natatory, the thoracic legs and gills appear, and, throughout a series
of moults, the pleopods and photophores become fully developed. The furcilia
passes through various numbers of moults both between and within species
and the rate of addition of functional parts varies, depending on environmental
conditions. Euphausia superba has the least number of furcilia stages of any
euphausiid (six) while species of Thysanoessa may have as many as 11 stages
(Mauchline & Fisher 1969).
Sheard (1953) described these complex larval phases of the life-cycle
in several species that happen to occur in New Zealand waters, including a
detailed description of those in Nyctyphanes australis. Typical of coastal species,
the number of larval instars and the sequence of addition of morphological
characters (the developmental pathways taken) in N. australis is variable, and
PHYLUM ARTHROPODA
CRUSTACEA
more so than in oceanic species. The final furcilia moults to the first adolescent
stage with little morphological change.
Food, predation, and parasitism
Krill are omnivorous, feeding on phytoplankton, zooplankton, and organic
detritus from bottom sediments. Species with highly fringed feeding limbs
use them to filter minute protozoans and algal plankton from the water. The
bristles effectively form a fine net to strain food from currents created by the
thoracic limbs and pleopods. Species with less setose appendages feed more on
zooplankton.
The anterior thoracic limbs can be held in such a way as to form a ‘food
basket’ between them and the mouthparts (Mauchline 1984). Bottom-feeding
krill employ two methods of collecting food. In one, the animal approaches the
bottom in a near-vertical position and, by beating the thoracic exopods, raises
into suspension sediment that is filtered by the mouthparts. In the second
method, the animal approaches the bottom at a flatter angle and ploughs up the
soft sediments with its antennae to form a lump, which it ‘sucks’ into the food
basket by a sudden lateral movement of the thoracic limbs. This method is also
used repeatedly as the animals swim, to trap planktonic prey such as copepods
or chaetognaths in the food basket.
Among New Zealand krill, ‘round-eye’ Euphausia, Nyctiphanes, and Thysanopoda species have more highly fringed feeding limbs than ‘bilobed-eye’ Nematobrachion, Nematosceles, Stylocheiron, and Thysanoessa species. In general, the
former group is omnivorous, consuming bottom detritus as well as small
plankton and non-living particles from the water column. The two large deepsea species Thysanopoda cornuta and T. egregia are also known to eat live prey,
having been found with copepods, arrow worms, and juvenile fish in their
stomachs (Brinton et al. 1999). Carnivory had been suspected in the latter group
of krill because bilobed eyes and elongated legs are thought to be adaptations
for the capture of live prey (Mauchline & Fisher 1969). The two large deep-sea
species Thysanopoda cornuta and T. egregia are also known to eat live prey, having
been found with copepods, arrow worms, and juvenile fish in their stomachs
(Brinton et al. 1999).
Nyctiphanes australis is the only one among the above species whose feeding
has been studied in New Zealand waters. Bradford (1972) found maximum
numbers of this species in Kaikoura waters underneath concentrations of
copepods, eating their faecal pellets. Blackburn (1980) listed diatoms, copepods,
and copepod faecal pellets in its diet and McClatchie et al. (1991a) also confirmed
omnivory in the species in Otago waters.
Dalley and McClatchie (1989) carried out a detailed study of the feeding
morphology of Nyctiphanes australis in Otago, and McClatchie et al. (1991a)
measured the spaces between setae of the food basket at 2–8 micrometres, the
finest of any euphausiid measured to that time. This suggested N. australis is
equipped to filter nanoplankton-sized particles. However, Dalley and McClatchie
(1989) also concluded that the species is an ‘opportunistic omnivore’ since it has
both a mandibular molar process typical of predators and a mandibular palp
and stomach armature characteristic of herbivores. Gut contents, measured
using a pigment fluorescence technique (McClatchie et al. 1991a), also revealed
substantial amounts of chlorophyll pigments from phytoplankton much larger
than nannoplankton, consumed directly, or secondarily in the gut contents of
prey. The swarming of N. australis in Otago Harbour also coincides with the
spring diatom bloom (McClatchie et al. 1991a).
Krill are eaten by a wide variety of cetaceans, fish, and birds. Mauchline
(1980) listed the euphausiid species, their major predators, and whether they
swarm or not, swarming being an important aspect of their consumption in large
numbers. Little appears to be known about predators of Euphausia longirostris but
five of the other six New Zealand Euphausia species that swarm are an important
Stylocheiron elongatum.
After Sars 1885
177
NEW ZEALAND INVENTORY OF BIODIVERSITY
constituent in the diets of baleen whales. Euphausia vallentini was reported by
Nemoto (1962b in Mauchline and Fisher 1969) to be eaten by fin and sei whales
in waters south of New Zealand. Among the six species of Stylocheiron, only S.
abbreviatum is reported as swarming, but all are known to be important food for
planktivorous and micronektonic fish. Being mesopelagic, S. maximum is also
found in the stomach contents of some demersal fish. Whales, planktivorous
fish, and seabirds all eat Thysanoessa gregaria when it swarms at the sea surface
but, while T. macrura has been found in whale stomachs, much less is known
about it as a food item. Nematosceles megalops swarms but both it and N. microps
apparently do not approach the surface and are preyed on by demersal and
planktivorous fish. Pelagic and midwater fish feed on Thysanopoda monacantha,
and whales and demersal fish on T. acutifrons.
Studies of feeding in New Zealand fish and seabirds have revealed that
Nyctiphanes australis plays an important role in their diets. Kahawai (Arripis
trutta) around Kaikoura depend on N. australis for much of their diet (Bradford
1972) and barracouta (Thyrsites atun) also eat this species (Bartle 1976). O’Driscoll
and McClatchie (1998) used side-scan radar to study schooling behaviour in
barracouta off Otago and came to the conclusion that ‘schooling of barracouta
seems to be a feeding strategy to exploit surface swarms of krill’. They also found
that jack mackerel (Trachurus murphyi) and slender tuna (Allothunnus fallai)
prey on N. australis. Blackburn (1980) reported that southern bluefin (Thunnus
thynnus maccoyii) and skipjack tuna (Katsuwonus pelamis), common in New
Zealand waters, eat N. australis off Australia. No doubt other pelagic fish prey
on this species, and Fenwick (1978) saw six different species of bottom-dwelling
fish attacking a swarm near The Snares.
With the exception of penguins, seabirds can exploit krill only at or near
the sea surface. Rockhopper penguin (Eudyptes chrysocome) stomachs have been
found with N. australis remains – mainly eyes, which seem to resist digestion
longer than other body parts (Te Papa unpubl. data). Many flying birds also
exploit this species, e.g. grey-faced petrels (Pterodroma macroptera), fairy prions
(Pachyptila turtur) (Bartle 1976), and, importantly, black-billed gulls (Larus
bulleri) (McClatchie et al. 1989). They are eaten at sea by red-billed gulls (Larus
novaehollandiae) but not by black-backed gulls (Larus dominicanus), which prefer
stranded krill (McClatchie et al. 1991b).
Krill are hosts to various parasites. Mauchline (1980) listed three types of
ectoparasites – ellobiopsid and apostome protozoans and dajid isopods. The
effects of ectoparasites on the host are not always obvious but it is thought that
they impair swimming, increase the risk of predation, and damage the cuticle,
allowing bacterial infections (McClatchie et al. 1990). Among krill species found
in New Zealand, Euphausia lucens, E. recurva, E. similis, E. vallentini, Nyctiphanes
australis, and Thysanoessa gregaria have been recorded as being infested with the
ellobiopsid protozoan Thalassomyces fagei (phylum Myzozoa) (Mauchline 1980).
Its precise life-history is not known, but T. fagei first appears under the upper
carapace of the host, sends a root-like structure down among the organs to gain
nourishment, then grows a ‘neck’, up through the carapace, that branches and
produces spores. Dajid isopods attach themselves to the cephalothorax of the
host. Among the krill recorded in New Zealand, dajids have been observed in
Nematosceles megalops, T. gregaria, and Stylocheiron longicorne. McClatchie et al.
(1990) discovered that a stalked pennate diatom also grows externally on N.
australis caught in Otago Harbour, the first record of such an infestation; 50–70%
of N. australis sampled in the Harbour were infested. The effects of the diatom
on the host were unclear but diatom chlorophyll introduced error into their
chlorophyll pigment fluorescence experiments on the krills’ diet.
Commercial exploitation and resource potential of krill
The publication by Nicol and Endo (1997) on the world’s krill fisheries is an
accessible and essential reference for anyone interested in the subject. These
178
PHYLUM ARTHROPODA
CRUSTACEA
authors listed six species of krill commercially harvested in various parts of the
world – Euphausia superba in the Antarctic Ocean, E. pacifica off Japan and British
Columbia, E. nana off southern Japan, Thysanoessa inermis off northern Japan and
in the Gulf of St Lawrence (eastern Canada), and T. raschi and Meganyctiphanes
norvegica also in the Gulf of St Lawrence. In 1997, the annual catch of krill for
human use was estimated at 160,000 tonnes, with E. superba the most important
species.
Japan is the major fishing nation of both Antarctic krill and northern species,
but Ukraine and Poland also have an important stake in the Antarctic fishery.
Russia, Korea, and Chile have also been involved at various times. Probably of
more interest to New Zealand is research carried out in Tasmania on the potential
for a fishery there for Nyctiphanes australis, since the species is abundant in New
Zealand coastal waters as well.
Human uses of krill include food, bait for sport fishing, aquarium food, and
aquaculture food, which is the major use. Krill are of high nutritional value and in
Japan are also used to add colour to fish flesh for human consumption. Like the
exploited species, N. australis has also been shown to have high nutritional value.
Krill contain a wide variety of biochemicals, some of possible pharmaceutical
value, and Nicol and Endo (1997) listed and discussed their properties and
potential uses. They also outlined conservation needs for krill. Current catch
rates are thought to be far below the potential for sustainable fishing but the
importance of krill in marine food-webs is enormous. The probable effects of
overfishing on the many bird, cetacean, and fish predators of krill was important
in setting the regulatory Convention on the Conservation of Antarctic Marine
Resources in 1980.
Scope for future work
New records of krill species found elsewhere can be expected in the New
Zealand region and there is a need to clarify the status of species ‘forms’ and
species groups. Compared to the northern Pacific and Atlantic Oceans there is a
lack of data on krill in the SW Pacific. Should a fishery for Nyctiphanes australis
prove commercially viable off Tasmania, investment in further research on this
and other species in New Zealand waters will probably follow.
Order Decapoda: Shrimps, lobsters, crabs, and kin
Decapods (‘10-footed’) are the most familiar crustaceans, numbering more than
10,000 living species worldwide – almost half the named species of Crustacea.
They occur in a great diversity of forms and habitats and some are highly
specialised. Most decapods are marine, living from above high tide to depths of
more than 5000 metres and at all levels of the ocean. Some live in fresh water
and on land but all land dwellers, including the forest crabs of tropical latitudes,
must have access to water to hatch their eggs and to drink. Decapods range in
size from tiny shrimps about a millimetre long to the largest of all arthropods,
the giant Japanese spider crab Macrocheira kaempferi with claws that can span
up to four metres. There are tiny crabs that live out their lives within coral galls
and the huge xanthid crab Pseudocarcinus gigas of southern Australia that reaches
15 kilograms in weight. While North American clawed lobsters are the heaviest
of all crustaceans, the largest rock (spiny) lobster is the packhorse rock lobster
Sagmariasus verreauxi of New Zealand and eastern Australia that can weigh 16
kilograms.
Behaviourally, some shrimps and prawns spend their whole lives swimming,
while others associate with various bottom habitats. Lobsters and crabs inhabit
all kinds of rocky or soft substrata, some bury themselves temporarily, and others
live in permanent burrows in mud and sand. Certain genera of squat lobsters
are found only on deep-sea branching corals, while small shrimps are often
closely associated with algae, adjusting their colours to blend in. A small number
Carcinologist Rick Webber with a historic
specimen of the large packhorse rock lobster
Sagmariasus verreauxi.
Te Papa Tongarewa
179
NEW ZEALAND INVENTORY OF BIODIVERSITY
Alvinocaris niwa, a hot-vent shrimp.
From Webber 2004
180
of shrimp species have become specialised fish cleaners and a few decapods
are confined to very circumscribed habitats such as coastal anchialine caves,
underwater geothermal vents and cold-water or hydrocarbon seeps, or are
specialised to live on decaying wood or whale bone.
The relationships of decapods with other orders of Malacostraca continue to
be argued as do relationships among decapod groups (e.g. Martin et al. 2009).
The classification followed here is that of De Grave et al. (2009). The traditional
separation of decapods into natants and reptants has no formal status but is
useful when discussing the ‘swimming’ and ‘crawling’ members of the order and
is used here informally.
The Decapoda is divided into two suborders, the Dendrobranchiata, which
includes the penaeoid and sergestoid prawns with gill lamellae divided into many
dendritic branches, and the Pleocyemata, including all remaining Decapoda,
whose gill lamellae are not dendritic (gills are lamellar in the caridean shrimps
and prawns, Brachyura and most Anomura; filamentous in crayfish, lobsters and
some dromiid crabs – see McLaughlin 1980 for description of gill types). The
Pleocyemata thus includes the majority of shrimp and prawn species as well as
freshwater crayfish, clawed, slipper and rock lobsters, true crabs and king crabs,
hermit crabs, and squat lobsters.
Along with all other members of the class Malacostraca, the decapod body
consists of five cephalic (head) somites (six if the eyes are taken as representing
a separate somite), eight thoracic, and six abdominal somites. Appendages of
the anterior three thoracic somites are modified as food-handling maxillipeds,
a principal diagnostic character of the Decapoda since other Crustacea have no
more than two pairs of maxillipeds, while the legs articulate with the five posterior
thoracic segments. In all decapods the cephalic and thoracic segments are fused,
and protected by a carapace that extends down each side of the cephalothorax
to enclose the gills and form branchial chambers. The carapace varies from more
or less cylindrical in shrimps, prawns, and lobsters to rounded and flattened
in crabs but it is the abdomen that has undergone the greatest modifications.
In the natants, the decapod abdomen is at its largest, most muscular, and least
flexible. It is substantial but proportionately smaller in the reptant lobsters and
their relatives, and able to be curved under the cephalothorax, but is reduced
to a flap normally held firmly beneath the cephalothorax, in crabs and crablike Anomura. Despite this variation, all but males of a few hermit-crab species
retain at least some abdominal pleopods. Pleopods provide propulsion in natant
forms and penis-like organs in male decapods, and in female Pleocyemata
remain large enough to carry eggs, even in the shell-inhabiting hermit crabs,
whose abdomens are soft and pleopod numbers reduced.
The chitinous integument (exoskeleton) of crustaceans is variously hardened
by the addition of calcium salts to increase its strength and rigidity. In crabs and
lobsters the skeleton is generally hard and well calcified, except of course at
the joints of appendages and abdominal segments, and most extreme in the
huge claws of lobsters and mature male crabs. But calcium also adds weight
and is therefore minimal in open-water shrimps and prawns. There is also little
calcification in burrowing forms, particularly the callianassid ‘ghost shrimps’,
which seldom if ever venture from their protective tunnels, and in hermit crabs
the claws and front end of the body are well calcified while the abdomen remains
membranous and flexible.
In decapods the sexes are usually separate, although protandry (in which
males change to females as they grow) occurs in a number of species and
protandric hermaphroditism (where male and female reproductive systems
remain functional after the female system develops) has been observed in a
shrimp genus. Mating involves the deposition of non-motile sperm, packaged
in spermatophores, either externally on the cuticular surface of the female, or
internally. Eggs are laid into the surrounding water by dendrobranchs but in the
Pleocyemata are retained by the female’s pleopods until hatching .
PHYLUM ARTHROPODA
Historical overview of studies on New Zealand Decapoda
Sydney Parkinson, artist on Cook’s first voyage to the South Pacific in 1769,
illustrated the spider crab now known as Notomithrax peronii from material
collected in New Zealand. Early settlers and explorers observed and collected
intertidal and shallow-water Crustacea (Yaldwyn 1957a) and Cook and his crews
traded ‘crayfish’ with Maori in the Bay of Plenty (Begg & Begg 1969), a hundred
years before the species Jasus edwardsii (Hutton, 1875) was formally described.
In the last half-century, major reviews of some New Zealand decapod
groups have appeared, summarising historical research on these taxa. Forest et
al. (2000) monographed the hermit crabs (Diogenidae, Paguridae, Parapaguridae,
and Pylochelidae). Their historical account documents an increasingly confused
taxonomy of these families in New Zealand, a problem not confined to the
hermits. Thirteen years earlier, McLay (1988) published his indispensable
book on New Zealand crabs and listed previous contributors to the group.
These included Melrose (1975) who reviewed the hitherto poorly known
Hymenosomatidae, Griffin (1966) who reviewed the majid spider crabs and
their research history, and Bennett (1964) who had himself monographed the
Brachyura and provided a critical history of contributions to the group. In two
unpublished theses,Yaldwyn (1954, 1959) detailed the history of contributions to
New Zealand shrimp and prawn systematics. Wear and Fielder (1985) outlined
the very brief history of local larval taxonomy in a monograph on New Zealand
brachyuran larvae, a publication that probably advanced knowledge of New
Zealand’s crab larvae beyond that of any other region.
The first decapod described from New Zealand is probably the shallowwater spider crab Notomithrax ursus (Herbst, 1788) collected on one of Cook’s
voyages. Halicarcinus planatus (Fabricius, 1775) may have been the first but McLay
(1988) considered this unlikely. No further descriptions of New Zealand material
appeared for 46 years (although 14 species now recorded in New Zealand were
described from other localities prior to 1834). The mid-1830s saw an increase in
taxonomic activity resulting from collections made during exploratory voyages
by ships from Europe and North America visiting the New Zealand region.
Several explorations of the region provided early knowledge of decapod
diversity. These included d’Urville’s first voyage to New Zealand (1826–29)
(decapods reported by H. Milne Edwards, e.g. 1834–1840); the U. S. Exploring
Expedition (1838–42) (decapods reported by Dana, e.g. 1853–55); HMS Erebus
and Terror (1839–43) (decapods reported by White, e.g. 1847); and the Austrian
frigate Novara (1857–59) (some decapods reported by Heller, e.g. 1868). Decapoda
from early exploratory work were first listed with the ‘Annulose Animals’ by
White and Doubleday (1843) in Dieffenbach’s Travels in New Zealand. The
1880s were the most significant decade of the 19th century in terms of additions
to the fauna. The 1874 French Mission de l’Île Campbell made collections from
Cook Strait, Stewart Island, and the subantarctic islands (decapods reported by
Filhol, e.g. 1886). HMS Challenger visited New Zealand on its round-the-world
journey (1873–76) and was the first to sample deep-water stations east and west
of the country and off the Kermadec Islands (Yaldwyn 1957). Bate (1881, 1888)
reported on the mostly meso- and bathypelagic natants, Henderson (1888) the
Anomura, and Miers (1886) the Brachyura. Miers (1876) also compiled a Catalog
of the Stalk- and Sessile-eyed Crustacea of New Zealand from the literature, museum
collections, and a collection borrowed from the New Zealand Government.
New Zealanders began to contribute to local decapod taxonomy with the
first publication of G. M. Thomson (1879b) describing two natant species.
Thomson went on to make an important contribution to New Zealand
crustacean studies, including revisions of the New Zealand hermit crabs (1898)
and natants. With Charles Chilton he provided a list of New Zealand decapods
for Hutton’s (1904) Index Faunae Novae Zealandiae. Chilton made a valuable
contribution to crustacean systematics in New Zealand in a career lasting more
than 40 years. Beginning in 1882 he dealt with a variety of reptants and natants,
CRUSTACEA
Spider crab Notomithrax ursus.
From Griffin 1966
181
NEW ZEALAND INVENTORY OF BIODIVERSITY
King crab Lithodes aotearoa.
From Ahyong 2010
182
from the Subantarctic to the Kermadec Islands and greatly increased knowledge
of their distributions. Chilton (1911c) reported on the New Zealand Government
Nora Niven Trawling Expeditions that covered most of New Zealand’s coastlines.
His 1910 paper on crustaceans from the Kermadec Islands, collected by Oliver
in 1908, remained the major reference to the Decapoda of these islands until the
21st century. The British Terra Nova expedition of 1911 sampled a single but very
valuable bottom station off Northland from which Borradaile (1916) described
brachyurans, hermit crabs, chirostylids, and natants. Decapods collected from
the Auckland and Campbell Islands by Mortensen’s Pacific Expedition of 1914–
16 were described by Stephensen (1927), and Balss (1929) reported on those
collected by the 1924 German Expedition to the Subantarctic Islands led by
Kohl-Larsen.
Foreign expeditions continue to visit New Zealand but the contribution of
local surveys has greatly increased since World War II, such as those organised
by university and museum researchers (e.g. Yaldwyn 1957) and the former
New Zealand Oceanographic Institute of the DSIR (incorporated into NIWA
since 1992). The Ministry of Fisheries’ Observer Programme, in which onboard
observers monitor commercial fish catches within the EEZ, has yielded a steady
flow of interesting decapods from deep water. In addition, NIWA vessels are
currently adding new and rare decapods taken in deep water, on and around
seamounts and other locations not previously sampled.
In the postwar period, crab systematics was advanced by the work of
Richardson (1949a,b) and Dell (e.g. 1960, 1963a,b, 1968a,b, 1971, 1972, 1974),
sometimes in collaboration (e.g. Richardson & Dell 1964; Dell et al. 1970). The
first recognition of lithodid king crabs in New Zealand waters came from the
identification of Paralomis zealandica (as Lithodes sp.) from Cook Strait by King
(1958), and, as deep-water investigations increased, five further species were
added (Yaldwyn & Dawson 1970; Dawson & Yaldwyn 1970, 1971, 1985; Dawson
1989; O’Shea et al. 1999), with the total New Zealand fauna now numbering
at least 13 species (Ahyong 2010). Schembri and McLay (1983) published an
annotated key to hermit crabs of the Otago region that, in the absence of any
similar publication, proved a particularly useful guide to identification until the
comprehensive review by Forest et al. (2000).
John Yaldwyn of the Dominion (later National) Museum published on
several decapod groups but his most extensive contribution concerned the New
Zealand shrimp and prawn fauna. In 1957, he described the Sergestidae of Cook
Strait, a significant contribution to this difficult group (Yaldwyn 1957b). He and
L. R. Richardson published keys to New Zealand’s natant decapods (Richardson
& Yaldwyn 1958), now outdated but still the only comprehensive guide available.
He added numerous new species to the fauna, notably those collected by
the Chatham Islands 1954 Expedition (Yaldwyn 1960) and from the National
Museum’s collection (Yaldwyn 1971), and published or contributed to numerous
other works (e.g. Yaldwyn 1954a,b, 1959, 1961, 1974; Yaldwyn & Dawson 1985).
Since 2000, the rate of publication on New Zealand decapod taxonomy has
increased Papers on brachyuran crabs have predominated, with the emphasis
on collections from the Kermadec Islands (e.g. Takeda & Webber 2006; McLay
2007; Ahyong 2008) and sea mounts and chemosynthetic habitats (Ahyong
2008). Reviews of the chirostylid squat lobsters (Schnabel 2009) and king crabs
(Ahyong 2010) added many new species.
It appears the first systematically collected and recorded New Zealand
collection of decapods (and other Crustacea) was that of Charles Chilton, who
deposited his material in the Canterbury Museum. Another collection of note
is that of A. W. B. Powell at the Auckland Institute and Museum, collected in
the 1930s and ‘40s. After World War II, the collection of Decapoda at the then
Dominion Museum increased steadily with the efforts of Moreland and Dell and
was continued at greater pace by Yaldwyn between 1959 and 1969 and by Webber
into the 1990s. This museum collection is particularly strong in offshore natants
PHYLUM ARTHROPODA
CRUSTACEA
and decapod larvae but has a wide coverage of New Zealand decapods as well as
some valuable material from Pacific Islands. A small collection made by Betty
Batham in the 1940s and `50s is housed at the Portobello Marine Laboratory of
Otago University. NIWA, Wellington, has a major collection of decapods, which
has become the fastest growing in New Zealand.
The New Zealand decapod fauna
Some 591 decapod species (492 living, ~99 fossil) are known from New Zealand,
not all of them formally named, and there are still more to be discovered. New
Zealand’s decapod fauna is generally considered depauperate compared to other
regions (Dell 1968a), given the extent of the EEZ over 30 degrees of latitude, the
exceptionally large area of continental shelf and slope, and the wide variety of
seafloor relief and ecological niches available. It is difficult to find comparable
areas but the numbers of New Zealand crabs have been compared with South
Australia by Dell (1968a) and with the Galápagos, Chile, eastern USA, China,
and Japan by Feldmann and McLay (1993). These comparisons certainly indicate
the limited nature of New Zealand’s crab fauna. This is more simply observed
in the lack of variety and number of crabs found on seashores or the small
number of locally caught crabs, shrimps, or lobsters in fish shops compared
with neighbouring Australia and many places further afield. It is generally felt
that this limited diversity of species has resulted from New Zealand’s isolation
from centres of diversity that might have acted as sources of recruitment. Dell
(1968a) suggested that New Zealand’s separation from Australia in the Early
Tertiary occurred before evolutionary radiation gave Australia its diverse fauna
but it is unclear why a similar process has not occurred in New Zealand. It is
reasonable to view most of New Zealand’s decapod taxa as depauperate but
there are exceptions – New Zealand is well represented by southern hemisphere
oceanic natants that live independently of shallow water and are less limited
by constraints on dispersal, but there is also a high diversity of hermit
crabs and some squat lobster genera and the two crab families Majidae and
Hymenosomatidae are also well represented.
Taxonomic knowledge of New Zealand’s present-day Decapoda is comprehensive for the hermit crabs and squat lobsters, and reasonably good for
coastal and shelf natants and the Brachyura, but not so for the thalassinids and
penaeoid and sergestoid shrimps and prawns. Present exploration of deep-sea
rocky habitats, notably the many seamounts in the New Zealand region, is rapidly
increasing our knowledge of decapods in these places. Geographically, the least
well-known areas are the Kermadec Islands (although knowledge of the shallowwater crab fauna is rapidly increasing), and much of the west coast of New Zealand.
Decapods are an important component of the luxury food market worldwide.
Despite New Zealand’s limited variety of edible species, some nevertheless
support very valuable fisheries, most notably the red rock lobster Jasus edwardsii.
Interest in developing new crustacean fisheries is growing, and considerable
research effort is now expended on ways of improving rock-lobster productivity
and quality through habitat enhancement, ongrowing of juveniles, and the
possibility of culturing.
The main collections of New Zealand decapods are held at the Museum of
New Zealand and NIWA, but considerable historic collections and the majority
of types are kept at the Natural History Museum (London) and the Muséum
National d’Histoire Naturelle in Paris. Other significant collections are located
in the Senckenburg Museum (Frankfurt) and the Australian Museum (Sydney),
while further important material resides in a number of other institutions,
notably Museum Victoria, the U. S. National Museum of Natural History, and
the National Science Museum in Tokyo. The largest type collection in the country
is housed at the Museum of New Zealand, where there are 202 lots, including
just 42 primary types. A smaller collection of types is held by NIWA and type
material is also kept at Auckland, Canterbury, and Otago Museums.
Endemic triangle crab
Eurynolambrus australis.
From Griffin 1966
183
NEW ZEALAND INVENTORY OF BIODIVERSITY
A total of 492 living decapod species have been recorded within New
Zealand’s EEZ (see end-chapter checklist). When the first Decapoda checklist was
compiled for Species 2000 New Zealand in 2002 the classification used was that
of Martin and Davis (2001). The greatest effect their revised classification had on
the hierarchy of New Zealand decapods was to increase the number of families
recognised locally, mainly by raising subfamilies to family status, especially in
the Brachyura. Since then, there has been less change in the classification of
shrimps and prawns and other non-brachyuran groups but changes continue
to be made in brachyuran families (e.g. Ng et al. 2008). New Zealand has 84 of
the 151 families of Martin and Davis (2001) although a large proportion of them
(43%) contain only one or two species (20 with only one species, 15 with two).
In contrast, the three most species-rich families contain 112 species, or almost a
quarter of the decapod fauna. Of these three, the Galatheidae has the greatest
number with 46 species, the Paguridae with 34 species and the Chirostylidae
with 33. The Chirostylidae also includes the most speciose New Zealand genus,
Uroptychus, with 27 named species. The largest natant family is the Oplophoridae
with 18 species, all named. Among the subfamilies raised to family in Martin
and Davis (2001) are those of the superfamily Majoidea (previously family
Majidae), which contains 33 species. Despite this division, however, the previous
subfamily Majinae (now the Majidae in the strict sense) contains 17 species,
almost as many as the largest New Zealand brachyuran family, Xanthidae (18
species).
Freshwater hymenosomatid
crab Amarinus lacustris.
From Melrose 1975
Native paddle crab Ovalipes catharus.
Shane Ahyong
184
Endemism
Of the 492 living New Zealand decapods known, 12 are unnamed or not yet
fully determined. The level of endemism is only ~30% (144 species). As might
be expected, endemism is lowest in pelagic offshore species and highest among
benthic and shallow-water forms. Thus all seven dendrobranch families (23
named species, two undetermined) contain no endemics at all and the four
pelagic carid families Nematocarcinidae, Oplophoridae, Pandalidae, and
Pasiphaeidae (44 species in total) include only one endemic species. New
Zealand’s dearth of nearshore pelagic natants in any of these groups is reflected
in this low endemism and, although an estimated 35 additional penaeoid and
sergestoid species may be anticipated for the fauna, few if any are likely to be
restricted to New Zealand waters. Subtract offshore natant groups from the
named decapods and the proportion of endemics rises. But lower endemism is
not characteristic of all natants – of the 471 named living New Zealand Decapoda,
97 are carid shrimps of which 30 (~31%) are endemic, the same proportion as
for the reptants alone, of which 106 (~31%) are confined to the New Zealand
region. Ten of the 253 New Zealand decapod genera are endemic, viz the
brachyurans Eurynolambrus, Halimena, Heterozius, Jacquinotia, Neohymenicus,
Neommatocarcinus, Nepinnotheres, Pteropeltarion, and Trichoplatus and the slipper
lobster genus Antipodarctus – all of which contain a single species. One family,
Belliidae, is endemic.
Most New Zealand species of Crangonidae and Palaemonidae are endemic,
as are both species of Spongicolidae, probably reflecting their close association
with hexactinellid sponges. There is also higher-than-average endemism of
Axiidea and Gebiidea (former Thalassinidea), Diogenidae, and Paguridae.
This is in contrast to the deeper-water hermit crabs of the Pylochelidae and
Parapaguridae, which each have only a single endemic species.
While the two freshwater parastacid crayfish Paranephrops planifrons and
P. zealandicus and the only freshwater shrimp Paratya curvirostris are endemic,
the freshwater hymenosomatid crab Amarinus lacustris is not, occurring also at
Norfolk and Lord Howe Islands and in southeastern Australia and Tasmania.
A number of rarely caught deep-sea species previously thought to be
endemic to New Zealand have been found in greater numbers and further afield,
particularly in southeast Australian waters (e.g. Lipkius holthuisi, Teratomaia
PHYLUM ARTHROPODA
CRUSTACEA
richardsoni). The apparent endemism and rarity of some deep-sea species
are probably the result of insufficient sampling. Endemism in New Zealand’s
second-largest crab family, Majidae, is rather low at 35% (six of 17 species) but
includes intertidal (e.g. Notomithrax peronii) and shelf/slope (e.g. Thacanophrys
filholi) taxa. Hymenosomatid crabs are well represented in New Zealand and 10
of the 14 species (71%) are also endemic. One of the non-endemics, Halicarcinus
innominatus, is thought to be of New Zealand origin but accidentally introduced
to Tasmania.
New Zealand’s two species of Pinnotheridae (pea crabs) are both endemic,
as might be expected of shallow-water associates of bivalve molluscs, but
endemism in the crab families Portunidae (paddle crabs) and Xanthidae is quite
low at less than 30%. Just three of 11 native portunids and three of 15 native
xanthids (all found only at the Kermadec Islands) are endemic. Portunids and
species of Varunidae tend to have long larval lives and some are able to travel
great distances as adults so that most species are distributed widely. Even New
Zealand’s only terrestrial decapod, Geograpsus grayi of the Kermadec Islands, is
widespread in the Indo-West Pacific.
Of New Zealand’s 132 endemic decapods, 14 are recorded from the Kermadec
Islands and nine are restricted there. Five are hermit crabs, all from moderately
deep water except Pagurixus kermadecensis, which is found in rock pools. Like a
number of other apparent endemics, the shrimp Stylodactylus discissipes is known
from only a single station at 1100 m depth and is likely to be more widespread.
Ecological studies
Paddle crabs (Ovalipes catharus) are numerous enough to comprise a
small fishery, encouraging investigation of marketing (Cameron 1984) and
reproductive biology (Haddon 1994, 1995; Haddon & Wear 1993). University
research has made a considerable contribution to decapod biology, particularly
that carried out over the years by Malcolm Jones and Colin McLay (Canterbury)
and Bob Wear (Wellington), with their students. The physiology of musculature,
haemolymph, locomotion, and eye function in shore crabs have been addressed
(e.g. Jones & Greenwood 1982; Bedford et al. 1991; Forster 1991; MeyerRochow & Reid 1994; Palmer & Williams 1993; Meyer-Rochow & Meha 1994;
Depledge & Lundebye 1996) as have the effects of low oxygen and varying pH
on freshwater shrimp (West et al. 1997; Dean & Richardson 1999). Feeding
studies of shore crabs were carried out (e.g. Wear & Haddon 1987; Creswell &
McLay 1990; Woods 199l; Woods & McLay 1994). Jones (1976, 1977, 1978, 1980,
1981), Jones and Winterbourn (1978), and Jones and Simons (1981, 1982, 1983)
undertook significant work on intertidal crabs of the Avon-Heathcote Estuary
and Kaikoura, and other ecological studies were made by McLay and McQueen
(1995), Palmer (1995), and Morrisey et al. (1999). Several papers on the behaviour
and associations of shore crabs have also appeared (e.g. Field 1990; Taylor 1991;
Chatterton & Williams 1994; Woods & McLay 1994; Woods 1995; Woods & Page
1999) and Berkenbush and Rowden (1998, 1999) studied population dynamics
and sediment turnover in the burrowing ghost shrimp Callianassa filholi.
Alien species
Interest in adventive species is growing rapidly in New Zealand (see Cranfield et
al. 1998 for a list of adventive decapods and the Ministry of Fisheries for details
of potential invaders (Marine Pest Identification Guide series)). Some decapods
have been introduced intentionally but mostly without success; this is probably
a good thing as some crab and lobster species are among the most destructive
of invaders. The first such introduction appears to have been of the Australian
penaeid prawn Melicertus canaliculatus (as Penaeus canaliculatus), released at
Nelson in 1892 and at the entrance to Otago Harbour in 1894 (Thomson 1922).
They were never seen again. Between 1906 and 1918, a more serious attempt
185
NEW ZEALAND INVENTORY OF BIODIVERSITY
Alien paddle crab Charybdis japonica.
Shane Ahyong
Projasus parkeri, a recent palinurid.
W. Richard Webber
186
was made to introduce the European lobster Homarus gammarus into New
Zealand. A similar project was undertaken with the European edible crab Cancer
pagurus between 1907 and 1914 (Thomson & Anderton 1921). Live crabs and
lobsters were imported from the United Kingdom and kept at the Portobello
Marine Fish-Hatchery in Otago Harbour. Several million crab larvae and more
than 750,000 lobster larvae were hatched and liberated in the harbour during
those years. Some young lobsters were reared for several years before release,
and mature adults of both species were also liberated but no trace of free-living
specimens of either species has been found in Otago or New Zealand waters
since.
There was a short-term attempt in the early 1990s to farm a ‘saltwater king
prawn’ from Hong Kong, probably the penaeid Fenneropenaeus chinensis, at
South Kaipara Heads. Like the H. gammarus and C. pagurus experiments this
also failed but in this case the stock was destroyed. So too was the entire stock
at a pond farm of the Western Australian crayfish or marron, Cherax tenuimanus,
at Warkworth, north of Auckland in the late 1980s and early 1990s (Hughes
1988; Lilly 1992). Fear of their escape into waterways led to this action but the
same problem does not occur with large palaemonid prawns farmed at Wairakei,
near Taupo. Here, Macrobrachium rosenbergii from South-east Asia and northern
Australia is successfully farmed in artificially heated water. This is drawn from
the Waikato River and warmed by a heat exchanger using hot-water runoff from
a geothermal power station nearby. Macrobrachium rosenbergii cannot breed or
survive in ambient New Zealand fresh waters.
Foreign decapods periodically appear accidentally in New Zealand,
apparently introduced in ships’ ballast water or on hulls. Some species disappear
but others threaten to become established and compete with the local biota.
The hymenosomatid crab Halicarcinus ovatus, normally found around western,
southern, and eastern Australia, was recorded just once at Port Chalmers, Otago,
by Filhol (1886) but has not been recorded in New Zealand since (Melrose 1975;
McLay 1988). In 1978, the small inachoidid spidercrab Pyromaia tuberculata,
originally from the Central American west coast but subsequently found in
other localities in the Pacific and Atlantic Oceans, was discovered in the Firth of
Thames (Webber & Wear 1981). It has become established but is not often found
and does not seem to be a major threat to endemic species.
In the early 1990s live specimens of three species of crab were found in a
ship’s sea chest at a Nelson slipway – Pilumnus minutus, Carupa tenuipes, and
Charybdis hellerii (Dodgshun & Coutts 1993). The significance of sea chests
(recesses in ship hulls housing the intakes of ballast water) as a mode of
introduction quickly became apparent. Pilumnus minutus is small and uncommon
and C. tenuipes tropical, and neither is likely to become established, but the Asian
and northern Indian Ocean portunid C. hellerii is a successful invader of the
eastern Mediterranean and western Atlantic from Florida to Brazil. It is unlikely
that C. hellerii could establish itself in New Zealand, except perhaps in the far
north, but a close relative has. First reported from Waitemata Harbour in 2001,
hundreds of Charybdis japonica, including egg-bearing females, have since been
caught, and it is also present in the Firth of Thames (Webber 2001; Smith et al.
2003). Almost as large, and far more aggressive than the paddlecrab Ovalipes
catharus, C. japonica is likely to exclude the local species from harbour and estuary
mouths but is unlikely to spread to open sand beaches or much further south,
as it is a warm-water species. Its behaviour in nets causes problems for flounder
fishers but if it remains in large-enough numbers, it may at least become a new
fishery.
Introductions have also occurred in the opposite direction. The small
hymenosomatid crab Halicarcinus innominatus and the larger pie-crust crab
Metacarcinus novaezelandiae were probably accidentally introduced to Tasmania
when Ostrea angasi was imported from New Zealand to enhance the oyster
fishery (Lucas 1980).
PHYLUM ARTHROPODA
New Zealand fossil Decapoda
The fossil decapod fauna comprises approximately 99 species, although only
56 of these are named unequivocally owing to the high proportion of small or
unique specimens or their often incomplete or fragmentary state. There are 48
named genera in 27 families, and six of the seven Recent reptant infraorders
(only Polychelida lacking), and only the Glypheidea (superorder Pleocyemata)
among the natants. Nineteen of the 58 Recent reptant families include fossil
species, with five families represented in New Zealand only by fossils. Some 22
fossil genera also occur in the present-day New Zealand fauna and four Recent
species are represented in the New Zealand fossil record, possibly six, should
fossil Ctenocheles cf. maorianus and Ommatocarcinus cf. Neommatocarcinus huttoni
prove indistinguishable from their living namesakes.
Although the fossil decapod fauna of 99 species is small relative to the
present-day fauna, recent research has revealed its significance to the origins
of decapods in New Zealand and in the South Pacific (Feldmann 2003). The
xanthid crab Tumidocarcinus tumidus was the first fossil decapod described
from New Zealand, but 94 years were to elapse before additional records were
published. Glaessner (1960) published his signal work on the New Zealand fossil
Decapoda, recognising 29 species in eight genera, including a new genus and 16
new species. Most of these were brachyurans (22 crabs in seven families) but
Glaessner also identified five callianassid ghost shrimps and three astacoidean
lobsters of the families Glypheidae and Mecochiridae. In addition, he described
the palinurid rock lobster Sagmariasus flemingi (as Jasus flemingi), the only fossil
yet discovered among the nine Recent species of non-stridulating Palinuridae
(Jasus, Projasus, and Sagmariasus species, all austral).
Glaessner’s (1960) work remains the most important contribution in terms
of numbers of taxa added to the fossil fauna, although subsequent work has
trebled the known fauna. Only three more new species were added to the fauna
during the 1960s and 1970s, but momentum and diversity then increased, with
nine new species described in the 1980s and 16 in the 1990s. Crabs predominate
among the new records, but several other new taxa have also been identified,
leading to fresh interpretations of their origins and relationships to Recent
forms. For example, New Zealand’s first fossil nephropid lobster, Metanephrops
motunauensis, was described from north Canterbury.
The first decapod added to the fauna by a New Zealand worker (Trichopeltarion
greggi) was also the first fossil species of the extant family Atelecyclidae (Dell
1969). The tymoloid family Torynommidae was erected by Glaessner (1980)
to contain several extinct Australasian crabs including two new New Zealand
species, and in the same paper Glaessner named three new species of raninids
for New Zealand. Hyden and Forest (1980) described the first, and so far the only
named, fossil hermit crab from New Zealand (Diacanthurus spinulimanus), and
the late Sir Charles Fleming (1981) described Miograpsus papaka, so far the only
fossil grapsid recorded from New Zealand.
The description of the squat-lobster-like anomuran Haumuriaegla glaessneri
was significant, both for the implications it had for the interpretation of
New Zealand’s fossil record and as the beginning of a major and continuing
contribution to New Zealand decapod palaeontology by its author (Feldmann
1984). Linuparus korura was the second palinurid added to the New Zealand fossil
fauna (Feldmann & Bearlin 1998) and Feldmann and Maxwell (1999) described
five more decapods – two raninids, two majids, and a single goneplacid, the first
New Zealand fossil of the genus Carcinoplax. At this point, a review of the fossil
decapods of New Zealand by Feldmann and Keyes (1992) appeared, listing all
previously published records, giving a detailed index of locality records and an
updated checklist of taxa, and tabulating their stratigraphic ranges in the Mesozoic
and Cenozoic. Some 81 decapods were recorded, although just 38 species were
named. Forty genera were recorded in 21 or 22 families, a considerable increase
from the eight genera in 11 families recognised by Glaessner (1960). Five more
CRUSTACEA
Native frog crab Notosceles pepeke.
From Yaldwyn & Dawson 2000
187
NEW ZEALAND INVENTORY OF BIODIVERSITY
Planktonic zoea larva of the
majid crab Jacquinotia edwardsii.
W. Richard Webber
188
new species were soon added to the fauna by Feldmann (1993), including the
first published record for New Zealand of the Calappidae (Calappilia maxwelli),
the first record of the genus Glyphea (G. stilwelli), and one further species in each
of the Holodromiidae, Torynommidae, and Majidae.
Feldmann and Keyes’ (1992) review and McLay’s (1988) survey of New
Zealand’s Recent crab fauna were closely followed by a substantial paper on the
paleogeographic history of the New Zealand Brachyura (Feldmann & McLay
1993). In their analysis, these authors compared New Zealand’s extant Brachyura
with that of other, mostly Pacific, regions and went on to identify significant
relationships not recognised previously between New Zealand’s Recent and fossil
faunas. A number of new taxa have come to light since these works, supporting
their observations.
The first recognition of the family Parastacidae in the fossil record (Paranephrops fordycei) was published from a single specimen found in Miocene
deposits of Central Otago (Feldmann & Pole 1994). Two further majids were
added to the fauna by McLay et al. (1995) and a new cancrid by Feldmann and
Fordyce (1996). The world’s first fossil lithodid (king) crab (Paralomis debodeorum)
was described only in the 1990s (Feldmann 1998), along with a glypheid lobster,
Glyphea christeyi (Feldmann & Maxwell 1999), both from Canterbury.
The origins of New Zealand’s decapod fauna are far from clear and continue to be debated, particularly because of fossil discoveries over the past 20
years in both New Zealand and Antarctica. Until the early 1980s it was believed
that New Zealand’s decapod fauna was primarily of Australian and Indo-Pacific
origin. Glaessner’s (1960) Tertiary material occurred no earlier than the middle
Eocene (45–50 million years ago). He considered the presence of Tumidocarcinus
in the middle Tertiary of Australia and in the Eocene and Miocene of New
Zealand as indicative of a ‘distinctive zoogeographical province’ and that
Australasian genera could be considered as originating in the ancient Tethys
Sea. Fleming (1962, 1979) also concluded that New Zealand decapods were
primarily of Tethyan origin and that typical New Zealand marine decapod
faunas had appeared since the Oligocene. In his analysis of the distribution
and composition of New Zealand’s extant Brachyura, Dell (1968a) found that
the strongest external elements in the present-day crab fauna are Australian
and Malayo-Pacific in practically equal strength, which also implies a Tethyan
origin.
The late Mesozoic H. glaessneri from North Canterbury was a shallow-water
marine species and the earliest known representative of the extant freshwater
anomuran family Aeglidae, which is confined to temperate latitudes of southern
South America. This discovery, and analysis of fossil and recent species of
Lyreidus (Raninidae), led Feldmann (1984, 1986, 1990) to believe that these and
other decapod genera had evolved in high-latitude southern waters rather than
originating in the Tethys. This occurred during the late Mesozoic prior to New
Zealand’s split from Australia and Australia’s split from Antarctica, which also
had a cool-temperate climate. Feldmann considered that species evolving along
this coast would be dispersed eastwards by the southern Pacific gyre but that this
would have discontinued with a cooling climate and the break of Australia from
the Antarctic, allowing the circumpolar current to develop.
Newman (1991), however, questioned this view and suggested that taxa like
the entirely austral Jasus species may have resulted by reliction (reduction in
range) following an amphitropical (northern as well as southern hemisphere)
distribution. He offered three hypotheses on how such southern hemisphere
endemism could have come about – centres of origin, dispersal to the southern
hemisphere, or vicariance (see Newman 1991).
This debate continues, with research on fossil decapods worldwide increasing
in recent years. Schweitzer (2001) has summarized decapod paleobiogeography
and the diverse literature on decapod fossils and their interpretation was
reviewed by Feldmann (2003).
PHYLUM ARTHROPODA
Decapod development
No discussion of decapod diversity would be complete without reference to their
larvae. The morphology of decapod developmental stages is an important aspect
of decapod systematics, and knowledge of larval biology and recruitment to
adult populations is essential to managing decapod fisheries.
Development in the great majority of Decapoda, both natants and reptants,
includes free-swimming planktonic larvae. In the penaeoid and sergestoid
(dendrobranch) shrimps and prawns, eggs are laid into the surrounding water
and tiny, motile nauplius larvae subsequently hatch into the plankton. All other
decapod groups (the Pleocyemata) retain their eggs attached to the pleopods
until larvae hatch. In the plankton, larvae grow through a series of instars until,
at the final moult, they metamorphose into a post-larva, an intermediate form
looking more or less like the adult but retaining the ability to swim. The role of
the post-larva is to relocate itself to the milieu of the adult phase where it again
moults to become a juvenile crab, lobster, shrimp, or prawn. Like their larvae,
shrimps and prawns are pelagic. The transition from larva through post-larva to
juvenile is less abrupt although the final larval moult is still marked in pelagic
species by a fundamental change in locomotion from using appendages of the
cephalothorax to propulsion by the abdominal appendages (pleopods).
Most decapod families have different though predictable numbers of larval
growth stages and a single post-larva during development, but a few groups
and species have either extended or abbreviated development. Some have even
eliminated free-swimming larval or post-larval phases altogether, with juveniles
hatching directly from the eggs. The number of larval stages relates to the duration
of the larval phase, and those species with abbreviated or direct development
usually occur in habitats where free-swimming larvae would be lost. Some of
these different strategies are exemplified by New Zealand Decapoda.
Larval decapods are of taxonomic interest because they differ morphologically
from adults. This is particularly so in benthic forms, which make up the majority
of decapod species and occupy very different habitats from their offspring. Pelagic
larvae have evolved their own adaptations to planktonic life, yet the medium
they frequent is in many ways more uniform than the variety of substrata or
depths occupied by the adult phase, which serves to emphasise the importance
to taxonomy of differences in larval features.
Limits to the use of larval features are more practical than theoretical,
however; while larvae caught in plankton can usually be attributed confidently to
higher taxa, incorrect identifications of genera and species based on morphology
are often made (e.g. McWilliam et al. 1995). The only foolproof method of
putting names to larvae caught in plankton is to hatch them from eggs of known
parentage or rear planktonic larvae through to identifiable adults. Since Vaughan
Thompson (1828) first put the provenance of decapod larvae beyond doubt by
observing larvae hatching (see Gurney 1942), rearing techniques have improved,
but maintaining ovigerous females and their delicate offspring in captivity, even
when robust berried females can be caught, is always difficult and sometimes
impossible. However, this impasse has begun to be resolved in the last few years
as molecular analysis has enabled more precise matching of adult and larval
forms. DNA analysis has even enabled the type species of some old larval genera
and species to be matched to the adults they correctly belong with (Palero et al.
2008).
New Zealand’s larval decapods, particularly the Brachyura, are comparatively
well known, thanks largely to the work of Robert Wear and his students
(1965–1985) at Victoria University in Wellington. Their efforts are summarised
in two particularly useful publications. One (Wear & Fielder 1985) consists
of a comprehensive illustrated atlas of all previously described New Zealand
brachyuran larvae, with keys and some new descriptions; the other (Wear 1985),
is an annotated list of all non-brachyuran New Zealand species whose larvae
had been described to that time. Prior to 1985, numerous authors published
CRUSTACEA
Megalopa larva of spider crab
Notomithrax minor.
From Webber & Wear 1981
189
NEW ZEALAND INVENTORY OF BIODIVERSITY
Final phyllosoma larval stage of the
rock lobster Jasus edwardsii.
From Kittaka et al. 2005
190
descriptions of New Zealand decapod larvae but only the more significant are
referred to here. Thomson and Anderson were the first New Zealanders to
describe the larvae of brachyurans of the region, hatched at Portobello marine
station. Prior to the 1960s, the most substantial contribution to New Zealand
larval taxonomy was made by Gurney (1924, 1936, 1942), who described eight
decapod species (in seven families) collected by the Terra Nova and Discovery
Expeditions. Webber (1979) described the developmental stages of eight majid
spider crabs, published later by Webber and Wear (1981). Larvae of 12 species
of carid shrimps, in the families Crangonidae, Hippolytidae, and Palaemonidae,
were described in detail by Packer (1983) who published a guide to these and
six other shallow-water shrimp species in 1985. Since then, the output of larval
taxonomy has slowed. Horn and Harms (1988) completed the larval description of
Halicarcinus varius; Lemaitre and McLaughlin (1992) described the megalopa of
the deep-water parapagurid Sympagurus dimorphus; the complete development
of the packhorse rock lobster Sagmariasus verreauxi was described by Kittaka et
al. (1997); and those of the red rock lobster Jasus edwardsii by Kittaka et al. (2005)
from lobsters cultured in Japan; Cuesta et al. (2001) re-examined the zoeas of
Cyclograpsus lavauxi, Hemigrapsus sexdentatus, and H. crenulatus; and detailed
descriptions of the phyllosomas and nisto of a slipper lobster Scyllarus sp. Z
(probably S. aoteanus) were published by Webber and Booth (2001).
Developmental stages of 94 species (21%) of New Zealand Decapoda have
been described, but a much greater proportion of higher taxa is represented by
this number. Descriptions of larvae, post-larvae, or both have been published
from 45 (54%) of the 84 families recorded from New Zealand. These percentages
reflect the high proportion of families containing only one species (larval
descriptions of single species account for 27 families) but it also indicates the
broad spectrum of decapods whose various larval forms are known to some
degree. Best documented are the Brachyura, with 22 of New Zealand’s 39 families
represented by larval descriptions. The remaining 17 families contain 54 of the
167 brachyuran species, while, in the larger families, 11 of 14 hymenosomatid
and five of 12 portunid species include larval descriptions.
Descriptions of all stages in the development of New Zealand’s crayfish and
lobsters were completed relatively recently, but commercial interest has now
generated considerable investment in research into all aspects of their biology.
The freshwater crayfish Paranephrops planifrons provides an example of direct
development in which there are no larval stages and crayfish hatch from the eggs
(Hopkins 1967). Young crayfish, with the cephalothorax packed with yolk, attach
themselves to the female’s pleopods and pass through three stages with the
third having exhausted its supply of yolk. Development in scampi (Metanephrops
challengeri) is not direct but apparently abbreviated. Wear (1976) found that while
larvae hatch as prezoeas the prezoeal cuticle is quickly shed and the single-stage
large zoea appears to last only two to three days or less before moulting to the
post-larva. Scampi zoeas are not found in surface plankton and have a restricted
ability to swim, which led Wear (1976) to suggest they are very short-lived and
settle as a post-larva soon after hatching.
At the other end of the scale are the palinurid and scyllarid lobsters. New
Zealand’s rock lobsters Jasus edwardsii and Sagmariasus verreauxi, and slipper
lobsters whose larval development is known (Ibacus alticrenatus and Scyllarus sp.
Z), are typical of the Palinuroidea in having a long-lived larval phase. Longest
of all is that of J. edwardsii, with 11 phyllosoma stages that can last more than
a year, perhaps as long as 24 months, in the plankton (Booth & Phillips 1994).
Sagmariasus verreauxi has a similar number of stages but of shorter duration
(up to a year) (Booth & Phillips 1994), I. alticranatus still shorter (4–6 months)
with seven stages (Atkinson & Boustead 1982), and Scyllarus sp. Z with 10
phyllosoma stages that probably have a duration as short as or shorter than
I. alticrenatus. Planktonic larval sampling has concentrated on J. edwardsii
because of its high economic value, but the incidental capture of phyllosomas
PHYLUM ARTHROPODA
of other species has enabled useful comparisons to be made. After hatching and
shedding the naupliosoma cuticle, early-stage phyllosomas drift out to sea. Most
sampled mid- to late-stage larvae of J. edwardsii appeared to become entrained
in the Wairarapa Eddy southeast of the North Island, while those of Scyllarus
sp. Z are found much closer to the North Island east and northeast coasts but
also in oceanic waters to the north and northwest of New Zealand (Webber
& Booth 2001). While mid- and late-stage J. edwardsii are rarely found inside
the continental-shelf break, all stages of Scyllarus sp. Z are found there in good
numbers, indicating that they go through larval development closer to shore.
This accords with the much shorter larval duration in the scyllarid species and
it is assumed that the widely scattered phyllosomas to the north and northwest
are lost. The distribution of adult Scyllarus sp. Z is confined to the northeast
coast of the North Island between Cape Maria van Diemen and Gisborne and
is completely overlapped by J. edwardsii, yet the larvae they produce become
distributed in different geographical areas. Phyllosomas have very limited ability to
swim horizontally but they can move vertically through the water column. Coupled
with changing phototactic responses during development, vertical mobility
enables larvae to exploit currents flowing in different directions at different depths,
a strategy that enables them to position themselves in water masses from which
they can return to the coast as post-larvae (Webber & Booth 2001).
CRUSTACEA
Rock lobster Jasus edwardsii.
W. Richard Webber
Commercial exploitation and resource potential of decapods
Studies of decapod biology and ecology have increased in the last half-century,
especially of commercially important species. Early surveys of fishing potential
included the southern spider crab Jacquinotia edwardsii (Ritchie 1970, 1971; Ryff
& Voller 1976), prawns in the Bay of Plenty in the 1970s, and experiments aimed
at culturing freshwater crayfish. As one of New Zealand’s most valuable fisheries,
Jasus rock lobsters are the subject of numerous and continuing studies. Their
movements and migratory behaviour have been investigated for more than 30
years (e.g. Street 1969, 1971, 1973, 1994; Annala 1981; McCoy 1983; Booth 1984,
1997; MacDiarmid 1991, 1994; MacDiarmid et al. 1991; Andrew & MacDiarmid
1991; Annala & Bycroft 1993; Kelly 1995; Babcock et al. 1999; Butler et al. 1999;
Kelly et al. 1999). Because rock lobsters have pelagic larvae and post-larvae,
research has been carried out on the ecology and recruitment of developmental
stages to adult populations (Booth 1979, 1986, 1994, 1995, 1997; Hayakawa et al.
1990; Booth & Grimes 1991; Booth et al. 1991; Booth & Stewart 1992; Booth &
Phillips 1994; Booth & Kittaka 1994; Booth et al. 1998, 2000; Nishida et al. 1995;
Chiswell & Booth 1999; Chiswell & Roemmich 1999). Rearing of New Zealand
lobster larvae has advanced greatly (Kittaka 1994a,b; Kittaka et al. 1997; Tong et
al. 1997, 2000a,b; Moss et al. 1999), while additional research on their biology
and fisheries has also appeared (e.g. Booth & Breen 1994; James & Tong 1998;
MacDiarmid & Butler 1999a,b). Genetic techniques have been employed to
improve Jasus species stock identities (Ovenden et al. 1992; Ovenden & Brasher
1994; Booth & Ovenden 2000). Allozyme variation has also been identified in
scampi populations around New Zealand.
Acknowledgements
Drs Paul Sagar (NIWA) and Wolfgang Zeidler (South Australian Museum)
provided information on amphipods (literature on amphipods as prey for birds,
and Hyperiidea, respectively). Dr Merlijn Jocqué (University of Leuven, Belgium)
checked the section on Mysidacea and added a new endemic species. Thanks
are due to Dr Bob McDowall (NIWA, Christchurch) for his constructive review
of the Amphipoda section. Drs Michael Ayress (Ichron, UK) and Kerry Swanson
(University of Canterbury, Christchurch) clarified aspects of ostracod taxonomy
for the checklist; John Simes provided information on pre-Tertiary fossils.
191
NEW ZEALAND INVENTORY OF BIODIVERSITY
Authors
Dr Shane T. Ahyong National Institute of Water & Atmospheric Research, Private Bag 14901,
Kilbirnie, Wellington, New Zealand [s.ahyong@niwa.co.nz] Hoplocarida
Dr Graham J. Bird
nz] Tanaidacea
8 Shotover Grove, Waikanae, Kapiti Coast 5036, New Zealand [zeuxo@clear.net.
Dr Janet M. Bradford-Grieve National Institute of Water & Atmospheric Research, Private Bag
14901, Kilbirnie, Wellington, New Zealand [j.grieve@niwa.co.nz] marine Copepoda, Branchiura,
Tantulocarida
Dr Niel L. Bruce Museum of Tropical Queensland, 70–102 Flinders Street, Townsville, Queensland
4810, Australia [niel.bruce@qm.qld.gov.au] Isopoda
Professor John S. Buckeridge School of Civil, Environmental and Chemical Engineering, RMIT
University, GPO Box 2476V, Melbourne, Victoria 3001, Australia
[john.buckeridge@rmit.edu.au] Cirripedia
Dr M. Anne Chapman Deceased. Formerly Department of Biological Sciences, Waikato University,
Private Bag 3105, Hamilton, New Zealand Freshwater crustacean ecology
Dr W. A. (Tony) Charleston 488 College Street, Palmerston North, New Zealand [charleston@
inspire.net.nz] Pentastomida
Mr Elliot W. Dawson Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New
Zealand [edawson@xtra.co.nz] Leptostraca, Syncarida
Mr Stephen H. Eagar School of Earth Sciences, Victoria University of Wellington,
P.O. Box 600, Wellington, New Zealand [stephen.eagar@paradise.net.nz] Ostracoda
Dr Graham D. Fenwick National Institute of Water & Atmospheric Research (NIWA), P.O. Box
8602, Christchurch, New Zealand [g.fenwick@niwa.co.nz] Amphipoda
Dr John D. Green 36 Paturoa Road, Titirangi, Waitakere, Auckland 0604, New Zealand [john.green@
worldnet.co.nz] Freshwater copepod ecology
Dr Ju-Shey Ho Department of Biological Sciences, California State University, Long Beach, 1250
Bellflower Boulevard, Long Beach, California 90840-3702, USA
[jsho@csulb.edu] Parasitic copepoda
Dr J. Brian Jones Fisheries WA, C/o Animal Health Lab., Agriculture WA, Locked Bag 4, Bentley
Delivery Centre, WA 6983, Australia [bjones@agric.wa.gov.au] Branchiura, parasitic Copepoda
Dr Kim Larsen CIIMAR, University of Porto, Rua dos Bragas n. 289, 4050-123 Porto, Portugal
[tanaids@hotmail.com] Tanaidacea
Dr Anne-Nina Lörz National Institute of Water & Atmospheric Research, Private Bag 14901,
Kilbirnie, Wellington, New Zealand [a.lorez@niwa.co.nz] Rhizocephala
Dr Jørgen Olesen Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100
Copenhagen, Denmark [J1Olesen@zmuc.ku.dk] Branchiopoda
Dr Gary C. B. Poore Museum Victoria, GPO Box 666E, Melbourne, Victoria 3001, Australia
[gpoore@museum.vic.gov.au] Isopoda
Dr Carlos E. F. Rocha Universidade de São Paulo, Departamento de Zoologia, Caixa Postale 11461,
CEP 05422 970, São Paulo, Brazil [cefrocha@usp.br] Copepoda: Oithonidae
Dr Russell J. Shiel Department of Environmental Biology, University of Adelaide, Adelaide, South
Australia 5005, Australia [russell.shiel@adelaide.edu.au] Freshwater Copepoda
Dr Les Watling Department of Zoology, University of Hawaii at Manoa, Honolulu, HI 96822, USA
[watling@hawaii.edu] Cumacea
Dr John B. J. Wells Department of Biological Sciences, Victoria University of Wellington, P.O. Box
600, Wellington, New Zealand [wellsjm@xtra.co.nz] Harpacticoida
Mr W. R. (Richard) Webber Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington
New Zealand [rickw@tepapa.govt.nz] Decapoda, Euphausiacea, Mysidacea
192
PHYLUM ARTHROPODA
CRUSTACEA
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PHYLUM ARTHROPODA
CRUSTACEA
Checklist of New Zealand living Crustacea
The following classification is based mostly on Martin and Davis (2001). All species are
to be regarded as marine unless indicated otherwise by habitat codes.
All species: A, adventive; B, brackish/estuarine; C, commensal; E, endemic; F,
freshwater; S, supralittoral; T, terrestrial; *, unpublished (new) record; ? after a genus
name or before a species name indicates uncertainty or a possible misidentification.
Endemic genera are underlined (first mention).
Notostraca: Hs, hypersaline environments.
Cirripedia: Letters in parentheses following new records indicate where material
is held, i.e. AUT (Earth and Oceanic Sciences Research Centre, Auckland University of
Technology); GNS (GNS Science, Lower Hutt); NIWA (National Institute of Water &
Atmosphere, Wellington); UA (Geology Department, University of Auckland).
Other groups, especially Copepoda: Habitat codes – Be, benthic; L, littoral; Sl,
sublittoral (to ca. 10 metres depth); Sh, shelf (ca. 10–200 metres depth); Ba, bathyal (>
200 metres depth); Bp, benthopelagic; Co, coastal; F, freshwater (including wells, as well
as species found in terrestrial mosses as they comprise an essentially aquatic habitat);
O, oceanic; P, parasitic; Pe, pelagic (planktonic); Ep, epipelagic; Me, mesopelagic; By,
bathypelagic; Ph, phytal (if marine, usually in algal and seagrass communities in the
littoral or sublittoral, but W indicates decaying or mollusc-bored wood, which may have
been dredged from depths up to 2000 metres. If freshwater, usually in algal or floweringplant communities but M indicates moss or liverwort and includes water courses and
damp terrestrial situations. Zoogeography codes: Ant, Antarctic; Ca, Campbell Island;
Ch, Chatham Islands; Sa, subantarctic; Sn, Snares Islands; Tr/St, tropical/subtropical; Tz,
transition zone; W, widespread.
Amphipoda: Families of the section Gammaridea sensu Barnard and Barnard (1983)
(Barnard’s 1969 family Gammaridae), follow Barnard and Barnard (1983) and Barnard
and Karaman (1991). Known unpublished amphipod taxa are not included in the list.
SUBPHYLUM CRUSTACEA
Class BRANCHIOPODA
Subclass PHYLLOPODA
Order ANOSTRACA
ARTEMIIDAE
Artemia franciscana Kellogg, 1906 Hs A?
Order NOTOSTRACA
TRIOPSIDAE
Lepidurus apus viridis Baird, 1850 F
Order DIPLOSTRACA
Suborder SPINICAUDATA
LIMNADIIDAE
Eulimnadia marplesi Timms & McLay, 2005 F E
Suborder CLADOCERA
Infraorder ANOMOPODA
BOSMINIDAE
Bosmina meridionalis Sars, 1904 F
CHYDORIDAE
Alona abbreviata Sars, 1896 F
Alona affinis s.l. (Leydig, 1860) F
Alona cambouei Guerne & Richard, 1893 F
Alona guttata s.l. Sars, 1862 F
Alona quadrangularis (Müller, 1785) F
Alona rectangula s.l. Sars, 1862 F
Armatolona macrocopa Sars, 1895 F
Camptocercus australis Sars, 1896 F
Camptocercus rectirostris Schödler, 1862 F
Chydorus sphaericus s.l. (Müller, 1785) F
Dunhevedia crassa King, 1853 F
Ephemeroporus barroisi s.l. (Richard, 1894) F
Graptoleberis testudinaria (Fischer, 1851) F
Leydigia ?australis Sars, 1885 F
Monospilus dispar Sars, 1861 F A?
Oxyurella tenuicaudis (Sars, 1862) F
Pleuroxus hastirostris Sars, 1904 F E
Pleuroxus helvenacus Frey, 1991 F E
Pleuroxus unispinus Henry, 1922 F
DAPHNIIDAE
Ceriodaphnia dubia Richard, 1895 F
Ceriodaphnia cf. pulchella Sars, 1862 F
Ceriodaphnia ?reticulata (Jurine, 1820) F
Daphnia carinata s.l. King, 1852 F
Daphnia dentifera Forbes, 1893 F A
Daphnia lumholtzi Sars, 1903 F
Daphnia obtusa Kurz, 1942 F
Scapholeberis kingi Sars, 1903 F
Simocephalus exspinosus (Koch, 1841) F
Simocephalus obtusatus (Thomson, 1894) F E
Simocephalus ?vetulus (Müller, 1776) F
ILYOCRYPTIDAE
Ilyocryptus sordidus s.l. (Lieven, 1848) F
MACROTHRICIDAE
Lathonura ?rectirostris (Müller, 1785) F
Macrothrix schauinslandi Sars, 1904 F
Pseudomoina lemnae (King, 1853) F
Streblocerus serricaudatus (Fischer, 1849) F
MOINIDAE
Moina australiensis Sars, 1896 F
Moina tenuicornis Sars, 1896 F
NEOTHRICIDAE
Neothrix armata Gurney, 1927
SAYCIIDAE
Saycia cooki novaezealandiae Frey, 1971 F E
SIDIDAE
Penilia avirostris Dana, 1852
Penilia pacifica Kraemer, 1895
Suborder ONYCHOPODA
PODONIDAE
Evadne nordmanni Loven, 1836
Evadne aspinosus Kraemer, 1895
Pleopis polyphaemoides (Leuckart, 1859)
Pleopis trisetosus Kraemer, 1895
Class CEPHALOCARIDA
Order BRACHYPODA
HUTCHINSONIELLIDAE
Chiltoniella elongata Knox & Fenwick, 1977 E
Class MAXILLOPODA
Subclass THECOSTRACA
Infraclass ASCOTHORACIDA
Order LAURIDA
SYNAGOGIDAE
Gen. et sp. indet. Te Papa Palmer 1997
Order DENDROGASTRIDA
DENDROGASTRIDAE
Dendrogaster argentinensis Grygier & Salvat, 1987
Dendrogaster otagoensis Palmer, 1997 E
Infraclass CIRRIPEDIA
Superorder ACROTHORACICA
Order PYGOPHORA
CRYPTOPHIALIDAE
Australophialus melampygos (Brandt, 1907) E
Superorder RHIZOCEPHALA
Order KENTROGONIDA
LERNAEODISCIDAE
Triangulus munidae Smith, 1906
PELTOGASTRIDAE
Boschmaia munidicola Reinhard, 1958
Briarosaccus callosus Boschma, 1930
Galatheascus babai Lützen, 1985
Peltogaster sp. Lörz et al. 2008 E
Tortugaster discoidalis Lützen, 1985 E
SACCULINIDAE
Sacculina sp. Brockerhoff, McLay & Kluza 2006
Order AKENTROGONIDA
THOMPSONIIDAE
?Thompsonia affinis Krüger, 1912
Thylacoplethus novaezealandiae Lützen, Glenner &
Lörz, 2009 E
INCERTAE SEDIS
Parthenopea vulcanophila Lützen, Glenner & Lörz,
211
NEW ZEALAND INVENTORY OF BIODIVERSITY
2009 E
Gen. et sp. indet. Lützen, Glenner & Lörz 2009
Superorder THORACICA
Order IBLIFORMES
IDIOIBLIDAE
Chaetolepas segmentata Studer, 1889 E
Chitinolepas spiritsensis Buckeridge & Newman,
2006 E
Idioibla idiotica (Batham, 1945) E
Order LEPADIFORMES
Suborder LEPADOMORPHA
LEPADIDAE
Alepas pacifica Pilsbry, 1907
Conchoderma auritum (Linné, 1767)
Conchoderma virgatum (Spengler, 1790)
Dosima fascicularis (Ellis & Solander, 1786)
Lepas anatifera Linné, 1758 A
Lepas australis Darwin, 1851
Lepas pectinata Spengler, 1793
Lepas testudinata Aurivillius, 1892
OXYNASPIDAE
Oxynaspis indica (Annandale, 1910)
Oxynaspis terranovae Totton, 1923 E
POECILASMATIDAE
Megalasma carinatum (Hoek, 1883)
Megalasma striatum (Hoek, 1883)
Poecilasma kaempferi (Darwin, 1851)
Trilasmis eburneum Hinds, 1883
Suborder HETERALEPADOMORPHA
ANELASMATIDAE
Anelasma squalicola Lovén, 1845*
HETERALEPADIDAE
Heteralepas japonica (Aurivillius, 1892)
Paralepas minuta (Philippi, 1836)
Paralepas quadrata (Aurivillius, 1894)
Order SCALPELLIFORMES
CALANTICIDAE
Calantica spinosa (Quoy & Gaimard, 1834) E
Calantica spinilatera Foster, 1979 E
Calantica villosa (Leach, 1824) E
Scillaelepas fosteri Newman, 1980 E
Scillaelepas studeri (Weltner, 1922)
Scillaelepas n. sp. 1* NIWA E
Scillaelepas n. sp. 2* NIWA E
Smilium acutum (Hoek, 1883)
Smilium zancleanum (Seguenza, 1876)
EOLEPADIDAE
Ashinkailepas kermadecensis Buckeridge, 2009 E
Vulcanolepas osheai (Buckeridge, 2000) E
SCALPELLIDAE
Alcockianum persona (Annandale, 1916)
Amigdoscalpellum costellatum (Withers, 1935)
Amigdoscalpellum vitreum (Hoek, 1883)
Anguloscalpellum pedunculatum (Hoek, 1883) E
Anguloscalpellum n. sp.* NIWA E
Arcoscalpellum trochelatum Foster, 1979 E
Arcoscalpellum affbricatum Foster, 1979 E
Arcoscalpellum pertosum Foster, 1979 E
Gymnoscalpellum intermedium (Hoek, 1883)
Verum novaezelandiae (Hoek, 1883)
Verum raccidium (Foster, 1979) E
Gen. indet. et n. spp. (2)* NIWA 2E
Order SESSILIA
Suborder VERRUCOMORPHA
VERRUCIDAE
Altiverruca galapagosa Zevina, 1978*
Altiverruca gibbosa (Hoek, 1883)
Altiverruca nitida (Hoek, 1883)*
Metaverruca recta (Aurivillius, 1898)
Metaverruca cf. defayeae Buckeridge, 1994*
212
Gen. nov. et n. sp.* J. Buckeridge E
Suborder BALANOMORPHA
ARCHAEOBALANIDAE
Acasta sp. *AUT
Notobalanus vestitus (Darwin, 1854) E
Solidobalanus auricoma (Hoek, 1913)
AUSTROBALANIDAE
Austrominius modestus (Darwin, 1854) E
Epopella kermadeca Foster, 1979 E
Epopella plicata (Gray, 1843) E
BALANIDAE
Amphibalanus amphitrite (Darwin, 1854) A
Amphibalanus variegatus (Darwin, 1854) A
Austromegabalanus nigrescens (Lamarck 1818)
Austromegabalanus psittacus (Molina, 1782)
Balanus trigonus Darwin, 1854
Notomegabalanus campbelli (Filhol, 1885) E
Notomegabalanus decorus (Darwin, 1854) E
Megabalanus tintinnabulum linzei (Foster, 1979)
BATHYLASMATIDAE
Bathylasma alearum (Foster, 1979)
Hexelasma gracilis Foster, 1981 E
Hexelasma nolearia (Foster, 1979) E
Mesolasma fosteri (Newman & Ross, 1971) E
Tetrachaelasma tasmanicum Buckeridge, 1999
CHIONELASMATIDAE
Chionelasmus crosnieri Buckeridge, 1998
CHTHAMALIDAE
Chamaesipho brunnea Moore, 1944 E
Chamaesipho columna (Spengler, 1790) E
CORONULIDAE
Coronula diadema (Linné, 1767)
Coronula reginae Darwin, 1854
Tubinicella major Lamarck, 1802
PACHYLASMATIDAE
Pachylasma auranticacum Darwin, 1854
Pachylasma scutistriata Darwin, 1854
PLATYLEPADIDAE
Platylepas hexastylos (Fabricus, 1798)
Stomatolepas elegans (Costa, 1838)
PYRGOMATIDAE
Cantellius septimus (Darwin, 1854)
Creusia spinulosa Leach, 1824
TETRACLITIDAE
Tesseropora rosea (Krauss, 1848)
Tetraclita aoranga Foster, 1979 E
Tetraclitella depressa Foster & Anderson, 1986 E
Subclass TANTULOCARIDA
DEOTERTHRIDAE
Deoterthron dentatum Bradford & Hewitt, 1980 P E
(ostracod host)
Doryphallophora aselloticola (Boxshall & Lincoln,
1983) P (isopod host)
Doryphallophora megacephala (Lincoln & Boxshall,
1983) P (isopod host) E
Subclass BRANCHIURA
Order ARGULOIDA
ARGULIDAE
Argulus japonicus Thiele, 1900 F P (fish host) A
Subclass PENTASTOMIDA
Order POROCEPHALIDA
LINGUATULIDAE
Linguatula serrata (Leuckart, 1860) T P (mammal) A
Subclass COPEPODA
Order CALANOIDA
ACARTIIDAE
Acartia danae Giesbrecht, 1889 Pe O Ep Tr/St
Acartia negligens Dana, 1849 Pe O Ep Tr
Acartia ensifera Brady, 1899 Pe Co Ep St E
Acartia jilletti Bradford, 1976 Pe Co Ep St E
Acartia simplex Sars, 1905 Pe Co Ep St E
AETIDEIDAE
Aetideus acutus Farran, 1929 Pe Ep Tr
Aetideus australis (Vervoort, 1957) Pe Ep Sa
Aetideus giesbrechti Cleve, 1904 Pe Ep Tr/St
Aetideus pseudarmatus Bradford, 1971 Pe Ep Tr
Aetideopsis tumorosa Bradford, 1969 Pe/BP Me Sa
Bradyidius capax Bradford-Grieve, 2003 Ba Bp
Bradyidius spinifer Bradford, 1969 Ba Bp
Chiridius molestus Tanaka, 1957 Pe Ep/Me Tr/St
Chiridius pacificus Brodsky, 1950 Pe By Tr/St
Chiridius poppei Giesbrecht, 1892 Pe Me Tr
Chirundina streetsii Giesbrecht, 1895 Pe Me Tr/St
Comantenna crassa Bradford, 1969 Ba Bp
Crassantenna comosa Bradford, 1969 Ba Bp
Crassantenna mimorostrata Bradford, 1969 Ba Bp
Euchriella amoena Giesbrecht, 1888 Pe Me Tr
Euchirella bitumida With, 1915 Pe Me Tr
Euchirella curticauda Giesbrecht, 1888 Pe Me Tr/St
Euchirella formosa Vervoort, 1949 Pe Me Tr/St
Euchirella latirostris Farran, 1929 Pe Me Sa
Euchirella messinensis indica Vervoort, 1949 Pe Me
Tr/St
Euchirella m. messinensis (Claus, 1863) Pe By Tr/St
Euchirella rostrata (Claus, 1866) Pe Me Tr/St/Sa
Euchirella rostromagna Wolfenden, 1911 Pe Me Sa/
Ant
Euchirella similis Wolfenden, 1911 Pe By Tr/St
Euchirella speciosa Grice & Hulsemann, 1968 Pe
Me Tr/St
Euchirella truncata Esterly, 1911 Pe Me Tr/St
Euchirella venusta Giesbrecht, 1888 Pe Me Tr/St
Gaetanus brevicornis Esterly, 1906 Pe By Tr/St
Gaetanus brevispinus (Sars, 1900) Pe By Tr/St
Gaetanus kruppii Giesbrecht, 1903 Pe By Tr/St
Gaetanus latifrons Sars, 1905 Pe By Tr/St
Gaetanus minor Farran, 1905 Pe Me Tr/St
Gaetanus minutus (Sars, 1907) Pe Me Tr/St
Gaetanus pileatus Farran, 1903 Pe By Tr/St
Gaetanus secundus Esterly, 1911 Pe Me Tr/St
Gaetanus tenuispinus (Sars, 1900) Pe MR Tr/St/Sa
Lutamator hurleyi Bradford, 1969 Ba Bp
Pseudeuchaeta brevicauda Sars, 1905 Pe By W
Pseudeuchaeta flexuosa Bradford, 1969 Ba Bp
Pseudeuchaeta magna Bradford, 1969 Ba Bp
Pseudochirella dentata (A. Scott, 1909) Pe By Tr/St
Pseudochirella mawsoni Vervoort, 1957 Pe BySt/ Sa/
Ant
Pseudochirella notacantha (Sars, 1905) Pe By Tr/St
Pseudochirella obesa Sars, 1920 Pe By Tr/St
Pseudochirella obtusa (Sars, 1905) Pe By Tr/St
Pseudotharybis brevispinus (Bradford, 1969) Ba Bp
Pseudotharybis dentatus (Bradford, 1969) Ba Bp
Pseudotharybis robustus (Bradford, 1969) Ba Bp
Pseudotharybis spinibasis (Bradford, 1969) Ba Bp
Sursamucro spinatus Bradford, 1969 Ba Bp
Undeuchaeta incisa Esterly, 1911 Pe By Tr/St
Undeuchaeta major Giesbrecht, 1888 Pe Me Tr/St
Undeuchaeta plumosa (Lubbock, 1856) Pe Me Tr/St
Valdiviella insignis Farran, 1908 Pe By Tr/St
ARIETELLIDAE
Arietellus aculeatus (T. Scott, 1894b) Pe Me Tr
Arietellus setosus Giesbrecht, 1892 Pe Me/By Tr
Campaneria latipes Ohtsuka, Boxshall & Roe, 1994
Ba Bp St
Paramisophria n. sp.* Bp Sh
Paraugaptiloides magnus (Bradford, 1974) Ba Bp St
Paraugaptilus ?buchani Wolfenden, 1904 Pe Me Tr
Scutogerulus pelophilus Bradford, 1969 Ba Bp St
AUGAPTILIDAE
Augaptilus longicaudatus (Claus, 1863) Pe Me Tr/St
Centraugaptilus horridus (Farran, 1908) Pe By Tr/ St
Euaugaptilus bullifer (Giesbrecht, 1889) Pe By Tr/
St/Sa
Euaugaptilus filigerus (Claus, 1963) Pe By T/St
Euaugaptilus hecticus (Giesbrecht, 1889) Pe Ep/
PHYLUM ARTHROPODA
Me Tr
Euaugaptilus humilis Farran, 1926 Pe By Tr
Euaugaptilus laticeps (Sars, 1905) Pe By Tr/St
Euaugaptilus longimanus (Sars, 1905) Pe By Tr
Euaugaptilus nodifrons (Sars, 1905) Pe By Tr/St/Sa
Euaugaptilus oblongus (Sars, 1905) Pe By Tr/St
Euaugaptilus palumbii (Giesbrecht, 1889) Pe Me Tr
Haloptilus acutifrons (Giesbrecht, 1892) Pe Me Tr/St
Haloptilus fons Farran, 1908 Pe Me/By Tr/St/Sa
Haloptilus longicornis (Claus, 1893) Pe Ep/Me Tr/
St/Sa
Haloptilus ornatus (Giesbrecht, 1892) Pe Ep/Me Tr/
St
Haloptilus oxycephalus (Giesbrecht, 1889) Pe Ep/ Me
Tr/St/Sa
Haloptilus spiniceps (Giesbrecht, 1892) Pe Ep/Me Tr
Pachyptilus eurygnathus (Sars, 1905) Pe By Tr/St
BATHYPONTIIDAE
Temorites elongata (Sars, 1905) Pe By W
CALANIDAE
Calanoides acutus (Giesbrecht, 1902) Pe Ep/Me Sa/
Ant
Calanoides macrocarinatus Brodsky, 1972 Pe Ep/
Me St
Calanus australis Brodsky, 1959 Pe Co Ep St/Sa
Calanus simillimus Giesbrecht, 1902 Pe Ep Sa
Canthocalanus pauper (Giesbrecht, 1888) Pe Ep Tr
Cosmocalanus darwinii (Lubbock, 1860) Pe Ep Tr
Mesocalanus tenuicornis (Dana, 1849) Pe Ep T/St/ Sa
Nannocalanus minor (Claus, 1863) Pe Ep Tr/St
Neocalanus gracilis Dana, 1849 Pe Ep Tr/St
Neocalanus tonsus (Brady, 1883) Pe Ep/Me St/Sa
CANDACIIDAE
Candacia bipinnata (Giesbrecht, 1888) Pe Ep/Me
Tr/St
Candacia cheirura Cleve, 1904 Pe Ep/Me St/Sa
Candacia ethiopica (Dana, 1849) Pe Ep/Me Tr
Candacia longimana (Claus, 1863) Pe Ep/Me Tr/St
Candacia pachydactyla (Dana, 1849) Pe Ep/Me St
Candacia tenuimana (Giesbrecht, 1888) Pe Me Tr/St
Paracandacia simplex (Giesbrecht, 1889) Pe Ep T/St
Paracandacia worthingtoni Grice, 1981 Pe Ep Tr
CENTROPAGIDAE
Boeckella delicata Percival, 1937 F Pe
Boeckella dilatata Sars, 1904 F Pe E
Boeckella hamata Brehm, 1928 F Pe E
Boeckella minuta Sars, 1896 F Pe A
Boeckella propinqua Sars, 1904 F Pe
Boeckella symmetrica Sars, 1908 F Pe A
Boeckella tanea Chapman, 1973 F Pe E
Boeckella triarticulata (Thomson, 1883) F Pe
Calamoecia lucasi Brady, 1906 F Pe
Centropages aucklandicus Krämer, 1895 Pe Co Ep
St E
Centropages bradyi Wheeler, 1900 Pe Me Tr/St
Centropages elegans Giesbrecht, 1895 Pe O Ep Tr
Centropages violaceus (Claus, 1863) Pe O Ep Tr
Gladioferens pectinatus (Brady, 1899) B Pe Ep St
Gladioferens spinosus Henry, 1919 B Pe Ep St
CLAUSOCALANIDAE
Clausocalanus arcuicornis (Dana, 1849) Pe Ep Tr/St
Clausocalanus brevipes Frost & Fleminger, 1968 Pe
Ep Sa
Clausocalanus ingens Frost & Fleminger, 1968 Pe Ep
Tr/St/Sa
Clausocalanus jobei Frost & Fleminger, 1968 Pe Ep
St
Clausocalanus laticeps Farran, 1929 Pe Ep Sa
Clausocalanus lividus Frost & Fleminger, 1968 Pe
Ep Tr/St
Clausocalanus parapergens Frost & Fleminger, 1968
Pe Ep Tr/St
Clausocalanus paululus Farran, 1926 Pe Ep Tr/St
Clausocalaus pergens Farran, 1926 Pe Ep St
Ctenocalanus vanus Giesbrecht, 1888 Pe Ep St
Drepanopus pectinatus Brady, 1883 Pe Ep Co Sa
DIAPTOMIDAE A
Sinodiaptomus valkanovi Kiefer, 1938 F Pe A
Skistodiaptomus pallidus (Herrick, 1879) F Pe A
EUCALANIDAE
Eucalanus hyalinus (Claus, 1866) Pe Ep/Me Tr/St
Pareucalanus langae (Fleminger, 1973) Pe Ep Tr
Pareucalanus sewelli (Fleminger, 1973) Pe Ep Tr/St
Rhincalanus gigas Brady, 1883 Pe Ep/Me Sa/Ant
Rhincalanus nasutus Giesbrecht, 1888 Pe Ep/Me St
Rhincalanus rostrifrons (Dana, 1852) Pe Ep Tr
Subeucalanus crassus (Giesbrecht, 1888) Pe Ep Tr/St
Subeucalanus longiceps (Matthews, 1925) Pe Ep Sa
Subeucalanus mucronatus (Giesbrecht, 1888) Pe Ep
Tr
EUCHAETIDAE
Euchaeta acuta Giesbrecht, 1892 Pe Ep Tr/St
Euchaeta media Giesbrecht, 1888 Pe Ep Tr/St
Euchaeta longicornis Giesbrecht, 1888 Pe Ep T/St
Euchaeta rimana Bradford, 1974 Pe Ep T/St
Euchaeta pubera Sars, 1907 Pe Ep T/St
Euchaeta spinosa Giesbrecht, 1892 Pe Me Tr
Pareuchaeta biloba Farran, 1929 Pe Me Sa/Ant
Pareuchaeta bisinuata (Sars, 1907) Pe By Tr/St
Pareuchaeta comosa Tanaka, 1958 Pe By Tr/St
Pareuchaeta exigua (Wolfenden, 1911) Pe By Tr/St
Pareuchaeta hansenii (With, 1915) Pe Me Tr/St
Pareuchaeta pseudotonsa (Fontaine, 1967) Pe By Tr/
St/Sa
Pareuchaeta sarsi (Farran, 1908) Pe By W
HETERORHABDIDAE
Disseta magna Bradford, 1971 Pe By St
Disseta palumbii Giesbrecht, 1889 Pe By Tr/St
Heterorhabdus abyssalis (Giesbrecht, 1889) Pe Me/
By St
Heterorhabdus austrinus Giesbrecht, 1902 Pe Me/
By Sa/Ant
Heterorhabdus caribbeanensis Park, 1970 Pe Me Tr
Heterorhabdus lobatus Bradford, 1971 Pe Me Tr
Heterorhabdus pacificus Brodsky, 1950 Pe By Tr/ St
Heterorhabdus papilliger (Claus, 1863) Pe Ep/me Tr
Heterorhabdus proximus Davis, 1949 Pe Me St
Heterorhabdus robustus Farran, 1908 Pe
Heterorhabdus spinifer Park, 1970 Pe Me Tr
Heterorhabdus spinifrons (Claus, 1863) Pe Me Tr/St
Heterohabdus spinosus Bradford 1971 Pe Me St
Heterostylites longicornis (Giesbrecht, 1889) Pe Me
Tr/St
LUCICUTIIDAE
Lucicutia bicornuta Wolfenden, 1905 Pe Ep/Me Tr/St
Lucicutia clausi (Giesbrecht, 1889) Pe Me Tr/St
Lucicutia curta Farran, 1905 Pe Me W
Lucicutia flavicornis (Claus, 1863) Pe Ep/Me Tr/St
Lucicutia cf. flavicornis, Bradford-Grieve, 1999 Pe
Ep/Me Tr/St
Lucicutia gemina Farran, 1926 Pe Ep/Me Tr
Lucicutia grandis (Giesbrecht, 1895) Pe By W
Lucicutia longiserrata (Giesbrecht, 1889) Pe By Tr
Lucicutia magna Wolfenden in Fowler, 1903 Pe By W
Lucicutia ovalis (Giesbrecht, 1889) Pe Ep/Me Tr
MECYNOCERIDAE
Mecynocera clausi Thompson, 1888 Pe Ep Tr/St
MEGACALANIDAE
Megacalanus longicornis Sars, 1925 Pe By W
METRIDINIDAE
Gaussia princeps T. Scott, 1894 Pe By Tr/St
Metridia brevicauda Giesbrecht, 1889 Pe Me/By Tr/
St
Metridia curticauda Giesbrecht, 1889 Pe Me/By W
Metridia lucens Boeck, 1865 Pe Ep/Me Tr/St/Sa
Metridia princeps Giesbrecht, 1892 Pe By W
Metridia venusta Giesbrecht, 1892 Pe Me/By Tr/ St
Pleuromamma abdominalis (Lubbock, 1856) Pe Me
Tr/St/Sa
Pleuromamma borealis (Dahl, 1893) Pe Me Tr/St/Sa
CRUSTACEA
Pleuromamma gracilis (Claus, 1863) Pe Me Tr/St
Pleuromamma piseki Farran, 1929 Pe Me Tr/St
Pleuromamma quadrungulata (Dahl, 1893) Pe Me
Tr/St/Sa
Pleuromamma robusta (Dahl, 1893) Pe Me Tr/St/Sa
Pleuromamma xiphias Giesbrecht, 1889 Pe Me Tr/St
NULLOSETIGERIDAE
Nullosetigera bidentatus (Brady, 1883) Pe Me W
Nullosetigera helgae (Farran, 1908) Pe Me/By W
PARACALANIDAE
Calocalanus longispinus Shmeleva, 1978 Pe Ep Tr/St
Calocalanus minutus Andronov, 1973 Pe Ep Tr/St
Calocalanus namibiensis Andronov, 1973 Pe Ep Tr/St
Calocalanus neptunus Schmeleva, 1965 Pe Ep Tr/St
Calocalanus pavo (Dana, 1849) Pe Ep Tr/St
Calocalanus plumulosus (Claus, 1863) Pe Ep T/St
Calocalanus styliremis Giesbrecht, 1888 Pe Ep Tr/St
Calocalanus tenuis Farran, 1926 Pe Ep Tr/St
Paracalanus aculeatus Giesbrecht, 1892 Pe Ep Tr/St
Paracalanus indicus Wolfenden, 1905 Pe Ep Tr/St
PHAENNIDAE
Cornucalanus chelifer (I.C. Thompson, 1903) Pe By
Tr/St
Onchocalanus cristatus (Wolfenden, 1904) Pe By T/St
Onchocalanus trigoniceps Sars, 1905 Pe By Tr/St
Neoscolecithrix cf. magna (Grice, 1972) Bp
Neoscolecithrix ornata Bradford-Grieve, 2001 Bp
Phaenna spinifera Claus, 1863 Pe Me T/St
Xanthocalanus penicillatus Tanaka, 1960 Pe By Tr/St
PONTELLIDAE
Calanopia aurivilli Cleve, 1901 Pe O Ep Tr
Labidocera cervi Krämer, 1895 Pe Co Ep St
Labidocera detruncata (Dana, 1849) Pe O Ep Tr
Pontella novaezelandiae Farran, 1929 Pe Co Ep St E
Pontella valida Dana, 1852 Pe O Ep Tr
Pontella whiteleggei Krämer, 1896 Pe O Ep Tr
Pontellina plumata (Dana, 1849) Pe O Ep Tr
Pontellopsis grandis (Lubbock, 1853) Pe O Ep Tr
PSEUDOCYCLOPIDAE
Pseudocyclops n. sp.* Bp Sh
SCOLECITRICHIDAE
Amallothrix arcuata (Sars, 1920) Pe By Tr/St
Amallothrix dentipes (Vervoort, 1951) Pe Me Sa/Ant
Amallothirx emarginata (Farran, 1905) Pe By Tr/St
Amallothrix gracilis (Sars, 1905) Pe By Tr/St
Amallothrix parafalcifer (Park, 1980) Pe By St
Amallothrix pseudopropinqua (Park, 1980) Pe By St
Amallothrix valida (Farran, 1908) Pe By W
Lophothrix frontalis Giesbrecht, 1895 Pe By Tr/St
Lophothrix latipes (T. Scott, 1894) Pe Me Tr
Scaphocalanus affinis (Sars, 1905) Pe By W
Scaphocalanus brevicornis (Sars, 1900) Pe Me Tr/St
Scaphocalanus curtus (Farran, 1926) Pe Ep Tr
Scaphocalanus echinatus (Farran, 1905) Pe Ep Tr/
St/Sa
Scaphocalanus longifurca (Giesbrecht, 1888) Pe Me
Tr/St
Scaphocalanus magnus (T. Scott, 1894) Pe By W
Scaphocalanus major (T. Scott, 1894) Pe Me Tr/St
Scaphoclanaus subbrevicornis (Wolfenden, 1911) Pe
Me W
Scolecithricella abyssalis (Giesbrecht, 1888) Pe Me
Tr/St
Scolecithricella dentata (Giesbrecht, 1892) Pe Me
Tr/St
‘Scolecithricella’ fowleri (Farran, 1926) Pe Me Tr
Scolecithricella minor (Brady, 1883) Pe Ep W
Scolecithricella ovata (Farran, 1905) Pe Me W
Scolecithricella schizosoma Park, 1980 Pe By Sa/Ant
Scolecithricella vittata (Giesbrecht, 1892) Pe Me Tr/
St
Scolecithrix bradyi Giesbrecht, 1888 Pe Ep Tr
Scolecithrix danae (Lubbock, 1856) Pe Ep Tr
Scopalatum sp. Bradford et al. 1983 Pe Me St
Scottocalanus helenae (Lubbock, 1856) Pe Me Tr/St
213
NEW ZEALAND INVENTORY OF BIODIVERSITY
Scottocalanus securifons (T. Scott, 1894) Pe By Tr/ St
Scottocalanus terranovae Farran, 1929 Pe By St
Scottocalanus thorii With, 1915 Pe By Tr/St
SPINOCALANIDAE
Spinocalanus longicornis Sars, 1900 Pe By W
Spinocalanus spinosus Farran, 1908 Pe By Tr
STEPHIDAE
Stephos angulatus Bradford-Grieve, 1999 Bp Sh E
Stephos hastatus Bradford-Grieve, 1999 BP Sh E
SULCANIDAE
Sulcanus conflictus Nicholls, 1945 B Pe Co Ep A?
TEMORIDAE
Temora turbinata (Dana, 1849) Pe Co Ep S/St
Temoropia minor Deevey, 1972 Pe By Tr
Gen. et sp. indet.* Bp Sh
THARYBIDAE
Tharybis inaequalis Bradford-Grieve, 2001 Ba Bp
Tharybis spp. (2)* Bp Sh
Undinella brevipes Farran, 1908 Pe Me Tr/St
Order CYCLOPOIDA
ASCIDICOLIDAE
Botryllophilus cf. banyulensis Brément, 1909*
Enteropsis onychophorus Schellenberg, 1922 P
(tunicates)
Haplostoma gibberum (Shellenberg, 1922) P
(tunicates)
Haplostomides otagoensis Ooishi, 2001 P (tunicates)
BOMOLOCHIDAE
Acanthocolax sp. Beresford 1991 P (fish)
Pseudoeucanthus australiensis Roubal, Armitage &
Rohde, 1983* P (fish)
Pseudoeucanthus uniserratus Wilson, 1913 P (fish)
Unicolax chrysophryenus Roubal, Armitage &
Rohde, 1983 P (fish)
CHITONOPHILIDAE
Cocculinika myzorama Jones & Marshall, 1986 P
(molluscs)
CHONDRACANTHIDAE
Acanthochondria incisa Shiino, 1955 P (fish)
Chondracanthodes radiatus Müller, 1777 P (fish)
Chondracanthus australis Ho, 1991 P (fish)
Chondracanthus distortus Wilson, 1922 P (fish)
Chondracanthus genypteri Thomson, 1890 P (fish)
Chondracanthus lotellae Thomson, 1890 P (fish)
Chondracanthus yanezi Atria, 1980 P (fish)
Mecaderochondria pilgrimi Ho & Dojiri, 1987 P
(fish)
Prochondracanthus platycephali Ho, 1975 P (fish)
Pseudochondracanthus chilomycteri (Thomson, 1890)
P (fish)
CLAUSIDIIDAE
Hemicyclops? n. sp., n. gen.? * Be C
Teredicola typicus Wilson, 1942 P (boring molluscs)
CORYCAEIDAE
Corycaeus agilis Dana, 1849* Pe Ep Tr/St
Corycaeus aucklandicus Kramer, 1895 Pe Ep Co E
Corycaeus clausi F. Dahl, 1894* Pe Ep Tr/St
Corycaeus crassiusculus Dana, 1849* Pe Ep Tr/St
Corycaeus flaccus Giesbrecht, 1891* Pe Ep Tr/St
Corycaeus furcifer Claus, 1863* Pe Ep Tr/St
Corycaeus latus Dana, 1849* Pe Ep Tr/St
Corycaeus limbatus Brady, 1883* Pe Ep Tr/St
Corycaeus longistylis Dana, 1849* Pe Ep Tr
Corycaeus speciosus Dana, 1849* Pe Ep Tr/St
Corycaeus typicus Krøyer, 1849* Pe Ep Tr
Farranula rostata (Claus, 1863)* Pe Ep S/St
CYCLOPIDAE
Abdiacyclops cirratus Karanovic, 2005 F E
Acanthocyclops robustus (Sars, 1863) F Be A?
Acanthocyclops vernalis (Fischer, 1853) F Pe
Cyclops? strennus strennus Fischer, 1851 P
Diacyclops bicuspidatus (Claus, 1857) F Be
Diacyclops bisetosus (Rehberg, 1880) F Be A?
Eucyclops serrulatus (Fischer, 1851) F Pe A?
214
Euryte? longicauda Philippi, 1843 Be
Goniocyclops silvestris Harding, 1958 F Ph E
Halicyclops? magniceps (Lilljeborg, 1853) B Be
Halicyclops? neglectus Kiefer, 1935 F/B Be/Pe
Macrocyclops albidus (Jurine, 1820) F Be
Mesocyclops? australiensis (Sars, 1908) F Pe
Mesocyclops? leuckarti (Claus, 1857) F Pe
Metacyclops monacanthus (Kiefer, 1928) B Pe E
Microcyclops? varicans Sars, 1863 F Be/Pe
Paracyclops chiltoni (Thomson, 1883) F Be
Paracyclops fimbriatus (Fischer, 1853) F/B Be A?
Paracyclops waiariki Lewis, 1974 F Be E
Tropocyclops? prasinus (Fischer, 1860) F Be/Pe
Zealandcyclops fenwicki Karanovic, 2005 F E
Zealandcyclops haywardi Karanovic, 2005 F E
ERGASILIDAE
Abergasilus amplexus Hewitt, 1978 B P (fish)
Paeonodes nemaformis Hewitt, 1969 F P (fish,
extinct?) E
Thersitina inopinata Percival, 1937 F Pe P (fish,
extinct?)
LERNAEIDAE
Lernaea cyprinacea Linnaeus, 1758 F P (fish) A
LICHOMOLGIDAE
Lichomolgidium tupuhiae Jones, 1975 C (molluscs)
Lichomolgus uncus Jones, 1976 C (molluscs)
MYTILICOLIDAE
Pseudomyicola spinosus (Raffaele & Monticelli,
1885) C (molluscs)
NOTODELPHYIDAE
Pygodelphys novaeseelandius (Shellenberg, 1922) C
(tunicates)
Doropygus globosus Jones, 1974 C (tunicates)
Doropygus louisae Jones, 1980 C (tunicates)
Doropygus platythorax Jones, 1974 C (tunicates)
Doropygus pulex Shellenberg, 1922 C (tunicates)
Doropygus spinosus Jones, 1980 C (tunicates)
Doropygus trisetosus Shellenberg, 1922 C (tunicates)
Ophioseides schellenbergi Jones, 1980 C (tunicates)
OITHONIDAE
Oithona atlantica Farran, 1908 Pe Ep St
Oithona nana Giesbrecht, 1892 Pe Ep Tr/St
Oithona plumifera Baird, 1843 Pe Ep Tr/St
Oithona similis Claus, 1866 Pe Ep W
ONCAEIDAE
Conaea rapax Giesbrecht, 1891 Pe Me W
Lubbockia aculeata Giesbrecht, 1891 Ep/Me Tr/St
Lubbockia squillimana Claus, 1863 Pe Ep/Me Tr/St
Oncaea antarctica Heron, 1977 Pe Ep/Me Sa/Ant
Oncaea conifera Giesbrecht, 1891 Pe Ep/Bap Tr/St
Oncaea derivata Heron & Bradford-Grieve, 1995 Pe
Me Tr/St
Oncaea englishi Heron, 1977 Pe Ep/Bap W
Oncaea furcula Farran, 1936 Pe Me Tr/St
Oncaea inflexa Heron, 1977 Pe Ep/Me Sa
Oncaea media Giesbrecht, 1891 Pe Ep/Me Tr/St
Oncaea mediterranea (Claus, 1863) Pe Ep/Me W
Oncaea quadrata Heron & Bradford-Grieve, 1995
Pe Ep/Me St
Oncaea redacta Heron & Bradford-Grieve, 1995 Pe
Ep/Me Tr
Oncaea scottodicarloi Heron & Bradford-Grieve,
1995 Pe Ep Tr/St
Oncaea similis Sars, 1918 Pe Ep/Me St
Oncaea venusta Philippe, 1843 Pe Ep/Me Tr/St
PHILICHTHYIDAE
Philichthys xiphiae Steenstrup, 1862 P (fish)
Sarcotaces sp. Avdeev & Avdeev 1975 P (fish)
SAPPHIRINIDAE
Copilia hendorffi Dahl, 1892* Pe Ep Tr/St
Copilia mirabilis Dana, 1849* Pe Ep/Me Tr/St
Copilia vitrea (Haeckel, 1864)* Pe Ep/Me Tr
Sapphirina angusta Dana, 1849* Pe Ep Tr/St
Sapphirina autonitens-sinuicauda Lehnhofer, 1929*
Pe Ep Tr/St
Sapphirina ovatolanceolata-gemma Lehnhofer, 1929*
Pe Ep Tr/St
Sapphirina intestinata Giesbrecht, 1891* Pe Ep T/St
Sapphirina iris Dana, 1849* Pe Ep Tr/St
Sapphirina opalina-darwini Lehnhofer, 1929* Pe
Ep Tr/St
Sapphirina sali Farran, 1929* Pe Ep St
Sapphirina scarlata Giesbrecht, 1891* Pe Ep T/St
THAMNOMOLGIDAE
Thamnomolgus eurycephalus Humes & Kiss, 2004 P
(black coral)
Order MORMONILLOIDA
Mormonilla phasma Giesbrecht, 1891* Pe
Order HARPACTICOIDA
AEGISTHIDAE
Aegisthus mucronatus Giesbrecht, 1891 Pe
AMEIRIDAE
Ameira minuta Boeck, 1864 Ph
Ameira parvula (Claus, 1866) Ph BeL
Ameira sp.* BeL
Ameiropsyllus (?) spp. (5)* BeL
Leptameira sp.* BeL
Nitocra fragilis Sars, 1905 Ch B Be
Nitocra sp. (2)* BeL
Parapseudoleptomesochra (?) sp.* BeL
Parevansula sp.* BeL
Psyllocamptus sp.* BeL
ANCORABOLIDAE
Laophontodes hamatus (Thomson, 1883) Ph E
Laophontodes whitsoni T. Scott, 1912 Ca Be
Paralaophontodes sp.* BeL
ARENOPONTIIDAE
Arenopontia sp.* BeL
CANTHOCAMPTIDAE
Antarctobiotus australis Lewis, 1972 F Ph(M) E
Antarctobiotus diversus Lewis, 1972 F Ph(M) E
Antarctobiotus elongatus Lewis, 1972 F Ph(M) E
Antarctobiotus exiguus Lewis, 1972 F Ph(M) E
Antarctobiotus ignobilis Lewis, 1972 F Ph(M) E
Antarctobiotus triplex Lewis, 1972 F Ph(M) E
Antarctobiotus n. sp.* F Ph(M)
Antipodiella chappuisi Brehm, 1928* F Ph(M)
Antipodiella n. spp. (3)* 3F Ph(M)
Attheyella (Chappuisiella) fluviatilis Lewis, 1972 F
Ph(M) E
Attheyella (C.) maorica (Brehm, 1928) F Ph(M) E
Attheyella (C.) rotoruensis Lewis, 1972 F Pe E
Attheyella (Delachauxiella) bennetti Brehm, 1927 F
Ph(M) E
Attheyella (D.) brehmi Kiefer, 1928 F Ph(M) E
Attheyella (D.) humidarum Lewis, 1972 F Ph(M) E
Attheyella (D.) stillicidarum Lewis, 1972 F Ph(M) E
Bryocamptus (Rheocamptus) pygmaeus (Sars, 1862)*
F Ph(M)
Bryocamptus (Echinocamptus) stouti Harding, 1958 T
(forest litter) E
Bryocamptus n. spp. (3)* 3F
Elaphoidella bidens coronata Sars, 1904 F BeL
Elaphoidella silvestris Lewis, 1972 F Ph(M) E
Elaphoidella sp.* F Be
Epactophanes richardi Mrázek, 1893 F Ph, Ph(M)
Loeflerella n. sp.* F Ph(M)
Mesochra flava Lang, 1933 Ph
Mesochra meridionalis Sars, 1905 B
Mesochra parva Thomson, 1946 B BeL BeSL
Mesochra pygmaea (Claus, 1863)* BeL
Mesochra spp. (2)* BeL
Gen. nov. (2) et n. spp. (7)* 7F
CANUELLIDAE
Brianola sp.* B BeL
CLETODIDAE
Enhydrosoma variabile Wells, Hicks & Coull, 1982
BeL BeSL E
PHYLUM ARTHROPODA
Enhydrosoma spp. (2)* BeL
Enhydrosomella spp. (2)* BeL
Stylicletodes longicaudatus (Brady & Robertson,
1880) Ph
Stylicletodes sp.* BeL
DACTYLOPUSIIDAE
Dactylopusia frigida T. Scott, 1912 Ph
Dactylopusia tisboides (Claus, 1863) Ph BeL BeSL
Diarthrodes cystoecus Fahrenbach, 1954 Ph
Diarthrodes novaezealandiae Thomson, 1882 Ph E
Diarthrodes sp.* Ph
Paradactylopodia brevicornis (Claus, 1866) Ph
Paradactylopodia trioculata Hicks, 1988 Ph(W) E
DARCYTHOMPSONIIDAE
Gen. nov. et n. sp. Huys & Gee in press* BeL
ECTINOSOMATIDAE
Arenosetella sp. * BeL
Ectinosoma melaniceps Boeck, 1864 Ca Ch BeL
Ectinosoma sp.* BeL
Glabrotelson spp. (3)* BeL
Halectinosoma hydrofuge Wells, Hicks & Coull, 1982
BeL E
Halectinosoma otakoua Wells, Hicks & Coull, 1982
BeL E
Halectinosoma spp. (3)* BeL
Kliella (?) sp.* BeL
Microsetella norvegica (Boeck, 1864) Pe Ep W
Microsetella rosea (Dana, 1848) Pe Ep W
Noodtiella sp. * BeL
HARPACTICIDAE
Harpacticus furcatus Lang, 1936 Ph
Harpacticus glaber Brady, 1899 Pe SL E
Harpacticus pulvinatus Brady, 1910 Ph
Harpacticus spp. (2)* Ph
Perissocope litoralis Lang, 1934 Ph E
Tigriopus angulatus Lang, 1933 Ca Sn Ph
Tigriopus raki Bradford, 1967 Ph E
Zaus sp.* Ph
Zausopsis contractus (Thomson, 1883) Ph E
Zausopsis mirabilis Lang, 1934 Ph E
LAOPHONTIDAE
Afrolaophonte sp.* BeL
Apolethon sp.* BeL
Folioquinpes chathamensis (Sars, 1905) B E
Harrietella simulans (T. Scott, 1894) Ph(W)
Heterolaophonte campbelliensis (Lang, 1934) Ca Ph
Heterolaophonte tenuispina (Lang, 1934) Ca Ph
Klieonychocamptoides sp.* BeL
Laophonte australasica Thomson, 1883 E
Laophonte cornuta Philippi, 1840 Ca Ph
Laophonte elongata barbata Lang, 1934 Ph
Laophonte inornata A. Scott, 1902 Ph
Laophonte lignosa Hicks, 1988 Ph(W) E
Laophonte sima Gurney, 1927 Ph
Laophonte spp. (2)* BeL
Onychocamptus mohammed (Blanchard & Richard,
1891) B
Paeudonychocamptus sp.* BeL
Paralaophonte aenigmaticum Wells, Hicks & Coull,
1982 BeL E
Paronychocamptus exiguus (Sars, 1905) B E
Paralaophonte meinerti (Brady, 1899) Ca Ph
Paralaophonte spp. (4)* BeL
Pseudolaophonte spp. (2)* BeL
Quinquelaophonte candelabrum Wells, Hicks &
Coull, 1982 BeL BeSL Ph E
Quinquelaophonte longifurcata (Lang, 1965) Ph
Quinquelaophonte sp.* BeL
Xanthilaophonte trispinosa (Sewell, 1940) BeL BeSL
LEPTASTACIDAE
Leptastacus sp.* BeL
LOURINIIDAE
Lourinia armata (Claus, 1866) Ph
MIRACIIDAE
Amonardia perturbata Lang, 1965 Ph
Amphiascoides nichollsi Lang, 1965 Ph
Amphiascoides sp.* BeL
Amphiascopsis cinctus (Claus, 1866) Ph
Amphiascopsis southgeorgiensis (Lang, 1936) Ph
Amphiascus waihonu (Hicks, 1986) Be (C?) E
Bulbamphiascus imus (Brady, 1872) Ph
Bulbamphiascus spp. (2)* BeL
Cladorostrata sp.* BeL
Delavalia spp. (3)* BeL
Helmutkunzia sp.* BeL
Macrosetella gracilis (Dana, 1847) Pe Ep Tr/St
Metamphiascopsis monardi (Lang, 1934) Ph E
Miscegenus heretaunga Wells, Hicks & Coull, 1982
BeL BeSL E
Miscegenus spp. (2)* BeL
Oculosetella gracilis (Dana, 1849) Pe Ep Tr/St
Pseudostenhelia sp.* BeL
Robertgurneya sp.* BeL
Robertsonia propinqua (T. Scott, 1893) Ph
Sarsamphiascus hirtus (Gurney, 1927) Ca Ph
Sarsamphiascus lobatus (Hicks, 1971)
Sarsamphiascus pacificus (Sars, 1905) Ch Ph
Sarsamphiascus tainui (Hicks, 1989) W E
Sarsamphiascus spp. (2)* BeL
Schizopera clandestina (Klie, 1924) B
Schizopera longicauda Sars, 1905 Ch B Be
Schizopera sp.* BeL
Stenhelia xylophila Hicks, 1988 Ph(W) E
Stenhelia sp. BeL
Teissierella (?) sp.* BeL
Typhlamphiascus unisetosus Lang, 1965 Ph
Typhlamphiascus sp.*BeL
NANNOPODIDAE
Gen. et sp. indet.* BeL
NORMANELLIDAE
Normanella incerta Lang, 1934 Ph E
ORTHOPSYLLIDAE
Orthopsyllus linearis (Claus, 1866) Ph
PARAMESOCHRIDAE
Apodopsyllus sp.* BeL
Diarthrodella sp.* BeL
Emertonia sp.* BeL
PARASTENHELIIDAE
Parastenhelia hornelli Thompson & A. Scott, 1903
BeL
Parastenhelia megarostrum Wells, Hicks & Coull,
1982 BeL BeSL E
Parastenhelia spinosa (Fischer, 1860) CaPh BeL
BeSL
Parastenhelia sp.* BeL
PELTIDIIDAE
Alteutha depressa (Baird, 1837) Ph
Alteutha novaezealandiae (Brady, 1899) Ph E
Alteuthoides kootare Hicks, 1986 C (sponges) E
Clytemnestra rostrata (Brady, 1883) Pe Ep/Me Tr/St
Clytemnestra scutellata Dana, 1848 Pe Ep/Me Tr/St
Eupelte regalis Hicks, 1971 Ph E
Neopeltopsis pectinipes Hicks, 1976 Ph E
PHYLLOGNATHOPODIDAE
Phyllognathopus viguieri (Maupas, 1892) F Ph(M)
Phyllognathopus volcanicus Barclay, 1969 F BeL BeS
Ph E
PORCELLIDIIDAE
Dilatatiocauda dilatatum (Hicks, 1971) Ph E
Porcellidium erythrum Hicks, 1971 Ph E
Porcellidium fulvum Thomson, 1883 Ph E
Porcellidium interruptum Thomson, 1883 Ph E
Porcellidium tapui Hicks & Webber, 1983 C E
(hermit crabs)
PSAMMOPSYLLIDAE
Psammopsyllus sp.* BeL
PSEUDOTACHIDIIDAE
Dactylopodella flava (Claus, 1866) Ph(W)
Dactylopodella janetae Hicks, 1989 Ph(W) E
Dactylopodella sp.* Ph
CRUSTACEA
Danielssenia sp.* Be L
Donsiella bisetosa Hicks, 1988 Ph(W) E
Paranannopus sp.* BeL
Pseudomesochra sp.* BeL
Pseudonsiella aotearoa Hicks, 1988 Ph(W) E
Xouthous intermedia (Lang, 1934) Ph E
Xouthous novaezealandiae (Thomson, 1882) Ph E
Xylora bathyalis Hicks, 1988 Ph(W) E
Xylora neritica Hicks, 1988 Ph(W)E
RHIZOTHRICIDAE
Rhizothricidae sp.* BeL
RHYNCHOTHALESTRIDAE
Rhynchothalestris campbelliensis Lang, 1934 Ph E
TACHIDIIDAE
Euterpina acutifrons (Dana, 1848) Pe Ep W
Geeopsis incisipes (Klie, 1913) B
Tachidius sp.* BeL
TEGASTIDAE
Syngastes clausii (Thomson, 1883) Ph E
TETRAGONICIPITIDAE
Phyllopodopsyllus minor (Thompson & A. Scott,
1903) Ph
Phyllopodopsyllus sp.* BeL
THALESTRIDAE
Flavia crassicornis Brady, 1899 E
Thalestris australis Brady, 1899 Ph? E
Thalestris ciliata Brady, 1899 Ph? E
TISBIDAE
Scutellidium armatum (Wiborg, 1964) Ph
Scutellidium idyoides (Brady, 1883) Ph?
Scutellidium macrosetum Branch, 1975 Ph
Scutellidium plumosum Brady, 1899 Ca Ph BeL
Scutellidium ringueleti Pallares, 1969 Ph
Tisbe furcata (Baird, 1837) Ch Ph
Tisbe gurneyi (Lang, 1934) Ph E
Tisbe holothuriae Humes, 1957 Ph
Tisbe sp.* Ph
Order SIPHONOSTOMATOIDA
ARTOTROGIDAE
Artotrogus gordoni Kim, 2009 E (bryozoan)
ASTEROCHERIDAE
Cecidomyzon conophorae Stock, 1981 P (coral) E
Cystomyzon dimerum Stock, 1981 P (coral) E
Oedomyzon tripodum Stock, 1981 P (coral) E
CANCERILLIDAE
Cancerilla neozelandica Stephensen, 1927 P
(brittlestars) E
CALIGIDAE
Caligus aesopus Wilson, 1921 P (fish)
Caligus bonito Wilson, 1905 P (fish)
Caligus brevis Shiino, 1954 P (fish)
Caligus buechlerae Hewitt, 1964 P (fish) E
Caligus coryphaenae Steenstrup & Lütken, 1861 P
(fish)
Caligus elongatus Nordmann, 1832 P (fish)
Caligus epidemicus Hewitt, 1971 P (fish)
Caligus kahawai Jones, 1988 P (fish) E
Caligus lalandei Barnard, 1948 P (fish)
Caligus longicaudatus Brady, 1899 P (fish) E
Caligus pelamydis Krøyer, 1863 P (fish)
Caligus productus Dana, 1852 P (fish) ?
Caligus sp. 1 Sharples & Evans 1995 P (fish)
Caligus sp. 2 Sharples & Evans 1995 P (fish)
Dentigryps sp.* P (fish)
Lepeophtheirus argentus Hewitt, 1963 P (fish) E
Lepeophtheirus crassus Wilson & Bere, 1936 P (fish)
Lepeophtheirus distinctus Hewitt, 1963 P (fish) E
Lepeophtheirus erecsoni Thomson, 1891 P (fish) E
Lepeophtheirus hastus Shiino, 1960 P (fish)
Lepeophtheirus heegaardi Hewitt, 1963 P (fish)
Lepeophtheirus histiopteridi Kazachenko, Korotaeva
& Kurochkin, 1972 P (fish) E
Lepeophtheirus nordmanni (Edwards, 1840) P (fish)
Lepeophtheirus polyprioni Hewitt, 1963 P (fish) E
215
NEW ZEALAND INVENTORY OF BIODIVERSITY
Lepeophtheirus scutiger Shiino, 1952 P (fish)
Lepeophtheirus sekii Yamaguti, 1936 P (fish)
Lepeophtheirus sp.* P (fish)
CECROPIDAE
Cecrops latreillei Leach, 1816 P (fish)
DICHELESTHIIDAE
Anthosoma crassum (Abildgaard, 1794) P (fish)
ENTOMOLEPIDAE
Entomolepis ovalis Brady, 1899 E
EUDACTALINIDAE
Eudactylina acanthii Scott, 1901 P (fish)
Jusheyus shogunus Deets & Benz, 1987 P (fish)
Nemesis lamma lamma Risso, 1826 P (fish)
Nemesis l. vermi Scott, 1929 P (fish)
Nemesis robusta (van Beneden, 1851) P (fish)
EURYPHORIDAE
Euryphorus brachypterus (Gerstaecker, 1853) P (fish)
Euryphorus nordmanni Milne-Edwards, 1840 P
(fish)
Gloiopotes huttoni (Thomson, 1890) P (fish)
HATSCHEKIIDAE
Congericola kabatai Hewitt, 1975 P (fish) E
Hatschekia conifera Yamaguti, 1939 P (fish)
Hatschekia crenata Hewitt, 1969 P (fish) E
Hatschekia pagrosomi Yamaguti, 1939 P (fish)
Hatschekia quadrata Hewitt, 1969 P (fish) E
Hatschekia squamata Jones & Cabral, 1990 P E (fish)
HERPYLLOBIIDAE
Herpyllobius rotundus Lutzen & Jones, 1976 P
(polychaete) E
KROYERIIDAE
Kroyeria carchariaeglauci Hesse, 1897* P (shark)
Kroyeria cf. lineata P (fish)
LERNAEOPODIDAE
Albionella sp.* P (fish)
Alella tarakihi Hewitt & Blackwell, 1987 P (fish) E
Brachiella thynni Cuvier, 1830 P (fish)
Brachiella sp.* P (fish)
Charopinus parkeri (Thomson, 1816) P (fish)
Clavella zini Kabata, 1979 P (fish) E
Clavella sp.* P (fish)
Clavellodes sp. Vooren & Tracey 1976 P (fish)
Clavellopsis sargi (Kurz, 1877) P (fish)
Dendrapta sp. Jones, 1988 P (fish)
Lernaeopoda musteli Thomson, 1890 P (fish) E
Lernaeopoda sp. *B. Jones unpubl. P (fish)
Naobranchia sp. Pilgrim 1985 P (fish)
Parabrachiella amphipacifica Ho, 1982 P (fish)
Parabrachiella insidiosa f. lageniformes (Heller, 1865)
P (fish)
Parabrachiella sp. Pilgrim 1985 P (fish)
Pseudocharopinus bicaudatus (Kroyer, 1837) P (fish)
Schistobrachia pilgrimi Kabata, 1988 P (fish) E
Vanbenedenia sp. P (fish)
LERNANTHROPIDAE
Aethon garricki Hewitt, 1968 P (fish) E
Aethon morelandi Hewitt, 1968 P (fish)
Aethon percis (Thomson, 1890) P (fish) E
Lernanthropus microlamini Hewitt, 1968 P (fish) E
Lernanthropus sp.* P (fish)
Sagum foliaceus (Goggio, 1905) P (fish)
NICOTHOIDAE
Rhizorhina seriolis Green, 1959 P (isopod) E
Sphaeronella bradfordae Boxshall & Lincoln, 1983 P
(isopod) E
Sphaeronella serolis Monod, 1930 P (isopod) E
Sphaeronellopsis littoralis Hansen, 1905 P (ostracod)
E
PANDARIDAE
Demoleus latus Shiino, 1954 P (fish)
Dinemoura latifolia Steenstrup & Lütken, 1861 P
(fish)
Dinemoura producta (Müller, 1785) P (fish)
Echthrogaleus denticulatus Smith, 1874 P (fish)
Echthrogaleus coleoptratus (Gúerin-Meneville, 1837)
216
P (fish)
Nesippus orientalis Heller, 1865 P (fish)
Nogagus borealis (Steenstrup & Lütken, 1861) P
(fish)
Pandarus bicolor Leach, 1816 P (fish)
Pandarus satyrus Dana, 1852 P (fish)
Perissopus dentatus Steenstrup & Lütken, 1861 P
(fish)
Phyllothyreus cornutus (Edwards, 1840) P (fish)
PENNELLIDAE
Cardiodectes bellotti (Richiardi, 1882) P (fish)
Pennella histiophori Thomson, 1890 P (fish)
Trifur lotellae Thomson, 1890 P (fish)
PSEUDOCYCNIDAE
Pseudocycnus appendicualatus Heller, 1868 P (fish)
SPHYRIIDAE
Lophoura laticervix Hewitt, 1964 P (fish)
Lophoura spp. *B. Jones unpubl. P (fish)
Periplexis antarcticensis Hewitt, 1965 P (fish)
Sphyrion laevigatum (Quoy & Gaimard, 1824) P
(fish)
Sphyrion lumpi (Kroyer, 1845) P (fish)?
Sphyrion quadricornis Gavevskaya & Kovaleva, 1984
P (fish)
Order MONSTRILLOIDA
MONSTRILLIDAE?
Monstrilla sp.* P
Class OSTRACODA
Order PALAEOCOPIDA
Suborder BEYRICHICOPIDA
PUNCIIDAE
Manawa staceyi Swanson, 1989 E
Manawa tryphena Hornibrook, 1949 E
Puncia novozealandica Hornibrook, 1949 E
Order PODOCOPIDA
Suborder PODOCOPINA
BAIRDIIDAE
Bairdoppilata kerryi Milau, 1993
Bairdoppilata villosa (Brady, 1880)
Bairdoppilata sp. Swanson 1979
Neonesidea amygdaloides (Brady, 1880)
Neonesidea crosskeiana (Brady, 1886)
Neonesidea fusca (Brady, 1880)
Neonesidea ovata (Bosquet, 1853)
Neonesidea sp. Ayress 1993
BYTHOCYPRIDIDAE
Orlovibairdia arcaforma (Swanson, 1979) E
Orlovibairdia aff. angulata (Brady, 1870)
Orlovibairdia aff. fumata (Brady, 1890)
Orlovibairdia sp. Swanson 1979
BYTHOCYTHERIDAE
Baltraella cf. peterroyi Yassini & Jones, 1995
Bythocythere arenacea Brady, 1880
Bythocythere bulba Swanson, 1979
Bythoceratina decepta Hornibrook, 1952
Bythoceratina edwardsoni Hornibrook, 1952
Bythoceratina fragilis Hornibrook, 1952
Bythoceratina hornibrooki Jellinek & Swanson, 2003
Bythoceratina maoria Hornibrook, 1952
Bythoceratina mestayerae Hornibrook, 1952
Bythoceratina powelli Hornibrook, 1952
Bythoceratina tuberculata Hornibrook, 1952
Bythoceratina utilazea Hornibrook, 1952
Microceratina quadrata Swanson, 1980
Miracythere novaspecta Hornibrook, 1952 E
Miracythere speciosa Jellinek & Swanson, 2003 E
CYPRIDIDAE
Candona aotearoa Chapman, 1963 F E
Candona inexpecta Chapman, 1963 F E
Candonocypris assimilis Sars, 1894 F
Candonocypris novaezelandiae (Baird in White &
Doubleday, 1843) F E
Cypretta turgida (Sars, 1896) F E
Cypretta viridis (Thomson, 1879) F
Cyprinotus flavescens Brady, 1898 F E
Cyprinotus sarsi Brady, 1898 F E
Cypris kaiapoiensis Chapman, 1963 F E
Diacypris thomsoni (Chapman, 1963) F E
Eucypris lateraria (King, 1855) F
Eucypris sanguineus (Chapman, 1963) F E
Eucypris virens (Jurine, 1820) F A
Herpetocypris pascheri Brehm, 1929 F E
Heterocypris incongruens (Rhamdohr, 1808) F E
Ilyodromus stanleyanus (King, 1855) F
Ilyodromus obtusus Sars, 1894 F E
Ilyodromus smaragdinus Sars, 1894 F
Ilyodromus subsriatus Sars, 1894 F E
Ilyodromus varrovillius (King, 1855) F
Mesocypris insularis (Chapman, 1963) F E
Paracypria tenuis (Sars, 1905) F
Potamocypris sp. Hornibrook, 1955 F
Scottia audax (Chapman, 1961) T E
CYPRIDOPSIDAE
Cypridopsis obstinata Barclay, 1968 F E
Cypridopsis vidua (Müller, 1776) F A
Pleisiocypridopsis jolleae (Chapman, 1963) F E
Prionocypris marplesi Chapman, 1963 F E
CYTHERALISONIDAE
Cytheralison fava (Hornibrook, 1952) E
Cytheralison tehutui Jellinek & Swanson, 2003 E
Cytheralison sp. Jellinek & Swanson 2003
Debissonia fenestrata Jellinek & Swanson, 2003 E
Debissonia pravacauda (Hornibrook, 1952) E
Debissonia sp. Jellinek & Swanson 2003
CYTHERIDAE
Loxocythere crassa Hornibrook, 1952
Loxocythere hornibrooki McKenzie, 1967
Loxocythere kingi Hornibrook, 1952
Loxocythere sp. Hornibrook 1952
CYTHERIDEIDAE
Cytheridea aoteana Hornibrook, 1952 E
Hemicytheridea mosaica Hornibrook, 1952
Pseudeucythere sp. Jellinek & Swanson 2003
Pseudocythere (Pseudocythere) caudata Sars, 1866
Pseudocythere (Plenocythere) fragilis Swanson, 1979
Rotundracythere gravepuncta Hornibrook, 1952
Rotundracythere cf. gravepunctata Hornibrook, 1952
Rotundracythere inaequa Hornibrook, 1952
Rotundracythere mytila Hornibrook, 1952
Rotundracythere nux Jellinek & Swanson, 2003 E
Rotundracythere rotunda Hornibrook, 1952
Rotundracythere subovalis Hornibrook, 1952
Rotundracythere sp. A Jellinek & Swanson 2003
Rotundracythere sp. B Jellinek & Swanson 2003
Rotundracythere sp. C Jellinek & Swanson 2003
Rotundracythere sp. D Jellinek & Swanson 2003
Rotundracythere sp. E Jellinek & Swanson 2003
CYTHERURIDAE
Aversovalva aurea Hornibrook, 1952
Aversovalva sp. Ayress 1995
Cytheropteron anisovalva Ayress, Correge, Passlow
& Whatley, 1996
Cytheropteron confusum (Hornibrook, 1952)
Cytheropteron curvicaudum Hornibrook, 1952
Cytheropteron dividentum (Hornibrook, 1952)
Cytheropteron dorsocorrugatum Ayress, Correge,
Passlow & Whatley, 1996
Cytheropteron fornix (Hornibrook, 1952)
Cytheropteron hikurangiensis Swanson & Ayress,
1999 E
Cytheropteron latiscalpum Hornibrook, 1952
Cytheropteron obtusalum Hornibrook, 1952
Cytheropteron terecaudum Hornibrook,
Cytheropteron vertex Hornibrook, 1952
Cytheropteron wellingtoniense Brady, 1880
Cytheropteron wellmani Hornibrook, 1952
Cytheropteron willetti Hornibrook, 1952
PHYLUM ARTHROPODA
Cytheropteron sp. Ayress 1993 ?Rec
Cytheropteron sp. Hartmann 1982
Cytherura clausi Brady, 1880
Eucytherura boomeri Ayress, Whatley, Downing &
Millson, 1995
Eucytherura calabra (Colalongo & Pasini, 1980)
Eucytherura multituberculata Ayress, Whatley,
Downing & Millson, 1995
Eucytherura? anoda Ayress, Whatley, Downing, &
Millson, 1995
Hemicytherura (Hemicytherura) aucklandica
Hornibrook, 1952
Hemicytherura (H.) delicatula Hornibrook, 1952
Hemicytherura (H.) fereplana Hornibrook, 1952
Hemicytherura (H.) gravis Hornibrook, 1952
Hemicytherura (H.) pandorae Hornibrook, 1952
Hemicytherura (H.) pentagona Hornibrook, 1952
Hemicytherura (H.) quadrazea Hornibrook, 1952
Hemicytherura (Kangarina) radiata (Hornibrook,
1952)
Microcytherura hornibrooki (McKenzie, 1967)*
Microcytherura (Elofsonia) sp. Hayward 1981
Oculocytheropteron acutangulum (Hornibrook, 1952)
Oculocytheropteron confusum (Hornibrook, 1952)
Oculocytheropteron improbum (Hornibrook, 1952)
Pterygocythere mucronalata (Brady, 1880)
Semicytherura arteria Swanson, 1979
Semicytherura cf. costellata (Brady, 1880)
Semicytherura hexagona (Hornibrook, 1952
Semicytherura sericava (Hornibrook, 1952
DARWINULIDAE
Penthesilenula aotearoa (Rossetti, Eagar & Martens,
1998) F E
Penthesilenula kohanga (Rossetti, Eagar & Martens,
1998) F E
Penthesilenula? repoa (Chapman, 1963) F E
Penthesilenula sphagna (Barclay, 1968) F E
ENTOCYTHERIDAE
Laccocythere aotearoa Hart & Hart, 1970 E
HEMICYTHERIDAE
Ambostracon pumila (Brady, 1880)
Aurila sp. Hartmann 1985
Bradleya arata (Brady, 1880)
Bradleya claudiae Jellinek & Swanson, 2003 E
Bradleya cupa Jellinek & Swanson, 2003
Bradleya deltoides Hornibrook, 1952
Bradleya dictyon (Brady, 1880)
Bradleya fenwicki Jellinek & Swanson, 2003
Bradleya glabra Jellinek & Swanson, 2003 E
Bradleya lordhowensis Whatley, Downing, Kesler &
Harlow, 1984
Bradleya opima Swanson, 1979
Bradleya pelasgica Whatley, Downing, Kesler &
Harlow, 1984
Bradleya cf. pelasgica Whatley, Downing, Kesler &
Harlow, 1984
Bradleya perforata Jellinek & Swanson, 2003
Bradleya pygmaea Whatley, Downing, Kesler &
Harlow, 1984
Bradleya reticlava Hornibrook, 1952
Bradleya silentium Jellinek & Swanson, 2003 E
Bradleya wyvillethomsoni (Brady, 1880)
Bradleya n. sp. ‘dictyon’ Hornibrook 1952
Bradleya (Quasibradleya) cuneazea Hornibrook,
1952
Harleya ansoni (Whatley, Moguilevsky, Ramos &
Coxill, 1998)
Harleya davidsoni Jellinek & Swanson, 2003 E
Harleya sp. Jellinek & Swanson 2003
Hemicythere brunnea (Brady, 1898)
Hemicythere foveolata (Brady, 1880)
Hemicythere fulvotincta (Brady, 1880)
Hemicythere kerguelensis (Brady, 1880)
Hemicythere munida Swanson, 1979
Hermanites andrewsi Swanson, 1979
Hermanites briggsi Swanson, 1979
Jacobella papanuiensis Swanson, 1979
Mutilus cf. pumilus (Brady, 1866)
Poseidonamicus major Benson, 1972
Poseidonamicus minor Benson, 1972
Poseidonamicus ocularis Whatley, Downing, Kesler
& Harlow, 1986
Poseidonomicus sp. Jellinek & Swanson 2003
Poseidonamicus spp. Ayress, Neil, Passlow &
Swanson 1997
Procythereis (Serratocythere) lytteltonensis Hartmann,
1982
Quadracythere biruga Hornibrook, 1952
Quadracythere mediaruga Hornibrook, 1952
Quadracythere radizea Hornibrook, 1952
Quadracythere truncula Hornibrook, 1952
Waiparacythereis joanae Swanson, 1969
ILYOCYPRIDIDAE
Ilyocypris fallax Brehm, 1929 F E
KRITHIDAE
Krithe antisawanensis Ishizaki, 1966
Krithe comma Ayress, Barrows, Passlow & Whatley,
1999
Krithe compressa (Seguenza, 1980)
Krithe dolichodeira Bold, 1946
Krithe marialusae Abate, Barra, Aiello & Bonaduce,
1993
Krithe minima Coles, Whatley & Moguilevsky, 1994
Krithe morkhoveni morkhoveni Bold, 1960
Krithe nitida Whatley & Downing, 1993 ?Rec
Krithe producta Brady, 1880
Krithe pseudocomma Ayress, Barrows, Passlow &
Whatley, 1999
Krithe reversa Bold, 1958
Krithe swansoni Milau, 1993
Krithe trinidadensis Bold, 1958
Krithe sp. Ayress, Neil, Passlow & Swanson 1997
Krithe sp. 2 Ayress, Barrows, Passlow & Whatley
1999
Parakrithe sp. Swanson 1979
LEGUMINOCYTHERIDIDAE
Triginglymus? sp. Hornibrook 1952
LEPTOCYTHERIDAE
Callistocythere dorsotuberculata Hartmann, 1979
Callistocythere innominata (Brady, 1898)
Callistocythere mosleyi (Brady, 1880)
Callistocythere murrayana (Brady, 1880)
Callistocythere neoplana Swanson, 1979 E
Callistocythere obtusa Swanson, 1979 E
Callistocythere puri McKenzie, 1967
Callistocythere n. sp. cf. crispata Hornibrook, 1952
Callistocythere sp. Hornibrook 1952
Cluthia australis Ayress & Drapala, 1996
Kangarina unispinosa Swanson, 1980
Leptocythere hartmanni (McKenzie, 1967)
Leptocythere lacustris De Deckker, 1981
Leptocythere swansoni Hartmann, 1982 E
Swansonella novaezealandica (Hartmann, 1982) E
Swansonella newbrightonensis Guise, 2002 E
LIMNOCYTHERIDAE
Gomphocythere duffi (Hornibrook, 1955) F
Gomphocythere problematica (Brehm, 1932) F
Kiwicythere anneari Martens, 1992 F E
Kiwicythere vulgaris (McKenzie & Swanson, 1981)
FE
Paralimnocythere vulgaris McKenzie & Swanson,
1981 F
LOXOCONCHIDAE
Loxoconcha anomala Brady, 1880
Loxoconcha parvifoveata Hartmann, 1980 A
Loxoconcha punctata Thomson, 1879
Loxoconcha suteri Hartmann, 1982
Loxoconcha tubmani Swanson, 1980
Loxoconcha sp. Swanson 1969
Loxoconcha sp. Hartmann 1982
CRUSTACEA
MACROCYPRIDIDAE
Macrocyprina campbelli Jellinek & Swanson, 2003 E
Macrocyprina sp. Swanson 1979
Macrocyprina sp. A Jellinek & Swanson 2003
Macrocyprina sp. B Jellinek & Swanson 2003
Macrocyprina sp. C Jellinek & Swanson 2003
Macrocypris decora (Brady, 1866)
Macrocypris tumida Brady, 1880 (doubtful)
Macrocypris sp. Hornibrook 1952
Macrocypris sp. Swanson 1979
Macrocypris sp. Ayress 1993
Macromckensiea cf. porcelica Whatley & Downing,
1983
Macromckenziea swansoni Maddocks, 1990 E
Macropyxis andreseni Jellinek & Swanson, 2003
Macropyxis sonneae Jellinek & Swanson, 2003 E
‘Macropyxis’ thiedei Jellinek & Swanson, 2003 E
Macropyxis sp. Jellinek & Swanson 2003
Macrosarisa sp. Jellinek & Swanson 2003
Macroscapha procera Jellinek & Swanson, 2003 E
Gen et sp. indet. Jellinek & Swanson 2003
NEOCYTHERIDEIDAE
Copytus novaezealandiae (Brady, 1898) E
Neocytherideis muehlenhardtae Hartmann, 1982 E
Pontocythere hedleyi (Chapman, 1906)
NOTODROMADIDAE
Newnhamia fenestrata King, 1855
PARACYPRIDIDAE
Paracypris bradyi McKenzie, 1967
Phylctenophora zealandica Brady 1880
Tasmanocypris sp. Morley & Hayward 2007
PARADOXOSTOMATIDAE
Paradoxostoma spp. Hornibrook 1952
Sclerochillus littoralis (Thomson, 1879)
Sclerochillus sp. a Swanson 1979
Sclerochillus sp. b Swanson 1979
Sclerochillus sp. c Swanson 1979
PARVOCYTHERIDAE
Hemiparvocythere lagunicola Hartmann, 1982
PECTOCYTHERIDAE
Keijia demissa (Brady, 1968)
Kotoracythere formosa Swanson, 1979
Mckenzieartia sp. Morley & Hayward 2007
Munseyella aequa Swanson, 1979
Munseyella brevis Swanson, 1979
Munseyella dedeckeri (Swanson, 1980)
Munseyella modesta, Swanson, 1979
Munseyella punctata Whatley & Downing, 1983
Munseyella tumida Swanson, 1979
Munseyella sp. 10 Hartmann, 1982
Parakeijia aff. thomi (Yassini & Mikulandra, 1989)
Swansonites aequa (Swanson, 1979)
PONTOCYPRIDIDAE
Argilloecia clavata Brady, 1880 E
Argilloecia eburnea Brady, 1880
Argilloecia aff. pusilla (Brady, 1880)
Argilloecia sp. Swanson 1979
Propontocypris cf. attenuata Brady, 1868)
Propontocypris cf. herdmani (Scott, 1905)
Propontocypris (Ekpontocypris) epicyrta Maddocks,
1969
Propontocypris (Propontocypris) sp. Swanson 1979
Propontocypris (Schedopontocypris?) sp. 3 Maddocks
1969
TRACHYLEBERIDIDAE
Abyssophilos ktis Jellinek & Swanson, 2003
Actinocythereis thomsoni (Hornibrook, 1952)
Ambocythere christineae Jellinek & Swanson, 2003
Ambocythere recta Jellinek & Swanson, 2003
Apatihowella (Apatihowella) rustica Jellinek &
Swanson, 2003 E
Apatihowella (A.) sp. Jellinek & Swanson 2003
Apatihowella (Fallacihowella) caligo Jellinek &
Swanson, 2003
Apatihowella (F.) sol Jellinek & Swanson, 2003
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NEW ZEALAND INVENTORY OF BIODIVERSITY
Arculacythereis sp. Morley & Hayward 2007
Cletocythereis rastromarginata (Brady, 1880)
Clinocthereis australis Ayress & Swanson, 1991
Cythereis finlayi Hornibrook, 1952
Cythereis incerta Swanson, 1979
Dutoitella suhmi (Brady, 1880)
Henryhowella dasyderma (Brady, 1880)
Glencoeleberis armata Jellinek & Swanson, 2003
Glencoeleberis cf. armata Jellinek & Swanson, 2003
Glencoeleberis occultata Jellinek & Swanson, 2003 E
Glencoeleberis thomsoni (Hornibrook, 1952)
Legitimocythere acanthoderma (Brady, 1880)
Legitimocythere aculeata Jellinek & Swanson, 2003
Legitimocythere castanea Jellinek & Swanson, 2003
Legitimocythere sp. A Jellinek & Swanson 2003
Legitimocythere sp. B Jellinek & Swanson 2003
Philoneptunus gigas Jellinek & Swanson, 2003 E
Philoneptunus gravizea Hornibrook, 1952
Philoneptunus neesi Jelinek & Swanson, 2003
Philoneptunus paeminosus Whatley, Millson &
Ayress, 1992
Philoneptunus paragravazea Whatley, Millson &
Ayress, 1992
Philoneptunus planaltus (Hornibrook, 1952)
Philoneptunus provocator Jellinek & Swanson, 2003
Ponticocythereis decora Swanson, 1979
Ponticocythereis militaris (Brady, 1866)
Rugocythereis reticulata Ayress, 1993
Taracythere ayressi Jellinek & Swanson, 2003
Taracythere rhinoceros Jellinek & Swanson, 2003 E
Taracythere ulcus Jellinek & Swanson, 2003
Taracythere venusta Jellinek & Swanson, 2003 E
Taracythere sp. Jellinek & Swanson 2003
Trachyleberis cf. clavigera (Brady, 1880)
Trachyleberis lytteltonsis Harding & SylvesterBradley, 1953
Trachyleberis melobesoides (Brady, 1866)
Trachyleberis rugibrevis (Hornibrook, 1952)
Trachyleberis scabrocuneata (Brady, 1898)
Trachyleberis scutigera (Brady, 1880)
Trachyleberis tetrica (Brady, 1880)
Trachyleberis zeacristata Hornibrook, 1952
XESTOLEBERIDIDAE
Foveoleberis sp. Jellinek & Swanson 2003
Microxestoleberis triangulata Swanson, 1980
Semixestoleberis taiaroaensis Swanson, 1979
Xestoleberis africana Brady, 1880
Xestoleberis atra (Thomson, 1879) E
Xestoleberis aff. chilensis austrocontinentalis
Hartmann, 1978
Xestoleberis compressa Brady, 1898
Xestoleberis cf. curta (Brady, 1865)
Xestoleberis foveolata Brady, 1880
Xestoleberis luxata Brady, 1898
Xestoleberis olivacea Brady, 1898
Xestoleberis margaretea Brady, 1865
Xestoleberis setigera Brady, 1880
Xestoleberis cf. trimaculata Hartmann, 1962
Xestoleberis sp. Hornibrook 1952
Xestoleberis sp. Swanson 1979
Xestoleberis sp. A Jellinek & Swanson 2003
Xestoleberis sp. B Jellinek & Swanson 2003
Xestoleberis sp. C Jellinek & Swanson 2003
INCERTAE SEDIS
Bisulcocythere novaezealandiae Ayress & Swanson,
1991 E
Saida torresi (Brady, 1880)*
Suborder PLATYCOPINA
CYTHERELLIDAE
Cytherella corpusculum Swanson, Jellinek, & Malz,
2003
Cytherella eburnea Brady, 1898 E
Cytherella hemipuncta Swanson, 1969
Cytherella hiatus Swanson, Jellinek & Malz, 2003
218
Cytherella intonsa Swanson, Jellinek & Malz, 2003
Cytherella lata Brady, 1880
Cytherella paranitida Whatley & Downing, 1983
Cytherella permutata Swanson, Jellinek & Malz,
2003
Cytherella plusminusve Swanson, Jellinek & Malz,
2003
Cytherella polita Brady, 1880
Cytherella pulchra Brady, 1880
Cytherella punctata Brady, 1880
Cytheretta sp. Morley & Hayward 2007
Cytherelloidea willetti Swanson, 1969* E
Cytherelloidea n. sp. van den Bold 1963
Grammcythella dyspnoea Swanson, Jellinek & Malz,
2003
Inversacytherella tanantia Swanson, Jellinek & Malz,
2003
Order MYODOCOPIDA
Suborder MYODOCOPINA
CYPRIDINIDAE
Bathyvargula walfordi Poulsen, 1963
Codonocera crueta Brady, 1902
Cypridina inermis (Müller, 1906)
Cypridinodes reticulata Poulsen, 1962 E
Cypridinodes concentrica Kornicker, 1979 E
Gigantocypris australis Poulsen, 1962 Pe
Gigantocypris danae Poulsen, 1962 Pe
Macrocypridina castanea (Brady, 1897) Pe
Metavargula iota Kornicker, 1975 E
Metavargula bradfordi Kornicker, 1979 E
Metavargula mazeri Kornickeri, 1979 E
Paracypridina aberrata Poulsen, 1962 E
Vargula ascensus Kornicker, 1979 E
Vargula stathme Kornicker, 1975 E
PHILOMEDIDAE
Euphilomedes agilis (Thomson, 1879)
Euphilomedes ferox Poulsen, 1962
Harbansus n. sp. Eagar 1995
Scleroconcha arcuata Poulsen, 1962 E
Scleroconcha sculpta (Brady, 1898) E
Scleroconcha flexilis (Brady, 1898) E
Scleroconcha wolffi Kornicker, 1975 E
CYLINDROLEBERIDIDAE
Bathyleberis oculata Kornicker, 1975 E
Cycloleberis bradyi Poulsen, 1965
Diasterope grisea (Brady, 1898) E
Dolasterope johansoni Poulsen, 1965 E
Leuroleberis zealandica (Baird, 1850) E
Parasterope pectinata Poulsen, 1965 E
Parasterope quadrata (Brady, 1898) E
Pasterope crinita Kornicker, 1975 E
Synasterope empoulseni Korniker, 1975 E
SARSIELLIDAE
Ancohenia n sp. Eagar 1995
Chelicopia tasmanensis Kornicker, 1981
Cymbicopia brevicostata Kornicker, 1975 E
Cymbicopia hanseni (Brady, 1898) E
Cymbicopia hispida (Brady, 1898) E
Cymbicopia zealandica (Poulsen, 1965) E
HALOCYPRIDIDAE
Archiconchoecia cuculata (Brady, 1802)
Archiconchoecia versicula (Deevey, 1978)
Conchoecia acuticostata Müller, 1906
Conchoecia amblypostha Müller, 1906
Conchoecia antipoda Müller, 1906
Conchoecia belgicae Müller, 1906
Conchoecia bispinosa Claus, 1890
Conchoecia brachyaskos Müller (1906)
Conchoecia chuni Müller 1906
Conchoecia ctenophora (Müller, 1906)
Conchoecia discorphora Müller, 1906
Conchoecia eltaninae Deevey, 1982
Conchoecia hyalophyllum Claus, 1890 Pe
Conchoecia loricata (Claus, 1894)
Conchoecia macrocheira Müller, 1906 Pe
Conchoecia magna Claus, 1874 Pe
Conchoecia major Müller, 1906
Conchoecia nasotuberculata Müller, 1906
Conchoecia parvidentata Müller, 1906 Pe
Conchoecia pusilla Müller, 1906
Conchoecia rhynchena Müller, 1906
Conchoecia serrulata laevis Brady, 1907
Conchoecia skogsbergi Iles, 1953
Conchoecia spinifera Clauss, 1890
Conchoecia subarcuata Claus, 1890 Pe
Conchoecia stigmata Müller, 1906
Conchoecia teretivalvata Iles, 1953
Conchoecia (Alaca) hettacra (Müller, 1906)
Conchoecia (A.) valdiviae (Müller, 1906)
Conchoecia (Conchoecilla) chuni (Müller, 1906)
Conchoecia (C.) daphnoides (Clauss, 1890)
Conchoecia (Conchoecissa) ametra (Müller, 1906)
Conchoecia (C.) imbricata (Brady, 1880)
Conchoecia (C.) symmetrica (Müller, 1906)
Conchoecia (Discoconchoecia) elegans Sars, 1865
Conchoecia (Obtusoecia) antarctica (Muller, 1906)
Conchoecia (Orthoconchoecia) haddoni Brady &
Norman, 1896
Conchoecia (Porroecia) spinirostris Claus, 1874
Conchoecia (P.) porrecta Claus, 1890
Conchoecia (Pseudoconchoecia) serrulata Claus 1874
Fellia cornuta (Müller, 1906) Pe
Fellia dispar (Müller, 1906) Pe
Halocypris inflata (Dana, 1849) Pe
Halocypris globosa (Claus, 1874) Pe
Suborder CLADOCOPINA
POLYCOPIDAE
Polycope sp. Swanson 1979
Polycopsis cf. loscobanosi Hartmann, 1959
Class MALACOSTRACA
Subclass PHYLLOCARIDA
Order LEPTOSTRACA
NEBALIIDAE
Nebalia longicornis G.M. Thomson, 1879
Nebaliella antarctica Thiele, 1904
Sarsinebalia sp. 1 Dahl 1990
Sarsinebalia sp. 2 Dahl 1990
PARANEBALIIDAE
Levinebalia fortunata (Wakabara, 1976)
Subclass HOPLOCARIDA
Order STOMATOPODA
BATHYSQUILLIDAE
Bathysquilla microps (Manning, 1961)
HEMISQUILLIDAE
Hemisquilla australiensis Stephenson, 1967
ODONTODACTYLIDAE
Odontodactylus brevirostris (Miers, 1884)
SQUILLIDAE
Oratosquilla oratoria (de Haan, 1844) A
Pterygosquilla schizodontia (Richardson, 1953)
TETRASQUILLIDAE
Acaenosquilla brazieri (Miers, 1880)
Heterosquilla tricarinata (Claus, 1871) E
Heterosquilla tridentata (Thomson, 1882) E
Subclass EUMALOCOSTRACA
Superorder SYNCARIDA
Order ANASPIDACEA
STYGOCARIDIDAE
Stygocaris townsendi Morimoto, 1977 F E
Stygocaris sp. 1 Morimoto 1977 F E
Stygocaris sp. 2 Morimoto 1977 F E
Stygocaris sp. Schminke 1980 F
Stygocarella pleotelson Schminke, 1980 F E
Stygocarella sp. Schminke 1973 F E
PHYLUM ARTHROPODA
Order BATHYNELLACEA
BATHYNELLIDAE
Bathynella sp. 1 Schminke 1971 F E
Bathynella sp. 2 Schminke 1971 F E
PARABATHYNELLIDAE
Atopobathynella compagana Schminke, 1973 F E
Hexabathynella aotearoae Schminke, 1973 F E
Notobathynella chiltoni Schminke, 1973 F E
Notobathynella hineoneae Schminke, 1973 F E
Notobathynella longipes Schminke, 1978 F E
Notobathynella sp. Schminke 1973 F E
Superorder PERACARIDA
Order LOPHOGASTRIDA
GNATHOPHAUSIIDAE
Gnathophausia elegans G.O. Sars, 1883
Gnathophausia zoea Willemoes-Suhm, 1875
Neognathophausia ingens (Dohrn, 1870)
Neognathophausia gigas (Willemoes-Suhm, 1875)
LOPHOGASTRIDAE
Lophogaster sp.* MNZ
Paralophogaster glaber Hansen, 1910
Order MYSIDA
MYSIDAE
Boreomysis rostrata Illig, 1906
Euchaetomera oculata Hansen, 1910
Euchaetomera typica G.O. Sars, 1884
Euchaetomera zurstrasseni (Illig, 1906)
Gastrosaccus australis W. Tattersall, 1923 E
Siriella denticulata (Thomson, 1880) E
Siriella thompsonii (H. Milne Edwards, 1837)
Tenagomysis chiltoni W. Tattersall, 1923 E
Tenagomysis longisquama Fukuoka & Bruce, 2005 E
Tenagomysis macropsis W. Tattersall, 1923 E
Tenagomysis novaezealandiae Thomson, 1900 E
Tenagomysis producta W. Tattersall, 1923 E
Tenagomysis robusta W. Tattersall, 1923 E
Tenagomysis scotti W. Tattersall, 1923 E
Tenagomysis similis W. Tattersall, 1923 E
Tenagomysis tenuipes W. Tattersall, 1918 E
Tenagomysis thomsoni W. Tattersall, 1923 E
PETALOPHTHALMIDAE
Petalophthalmus sp.* MNZ
Order AMPHIPODA
Suborder INGOLFIELLIDEA
INGOLFIELLIDAE
“Pseudoingolfiella” sp. a Schminke & Noodt 1968
“Pseudoingolfiella” sp. b Schminke & Noodt 1968
Suborder GAMMARIDEA
AMARYLLIDAE
Amaryllis macrophthalma Haswell, 1880
AMPELISCIDAE
Ampelisca albedo Barnard, 1961 E
Ampelisca chiltoni Stebbing, 1888 E
Byblisoides esferis Barnard, 1961 E
Haploops decansa Barnard, 1961 E
AMPHILOCHIDAE
Amphilochus filidactylus Hurley, 1955 E
Amphilochus marionis? Stebbing, 1888
Amphilochus opunake Barnard, 1972 E
Gitanopsis desmondi Barnard, 1972 E
Gitanopsis kupe Barnard, 1972 E
Gitanopsis squamosa (Thomson, 1880)
AMPITHOIDAE
Ampithoe hinatore Barnard, 1972 E
Ampithoe sp. Barnard 1972 E
Parampithoe aorangi (Barnard, 1972) E
Pseudopleonexes lessoniae (Hurley, 1954) E
AORIDAE
Aora maculata (Thomson, 1879)
Aora typica Kroyer, 1845
Aora sp. Barnard 1972
Camacho bathyplous Stebbing, 1888
Camacho nodderi Coleman & Lörz, 2010 E
Haplocheira barbimana (Thomson, 1879)
Haplocheira lendenfeldi Chilton, 1884 E
Lembos? sp. No. 1 Barnard 1972
Lembos? sp. No. 3 Barnard 1972
Lembos? sp. No. 4 Barnard 1972
Meridiolembos acherontis (Myers, 1981) E
Meridiolembos hippocrenes (Myers, 1981) E
Meridiolembos pertinax (Myers, 1981) E
Microdeutopus apopo Barnard, 1972 E
CAPRELLIDAE
Caprella equilibra Say, 1818
Caprella manneringi McCain, 1979 E
Caprella mutica Schurin, 1935 A
Caprellina longicollis (Nicolet, 1849)
Caprellaporema subantarctica Guerra-García, 2003 E
Caprellinoides mayeri (Pfeffer, 1888)
Pseudaeginella campbellensis Guerra-García, 2003 E
Pseudoprotomima hurleyi McCain, 1969 E
CEINIDAE
Ceina egregia (Chilton, 1883) E
Taihape karori Barnard, 1972 E
Waitomo manene Barnard, 1972 E
CHELURIDAE
Chelura terebrans Philippi, 1839 A
CHEVALIIDAE
Chevalia sp. Ahyong
CHILTONIIDAE
Chiltonia enderbyensis Hurley, 1954 F E
Chiltonia mihiwaka (Chilton, 1898) F E
Chiltonia minuta Bousfield, 1964 ?F E
Chiltonia rivertonensis Hurley, 1954 F E
COLOMASTIGIDAE
Colomastix magnirama Hurley, 1954 E
Colomastix subcastellata Hurley, 1954 E
COROPHIIDAE
Apocorophium acutum Chevreux, 1908 A
Monocorophium acherusicum (Costa, 1857) A
Monocorophium insidiosum (Crawford, 1937) A
Monocorophium sextonae (Crawford, 1937) A
Paracorophium brisbanensis Chapman, 2002 B A
Paracorophium excavatum (Thomson, 1884) F B E
Paracorophium lucasi Hurley, 1954 F B E
CYAMIDAE
Cyamus balaenopterae Barnard, 1931
Cyamus boopis Lutken, 1873
Cyamus erraticus Roussel de Vauzeme, 1834
Cyamus gracilis Roussel de Vauzeme, 1834
Cyamus ovalis Roussel de Vauzeme, 1834
Isocyamus delphini Guerin-Meneville, 1837
Neocyamus physeteris (Pouchet, 1888)
Scutocyamus antipodensis Lincoln & Hurley, 1980 E
CYPHOCARIDIDAE
Cyphocaris anonyx Boeck, 1871
Cyphocaris richardi Chevreux, 1905
CYPROIDEIDAE
Neocyproidea otakensis (Chilton, 1900) E
Neocyproidea pilgrimi Hurley, 1955 E
Peltopes peninsulae (Hurley, 1955) E
Peltopes productus K.H. Barnard, 1930 E
DEXAMINIDAE
Atylus reductus (K.H. Barnard, 1930) E
Atylus taupo Barnard, 1972 E
Guernea timaru Barnard, 1972 E
Lepechinella sucia Barnard, 1961
Lepechinella wolffi Dahl, 1959 E
Paradexamine barnardi Sheard, 1938 E
Paradexamine houtete Barnard, 1972 E
Paradexamine muriwai Barnard, 1972 E
Paradexamine pacifica (Thomson, 1879) E
Paradexamine sp. Barnard 1972 E
Polycheria obtusa Thomson, 1882 E
Syndexamine carinata Chilton, 1914 E
DOGIELINOTIDAE
CRUSTACEA
Allorchestes compressa Dana, 1852
‘Allorchestes compressus’ Bousfield 1964 F? E
Allorchestes novizealandiae Dana, 1852 F E
ENDEVOURIDAE
Ensayara iara Lowry & Stoddart, 1983 E
Ensayara kermadecensis Kilgallen, 2009 E
Ensayara ursus Kilgallen, 2009 E
EOPHLIANTIDAE
Bircenna fulva Chilton, 1884 E
Bircenna macayai Lörz, Kilgallen & Thiel, 2009 E
Cylindryllioides kaikoura Barnard, 1972 E
Wandelia wairarapa Barnard, 1972 E
EPIMERIIDAE
Epimeria bruuni Barnard, 1961 E
Epimeria glaucosa Barnard, 1961 E
Epimeria horsti Lörz, 2008 E
Epimeria norfanzi Lörz, 2010
Epimeriella victoria Hurley, 1957 E
EUSIRIDAE
Atyloella moke Barnard, 1972 E
Bathyschraderia magnifica Dahl, 1959 E
Eusiroides monoculoides (Haswell, 1880)
Eusirus antarcticus Thomson, 1880
Gondogeneia bidentata (Stephensen, 1927)
Gondogeneia danai (Thomson, 1879) E
Gondogeneia rotorua Barnard, 1972 E
Gondogeneia subantarctica (Stephensen, 1938) E
Gondogeneia sp. Chilton 1909 E
Oradarea novaezealandiae (Thomson, 1879) E
Paramoera aucklandica (Walker, 1908) E
Paramoera chevreuxi (Stephensen, 1927) E
Paramoera fasciculata (Thomson, 1880) E
Paramoera fissicauda? (Dana, 1852)
Paramoera rangatira Barnard, 1972 E
Paramoera sp. Barnard 1972 E
Paramoera sp. Barnard 1972 F E
Prostebbingia? levis (Thomson, 1879) E
Regalia fascicularis Barnard, 1930 E
Rhachotropis chathamensis Lörz, 2010 E
Rhachotropis delicata Lörz, 2010 E
Rhachotropis levantis Barnard, 1961 E
Schraderia serraticauda (Stebbing, 1888)
Whangarusa translucens (Chilton, 1884) E
EXOEDICEROTIDAE
Patuki breviuropodus Cooper & Fincham, 1974 E
Patuki roperi Fenwick, 1983 E
HADZIIDAE
Zhadia subantarctica Lowry & Fenwick, 1983 E
HYALIDAE
Apohyale hirtipalma (Dana, 1852)
Apohyale media (Dana, 1853)
Apohyale novaezealandiae (Thomson, 1879) E
Protohyale (Protohyale) campbellica (Filhol, 1885) E
Protohyale (Boreohyale) grenfelli Chilton, 1916 E
Protohyale (B.) maroubrae Stebbing, 1899
Protohyale (B.) rubra (Thomson, 1879)
Hyale sp. Thomson 1899
IPHIMEDIIDAE
Amathillopsis grevei Barnard, 1961
Anisoiphimedia haurakiensis (Hurley, 1954) E
Curidia knoxi Lowry & Myers, 2003 E
Epimeria bruuni Barnard, 1961 E
Epimeria glaucosa Barnard, 1961 E
Epimeriella victoria Hurley, 1957 E
Iphimedia spinosa (Thomson, 1880) E
Labriphimedia hinemoa (Hurley, 1954) E
ISAEIDAE
Gammaropsis chiltoni (Thomson, 1897) E
Gammaropsis crassipes (Haswell, 1881)
Gammaropsis haswelli (Thomson, 1897)
Gammaropsis kermadeci (Stebbing, 1888) E
Gammaropsis longimana (Chilton, 1884) E
Gammaropsis tawahi Barnard, 1972 E
Gammaropsis thomsoni Stebbing, 1888
Gammaropsis typica (Chilton, 1884) E
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NEW ZEALAND INVENTORY OF BIODIVERSITY
Gammaropsis sp. Barnard 1972 E
Pagurisaea schembrii Moore, 1983 E
Photis brevicaudatus Norman, 1867
Photis nigrocula Lowry, 1979 E
Photis phaeocula Lowry, 1979 E
Photis sp. Barnard 1972 E
ISCHYROCERIDAE
Ericthonius pugnax (Dana, 1852) A
Ischyrocerus longimanus (Haswell, 1880)
Jassa alonsoae Conlan, 1990
Jassa fenwicki Conlan, 1990
Jassa hartmannae Conlan, 1990 E
Jassa justi Conlan, 1990
Jassa marmorata Conlan, 1990
Jassa slatteryi Conlan, 1990
Notopoma fallohidea (Lowry, 1981) E
Notopoma harfoota (Lowry, 1981) E
Notopoma stoora (Lowry, 1981) E
Parajassa andromedae Moore, 1985 E
Runanga coxalis Barnard, 1961 E
Runanga wairoa McCain, 1969 E
Ventojassa frequens (Chilton, 1883) E
KAMAKIDAE
Aorcho delgadus Barnard, 1961
LEUCOTHOIDAE
Leucothoe trailli Thomson, 1882 E
LILJEBORGIIDAE
Liljeborgia aequabilis Stebbing, 1888
Liljeborgia akaroica Hurley, 1954
Liljeborgia barhami Hurley, 1954 E
Liljeborgia dubia (Haswell, 1880)
Liljeborgia hansoni Hurley, 1954 E
LYSIANASSIDAE
Acheronia pegasus Lowry, 1984 E
Acontiostoma marionis Stebbing, 1888
Acontiostoma tuberculata Lowry & Stoddart, 1983 E
Acontiostoma sp.
Ambasiopsis robustus Barnard, 1961 E
Bruunosa bruuni (Dahl, 1959) E
Cheirimedon cansada (Barnard, 1961)
Eurythenes gryllus (Lichtenstein, 1822)
Hippomedon antitemplado Barnard, 1961 E
Hippomedon concolor Barnard, 1961 E
Hippomedon hake Lowry & Stoddart, 1983 E
Hippomedon hurleyi Kilgallen, 2009 E
Hippomedon incisus K.H. Barnard, 1930 E
Hippomedon iugum Kilgallen, 2009 E
Hippomedon kergueleni (Miers, 1875)
Hippomedon tasmanicus Barnard, 1961 E
Hirondella dubia Dahl, 1959 E
Kakanui punui Lowry & Stoddart, 1983 E
Lepidecreella bidens (Barnard, 1930) E
Lysianopsis tieke Lowry & Stoddart, 1983 E
Ocosingo fenwicki Lowry & Stoddart, 1983 E
Orchomene aahu Lowry & Stoddart, 1983 E
Orchomenella cavimanus (Stebbing, 1888)
Paracentromedon? manene (Lowry & Stoddart, 1983)
E
Paracentromedon? matikuku (Lowry & Stoddart,
1983) E
Paracentromedon? whero (Fenwick, 1983) E
Paralicella similis Birnstein & Vinogradov, 1960
Parawaldeckia angusta Lowry & Stoddart, 1983 E
Parawaldeckia dabita Lowry & Stoddart, 1983 E
Parawaldeckia hirsuta Lowry & Stoddart, 1983 E
Parawaldeckia karaka Lowry & Stoddart, 1983 E
Parawaldeckia kidderi Lowry & Stoddart, 1983
Parawaldeckia parata Lowry & Stoddart, 1983 E
Parawaldeckia pulchra Lowry & Stoddart, 1983 E
Parawaldeckia stephenseni Hurley & Cooper, 1974 E
Parawaldeckia suzae Lowry & Stoddart, 1983 E
Parawaldeckia thomsoni (Stebbing, 1906) E
Parawaldeckia vesca Lowry & Stoddart, 1983 E
Pseudambasia rossii Stephensen, 1927 E
Schisturella abyssi tasmanensis (Barnard, 1961) E
220
Stomacontion hurleyi Lowry & Stoddart, 1983 E
Stomacontion pungapunga Lowry & Stoddart, 1983 E
Stomacontion sp.
Tryphosella moana Kilgallen, 2009 E
Tryphosella serans Lowry & Stoddart, 1983 E
Valettiopsis multidentata Barnard, 1961 E
MELITIDAE
Ceradocopsis macracantha Lowry & Fenwick, 1983 E
Ceradocopsis carnleyi (Stephensen, 1927) E
Ceradocopsis peke Barnard, 1972 E
Ceradocus chiltoni Sheard, 1939 E
Ceradocus rubromaculatus haumuri Barnard, 1972
Elasmopus bollonsi Chilton, 1915
Elasmopus neglectus Chilton, 1915 E
Elasmopus wahine Barnard, 1972 E
Gammarella hybophora Lowry & Fenwick, 1983 E
Hoho hirtipalma (Barnard, 1972) E
Linguimaera tias Krapp-Schickel, 2003
Maera incerta Chilton, 1883 E
Maera spp. Barnard 1972
Mallacoota nanaui Myers, 1985
Melita awa Barnard, 1972 B E
Melita festiva (Chilton, 1884)
Melita inaequistylis Dana, 1852 E
Melita? solada Barnard, 1961 E
Melita sp. Barnard 1972 E
Micramaera tepuni (Barnard, 1972) E
Parapherusa crassipes (Haswell, 1880)
Tagua aporema Lowry & Fenwick, 1983 E
MELPHIDIPPIDAE
Horniella whakatane (Barnard, 1972) E
NIHOTUNGIDAE
Nihotunga noa Barnard, 1972 E
OCHLESIDAE
Curidia knoxi Lowry & Myers, 2003 E
OEDICEROTIDAE
Bathymedon neozelanicus Barnard, 1930 E
Carolobatea novaezealandiae Chilton, 1909
Lopiceros forensia Barnard, 1961 E
Monoculodes abacus Barnard, 1961 E
Oediceroides apicalis Barnard, 1931
Oediceroides limpieza Barnard, 1961 E
Oediceroides microcarpa Barnard, 1930 E
Oediceroides wolffi Barnard, 1961
PARACALLIOPIIDAE
Paracalliope fluviatilis (Thomson, 1879) F E
Paracalliope karitane Barnard, 1972 F E
Paracalliope novizealandiae (Dana, 1853) E
PARACRANGONYCTIDAE E
Paracrangonyx compactus (Chilton, 1882) F E
Paracrangonyx winterbourni Fenwick, 2001 F E
Pseudoingolfiella Morimotoi Grosso, Peralta & Ruffo,
2006 F E
PARALEPTAMPHOPIDAE E
Paraleptamphopus caeruleus (Thomson, 1885) F E
Paraleptamphopus subterraneus (Chilton, 1882) F E
Paraleptamphopus spp. (10) 10E G. D. Fenwick
Ringanui koonuiroa Fenwick, 2006 F E
Ringanui toonuiiti Fenwick, 2006 F E
Gen. nov. (~10) et n. spp. (~20) ~ 20E G. D.
Fenwick
PARDALISCIDAE
Arculfia trago Barnard, 1961 E
Halice macronyx (Stebbing, 1888)
Halice secunda (Stebbing, 1888)
Halice sublittoralis Lowry, 1979 E
Halicoides tambiella Barnard, 1961 E
Pardaliscoides longicaudatus Dahl, 1959 E
Princaxelia abyssalis Dahl, 1959
PHLIANTIDAE
Iphinotus typicus (Thomson, 1882) E
PHOXOCEPHALIDAE
Booranus? spinibasus (Cooper, 1974) E
Cephaloxoides keppeli (Barnard & Drummond,
1978) E
Cephalophoxus regium (Barnard, 1930) E
Harpiniopsis nadania (Barnard, 1961) E
Joubinella traditor Pirlot, 1932
Palabriaphoxus palabria Barnard, 1961 E
Parajoubinella concinna Gurjanova, 1977 E
Paraphoxus? pyripes Barnard, 1930 E
Protophoxus australis Barnard, 1930
Ringaringa littoralis (Cooper & Fincham, 1974) E
Synphoxus novaezelandicus Gurjanova, 1980 E
Torridoharpinia hurleyi (Barnard, 1958) E
Trichophoxus capillatus Barnard, 1930 E
Waitangi rakiura (Cooper & Fincham, 1974) E
Waitangi? brevirostris Fincham, 1977 E
Waitangi? chelatus (Cooper, 1974) E
Wildus waipiro (Barnard, 1972) E
PHREATOGAMMARIDAE E
Phreatogammarus fragilis (Chilton, 1882) F E
Phreatogammarus helmsi Chilton, 1918 F E
Phreatogammarus propinquus Chilton, 1907 F E
Phreatogammarus waipoua Chapman, 2003 F E
PLATYISCHNOPIDAE
Otagia neozelanicus (Chilton, 1987) E
PODOCERIDAE
Podocerus cristatus (Thomson, 1879) E
Podocerus karu Barnard, 1972 E
Podocerus manawatu Barnard, 1972 E
Podocerus sp. Chilton, 1926
Podocerus wanganui Barnard, 1972 E
RAKIROIDAE E
Rakiroa rima Lowry & Fenwick, 1982 E
SCOPELOCHEIRIDAE
Scopelocheirus? schellenbergi Bernstein &
Vinogradov, 1958
SEBIDAE
Seba typica (Chilton, 1884)
STEGOCEPHALIDAE
Andaniotes corpulentus (Thomson, 1882)
Euandandania gigantea (Stebbing, 1888)
Phippsiella nipoma Barnard, 1961
Stegosoladidus simplex (Barnard, 1930) E
Tetradeion crassum (Chilton, 1883) E
STENOTHOIDAE
Mesoproboloides? excavata Fenwick, 1977 E
Parathaumatelson nasicum (Stephensen, 1927) E
Probolisca ovata (Stebbing, 1888)
Raukumara rongo (Barnard, 1972) E
Stenothoe aucklandicus Stephensen, 1927 E
Stenothoe gallensis Walker, 1904 A
Stenothoe moe Barnard, 1972 E
Stenothoe valida? Dana, 1853
STILIPEDIDAE
Alexandrella mixta (Nicholls, 1938)
Stilipes sanguineus (Hurley, 1954) E
SYNOPIIDAE
Syrrhoe affinis? Chevreux, 1908
TALITRIDAE
Arcitalitrus dorrieni (Hunt, 1925) T A
Arcitalitrus sylvaticus (Haswell, 1880) T A
Austroides sp. Fenwick & Webber 2008 T
Bellorchestia quoyana (Milne-Edwards, 1840) S E
Bellorchestia spadix Hurley, 1956 S E
Bellorchestia tumida Thomson, 1885 S E
Kanikania improvisa (Chilton, 1909) T E
Kanikania motuensis Duncan, 1994 T E
Kanikania rubroannulata (Hurley, 1957) T E
Makawe hurleyi (Duncan, 1968) T E
Makawe insularis (Chilton, 1909) T E
Makawe maynei (Chilton, 1909) T E
Makawe otamatuakeke Duncan, 1994 T E
Makawe parva (Chilton, 1909) T E
Makawe waihekensis Duncan, 1994 T E
Makawe sp. A Fenwick & Webber 2008 T E
Makawe sp. B Fenwick & Webber 2008 T E
Makawe sp. C Fenwick & Webber 2008 T E
Notorchestia aucklandiae (Bate, 1862) S E
PHYLUM ARTHROPODA
Orchestia? recens (Thomson, 1884) F E
Orchestia? sp. A Hurley, 1975 F E
Orchestia? sp. B Hurley, 1975 F E
Parorchestia ihurawao Duncan, 1994 T E
Parorchestia lesliensis (Hurley, 1957) T E
Parorchestia longicornis (Stephensen, 1938) T E
Parorchestia tenuis (Dana, 1852) T E
Protorchestia campbelliana (Bousfield, 1964) T E
Puhuruhuru aotearoa Duncan, 1994 T E
Puhuruhuru patersoni (Stephensen, 1938) T E
Puhuruhuru sp. Fenwick & Webber 2008 T E
Talitroides topitotum (Burt, 1934) T A
Tara hauturu Duncan, 1994 T E
Tara simularis (Hurley, 1957) T E
Tara sinbadensis (Hurley, 1957) T E
Tara sylvicola (Dana, 1852) T E
Tara taranaki Duncan, 1994 T E
Tara sp. A Fenwick & Webber 2008 T E
Tara sp. B Fenwick & Webber 2008 T E
Transorchestia bollonsi (Chilton, 1909) S E
Transorchestia chathamensis (Hurley, 1956) S E
Transorchestia cookii Filhol, 1885 S E
Transorchestia dentata (Filhol, 1885) S E
Transorchestia kirki (Hurley, 1956) S E
Transorchestia miranda (Chilton, 1916) S E
Transorchestia serrulata (Dana, 1852) S E
Transorchestia telluris (Bate, 1862) S E
Waematau kaitaia Duncan, 1994 T E
Waematau manawatahi Duncan, 1994 T E
Waematau muriwhenua Duncan, 1994 T E
Waematau reinga Duncan, 1994 T E
Waematau unuwhao Duncan, 1994 T E
URISTIDAE
Abyssorchomene abyssorum (Stebbing, 1888)
Galathella galatheae (Dahl, 1959) E
Galathella solivagus Kilgallen, 2009 E
UROTHOIDAE
Carangolia puliciformis Barnard, 1961 E
Urothoe elizae Cooper & Fincham, 1974 E
Urothoe wellingtonensis Cooper, 1974 E
Urothoides lachneessa (Stebbing, 1888)
Suborder HYPERIIDEA
ARCHAEOSCINIDAE
Archaeoscina steenstrupi (Bovallius, 1885)
Paralanceola wolffi Zeidler, 2006
BRACHYSCELIDAE
Brachyscelus crusculum Bate, 1861
Brachyscelus rapacoides Stephensen, 1925
Brachyscelus rapax (Claus, 1871)
CHUNEOLIDAE
Chuneola paradoxa Woltereck, 1909
CYLLOPIDAE
Cyllopus magellanicus Dana, 1853
CYSTISOMATIDAE
Cystisoma fabricii Stebbing, 1888
Cystisoma magna (Woltereck, 1903)
Cystisoma pellucida (Willemoes-Suhm, 1873)
DAIRELLIDAE
Dairella californica (Bovallius 1887)
HYPERIIDAE
Hyperia gaudichaudii Milne-Edwards, 1840
Hyperia spinigera Bovallius, 1889
Hyperiella antarctica Bovallius, 1887
Hyperoche mediterranea Senna, 1908
Hyperoche medusarum (Kroyer, 1838)
Lestrigonus schizogeneios (Stebbing, 1888)
Themisto australis (Stebbing, 1888)
Themisto gaudichaudi Guerin, 1825
IULOPIDIDAE
Iulopis loveni Bovallius, 1887
LANCEOLIDAE
Lanceola clausi Bovallius, 1885
Lanceola grunneri Zeidler, 2009
Lanceola intermedia Vinogradov, 1960
Lanceola longidactyla Vinogradov, 1964
Lanceola loveni (Bovallius, 1885)
Lanceola pacifica Stebbing, 1888
Lanceola sayana Bovallius, 1885
Lanceola serrata Bovallius, 1885
Scypholanceola aestiva (Stebbing, 1888)
LESTRIGONIDAE
Hyperietta luzoni (Stebbing, 1888)
Hyperietta vosseleri (Stebbing, 1904)
Hyperioides longipes Chevreux, 1900
Hyperionyx macrodactylus (Stephensen, 1924)
LYCAEIDAE
Lycaea nasuta Claus, 1879
Lycaea pachypoda (Claus, 1879)
Lycaea pulex Marion, 1874
Simorhynchotus antennarius (Claus, 1871)
LYCAEOPSIDAE
Lycaeopsis themistoides Claus, 1879
Lycaeopsis zamboangae (Stebbing, 1888)
MEGALANCEOLIDAE
Megalanceola stephenseni (Chevreux, 1920)
MICROPHASMIDAE
Microphasma agassizi Woltereck, 1909
MIMONECTIDAE
Mimonectes gaussi (Woltereck, 1904)
OXYCEPHALIDAE
Calamorhynchus pellucidus Streets, 1878
Leptocotis tenuirostris (Claus, 1871)
Oxycephalus piscator Milne-Edwards, 1830
Streetsia challengeri Stebbing, 1888
Streetsia porcella (Claus, 1879)
PARAPHRONIMIDAE
Paraphronima crassipes Claus, 1879
Paraphronima gracilis Claus, 1879
PARASCELIDAE
Parascelus edwardsi Claus, 1879
PHRONIMIDAE
Phronima atlantica Guérin-Menéville, 1836
Phronima sedentaria (Forsskål, 1775)
Phronimella elongata (Claus, 1862)
PHROSINIDAE
Anchylomera blossevillei Milne-Edwards, 1830
Phrosina semilunata Risso, 1822
Primno macropa Guérin-Menéville, 1836
PROLANCEOLIDAE
Prolanceola vibiliformis Woltereck, 1907
PLATYSCELIDAE
Amphithyrus bispinosus Claus, 1879
Hemityphis tenuimanus Claus, 1879
Paratyphis parvus Claus, 1887
Paratyphis spinosus Spandl, 1924
Platyscelus armatus (Claus, 1879)
Platyscelus ovoides (Risso, 1816)
Platyscelus serratulus Stebbing, 1888
Tetrathyrus arafurae Stebbing, 1888
Tetrathyrus forcipatus Claus, 1879
PRONOIDAE
Eupronoe maculata Claus, 1879
Eupronoe minuta Claus, 1879
Paralycaea gracilis Claus, 1879
Parapronoe campbelli Stebbing, 1888
Parapronoe crustulum Claus, 1879
Parapronoe parva Claus, 1879
Pronoe capito Guérin-Menéville, 1836
SCINIDAE
Acanthoscina acanthodes (Stebbing, 1895)
Scina borealis (G.O. Sars, 1882)
Scina crassicornis (Fabricius, 1775)
Scina curvidactyla Chevreux, 1914
Scina pusilla Chevreux, 1919
Scina tullbergi (Bovallius, 1885)
Scina wagleri abyssalis Vinogradov, 1957
TRYPHANIDAE
Tryphana malmi Boeck, 1871
VIBILIIDAE
CRUSTACEA
Vibilia antarctica Stebbing, 1888
Vibilia armata Bovallius, 1887
Vibilia borealis Bate & Westwood, 1868
Vibilia caeca Bulycheva, 1955
Vibilia chuni Behning & Woltereck, 1912
Vibilia cultripes Vosseler, 1901
Vibilia gibbosa Bovallius, 1887
Vibilia longicarpus Behning, 1913
Vibilia propinqua Stebbing, 1888
Vibilia pyripes Bovallius, 1887
Vibilia robusta Bovallius, 1887
Vibilia stebbingi Behning & Woltereck, 1912
Vibilia viatrix Bovallius, 1887
Order ISOPODA
Suborder ASELLOTA
ACANTHASPIDIDAE
Mexicope sushara Bruce, 2004 E
Acanthaspidia sp. E
DENDROTIIDAE
Acanthomunna proteus Beddard, 1886 E
Dendromunna mirabile Wolff, 1962 E
DESMOSOMATIDAE
Chelator spp. (3) N. Bruce 2008
Desmosoma sp. N. Bruce 2008
Eugerda sp. N. Bruce 2008
Eugerdella spp. (2) N. Bruce 2008
Mirabilicoxa sp. N. Bruce 2008
Prochelator tupuhi Brix & Bruce, 2008 E
HAPLONISCIDAE
Chauliodoniscus tasmanaeus Lincoln, 1985 E
Haploniscus kermadecensis Wolff, 1962 E
Haploniscus piestus Lincoln, 1985 E
Haploniscus miccus Lincoln, 1985 E
Haploniscus saphos Lincoln, 1985 E
Haploniscus silus Lincoln, 1985 E
Haploniscus tangaroae Lincoln, 1985 E
Hydroniscus lobocephalus Lincoln, 1985 E
Mastigoniscus pistus Lincoln, 1985 E
JANIRIDAE
Heterias n. sp. Scarsbrook et al. 2003 E
Iais californica (Richardson, 1904)
Iais pubescens (Dana, 1852)
Ianiropsis neglecta (Chilton, 1909) E
Iathrippa longicauda (Chilton, 1884) E
Iathrippa sp. NIWA N. Bruce
Mackinia sp. Scarsbrook et al. 2003
ISCHNOMESIDAE
Ischnomesus anacanthus Wolff, 1962 E
Ischnomesus birsteini Wolff, 1962 E
Ischnomesus bruuni Wolff, 1956 E
Ischnomesus spaercki Wolff, 1956 E
Mixomesus pellucidus Wolff, 1962 E
JOEROPSIDIDAE
Joeropsis neozealanica Chilton, 1892 E
Joeropsis palliseri Hurley, 1957 E
Joeropsis spp. (2) 2E
MUNNIDAE
Echinomunna sp. E
Munna neozelanica Chilton, 1892 E
Munna spp. (4) 4E
Uromunna schauinslandi (Sars, 1905) E
MUNNOPSIDIDAE
Bathybadistes andrewsi Merrin, Malyutina & Brandt,
2009
Disconectes madseni (Wolff, 1956) E
Echinozone n. sp. E
Epikopais mystax Merrin, 2009 E
Eurycope galatheae Wolff, 1956 E
Eurycope gibberifrons Wolff, 1962 E
Hapsidohedra aspidophora (Wolff, 1962) E
Ilyarachna kermadecensis Wolff, 1962 E
Ilyarachna n. spp. (7) 7E
Munneurycope harrietae Wolff, 1962 E
Munneurycope menziesi Wolff, 1962 E
221
NEW ZEALAND INVENTORY OF BIODIVERSITY
Munnopsis gracilis Beddard, 1886 E
Notopais euaxos Merrin & Bruce, 2006 E
Notopais zealandica Merrin, 2004 E
Paropsurus giganteus Wolff, 1962
Pseudarachna nohinohi Merrin, 2006 E
Storthyngura benti Wolff, 1956 E
Vanhoeffenura abyssalis Wolff, 1962 E
Vanhoeffenura furcata Wolff, 1956 E
Vanhoeffenura kermadecensis Wolff, 1962 E
Vanhoeffenura novaezelandiae (Beddard, 1885) E
Sursumura affinis Malyutina, 2004
PARAMUNNIDAE
Allorostrata n. sp. NIWA N. Bruce E
Austronanus aucklandensis Just & Wilson, 2006
Austronanus sp. A Just & Wilson 2006
Omanana serraticoxa Just & Wilson, 2004 E
‘Paramunna serrata’ sensu Stephenson 1927 E
Paramunna snaresi Just & Wilson, 2004 E
Spiculonana petraea Just & Wilson, 2004 E
Spiculonana platysoma Just & Wilson, 2004 E
Sporonana concavirostra Just & Wilson, 2004 E
Sporonana litoralis Just & Wilson, 2004 E
Gen. nov. 1 N. Bruce 2008 E
Gen. nov. 2 N. Bruce 2008 E
PSEUDOJANIRIDAE
Schottea taupoensis Serov & Wilson, 1999 E
Schottea n. sp. E
SANTIIDAE
Halacarsantia uniramea (Menzies & Miller, 1955) E
Kuphomunna n. sp. NIWA N. Bruce E
Santia hispida (Vanhöffen, 1914)
Santia n. spp. (2) 2E
STENETRIIDAE
Protallocoxa abyssale (Wolff, 1962) E
Stenetrium fractum Chilton, 1884 E
Suborder PHREATOICIDEA
PHREATOICIDAE
Neophreatoicus assimilis (Chilton, 1894) F E
Notamphisopus benhami Nicholls, 1944 F E
Notamphisopus dunedinensis (Chilton, 1906) F E
Notamphisopus flavius Nicholls, 1944 F E
Notamphisopus kirkii (Chilton, 1906) F E
Notamphisopus littoralis Nicholls, 1944 F E
Notamphisopus percevali Nicholls, 1944 F E
Phreatoicus orarii Nicholls, 1944 F E
Phreatoicus typicus Chilton, 1883 F E
Suborder CYMOTHOIDA
AEGIDAE
Aega komai Bruce, 1996
Aega monophthalam Johnston, 1834
Aega semicarinata Miers, 1875
Aega stevelowei Bruce, 2009
Aega urotoma Barnard, 1914
Aegapheles alazon (Bruce, 2004)
Aegapheles birubi (Bruce, 2004)
Aegapheles copidis Bruce, 2009
Aegapheles hamiota (Bruce, 2004)
Aegapheles mahana Bruce, 2009 E
Aegapheles rickbruscai (Bruce, 2004)
Aegapheles umpara (Bruce, 2004)
Aegiochus coroo (Bruce, 1983)
Aegiochus gordoni Bruce, 2009 E
Aegiochus insomnis Bruce, 2009 E
Aegiochus kakai Bruce, 2009 E
Aegiochus kanohi Bruce, 2009
Aegiochus laevis (Studer, 1883)
Aegiochus nohinohi Bruce, 2009
Aegiochus piihuka Bruce, 2009
Aegiochus riwha Bruce, 2009
Aegiochus tara Bruce, 2009
Aegiochus vigilans (Haswell, 1881)
Aegiochus sp. Bruce 2009
Epulaega derkoma Bruce, 2009
222
Epulaega fracta (Hale, 1940)
Rocinela bonita Bruce, 2009 E
Rocinela garricki Hurley, 1957 E
Rocinela leptopus Bruce, 2009 E
Rocinela pakari Bruce, 2009 E
Rocinela resima Bruce, 2009 E
Rocinela runga Bruce, 2009 E
Rocinela satagia Bruce, 2009 E
Rocinela sp. Bruce 2009
Syscenus latus Richardson, 1909 Pe
Syscenus springthorpei Bruce, 1997 Pe
Syscenus sp. Bruce 2009
ANTHURIDAE
Haliophasma novaezelandiae Wägele, 1985 E
Haliophasma platytelson Wägele, 1985 E
Quantanthura pacifica Wägele, 1985 E
Quantanthura raoulia Poore & Lew Ton, 1986 E
Mesanthura affinis (Chilton, 1883) E
ANUROPIDAE
Anuropus novaezealandiae Jansen, 1981 Pe E
Anuropus sp. N. Bruce 2008
BOPYRIDAE
Athelges lacertosi Pike, 1961 E
Eophrixus shojii Shiino, 1941
Gigantione pikei Page, 1985 E
Gyge angularis Page, 1985 E
Hemiarthrus nematocarcini Stebbing, 1914
Pleurocryptella infecta Nierstrasz & Brender à
Brandis, 1923
Pseudione affinis (Sars, 1882)
Pseudione hayi Nierstrasz & Brender à Brandis,
1931 E
Pseudione hyndmanni (Bate & Westwood, 1868)
Pseudione murawaiensis Page, 1985 E
Pseudione pontocari Page, 1985 E
Pseudostegias otagoensis Page, 1985 E
Rhopalione atrinicolae Page, 1985 E
CIROLANIDAE
Cirolana canaliculata Tattersall, 1921 E
Cirolana kokoru Bruce, 2004 E
Cirolana quechso Bruce, 2004 E
Cirolana quadripustulata Hurley, 19571 E
Cirolana n. spp. (5) 5E
Eurydice subtruncata Tattersall, 1921 E
Eurylana arcuata (Hale, 1925) E
Eurylana cooki (Filhol, 1885) E
Metacirolana caeca (Hansen, 1916) Pe
Metacirolana japonica (Hansen, 1890)
Natatolana amplocula Bruce, 1986
Natatolana aotearoa Keable, 2006 E
Natatolana honu Keable, 2006 E
Natatolana narica (Bowman, 1971) E
Natatolana paranarica Keable, 2006 E
Natatolana pellucida (Tattersall, 1921)
Natatolana rekohu Bruce, 2003 E
Natatolana rossi (Miers, 1876) E
Natatolana n. spp. (3) 3E
Pseudaega melanica Jansen, 1978 E
Pseudaega punctata Thomson, 1884 E
Pseudaega quarta Jansen, 1978 E
Pseudaega secunda Jansen, 1978 E
Pseudaega tertia Jansen, 1978 E
CRINONISCIDAE
Crinoniscus cephalatus Hosie, 2008 E
Crinoniscus politosummus Hosie, 2008 E
CYMOTHOIDAE
Ceratothoa imbricata (Fabricius, 1775)
Ceratothoa lineatus (Miers, 1876) E
Ceratothoa trillesi (Avdeev, 1979) E
Elthusa neocytta (Avdeev, 1975)
Elthusa propinqua (Richardson, 1904)
Elthusa raynaudii (Milne Edwards, 1840)
Mothocya ihi Bruce, 1986 E
Nerocila orbignyi (Guérin-Menéville, 1832)
EXPANATHURIDAE
Eisothistos adlateralis Knight-Jones & Knight-Jones,
2002 E
Heptanthura novaezealandiae Kensley, 1978 E
Rhiganthura spinosa Kensley, 1978 E
GNATHIIDAE
Bathygnathia tapinoma Cohen & Poore, 1994 E
Bathygnathia vollenhovia Cohen & Poore, 1994
Caecognathia akaroensis (Monod, 1926) E
Caecognathia nieli Svavarsson, 2005 E
Caecognathia pacifica (Monod, 1926) E
Caecognathia polythrix (Monod, 1926) E
Caecognathia regalis (Monod, 1926) E
Caecognathia sifae Svarvarsson, 2005 E
Caecognathia n. sp. E
Eunognathia n. sp. E
Gnathia brachyuropus Monod, 1926
Thaumastognathia diceros Monod, 1926 E
HEMIONISCIDAE
Scalpelloniscus nieli Hosie, 2008 E
Scalpelloniscus cf. penicillatus Grygier, 1981
Scalpelloniscus vomicus Hosie, 2008
HYSSURIDAE
Kupellonura proberti Wägele, 1985 E
LEPTANTHURIDAE
Albanthura rotunduropus Wägele, 1985 E
Albanthura stenodactyla Wägele, 1985 E
Bullowanthura crebrui Wägele, 1985 E
Cruregens fontanus Chilton, 1882 F E
Leptanthura chiltoni (Beddard, 1886) E
Leptanthura exilis Wägele, 1985 E
Leptanthura profundicola Wägele, 1985 E
Leptanthura truncatitelson Wägele, 1985 E
Psittanthura egregia Wägele, 1985 E
PARANTHURIDAE
Califanthura rima (Poore, 1981) E
Paranthura flagellata (Chilton, 1882) E
Paranthura longa Wägele, 1985 E
TRIDENTELLIDAE
Tridentella acheronae Bruce, 1988 E
Tridentella rosemariae Bruce, 2002 E
Tridentella tangaroae Bruce, 1988 E
Tridentella n. sp.
Suborder LIMNORIIDEA
LIMNORIIDAE
Limnoria convexa Cookson, 1991 E
Limnoria hicksi Schotte, 1989 E
Limnoria loricata Cookson, 1991 E
Limnoria quadripunctata Holthuis, 1949
Limnoria reniculus Schotte, 1989 E
Limnoria rugosissima Menzies, 1957
Limnoria segnis Chilton, 1883 E
Limnoria stephenseni Menzies, 1957 E
Limnoria tripunctata Menzies, 1951
Suborder SPHAEROMATIDEA
PLAKARTHRIIDAE
Plakarthrium typicum Chilton, 1883 E
SEROLIDAE
Acutiserolis sp. Poore & Storey 2009
Brucerolis brandtae Storey & Poore, 2009 E
Brucerolis howensis Storey & Poore, 2009 E
Brucerolis hurleyi Storey & Poore, 2009 E
Brucerolis osheai Storey & Poore, 2009 E
Myopiarolis bicolor (Bruce, 2008) E
Myopiarolis carinata (Bruce, 2008) E
Myopiarolis n. spp. (7) 7E
Spinoserolis latifrons (Miers, 1875) E
SPHAEROMATIDAE
Amphoroidea falcifer Thomson, 1879 E
Amphoroidea longipes Hurley & Jansen, 1977 E
Amphoroidea media Hurley & Jansen, 1971 E
Benthosphaera guaware Bruce, 1994
Bilistra cavernicola Sket & Bruce, 2004 F E
Bilistra millari Sket & Bruce, 2004 F E
Bilistra mollecopulans Sket & Bruce, 2004 F E
PHYLUM ARTHROPODA
Cassidina typa Milne Edwards, 1840 E
Cassidinopsis admirabilis Hurley & Jansen, 1977 E
Cerceis trispinosa (Haswell, 1882)
Cilicaea angustispinata Hurley & Jansen, 1977 E
Cilicaea caniculata (Thomson, 1879) E
Cilicaea dolorosa Hurley & Jansen, 1977 E
Cilicaea tasmanensis Hurley & Jansen, 1977 E
Cilicaeopsis n. sp. N. Bruce 2008 E
Cymodoce allegra Hurley & Jansen, 1977 E
Cymodoce australis Hodgson, 1902 E
Cymodoce convexa Miers, 1876 E
Cymodoce hamata Stephensen, 1927 E
Cymodoce hodgsoni Tattersall, 1921 E
Cymodoce iocosa Hurley & Jansen, 1977 E
Cymodoce penserosa Hurley & Jansen, 1977 E
Cymodocella capra Hurley & Jansen, 1977 E
Cymodocella egregia (Chilton, 1892) E
Cymodocella tubicauda Pfeffer, 1887
Cymodopsis impudica Hurley & Jansen, 1977 E
Cymodopsis sphyracephalata Hurley & Jansen, 1977
E
Cymodopsis torminosa Hurley & Jansen, 1977 E
Dynamenoides decima Hurley & Jansen, 1977 E
Dynamenoides vulcanata Hurley & Jansen, 1977 E
Dynamenopsis varicolor Hurley & Jansen, 1971 E
Exosphaeroma chilense (Dana, 1853)
Exosphaeroma echinense Hurley & Jansen, 1977 E
Exosphaeroma falcatum Tattersall, 1921 E
Exosphaeroma gigas (Leach, 1818)
Exosphaeroma montis (Hurley & Jansen, 1977) E
Exosphaeroma obtusum (Dana, 1853) E
Exosphaeroma planulum Hurley & Jansen, 1971 E
Exosphaeroma waitemata Bruce, 2005 E
Exosphaeroma n. sp. N. Bruce E
Ischyromene condita (Hurley & Jansen, 1977) E
Ischyromene cordiforaminalis (Chilton, 1883) E
Ischyromene hirsuta (Hurley & Jansen, 1971) E
Ischyromene huttoni (Thomson, 1879) E
Ischyromene insulsa (Hurley & Jansen, 1977) E
Ischyromene kokotahi Bruce, 2006 E
Ischyromene mortenseni (Hurley & Jansen), 1977 E
Isocladus armatus (Milne Edwards, 1840) E
Isocladus calcareus (Dana, 1853) E
Isocladus dulciculus Hurley & Jansen, 1977 E
Isocladus inaccuratus Hurley & Jansen, 1977 E
Isocladus reconditus Hurley & Jansen, 1977 E
Isocladus spiculatus Hurley & Jansen, 1977 E
Makarasphaera amnicosa Bruce, 2005 F E
Pseudosphaeroma callidum Hurley & Jansen, 1977 E
Pseudosphaeroma campbellensis Chilton, 1909
Scutuloidea kutu Stephenson & Riley, 1996 E
Scutuloidea maculata Chilton, 1883 E
Sphaeroma laurensi Hurley & Jansen, 1977 E
Sphaeroma quoianum Milne Edwards, 1840
Syncassidina aestuaria Baker, 1929 A?
INCERTAE SEDIS
Paravireia typica Chilton, 1925 E
Paravireia pistus Jansen, 1973 E
Suborder VALVIFERA
ANTARCTURIDAE
Caecarcturus quadraspinosus Schultz, 1981 E
Chaetarcturus myops (Beddard, 1886) E
ARCTURIDAE
Neastacilla antipodea Poore, 1981 E
Neastacilla fusiformis (Hale, 1946) E
Neastacilla levis (Thomson & Anderton, 1921) E
Neastacilla tattersalli Lew Ton & Poore, 1986 E
Neastacilla tuberculata (Thomson, 1879) E
Neastacilla spp. (4) N. Bruce 2008
AUSTRARCTURELLIDAE
Dolichiscus opiliones (Schultz, 1981) E
Austrarcturella galathea Poore & Bardsley, 1992 E
Pseudarcturella chiltoni Tattersall, 1921 E
Pseudarcturella crenulata Poore & Bardsley, 1992 E
CHAETILIIDAE
Macrochiridothea uncinata Hurley & Murray, 1968 E
Maoridotea naylori Jones & Fenwick, 1978 E
Maoridotea n. sp. N. Bruce E
HOLOGNATHIDAE
Cleantis tubicola (Thomson, 1885) E
Holognathus karamea Poore & Lew Ton, 1990 E
Holognathus stewarti (Filhol, 1885) E
IDOTEIDAE
Austridotea annectens Nicholls, 1937 F E
Austridotea benhami Nicholls, 1937 F E
Austridotea lacustris (Thomson, 1879) F E
Batedotea elongata (Miers, 1876)
Euidotea durvillei Poore & Lew Ton, 1993 E
Idotea? festiva Chilton, 1881 E
Idotea metallica Bosc, 1802
Paridotea ungulata Pallas, 1772
PSEUDIDOTHEIDAE
Pseudidothea richardsoni Hurley, 1957 E
Suborder ONISCIDEA
Infraorder LIGIAMORPHA
ACTAECIIDAE
Actaecia euchroa Dana, 1853 T E
Actaecia opihensis Chilton, 1901 T E
ARMADILLIDAE
Acanthodillo spinosus (Dana, 1853) T E
Coronadillo hamiltoni (Chilton, 1901) T E
Coronadillo milleri (Chilton, 1917) T E
Coronadillo suteri (Chilton, 1915) T E
Cubaris ambitiosa (Budde-Lund, 1885) T E
Cubaris minima Vandel, 1977 T E
Cubaris murina Brandt, 1833 T A
Cubaris tarangensis (Budde-Lund, 1904) T E
Merulana chathamensis (Budde-Lund, 1904) T E
Sphaerilloides? antipodum Vandel, 1977 T E
Sphaerilloides? invisibilis Vandel, 1977 T E
Sphaerilloides? macmahoni (Chilton, 1901) T E
Sphaerilloides? minimus Vandel, 1977 T E
Sphaerilloides? rugulosus (Miers, 1876) T E
Sphaerilloides? tuberculatus Vandel, 1977 T E
Spherillo bipunctatus Budde-Lund 1904 T E
Spherillo brevis Budde-Lund, 1904 T E
Spherillo danae Heller, 1865 T E
Spherillo inconspicuus (Miers, 1876) T E
Spherillo marginatus Budde-Lund, 1904 T E
Spherillo monolinus Dana, 1853 T E
Spherillo rufomarginatus Budde-Lund, 1904 T E
Spherillo setaceus Budde-Lund, 1904 T E
Spherillo speciosus (Dana, 1853) T E
Spherillo squamatus Budde-Lund, 1904 T E
Reductoniscus watti Vandel, 1977 T E
ARMADILLIDIIDAE
Armadillidium vulgare (Latreille, 1804) T A
LIGIIDAE
Ligia exotica Roux, 1828 T
Ligia novizealandiae Dana, 1853 T E
ONISCIDAE
Phalloniscus armatus Bowley, 1935 T E
Phalloniscus bifidus Vandel, 1977 T E
Phalloniscus bowleyi Vandel, 1977 T E
Phalloniscus chiltoni Bowley, 1935 T E
Phalloniscus cooki (Filhol, 1885) T E
Phalloniscus forsteri Vandel, 1977 T E
Phalloniscus kenepurensis (Chilton, 1901) T E
Phalloniscus lamellatus Vandel, 1977 T E
Phalloniscus minimus Vandel, 1977 T E
Phalloniscus montanus Vandel, 1977 T E
Phalloniscus occidentalis Vandel, 1977 T E
Phalloniscus propinquus Vandel, 1977 T E
Phalloniscus punctatus (Thomson, 1879) T E
PHILOSCIIDAE
Adeloscia dawsoni Vandel, 1977 T E
Okeaninoscia oliveri (Chilton, 1911) T E
Papuaphiloscia hurleyi Vandel, 1977 T
CRUSTACEA
Paraphiloscia brevicornis (Budde-Lund, 1912) T E
Paraphiloscia fragilis (Budde-Lund, 1904) T E
Philoscia novaezealandiae Filhol, 1885 T E
Philoscia pubescens (Dana, 1853) T E
Stephenoscia bifrons Vandel, 1977 T E
PORCELLIONIDAE
Porcellio scaber Latreille, 1804 T A
Porcellionides pruinosus (Brandt, 1833) T A
SCYPHACIDAE
Deto aucklandiae (Thomson, 1879) T E
Deto bucculenta (Nicolet, 1849) T
Scyphax ornatus Dana, 1853 T E
Scyphoniscus magnus Chilton, 1909 T E
Scyphoniscus waitatensis Chilton, 1901 T E
STYLONISCIDAE
Notoniscus australis (Chilton, 1909) T E
Notoniscus helmsii (Chilton, 1901) T E
Styloniscus commensalis (Chilton, 1910) T E
Styloniscus kermadecensis (Chilton, 1911) T E
Styloniscus magellanicus Dana, 1853 T
Styloniscus otakensis Chilton, 1901 T E
Styloniscus phormianus (Chilton, 1901) T E
Styloniscus thomsoni (Chilton, 1885) T E
Styloniscus phormianus (Chilton, 1901) T E
Styloniscus thomsoni (Chilton, 1885) T E
TRACHELIPODIDAE
Nagurus nanus (Budde-Lund, 1908) T A
TRICHONISCIDAE
Haplophthalmus danicus Budde-Lund, 1885 T A
Infraorder TYLOMORPHA
TYLIDAE
Tylos neozelanicus Chilton, 1901 T E
Order TANAIDACEA
Suborder APSEUDOMORPHA
APSEUDIDAE
Apseudes larseni Knight & Heard, 2006 E
Apseudes meridionalis Richardson, 1912*
Apseudes spectabilis Studer, 1883*
Apseudes spp. (9)
Gollumudes spp. (2?) NIWA G. Bird
Leviapseudes galatheae Wolff, 1956* E
Leviapseudes segonzaci Bacescu, 1981*
Spinosapseudes setosus (Lang, 1968) E
Taraxapseudes diversus (Lang, 1968)*
METAPSEUDIDAE
Apseudomorpha timaruvia (Chilton, 1882) E
Cyclopoapseudes latus (Chilton, 1883) E
Metapseudes aucklandiae Stephensen, 1927 E
Synapseudes n. spp. (2)*
PAGURAPSEUDIDAE
Pagurapseudes? sp.*
SPHYRAPIDAE
Kudinopasternakia dispar (Lang, 1968)*
INCERTAE SEDIS
Gen. et sp. indet. NIWA J. Sieg/G. Bird
Suborder NEOTANAIDOMORPHA
NEOTANAIDAE
Herpotanais kirkegaardi Wolff, 1956
Neotanais barfoedi Wolff, 1956
Neotanais hadalis Wolff, 1956
Neotanais mesostenoceps Gardiner, 1975*
Neotanais robustus Wolff, 1956
Neotanais vemae Gardiner, 1975*
Neotanais sp. NIWA G. Bird
Suborder TANAIDOMORPHA
AGATHOTANAIDAE
Agathotanais spinipoda Larsen, 1999*
Paragathotanais sp. NIWA G. Bird*
Paranarthrura fortispina Sieg, 1986*
Paranarthrura meridionalis Sieg, 1986*
Paranarthrura spp. (2)*
223
NEW ZEALAND INVENTORY OF BIODIVERSITY
ANARTHRURIDAE
Siphonolabrum sp. NIWA G. Bird
Gen. et spp. indet. (2) NIWA G. Bird
COLLETTEIDAE
Collettea cylindratoides Larsen, 1999*
Leptognathiella spp. (2) NIWA G. Bird
Libanius sp. NIWA G. Bird
Macrinella spp. (2?) NIWA G. Bird
LEPTOCHELIIDAE
Konarus sp. G. Bird
Leptochelia mirabilis Stebbing, 1905
LEPTOGNATHIIDAE
Leptognathia spp. (>3)*
NOTOTANAIDAE
Nototanais sp. G. Bird Ca
PARATANAIDAE
Bathytanais spp. (2) NIWA G. Bird
Paratanais oculatus (Vanhoeffen, 1914) B
Paratanais tenuis (G.M.Thomson, 1880) E
Paratanais sp.* Auckland Is.
Paratanais spp. (3)*
PSEUDOTANAIDAE
Akanthinotanais sp. NIWA G. Bird
Cryptocopoides arcticus (Hansen, 1886)
Cryptocopoides sp. NIWA G. Bird
Mystriocentrus sp. NIWA G. Bird
Pseudotanais nordenskioldi (Sieg, 1977)
Pseudotanais spp. (3)*
TANAELLIDAE
Araphura spp. (2) NIWA G. Bird
Araphuroides sp. NIWA G. Bird
Arthrura monocanthus (Vanhoeffen, 1914) n. comb.*
Tanaella forcifera (Lang, 1968)*
Tanaella spp. (4) NIWA G. Bird
TANAIDAE
Pancoloides litoralis (Vanhöffen, 1914)*
Pancoloides sp.* NIWA G. Bird
Sinelobus stanfordi (Richardson, 1901) F B C (sponge)
Synaptotanais sp. NIWA G. Bird
Tanais sp.*
Zeuxo novaezealandiae (Thomson, 1879) E
Zeuxo phytalensis Sieg, 1980*
Zeuxoides aka Bird, 2008 E
Zeuxoides helleri Sieg, 1980*
Zeuxoides ohlini (Stebbing, 1914)*
Zeuxoides pseudolitoralis Sieg, 1980*
Zeuxoides rimuwhero Bird, 2008 E
Zeuxoides sp.*
TYPHLOTANAIDAE
Hamatipeda spp. (2) NIWA G. Bird
Larsenotanais sp. NIWA G. Bird
Meromonakanatha sp. NIWA G. Bird
Paratyphlotanais sp. NIWA G. Bird
Typhlotanais greenwichensis Shiino, 1970*
Typhlotanais spp. (10)*
INCERTAE SEDIS
Akanthophoreus spp. (2) NIWA G. Bird
Chauliopleona spp. (2) NIWA G. Bird
Exspina typica Lang, 1968 C (holothurian)
Mirandotanais vorax Kussakin & Tzareva, 1974*
Stenotanais sp. NIWA G. Bird
Tanaopsis spp. (2) NIWA G. Bird
Order CUMACEA
BODOTRIIDAE
Apocuma n. sp. 1 B E
Bathycuma longirostre Calman, 1905 B
Cyclaspis argus Zimmer, 1902 E
Cyclaspis coelebs Calman, 1907 E
Cyclaspis elegans Calman, 1907 E
Cyclaspis laevis Thomson, 1892
Cyclaspis similis Calman, 1907
Cyclaspis tasmanica Jones, 1969 B E
Cyclaspis thomsoni Calman, 1907
Cyclaspis triplicata Calman, 1907 E
224
Cyclaspis n. sp. 1 B E
Cyclaspis n. sp. 2 B E
Cyclaspis n. sp. ?3 E
Gaussicuma scabra Jones, 1969 B E
Gaussicuma n. sp. 1 B E
Pomacuma australiae (Zimmer, 1921)
DIASTYLIDAE
Colurostylis castlepointensis Gerken & Lörz, 2007 E
Colurostylis lemurum Calman, 1917 E
Colurostylis longicauda Jones, 1963 E
Colurostylis pseudocuma Calman, 1911 E
Colurostylis stenocuma Lomakina, 1968 E
Diastylis acuminata Jones, 1960 E
Diastylis delicata Jones, 1969 B E
Diastylis insularum (Calman, 1908) E
Diastylis neozelanica Thomson, 1892 E
Diastylopsis crassior Calman, 1911 E
Diastylopsis elongata Calman, 1911 E
Diastylopsis thileniusi (Zimmer, 1902) E
Leptostylis profunda Jones, 1969 E
Leptostylis recalvastrata Hale, 1945
Makrokylindrus? mersus Jones, 1969 B E
Makrokylindrus neptunius Jones, 1969 E (abyssal)
Makrokylindrus sp. 1 B E
Paradiastylis? bathyalis Jones, 1969 E
Vemakylindrus sp. 1 E
GYNODIASTYLIDAE
Allodiastylis acanthanasillos Gerken, 2001 E
Axiogynodiastylis fimbriata Gerken, 2001 B E
Axiogynodiastylis kopua Gerken, 2001 E
Gynodiastylis carinata Calman, 1911 E
Gynodiastylis koataata Gerken, 2001 E
Gynodiastylis milleri Jones, 1963 E
Litogynodiastylis laevis (Calman, 1911) E
LAMPROPIDAE
Hemilamprops pellucidus Zimmer, 1908 S B
Hemilamprops ?n. sp. 1 E
Hemilamprops n. sp. 2 B E
Mesolamprops sp. B E
Paralamprops sp. 1 B E
Paralamprops sp. 2 B E
Paralamprops? sp. 3 B E
Paralamprops? sp. 4 B E
Watlingia cassis Gerken, 2010 E
Watlingia chathamensis Gerken, 2010 E
LEUCONIDAE
Eudorella hurleyi Jones, 1963 E
Eudorella truncatula (Bate, 1856) ?A
Eudorellopsis resima Calman, 1907 E
Hemileucon comes Calman, 1907 E
Hemileucon uniplicatus Calman, 1907 E
Heteroleucon akaroensis Calman, 1907 E
Leucon (Alytoleucon) sp. B E
Leucon (Crymoleucon) heterostylis Calman, 1907 E
Leucon (C.) sp. B E
Leucon (Epileucon) latispina Jones, 1963 E
Leucon (?n. subgen.) sp. B E
Paraleucon suteri Calman, 1907 E
NANNASTACIDAE
Campylaspis inornata Jones, 1969 B E
Campylaspis rex Gerken & Ryder, 2002 B E
Campylaspis sp. 2 B E
Campylaspis sp. 3 B E
Campylaspis sp. 4 B E
Campylaspis sp. 5 B E
Procampylaspis sp. 1 B E
Procampylaspis sp. 2 B E
Scherocumella pilgrimi (Jones, 1963) E
Styloptocuma sp. 1 B E
Gen. nov. et n. sp. B
Order EUPHAUSIACEA
EUPHAUSIIDAE
Euphausia longirostris Hansen, 1908
Euphausia lucens Hansen, 1905
Euphausia recurva Hansen, 1905
Euphausia similis G.O. Sars, 1883
Euphausia s. armata Hansen, 1911
Euphausia spinifera G.O. Sars, 1883
Euphausia vallentini Stebbing, 1900.
Nematobrachion flexipes (Ortmann, 1893)
Nematosceles megalops G.O. Sars, 1883
Nematosceles microps G.O. Sars, 1883
Nyctiphanes australis G.O. Sars, 1883
Stylocheiron abbreviatum G.O.Sars, 1883
Stylocheiron elongatum G.O. Sars, 1883
Stylocheiron longicorne G.O. Sars, 1883
Stylocheiron maximum Hansen, 1908
Stylocheiron suhmi G.O. Sars, 1883
Thysanoessa gregaria G.O. Sars, 1883
Thysanoessa macrura G.O. Sars, 1883
Thysanopoda acutifrons Holt & Tattersall, 1905
Thysanopoda obtusifrons G.O. Sars, 1883
Order DECAPODA
Suborder DENDROBRANCHIATA
ARISTEIDAE
Aristaeomorpha foliacea (Risso, 1826)
Aristaeopsis edwardsiana (Johnson, 1867)
Aristeus semidentatus Bate, 1881
Austropenaeus cf. nitidus (Barnard, 1947)
BENTHESICYMIDAE
Benthesicymus cereus Burkenroad, 1936
Benthesicymus investigatoris Alcock & Anderson,
1899
Gennadas capensis Calman, 1925 Pe
Gennadas gilchristi Calman, 1925 Pe
Gennadas incertus (Balss, 1927)
Gennadas kempi Stebbing, 1914 Pe
Gennadas tinayrei Bouvier, 1906 Pe
LUCIFERIDAE
Lucifer typus H. Milne Edwards, 1837 Pe
PENAEIDAE
Funchalia villosa (Bouvier, 1905) Pe
Funchalia woodwardi Johnson, 1867 Pe
SERGESTIDAE
Sergestes arcticus Kröyer, 1855 Pe
Sergestes disjunctus Burkenroad, 1940 Pe
Sergestes index Burkenroad, 1940 Pe
Sergestes cf. seminudus Hansen, 1919 Pe
Sergia japonica (Bate, 1881) Pe
Sergia kroyeri (Bate, 1881) Pe
Sergia potens (Burkenroad, 1940) Pe
SICYONIIDAE
Sicyonia inflexa (Kubo, 1940)*
Sicyonia truncata (Kubo, 1949)
SOLENOCERIDAE
Haliporoides sibogae (de Man, 1907)
Hymenopenaeus obliquirostris (Bate, 1881)
Solenocera comata Stebbing 1915
Infraorder CARIDEA
ALPHEIDAE
Alpheopsis garricki Yaldwyn, 1971 E
Alpheus euphrosyne richardsoni Yaldwyn, 1971 E
Alpheus hailstonei Coutière, 1905
Alpheus novaezealandiae Miers, 1876
Alpheus socialis Heller, 1865
Athanas indicus Coutière, 1903
Betaeopsis aequimanus (Dana, 1852) E
ALVINOCARIDIDAE
Alvinocaris alexander Ahyong, 2009 E
Alvinocaris longirostris Kikuchi & Ohta, 1995
Alvinocaris niwa Webber, 2004 E
Nautilocaris saintlaurentae Komai & Segonzac, 2004
ATYIDAE
Paratya curvirostris (Heller, 1862) F E
CAMPYLONOTIDAE
Campylonotus rathbunae Schmitt, 1926
CRANGONIDAE
PHYLUM ARTHROPODA
Aegaeon lacazei (Gourret, 1888)
Metacrangon knoxi (Yaldwyn, 1960) E
Metacrangon richardsoni (Yaldwyn, 1960) E
Philocheras acutirostratus (Yaldwyn, 1960) E
Philocheras australis (Thomson, 1879) E
Philocheras chiltoni (Kemp, 1911) E
Philocheras hamiltoni (Yaldwyn, 1971) E
Philocheras pilosoides (Stephensen, 1927) E
Philocheras quadrispinosus (Yaldwyn, 1971) E
Philocheras yaldwyni (Zarenkov, 1968) E
Parapontophilus junceus Bate, 1888 E
Prionocrangon curvicaulis Yaldwyn, 1960
DISCIADIDAE
Discias cf. exul Kemp, 1920
HIPPOLYTIDAE
Alope spinifrons (H. Milne Edwards, 1837) E
Bathyhippolyte yaldwyni Hayashi & Miyake, 1970 E
Hippolyte bifidrostris (Miers, 1876) E
Hippolyte multicolorata Yaldwyn, 1971 E
Lebbeus cristatus Ahyong, 2009 E
Lebbeus wera Ahyong, 2009 E
Leontocaris alexander Poore, 2009
Leontocaris amplectipes Bruce, 1990
Leontocaris yarramundi Taylor & Poore, 1998
Lysmata morelandi (Yaldwyn, 1971)
Lysmata trisetacea (Heller, 1861)
Lysmata vittata (Stimpson, 1860)
Merhippolyte chacei Kensley, Tranter & Griffin, 1987
Nauticaris marionis Bate, 1888
Tozeuma novaezealandiae Borradaile, 1916 E
GLYPHOCRANGONIDAE
Glyphocrangon caeca Wood-Mason & Alcock, 1891
Glyphocrangon lowryi Kensley, Tranter & Griffin,
1987
Glyphocrangon regalis Bate, 1888
Glyphocrangon sculpta (Smith, 1882)
NEMATOCARCINIDAE
Lipkius holthuisi Yaldwyn, 1960
Nematocarcinus cf. exilis (Bate, 1888) ZMUC
Nematocarcinus gracilis Bate, 1888
Nematocarcinus hiatus Bate, 1888
Nematocarcinus longirostris Bate, 1888
Nematocarcinus novaezealandicus Burukovsky, 2006
Nematocarcinus serratus Bate, 1888
Nematocarcinus undulatipes Bate, 1888
Nematocarcinus webberi Burukovsky, 2006
Nematocarcinus yaldwyni Burukovsky, 2006
OGYRIDIDAE
Ogyrides delli Yaldwyn, 1971
OPLOPHORIDAE
Acanthephyra brevirostris Smith, 1885 Pe
Acanthephyra eximia Smith, 1884 Pe
Acanthephyra pelagica (Risso, 1816) Pe
Acanthephyra quadrispinosa Kemp, 1939 Pe
Acanthephyra smithi Kemp, 1939 Pe
Ephyrina figueirai Crosnier & Forest, 1973 Pe
Heterogenys microphthalma (Smith, 1885) Pe
Hymenodora glacialis (Buchholz, 1874) Pe
Janicella spinicauda (A. Milne Edwards, 1883) Pe
Kemphyra corallina (A. Milne Edwards, 1883) Pe
Meningadora mollis Smith, 1882 Pe
Meningadora vesca (Smith, 1886) Pe
Notostomus auriculatus Barnard, 1950 Pe
Notostomus japonicus Bate, 1888 Pe
Oplophorus novaezeelandiae de Man, 1931 Pe
Oplophorus spinosus (Brullé, 1839) Pe
Systellaspis debilis (A. Milne Edwards, 1881) Pe
Systellaspis pellucida (Filhol, 1885) Pe
PALAEMONIDAE
Hamiger novaezealandiae (Borradaile, 1916) E
Palaemon affinis H. Milne Edwards, 1937 E
Periclimenes fenneri Bruce, 2005
Periclimenes tangeroa Bruce, 2005
Periclimenes yaldwyni Holthuis, 1959 E
PANDALIDAE
Chlorotocus novaezealandiae (Borradaile, 1916)
Heterocarpus laevigatus Bate, 1888
Notopandalus magnoculus (Bate, 1888) E
Plesionika costelloi (Yaldwyn, 1971)
Plesionika martia (A.Milne Edwards, 1883)
Plesionika spinipes Bate, 1888
PASIPHAEIDAE
Alainopasiphaea australis (Hanamura, 1989)
Eupasiphae gilesii (Wood-Mason, 1892) Pe
Parapasiphae compta Smith, 1884 Pe
Parapasiphae sulcatifrons Smith, 1884 Pe
Pasiphaea barnardi Yaldwyn, 1971 Pe
Pasiphaea burukovskyi Wasmer, 1992 Pe
Pasiphaea grandicula Burukovsky, 1976 Pe
Pasiphaea notosivado Yaldwyn, 1971 Pe
Pasiphaea tarda Kröyer, 1845 Pe
Psathyrocaris infirma Alcock & Anderson, 1894 Pe
PROCESSIDAE
Processa moana Yaldwyn, 1971 E
RHYNCHOCINETIDAE
Rhynchocinetes balssi Gordon, 1936
Rhynchocinetes ikatere Yaldwyn, 1971 E
STYLODACTYLIDAE
Stylodactyloides crosnieri Cleva, 1990
Stylodactylus discissipes Bate, 1888 E
Suborder PLEOCYEMATA
Infraorder STENOPODIDEA
SPONGICOLIDAE
Spongicoloides novaezelandiae Baba, 1980 E
Spongiocaris yaldwyni Bruce & Baba, 1973 E
STENOPODIDAE
Stenopus hispidus (Olivier, 1811)
Infraorder ASTACIDEA
NEPHROPIDAE
Metanephrops challengeri (Balss, 1914) E
Nephropsis suhmi Bate, 1888
PARASTACIDAE
Paranephrops planifrons White, 1842 F E
Paranephrops zealandicus (White, 1847) F E
Infraorder AXIIDEA
AXIIDAE
Axius cf. werribee (Poore & Griffin, 1979) MNZ
Calocarides vigila Sakai, 1992 E
Calocaris isochela Zarenkov, 1898 E
Dorphinaxius kermadecensis (Chilton, 1911)
Eiconaxius kermadeci Bate, 1888 E
Eiconaxius parvus Bate, 1888
Eucalastacus torbeni Sakai, 1992 E
Spongiaxius novaezealandiae (Borradaile, 1916) E
CALLIANASSIDAE
Corallianassa articulata (Rathbun, 1906)
Corallianassa cf. collaroy (Poore & Griffin, 1979)
MNZ
‘Callianassa’ filholi (A. Milne Edwards, 1879) E
Vulcanocalliax sp. E
CTENOCHELIDAE
Ctenocheles maorianus Powell, 1949 E
Infraorder GEBIIDEA
LAOMEDIIDAE
Jaxea novaezealandiae Wear & Yaldwyn, 1966 E
UPOGEBIIDAE
Acutigebia danai (Miers, 1876) E
Upogebia hirtifrons (White, 1847) E
Infraorder ACHELATA
PALINURIDAE
Jasus edwardsii (Hutton, 1875)
Sagmariasus verreauxi (H. Milne Edwards, 1851)
Projasus parkeri (Stebbing, 1902)
Infraorder POLYCHELIDA
CRUSTACEA
POLYCHELIDAE
Pentacheles laevis Bate, 1878
Pentacheles validus A. Milne Edwards, 1880
Polycheles enthrix (Bate, 1878)
Polycheles kermadecensis (Sund, 1920)
Stereomastis nana (Smith, 1884)
Stereomastis sculpta (Smith, 1880)
Stereomastis suhmi Bate, 1878
Stereomastis surda (Galil, 2000)
Willemoesia pacifica Sund, 1920
SCYLLARIDAE
Antarctus mawsoni (Bage, 1938)
Antipodarctus aoteanus (Powell, 1949) E
Arctides antipodarum Holthuis, 1960
Ibacus alticrenatus Bate, 1888
Ibacus brucei Holthuis, 1977
Scyllarides haanii (de Haan, 1841)
Infraorder ANOMURA
ALBUNEIDAE
Albunea microps Miers, 1878
CHIROSTYLIDAE
Chirostylus novaecaledoniae Baba, 1991
Eumunida pacifica Gordon, 1930
Gastroptychus novaezelandiae (Baba, 1974)
Gastroptychus rogeri (Baba, 2000)
Uroptychodes epigaster Baba, 2004
Uroptychodes spinimarginatus (Henderson, 1885)
Uroptychus alcocki Ahyong & Poore, 2004
Uroptychus australis (Henderson, 1885)
Uroptychus bicavus Baba & de Saint Laurent, 1992
Uroptychus cardus Ahyong & Poore, 2004
Uroptychus empheres Ahyong & Poore, 2004
Uroptychus flindersi Ahyong & Poore, 2004
Uroptychus gracilimanus (Henderson, 1885)
Uroptychus kaitara Schnabel, 2009
Uroptychus latus Ahyong & Poore, 2004
Uroptychus longicheles Ahyong & Poore, 2004
Uroptychus longvae Ahyong & Poore, 2004
Uroptychus maori Borradaile, 1916 E
Uroptychus multispinosus Ahyong & Poore, 2004
Uroptychus novaezealandiae Borradaile, 1916 E
Uroptychus paku Schnabel, 2009
Uroptychus paracrassior Ahyong & Poore, 2004
Uroptychus pilosus Baba, 1981
Uroptychus politus (Henderson, 1885) E
Uroptychus raymondi Baba, 2000
Uroptychus rutua Schnabel, 2009
Uroptychus scambus Benedict, 1902
Uroptychus spinirostris (Ahyong & Poore, 2004)
Uroptychus thermalis Baba & de Saint Laurent, 1992
Uroptychus toka Schnabel, 2009
Uroptychus tomentosus Baba, 1975 E
Uroptychus webberi Schnabel, 2009
Uroptychus yaldwyni Schnabel, 2009
DIOGENIDAE
Calcinus imperialis Whitelegge, 1901
Cancellus frontalis Forest & McLaughlin, 2000 E
Cancellus laticoxa Forest & McLaughlin, 2000 E
Cancellus rhynchogonus Forest & McLaughlin, 2000
E
Cancellus sphraerogonus Forest & McLaughlin,
2000 E
Dardanus arroser (Herbst, 1796)
Dardanus hessii (Miers, 1884)
Paguristes barbatus (Heller, 1862) E
Paguristes pilosus (H. Milne Edwards, 1836) E
Paguristes setosus (H. Milne Edwards, 1848) E
Paguristes subpilosus Henderson, 1888 E
GALATHEIDAE
Agononida incerta (Henderson, 1888)
Agononida marini (Macpherson, 1994)
Agononida nielbrucei Vereshchaka, 2005 E
Agononida procera Ahyong & Poore, 2004
Agononida squamosa (Henderson, 1885)
225
NEW ZEALAND INVENTORY OF BIODIVERSITY
Allogalathea elegans (Adams & White, 1848)
Galathea whiteleggii Grant & McCulloch, 1906
Galacantha quiquei Macpherson, 2007
Galacantha rostrata A. Milne Edwards, 1880
Leiogalathea laevirostris (Balss, 1913)
Munida acacia Ahyong, 2007
Munida chathamensis Baba, 1974 E
Munida collier Ahyong, 2007
Munida eclepsis Macpherson, 1994
Munida erato Macpherson, 1994
Munida endeavourae Ahyong & Poore, 2004
Munida exilis Ahyong, 2007
Munida gracilis Henderson, 1885 E
Munida gregaria (Fabricius, 1793)
Munida icela Ahyong, 2007
Munida isos Ahyong & Poore, 2004
Munida kapala Ahyong & Poore, 2004
Munida notata Macpherson, 1994
Munida psylla Macpherson, 1994
Munida notialis Baba, 2005
Munida rubrimana Ahyong, 2007
Munida spinicruris Ahyong & Poore, 2004
Munida zebra Macpherson, 1994
Munidopsis antonii (Filhol, 1884)
Munidopsis bractea Ahyong, 2007
Munidopsis comarge Taylor, Ahyong & Andreakis,
2010
Munidopsis kaiyoae Baba, 1974 E
Munidopsis marginata (Henderson, 1885)
Munidopsis maunga Schnabel & Bruce, 2006
Munidopsis papanui Schnabel & Bruce, 2006
Munidopsis proales Ahyong & Poore, 2004
Munidopsis cf. serricornis (Lovén, 1852)
Munidopsis tasmaniae Ahyong & Poore, 2004
Munidopsis treis Ahyong & Poore, 2004
Munidopsis valdiviae (Balss, 1913)
Munidopsis victoriae Baba & Poore, 2002
Onconida alaini Baba & de Saint Laurent, 1996
Paramunida antipodes Ahyong & Poore, 2004
Phylladiorhynchus integrirostris (Dana, 1852)
Phylladiorhynchus pusillus (Henderson, 1885)
Tasmanida norfolkae Ahyong, 2007
LITHODIDAE
Lithodes aotearoa Ahyong, 2010 E
Lithodes jessica Ahyong, 2010
Lithodes macquariae Ahyong, 2010
Lithodes robertsoni Ahyong, 2010 E
Neolithodes brodiei Dawson & Yaldwyn, 1970
Neolithodes bronwynae Ahyong, 2010
Paralomis dawsoni Macpherson, 2001
Paralomis echidna Ahyong, 2010
Paralomis hirtella Saint Laurent & Macpherson,
1997
Paralomis poorei Ahyong, 2010
Paralomis staplesi Ahyong, 2010
Paralomis webberi Ahyong, 2010 E
Paralomis zealandica Dawson & Yaldwyn, 1971 E
PAGURIDAE
Bathypaguropsis cruentus de Saint Laurent &
McLaughlin, 2000 E
Bathypaguropsis yaldwyni McLaughlin, 1994
Catapagurus spinicarpus de Saint Laurent &
McLaughlin, 2000 E
Diacanthurus ecphyma McLaughlin & Forest, 1997
Diacanthurus rubricatus (Henderson, 1888) E
Diacanthurus spinulimanus (Miers, 1876) E
Lophopagurus (Australeremus) cookii (Filhol, 1883) E
Lophopagurus (A.) cristatus (H. Milne Edwards,
1836) E
Lophopagurus (A.) eltaninae (McLaughlin & Gunn,
1992) E
Lophopagurus (A.) kirkii (Filhol, 1883 E
Lophopagurus (A.) laurentae (McLaughlin & Gunn,
1992) E
Lophopagurus (A.) stewarti (Filhol, 1883) E
226
Lophopagurus (A.) triserratus (Ortmann, 1892)
Lophopagurus (Lophopagurus) foresti McLaughlin &
Gunn, 1992 E
Lophopagurus (L.) lacertosus (Henderson, 1888) E
Lophopagurus (L.) ?nanus (Henderson, 1888)
Lophopagurus (L.) nodulosus McLaughlin & Gunn,
1992 E
Lophopagurus (L.) pumilis de Saint Laurent &
McLaughlin, 2000 E
Lophopagurus (L.) thompsoni (Filhol, 1885) E
Michelopagurus? sp. E
Pagurixus hectori (Filhol, 1883) E
Pagurixus kermadecensis de Saint Laurent &
McLaughlin, 2000 E
Pagurodes inarmatus Henderson, 1888
Pagurojacquesia polymorpha (de Saint Laurent &
McLaughlin, 1999)
Pagurus albidianthus de Saint Laurent &
McLaughlin, 2000 E
Pagurus iridocarpus de Saint Laurent & Mclaughlin,
2000 E
Pagurus novizealandiae (Dana, 1852) E
Pagurus sinuatus (Stimpson, 1858)
Pagurus traversi (Filhol, 1885) E
Porcellanopagurus chiltoni de Saint Laurent &
McLaughlin, 2000
Porcellanopagurus edwardsi Filhol, 1885 E
Porcellanopagurus filholi de Saint Laurent &
McLaughlin, 2000
Porcellanopagurus tridentatus Whitelegge, 1900
Propagurus deprofundis (Stebbing, 1924)
PARAPAGURIDAE
Oncopagurus sp. E
Paragiopagurus diogenes (Whitelegge, 1900)
Paragiopagurus hirsutus (de Saint Laurent, 1972)
Parapagurus abyssorum (Filhol, 1885)
Parapagurus bouvieri Stebbing, 1910
Parapagurus latimanus Henderson, 1888
Parapagurus richeri Lemaitre, 1999
Sympagurus dimorphus (Studer, 1883)
Sympagurus papposus Lemaitre, 1996
PORCELLANIDAE
Pachycheles pisoides (Heller, 1865)
Petrocheles spinosus (Miers, 1876) E
Petrolisthes elongatus (H. Milne Edwards, 1837)
Petrolisthes lamarckii (Leach, 1820)
Petrolisthes novaezelandiae Filhol, 1885 E
PYLOCHELIDAE
Cheiroplatea pumicicola Forest, 1987
Pylocheles mortensenii Boas, 1926
Trizocheles brachyops Forest & de Saint Laurent,
1987
Trizocheles perplexus Forest, 1987 E
Trizocheles spinosus (Henderson, 1888)
Trizocheles pilgrimi Forest & McLaughlin, 2000
Infraorder BRACHYURA
AETHRIDAE
Actaeomorpha erosa Miers, 1877
ATELECYCLIDAE
Pteropeltarion novaezelandiae Dell, 1972 E
Trichopeltarion fantasticum Richardson & Dell,
1964 E
Trichopeltarion janetae Ahyong, 2008
BELLIIDAE E
Heterozius rotundifrons A. Milne Edwards, 1867 E
BYTHOGRAEIDAE
Gandalfus puia McLay, 2007
CALAPPIDAE
Mursia australiensis Campbell, 1971
Mursia microspina Davie & Short, 1989
CANCRIDAE
Glebocarcinus amphioetus (Rathbun, 1898) A
Metacarcinus novaezelandiae (Hombron &
Jacquinot, 1846)
Romaleon gibbulosus (Rathbun, 1898) A
CRYPTOCHIRIDAE
Cryptochirus coralliodytes Heller, 1861
CYMONOMIDAE
Cymonomus aequilonius Dell, 1971 E
Cymonomus bathamae Dell, 1971 E
Cymonomas clarki Ahyong, 2008 E
DROMIIDAE
Cryptodromiopsis unidentata (Rüppell, 1830)
Metadromia wilsoni (Fulton & Grant, 1902)
Tumidodromia dormia (Linnaeus, 1763)
DYNOMENIDAE
Dynomene pilumnoides Alcock, 1900
Metadynomene tanensis (Yokoya, 1933)
EPIALTIDAE
Huenia heraldica (de Haan, 1839)
Leptomaia tuberculata Griffin & Tranter, 1986
Oxypleurodon wanganella Webber & Richer de
Forges, 1995 E
Rochinia ahyongi McLay, 2009 E
Rochinia riversandersoni (Alcock, 1895)
ERIPHIIDAE
Bountiana norfolcensis (Grant & McCulloch, 1907)
ETHUSIDAE
Ethusina castro Ahyong, 2008 E
Ethusina rowdeni Ahyong, 2008 E
GERYONIDAE
Chaceon bicolor Manning & Holthuis, 1989
Chaceon yaldwyni Manning, Dawson & Webber,
1990 E
GONEPLACIDAE
Goneplax marivenae Komatsu & Takeda, 2004
Neommatocarcinus huttoni (Filhol, 1886) E
Pycnoplax meridionalis (Rathbun, 1923)
Pycnoplax victoriensis (Rathbun, 1923)
Thyroplax truncata Castro, 2007
GRAPSIDAE
Geograpsus grayi (H. Milne Edwards, 1853) T
Leptograpsus variegatus (Fabricius, 1793)
Pachygrapsus minutus A. Milne Edwards, 1873
Planes major (MacLeay, 1838)
Planes marinus Rathbun, 1914
HOMOLIDAE
Dagnaudus petterdi (Grant, 1905)
Homola orientalis Henderson, 1888
Homola ranunculus Guinot & Richer de Forges,
1995
Homolochunia kullar Griffin & Brown, 1976
Yaldwynopsis spinimanus (Griffin, 1965)
HOMOLODROMIIDAE
Dicranodromia delli Ahyong, 2008 E
Dicranodromia spinulata Guinot, 1995
Homolodromia kai Guinot, 1993
HYMENOSOMATIDAE
Amarinus lacustris (Chilton, 1882) F
Elamena longirostris Filhol, 1885 E
Elamena momona Melrose, 1975 E
Elamena producta Kirk, 1879 E
Halicarcinus cookii (Filhol, 1885) E
Halicarcinus innominatus Richardson, 1949
Halicarcinus ovatus Stimpson, 1858
Halicarcinus planatus (Fabricius, 1775)
Halicarcinus tongi Melrose, 1975 E
Halicarcinus varius (Dana, 1851) E
Halicarcinus whitei (Miers, 1876) E
Halimena aotearoa Melrose, 1975 E
Hymenosoma depressum Hombron & Jacquinot,
1846 E
Neohymenicus pubescens (Dana, 1851) E
INACHIDAE
Achaeus akanensis Sakai, 1938
Achaeus curvirostris (A. Milne Edwards, 1873)
Achaeus kermadecensis Webber & Takeda, 2005 E
Cyrtomaia cornuta Richer de Forges & Guinot, 1988
Cyrtomaia lamellata Rathbun, 1906
PHYLUM ARTHROPODA
Dorhynchus ramusculus (Baker, 1906)
Platymaia maoria Dell, 1963
Platymaia wyvillethomsoni Miers, 1886
Trichoplatus huttoni A. Milne Edwards, 1876 E
Vitjazmaia latidactyla Zarenkov, 1994
INACHOIDIDAE
Pyromaia tuberculata (Lockington, 1877) A
LATREILLIIDAE
Eplumula australiensis (Henderson, 1888)
Latreillia metanesa Williams, 1982
LEUCOSIIDAE
Bellidilia cheesmani (Filhol, 1886) E
Ebalia humilis Takeda, 1977
Ebalia jordani Rathbun, 1906
Ebalia tuberculosa (A. Milne Edwards, 1873)
Ebalia webberi Komatsu & Takeda, 2007 E
Merocryptus lambriformis A. Milne Edwards, 1873
Tanaoa distinctus (Rathbun, 1893)
Tanaoa pustulosus (Wood-Mason in Wood-Mason
& Alcock, 1891)
MACROPHTHALMIDAE
Macrophthalmus (Hemiplax) hirtipes (Jacquinot in
Hombron & Jacquinot, 1846) E
MAJIDAE
Eurynolambrus australis H. Milne Edwards & Lucas,
1841 E
Eurynome bituberculata Griffin, 1964 E
Jacquinotia edwardsii (Jacquinot, 1853) E
Leptomithrax australis (Jacquinot, 1853) E
Leptomithrax garricki Griffin, 1966 E
Leptomithrax longimanus Miers, 1876
Leptomithrax longipes (Thomson, 1902)
Leptomithrax tuberculatus mortenseni Bennett, 1964
Naxia spinosa (Hess, 1865)
Notomithrax minor (Filhol, 1885)
Notomithrax peronii (H. Milne Edwards, 1834) E
Notomithrax spinosus (Miers, 1879)
Notomithrax ursus (Herbst, 1788)
Prismatopus filholi (A. Milne Edwards, 1876) E
Prismatopus goldsboroughi (Rathbun, 1906)
Schizophroida hilensis (Rathbun, 1906)
Teratomaia richardsoni (Dell, 1960)
MATHILDELLIDAE
Intesius richeri Crosnier & Ng, 2004
Mathildella mclayi Ahyong, 2008 E
Neopilumnoplax nieli Ahyong, 2008
OCYPODIDAE
Ocypode pallidula Jacquinot in Hombron &
Jacquinot, 1846
OZIIDAE
Ozius truncatus H. Milne Edwards, 1834
PALICIDAE
Pseudopalicus declivis Castro, 2000
Pseudopalicus oahuensis (Rathbun, 1906)
Pseudopalicus undulatus Castro, 2000
PARTHENOPIDAE
Actaeomorpha erosa Miers, 1877
Garthambrus allisoni (Garth, 1992)
Garthambrus tani Ahyong, 2008
Platylambrus constrictus (Takeda & Webber, 2007)
PILUMNIDAE
Actumnus griffini Takeda & Webber, 2006 E
Pilumnopeus serratifrons (Kinahan, 1856)
Pilumnus fimbriatus H. Milne Edwards, 1834
Pilumnus lumpinus Bennett, 1964 E
Pilumnus novaezelandiae Filhol, 1886 E
PINNOTHERIDAE
Nepinnotheres atrinicola (Page, 1983) E
Nepinnotheres novaezelandiae (Filhol, 1885) E
PLAGUSIIDAE
Miersiograpsus australiensis Türkay, 1978
Percnon planissimum (Herbst, 1804)
Plagusia chabrus (Linnaeus, 1758)
Plagusia dentipes de Haan, 1835
Plagusia squamosa (Herbst, 1790)
PORTUNIDAE
Caphyra acheronae Takeda & Webber, 2006 E
Charybdis japonica (A. Milne Edwards, 1861) A
Liocarcinus corrugatus (Pennant, 1777)
Nectocarcinus antarcticus (Jacquinot, 1853) E
Nectocarcinus bennetti Takeda & Miyake, 1969 E
Ovalipes catharus (White, 1843)
Ovalipes elongatus Stephenson & Rees, 1968
Ovalipes molleri (Ward, 1933)
Portunus pelagicus (Linnaeus, 1766)
Scylla serrata (Forskål, 1775)
Thalamita danae Stimpson, 1858
Thalamita macrops Montgomery, 1931
RANINIDAE
Lyreidus tridentatus de Haan, 1841
Notosceles pepeke Yaldwyn & Dawson, 2000 E
TRAPEZIIDAE
Calocarcinus africanus Calman, 1909
Trapezia cymodoce (Herbst, 1801)
CRUSTACEA
Trapezia guttata Rüppell, 1830
Trapezia septata Dana, 1852
VARUNIDAE
Austrohelice crassa (Dana, 1851) E
Cyclograpsus insularum Campbell & Griffin, 1966
Cyclograpsus lavauxi H. Milne Edwards, 1853 E
Hemigrapsus crenulatus (H. Milne Edwards, 1837)
Hemigrapsus sexdentatus (H. Milne Edwards, 1837) E
XANTHIDAE
Antrocarcinus petrosus Ng & Chia, 1994
Banareia armata A. Milne Edwards, 1869
Banareia banareias (Rathbun, 1911)
Euryxanthops chiltoni Ng & McLay, 2007 E
Gaillardiellus bathus Davie, 1997
Gaillardiellus rueppelli (Krauss, 1843)
Leptodius nudipes (Dana, 1852)
Liomera yaldwyni Takeda & Webber, 2006 E
Lybia leptochelis (Zehntner, 1894)
Medaeops serenei Ng & McLay, 2007 E
Miersiela haswelli (Miers, 1886)
Nanocassiope sp. Takeda & Webber 2006
Pilodius nigrochrinitus Dana, 1852
Platypodia delli Takeda & Webber, 2006 E
Pseudoliomera helleri (A. Milne Edwards, 1865)
Serenius actaeoides (A. Milne Edwards, 1834)
Xanthias dawsoni Takeda & Webber, 2006 E
Xanthias lamarckii (H. Milne Edwards, 1834)
XENOGRAPSIDAE
Xenograpsus ngatama McLay, 2007 E
Synonyms or possible synonyms in cyclopoid
Copepoda
Diacyclops crassicaudoides (Kiefer, 1928) = D.
bisetosus (Rehberg, 1880)
Eucyclops (Eucyclops) serrulatus (Fischer, 1851) (=
Cyclops novaezealandiae Thomson, 1879)
?Euryte longicauda Philippi, 1843 (= Thorellia
brunnae Boeck, 1864)
?Cyclops strennus strennus Fischer, 1851 (= C. ewarti
Brady, 1888)
Diacyclops bicuspidatus (Claus, 1857) (= Cyclops
gigas, Thomson, 1883)
?Halicyclops magniceps (Lilljeborg, 1853) (= ?C.
aequorus, Thomson, 1883)
?Macrocyclops distinctus (Richard, 1887) = M.
albidus (Jurine, 1820)
?Mesocyclops australiensis (Sars, 1908) (= ?M.
leuckarti)
Checklist of New Zealand fossil Crustacea
Letters in parentheses following new records indicate where material is held, i.e. AUT
(Earth and Oceanic Sciences Research Centre, Auckland University of Technology); GNS
(Institute of Geological and Nuclear Sciences, Lower Hutt); NIWA (National Institute of
Water and Atmospheric Sciences, Wellington); UA (Geology Department, University of
Auckland). Stratigraphic ranges, using abbreviations for New Zealand stages (Cooper
2004), follow each fossil species listing.
SUBPHYLUM CRUSTACEA
Class MAXILLOPODA
Infraclass CIRRIPEDIA
Superorder ACROTHORACICA
Order PYGOPHORA
CRYPTOPHIALIDAE
Australophialus? sp. nov.* Po-Pl (AUT) E
Gen. et sp. indet..* Po-Pl (UoA)
INCERTAE SEDIS
Zapfella sp.* Bm (GNS)
Zapfella? sp.* Ko (UoA)
Superorder RHIZOCEPHALA
Order KENTROGONIDA
SACCULINIDAE?
Gen. et sp. indet. Feldmann 1998 Mio
Pristinolepas haurakiensis (Buckeridge, 1983) Lw-Po E
Pristinolepas pakaurangiensis (Buckeridge, 1983)
Po-Pl E
Pristinolepas waikatoica (Buckeridge, 1983) Ld-Lw E
Pristinolepas n. sp. Ar E
Superorder THORACICA
Order LEPADIFORMES
LEPADIDAE
Lepas ?australis Darwin, 1851 Qu
Lepas clifdenica Buckeridge, 1983 Sl-Tt E
Lepas moturoaensis Maxwell, 1968 Po E
Pristinolepas harringtoni (Laws, 1948) Lw-Pl E
Order SCALPELLIFORMES
ARCOSCALPELLIDAE
Anguloscalpellum complanatum (Withers, 1924)
Lwh-Ld E
Anguloscalpellum cf. complanatum (Withers, 1924)
Po E
Anguloscalpellum crassiforme Buckeridge, 1983 Lwh
227
NEW ZEALAND INVENTORY OF BIODIVERSITY
E
Anguloscalpellum euglyphum (Withers, 1924) LwhLd E
Anguloscalpellum grantmackiei Buckeridge, 1983
Po-Sw E
Anguloscalpellum? striatulum (Withers, 1924) LwhLd E
Anguloscalpellum ungulatum (Withers, 1913) LwhSw E
CALANTICIDAE
Calantica spinilatera Foster, 1979 Ww-Rec E
Cretiscalpellum cf. glabrum (Roemer, 1841) Uk
Cretiscalpellum? sp. nov.* Cn (GNS) E
Cretiscalpellum? sp. Buckeridge 1983 Mp-Dt
Euscalpellum egmontense Buckeridge, 1983 Ww E
Pachyscalpellum cramptoni Buckeridge, 1991 Mp
Pachyscalpellum debodae Buckeridge, 1999 Mh E
Scillaelepas arguta (Withers, 1924) Lwh-Ld E
Scillaelepas? pittensis Buckeridge, 1984 Ab-Ar E
Scillaelepas cf. studeri (Weltner, 1922) Ab-Ar
Scillaelepas waitemata Buckeridge, 1983 Lw-Po E
Smilium calanticoideum Buckeridge, 1983 Dw-Dm
Smilium chathecum Buckeridge, 1984 Pl E
Smilium subplanum (Withers, 1913) Lw-Po E
Zeascalpellum crassum Buckeridge, 1983 Dm-Ab E
Gen. nov. et n. sp.* Mh-Dt (GNS) E
Gen. et sp. indet. Buckeridge 1983 Mp-Mh
EOLEPADIDAE
Eolepas? novaezelandiae Buckeridge 1983 Ce E
ZEUGMATOLEPADAE
Zeugmatolepas? sp. Buckeridge 1983 Kh
Order SESSILIA
Suborder VERRUCOMORPHA
VERRUCIDAE
Metaverruca recta (Aurivillius, 1898) Po-Rec
Verruca nuciformis Buckeridge, 1983 Dm-Po E
Verruca sauria Buckeridge, 2010 Mh E
Verruca tasmanica chatheca Buckeridge, 1983 DwDm E
Verruca t. tasmanica Buckeridge, 1983 Lwh
Suborder BALANOMORPHA
ARCHAEOBALANIDAE
Armatobalanus motuketeketeensis Buckeridge, 1983
Po E
Armatobalanus? sp. Buckeridge 1983 Po E
Striatobalanus zelandicus (Withers, 1924) Sl-Tt E
Notobalanus vestitus (Darwin, 1854) Lw-Rec E
Palaeobalanus lornensis Buckeridge, 1983 Ab-Ak E
Palaeobalanus? waihaoensis Buckeridge, 1983 Ab E
Tasmanobalanus acutus acutus (Withers, 1924)
Pl-Sw E
Tasmanobalanus a. clifdensensis Buckeridge, 1983
Sc E
Tasmanobalanus a. convexus Buckeridge, 1983 Pa E
Tasmanobalanus grantmackiei Buckeridge, 1983
Sw-Ww E
Zullobalanus everetti (Buckeridge, 1983) Lwh E
Zullobalanus novozelandicus (Buckeridge, 1983)
Ld-Lw E
AUSTROBALANIDAE
Austrobalanus imperator aotea Buckeridge, 1983
Ld-Po E
Austrobalanus macdonaldensis Buckeridge, 1983
Lwh E
Epopella eoplicata Buckeridge, 1983 Po E
Epopella cf. plicata Gray, 1843* Wp (AUT) E
Protelminius pomahakensis (Buckeridge, 1984) Ld E
BATHYLASMATIDAE
Bathylasma aucklandicum (Hector, 1888) Lw-Ww E
Bathylasma rangatira Buckeridge, 1983 Dt-Dm E
BALANIDAE
Amphibalanus variegatus (Darwin, 1854) Ww-Rec
Fistulobalanus kondakovi (Tarasov & Zevina, 1957)
228
?Wn
Fosterella chathamensis Buckeridge, 1983 Wo-Wn E
Fosterella tubulatus (Withers, 1924) Wo-Wn E
Notomegabalanus decorus argyllensis (Buckeridge,
1983) Wn-Qu E
Notomegabalanus miodecorus (Buckeridge, 1983)
Sw-Ww E
CHIONELASMATIDAE
Chionelasmus darwini (Pilsbry, 1907) Ak-Rec
CHTHAMALIDAE
Chamaesipho brunnea Moore, 1944 Po-Rec E
CORONULIDAE
Coronula aotea Fleming, 1959 Ww-Wm E
Coronula diadema (Linné, 1767) Wn-Rec
Coronula intermedia Buckeridge, 1983 Wn E
PACHYLASMATIDAE
Eolasma maxwelli Buckeridge, 1983 Dw-Dm E
Pachylasma distortum Buckeridge, 1983 Lwh E
Pachylasma? southlandicum Buckeridge, 1983 Ld-Po
E
Pachylasma veteranum Buckeridge, 1983 Dt-Dm E
Pachylasma sp.* Wp (AUT)
Waikalasma juneae Buckeridge, 1983 Po-Pl E
TETRACLITIDAE
Tesseroplax? maorica Buckeridge, 1983 Lw-Po E
Tesseropora cf. pacifica (Pilsbry, 1928) Po
Tetraclitella nodicostata Buckeridge, 2008 Lw-Po
Class OSTRACODA
All the marine Tertiary species may be regarded as
endemic.
Order ARCHAEOCOPIDA
Gen. et spp. indet. (2) Simes 1977 LPz
Order PALAEOCOPIDA
Suborder BEYRICHICOPIDA
PUNCIIDAE
Puncia goodwoodensis Hornibrook, 1963 Pl E
Order PODOCOPIDA
Suborder PODOCOPINA
BAIRDIIDAE
Bairdia canterburyensis Swanson, 1969 Pl E
Bairdoppilata kerryi Milau, 1993 Po-Rec
Bairdoppilata cf. austracretacea (Bate, 1972) Mh
Bairdoppilata sp. 5052 Dingle 2009 Mh
Neonesidea australis (Chapman, 1914) Ak-Lw
Neonesidea chapmani Whatley & Downing, 1983
Ak-Lw
Neonesidea waitematanensis Milau, 1993 Po E
Neonesidea sp. Ayress 1993 Ab-Rec
BYTHOCYPRIDIDAE
Bythocypris sudaustralis McKenzie, Reyment &
Reyment, 1991 Ak
Bythocypris cf. sudaustralis McKenzie, Reyment &
Reyment, 1991 Mh
Bythocypris cf. chapmani Neale, 1975 Mh
Bythocypris sp. Ayress, 1993 Lwh-Lw
BYTHOCYTHERIDAE
Abyssobythere inequivalva Ayress, Correge, Passlow
& Whatley, 1996 Wc
Bythoceratina decepta Hornibrook, 1952 Wc-Rec
Bythoceratina cf. dubia (Müller, 1908) Ak
Bythoceratina edwardsoni Hornibrook, 1952 Wc-Rec
Bythoceratina maoria Hornibrook, 1952 Sc-Rec
Bythoceratina mestayerae Hornibrook, 1952 Pl-Rec
Bythoceratina powelli Hornibrook, 1952 Ar-Rec
Bythoceratina robusta Milau, 1993 Po
Bythoceratina utilazea Hornibrook, 1952 Pl-Rec
Bythoceratina sp. Ayress 1993 Ld-Lw
Miracythere novaspecta Hornibrook, 1952 Lw-Rec E
Neobuntonia oneroaensis Milau, 1993 Po
Pseudeucythere biplana Ayress, 1995 Ak-Wc
Vitjasiella duplicispina Avress, 1993 Lw-Pl
Vitjasiella ferox (Hornibrook, 1952) Ab-Wc
CYPRIDIDAE
Candona sp. Hornibrook 1955 Wc F
Candonocypris assimilis Sars, 1894 Wc-Rec F
Cypretta viridis (Thomson, 1879) Wc-Rec F
Cypris sp. Hornibrook 1955 Wc F
Heterocypris ciliata (Thomson, 1879) Wc-Rec F
Heterocypris incongruens (Rhamdohr, 1808) Wc-Rec
FE
Ilyodromus stanleyanus (King, 1855) Wc-Rec F
CYTHERALISONIDAE
Cytheralison amiesi Hornibrook, 1953 Lwh-Ld
Cytheralison fava Hornibrook, 1952 Ab-Rec
Cytheralison parafava Ayress, 1993 Ld-Lw
Cytheralison spinosa Ayress, 1993 Ld-Lw
Cytheralison sp. Ayress 1995 Ak
Debissonia hornibrooki Ayress, 2003 Ld-Lw
Debissonia pravacauda (Hornibrook, 1952) Dm-Rec
CYTHERIDAE
Chejudocythere cf. higashikawai Ishizaki, 1981 Ak
Cythere allanthomsoni Chapman, 1926 Sw
Loxocythere crassa Hornibrook, 1952 Po-Rec
Loxocythere kingi Hornibrook, 1952 Pl-Rec
CYTHERIDEIDAE
Cytheridea aoteana Hornibrook, 1952 Wc-Rec E
Cytheridea symmetrica Swanson, 1969 Pl
Cytheridea (Clithrocytheridea) marwicki Hornibrook,
1953 Pl
Hemicytheridea mosaica Hornibrook, 1952 Dm-Rec
Eucythere sulcocostatula Ayress, 1995 Ak-Wc
Eucythere parapubera Whatley & Downing, 1983
Lwh-Ld
Eucythere cf. parapubera Whatley & Downing, 1983
Ak
Eucythere sp. Ayress 1995 Ak-Lw
Eucythere sp. 1 Ayress 1993 Lwh-Lw
Rostrocytheridea pukehouensis Dingle, 2009 E Mh
Rostrocytheridea aff. allaruensis? Krömmelbein,
1975 Cn
Rostrocytheridea? sp. 4992 Dingle 2009 Mh
Rotundracythere gravepuncta Hornibrook, 1952
Ar-Rec
Rotundracythre inaequa Hornibrook, 1952 Wc-Rec
Rotundracythere mytila Hornibrook, 1952 Ld-Rec
Rotundracythere rotunda Hornibrook, 1952 Ar-Rec
Rotundracythere subovalis Hornibrook, 1952 Ar-Rec
Pseudocythere (Pseudocythere) caudata Sars, 1866
Ld-Lw
Pseudocythere (P.) caudata Sars, 1866 Lw-Rec
CYTHEROMATIDAE
Malibaricythere oceanica Yassini & Jones, 1995 Lw
Paracytheroma stilwelli Ayress, 1990 Ld-Pl
Paracytheroma convexa Milau, 1993 Po
Pellucistoma coombsi Ayress, 1990 Ak-Pl
Pellucistoma fordycei Ayress, 1990 Ak-Pl
CYTHERURIDAE
Aversovalva aurea Hornibrook, 1952 Ab-Rec
Aversovalva pteroalata Ayress, 1993 Lwh-Ld n. nud.
Cytheropteron anisovalva Ayress, Correge, Passlow
& Whatley, 1996 Ar-Rec
Cytheropteron cuneatum Ayress, 1996 Ak
Cytheropteron confusum (Hornibrook, 1952) LwhRec
Cytheropteron crassicutum Ayress, 1998 Po-Wn
Cytheropteron curvicaudum Hornibrook, 1952
Lwh-Rec
Cytheropteron dividentum (Hornibrook, 1952)
Lwh-Rec
Cytheropteron dorsocorrugatum Ayress, Correge,
Passlow & Whatley, 1996 Wc
Cytheropteron fornix (Hornibrook, 1952) Ab-Rec
Cytheropteron obtusalum Hornibrook, 1952 Ar-Rec
Cytheropteron planalatum Guernet, 1985 Ak-Po
Cytheropteron terecaudum Hornibrook, 1952 Pl-Rec
Cytheropteron testudo Sars, 1869 Ak-Ar
Cytheropteron vertex Hornibrook, 1952 Wn-Rec
PHYLUM ARTHROPODA
Cytheropteron wellmani Hornibrook, 1952 Mp-Rec
Cytheropteron willetti Hornibrook, 1952 Wo-Rec
Cytheropteron sp. Ayress 1993 Ab-?Rec
Cytheropteron sp. Ayress 1995 Ak
Cytheropteron sp. 1 Ayress 1993 Lwh-Lw
Cytheropteron sp. 1 Ayress 1996 Ar-Lw
Cytheropteron sp. 2 Ayress 1993 Lwh-Ld
Cytheropteron sp. 2 Ayress, 1996 Ak
Cytheropteron sp. 3 Ayress 1993 Lwh-Ld
Eocytheropteron? sp. Ayress 1993 Ld-Lw
Cytherura clausi Brady, 1880 Pl-Rec
Cytherura nonspinosa Ayress, 1996 Ak
Eucytherura boomeri Ayress, Whatley, Downing, &
Millson, 1995 Wq
Eucytherura calabra (Colalongo & Pasini, 1980)
Ak-Rec
Eucytherura downingae Ayress, Whatley, Downing,
& Millson, 1995 Wc
Eucytherura elegantula Ayress, Whatley, Downing,
& Millson, 1995 Ab
Eucytherura pacifica Ayress, Whatley, Downing, &
Millson, 1995 Lw-Wc
Eucytherura tumida Ayress, Whatley, Downing, &
Millson, 1995 Wo-Wc (homonym of E. tumida
Bonnema, 1941)
Eucytherura bakeri Hornibrook, 1952 Po-Pl
Eucytherura batalaria Ayress, Whatley, Downing, &
Millson, 1995 Lwh-Wc
Eucytherura multituberculata Ayress, Whatley,
Downing, & Millson, 1995 Wo-Rec
Eucytherura sp. Ayress 1993 Ld
Eucytherura sp. 1 Ayress 1993 Ld-Lw
Eucytherura sp. 1 Ayress 1995 Ak
Eucytherura sp. 2 Ayress 1993 Ld
Eucytherura sp. 2 Ayress 1995 Ak
Eucytherura sp. 2 Ayress, Whatley, Downing, &
Millson 1995 Wo
Eucytherura? polydictyota Ayress, Whatley,
Downing, & Millson, 1995 Wc
Hemicytherura (Hemicytherura) aucklandica
Hornibrook, 1952 Lw-Rec
Hemicytherura (H.) delicatula Hornibrook, 1952
Lwh-Rec
Hemicytherura (H.) fereplana Hornibrook, 1952
Ak-Rec
Hemicytherura (H.) gravis Hornibrook, 1952 Ak-Rec
Hemicytherura (H.) quadrazea Hornibrook, 1952
Lwh-Rec
Hemicytherura sp. Ayress 1993 Ld-Lw
Hemicytherura (Kangarina) radiata (Hornibrook,
1952) Ak-Rec
Hemiparacytheridae leopardina Ayress, Whatley,
Downing & Millson, 1995 Wo
Hemiparacytheridea mediopunctata Ayress, Whatley,
Downing & Millson, 1995 Wo-Wc
Hemiparacytheridae vanharteni Ayress, Whatley,
Downing & Millson, 1995 Wc
Malabaricythere oceanica Yassini & Jones, 1995 Lw
Microcytherura alata Ayress, 1993 Lw n. nud.
Microcytherura sp. Ayress 1993 Lwh-Lw
Microcytherura haywardi Milau, 1993 Po
Microcytherura sp. Ayress 1993 Lwh-Lw
Microcytherura sp. 1 Ayress 1996 Ak-Ar
Microcytherura sp. 2 Ayress 1996 Ak-Ar
Oculocytheropteron aff. abyssorum (Brady, 1880) Ak
Oculocytheropteron acutangulum (Hornibrook, 1952)
Lwh-Rec
Oculocytheropteron australopunctatarum McKenzie,
Reyment & Reyment 1991 Ak
Oculocytheropteron confusum (Hornibrook, 1952)
Lwh-Rec
Oculocytheropteron ferrieri Milau, 1993 Po
Oculocytheropteron grantmackei Milau, 1993 Lw-Po
Oculocytheropteron improbum (Hornibrook, 1952)
Ak-Rec
Oculocytheropteron microfornix Whatley &
Downing, 1983 Ak
Oculocytheropteron paratinctum Ayress, 1996 Ak
Oculocytheropteron waihoensis Ayress, 1996 Ak
Oculocytheropteron sp. Ayress 1993 Lwh-Lw
Paracytheridea sp. Ayres, 1993 Ld-Lw
Pedicythere ?australis Neale, 1975 Ak
Pelecocythere? sp. 5042 Dingle 2009 Mh
Semicytherura arteria Swanson, 1979 Ak-Rec
Semicytherura coeca Ciampo, 1980 Ak-Lw
Semicytherura cf. costellata (Brady, 1880) Ak-Rec
Semicytherura eocenica Ayress, 1996 Ak-Ar
Semicytherura hexagona (Hornibrook, 1952) WnRec
Semicytherura okinawaensis Nohara, 1987 Ak
Semicytherura sericava (Hornibrook, 1952) Pl-Rec
Semicytherura sp. Ayress 1993 Ld-Lw
Semicytherura sp. 1 Ayress 1996 Ak
Semicytherura sp. 2 Ayress 1996 Ak
HEMICYTHERIDAE
Ambostracon sp. Ayress 1993 Lw
Ambostracon fredbrooki Milau, 1993 Po
Ambostracon (Patagonacythere) elongata Milau, 1993
Po
Bradleya arata (Brady, 1880) Wn-Rec
Bradleya clifdenensis Hornibrook, 1952 Ld-Pl
Bradleya dictyon (Brady, 1880) Dm-Rec
Bradleya kaiata Hornibrook, 1953 Ab-Ar
Bradleya opima Swanson, 1979 Ak-Rec
Bradleyla pakaurangia Hornibrook, 1952 Pl
Bradleya proarata Hornibrook, 1952 Ar-Lw
Bradleya pygmaea Whatley, Downing, Kesler &
Harlow, 1984 Mio-Rec
Bradleya reticlava Hornibrook, 1952 Ld-Rec
Bradleya semiarata Hornibrook, 1952 Pl
Bradleya (Quasibradleya) cuneazea Hornibrook,
1952 Ar-Rec
Bradleya (Q.) dictyonites Benson, 1972 Ak-Lw
Bradleya sp. Ayress 1993 Ab-Lwh
Bradleya sp. Ayress, 1993 Ld-Lw
Caudites impostor Hornibrook, 1953 Dh-Ab
Caudites cf. scopulicolus Hartmann, 1981
Hemicythere hornibrooki Swanson, 1969 Pl
Hemicythere munida Swanson, 1979 Ak-Rec
Hermanites andrewsi Swanson, 1979 Ld-Rec
Hermanites ?briggsi Swanson, 1979 Ak
Hermanites rectidorsa Milau, 1993 Po
Hermanites spinosa Milau, 1993 Po
Jacobella sp. Ayress 1995 Ak
Jugosocythereis reticulospinosa Ayress, 1993 Lwh-Lw
n. nud.
Limburgina quadrazea (Hornibrook, 1952) Dm-Ld
Patagonocythere tricostata Hartmann 1962 Ak
Patagonacythere waihaoensis Ayress, 1995 Ak
Patagonacythere parvitenuis (Hornibrook, 1953)
Ak-Ar
Poseidonamicus spp. Ayress, Neil, Passlow &
Swanson, 1997 Wc-Rec
Quadracythere alatazea Hornibrook, 1952 Pl-Sw
Quadracythere biruga Hornibrook, 1952 Ld-Rec
Quadracythere chattonensis Hornibrook, 1953
Ld-Lw
Quadracythere claremontensis Swanson, 1969 Pl
Quadracythere clavala Hornibrook, 1952 Lw-Sc
Quadracythere clifdenensis Hornibrook, 1952 Ak-Sl
Quadracythere longazea Hornibrook, 1952 Lwh-Sw
Quadracythere mediaplana Hornibrook, 1952 Po-Pl
Quadracythere mediaruga Hornibrook, 1952 Ak-Rec
Quadracythere planazea Hornibrook, 1952 Ld-Sl
Quadracythere radizea Hornibrook, 1952 Dm-Pl
Urocythereis opima Swanson, 1969 Lwh-Pl
Waiparacythereis caudata Swanson, 1969 Pl
Waiparacythereis decora Swanson, 1969 Pl
Waiparacythereis joanae Swanson, 1969 Pl-Rec
Waiparacythereis sp. Ayress 1993 Lwh
CRUSTACEA
KRITHIDAE
Krithe antisawanensis Ishizaki, 1966 Sl-Rec
Krithe comma Ayress, Barrows, Passlow & Whatley,
1999 Sl-Rec
Krithe compressa (Seguenza, 1980) Sw-Rec
Krithe dolichodeira Bold, 1946 Sw-Rec
Krithe marialusae Abate, Barra, Aiello & Bonaduce,
1993 Tt-Rec
Krithe minima Coles, Whatley & Moguilevsky, 1994
Lw-Rec
Krithe morkhoveni morkhoveni Bold, 1960 Wo-Rec
Krithe nitida Whatley & Downing, 1993 Ak-?Rec
Krithe pseudocomma Ayress, Barrows, Passlow &
Whatley, 1999 Lw-Rec
Krithe reversa Bold, 1958 Tk-Rec
Krithe swansoni Milau, 1993 Po-Rec
Krithe triangularis Ayress, Barrows, Passlow &
Whatley, 1999 Wc
Krithe trinidadensis Bold, 1958 Ww-Rec
Krithe sp. Ayress 1993 Lwh-Lw
Krithe sp. Ayress 1995 Ak
Krithe sp. 1 Ayress, Barrows, Passlow & Whatley
1999 Wn
Krithe sp. 2 Ayress, Barrows, Passlow & Whatley
1999 Lw-Rec
Krithe sp. 5055 Dingle 2009 Mh
Krithe sp. 5056 Dingle 2009 Mh
Krithe sp. 5079 Dingle 2009 Mh
Parakrithe sp. Ayress 1993 Lwh-Lw
Parakrithella lethiersi Milau, 1993 Po
LEGUMINOCYTHERIDIDAE
Triginglymus? hobsonensis Milau, 1993 Po
LEPTOCYTHERIDAE
Bisulcocythere campbelli Ayress & Swanson, 1991
Sw
Bisulcocythere compressa Ayress & Swanson, 1991
Po-Sw
Bisulcocythere eocenica Ayress & Swanson, 1991 Ak
Bisulcocythere micropunctata Ayress & Swanson,
1991 Lwh-Pl
Bisulcocythere novaezealandiae Ayress & Swanson,
1991 Pl-Rec
Callistocythere hanai Swanson, 1969 Pl
Callistocythere kaiata (Hornibrook, 1953) Ar-Ar
Callistocythere mansari Milau, 1993 Po
Cluthia antiqua Ayress & Drapala, 1996 Ak-Ar
Cluthia australis Ayress & Drapala, 1996 Wn-Rec
Cluthia micra Ayress & Drapala, 1996 Pl
Cluthia novaezealandiae Ayress & Drapala, 1996 Wn
Cluthia sp. Ayress 1993 Ld-Lw
Leptocythere sp. Ayress 1993 Ld-Lw
Leptocythere sp. Ayress 2006 Lw-Po
Leptocythere sp. Milau 1993 Po
Vandiemencythere phleboides Ayress & Warne, 1993
Ak-Lw
LIMNOCYTHERIDAE
Gomphocythere duffi (Hornibrook, 1955) Wc-Rec F
Limnocythere mowbrayensis Chapman, 1914 Wc F
Paralimnocythere vulgaris McKenzie & Swanson,
1981 Qu-Rec F
LOXOCONCHIDAE
Kuiperiana juglandica Ayress, 1993 Pl
Kuiperiana cf. lindsayi McKenzie, Reyment &
Reyment, 1991) Ak
Loxoconcha abrupta Hornibrook, 1952 Ld-Sw
Loxoconcha propunctata Hornibrook, 1952 Pl
Loxoconcha punctata Thomson, 1879 Ak-Rec
Loxoconcha sp. Milau 1969 Po
Microloxoconcha sp. Ayress 1995 Ak
Microloxoconcha sp. Ayress 1995 Ak
Palmoconcha juglandis Ayress, 1993 Lwh-Lw
Sagmatocythere carboneli Milau, 1993 Ak-Po
MACROCYPRIDIDAE
Macrocypris sp. Ayress 1993 Lwh-Lw
Macropyxis? sp. Ayress 2006 Lwh-Po
229
NEW ZEALAND INVENTORY OF BIODIVERSITY
Macroscapha? sp. Ayress 1995 Ak
NEOCYTHERIDEIDIDAE
Copytus pseudoelongatus Ayress, 1995 Ak
Copytus sp. Ayress 1993 Ld-Lw
Neocytherideis mediata Swanson, 1969 Ld-Pl
Neocytherideis reticulata Ayress, 1995 Ak-Lw
Pontocythere hedleyi (Chapman, 1906) Ak-Rec
NOTODROMADIDAE
Newnhamia fenestrata King, 1855 Wc-Rec
PARACYPRIDIDAE
Aglaia? praecox Chapman, 1926 Ld.
Paracypris eocuneata (Hornibrook, 1953) Ab-Lwh
Paracypris sp. 5040 Dingle 2009 Mh
Paracypris? sp. 5080 Dingle 2009 Mh
Phylctenophora zealandica Brady 1880 Ld-Rec
PARADOXOSTOMATIDAE
Cytherois parallella Milau, 1993 Po
Paracytherois cf. gracilis (Chapman, 1915) Ak
Paracytherois sp. Ayress 1993 Ld
PECTOCYTHERIDAE
Ameghinocythere eagari Dingle, 2009 Mh
Ameghinocythere? sp. 5078 Dingle 2009 Mh
Keijia? hornibrooki Milau, 1993 Po
Keijia sp. Ayress 2006 Po
Munseyella brevis Swanson, 1979 Ld-Rec
Munseyella dunoona McKenzie, Reyment &
Reyment, 1993 Ak
Munseyella modesta, Swanson, 1979 Ak-Rec
Munseyella pseudobrevis Ayress, 1995 Ak
Munseyella rectangulata Swanson, 1969 Pl
Munseyella cf. splendida Whatley & Downing, 1983
Ld-Lw
Swansonites aequa (Swanson, 1979) Ld-Rec E
Swansonites intermedia Milau, 1993 Po E
PONTOCYPRIDIDAE
Argilloecia acuticadata Whatley & Downing, 1983
Ak
Argilloecia australomiocenica Whatley & Downing,
1983 Ak
Argillaocia krithiformae Whatley & Downing, 1983
Ak
Argilloecia pusilla (Brady, 1880) Lwh-Lw
Australoecia sp. Ayress 1995 Ak-Lwh
Maddocksella argilloeciaformis (Whatley &
Downing, 1883) Ak
Maddocksella tumefacta (Chapman, 1914) Lwh-Lw
Maddocksella sp. 5047 Dingle 2009 Mh
Pontocypria sp. Ayress 1993 Lw
Propontocypris cf. herdmani (Scott, 1905) Ab-Rec
PROGONOCYTHERIDAE
Majungaella waiparaensis Dingle, 2009 E Mh
Majungaella wilsoni Dingle, 2009 E Mh
Majungaella sp. 4978 Dingle 2009 Mh
Parahystricocythere ericea Dingle, 2009 E Mh
Parahystricocythere sp. 5070 Dingle 2009 Mp
ROCKALLIIDAE
Arcacythere chapmani Hornibrook, 1952 Mp-Sw
Arcacythere aff. chapmani Hornibrook, 1952 LwhLw
Arcacythere eocenica (Whatley et al, 1980) Ak
SCHIZOCYTHERIDAE
Apateloschizocythere? colleni Dingle, 2009 Cn
TRACHYLEBERIDIDAE
Abyssocythere sp. Ayress 1993 Ld-Lw
Abyssophilos leptodictyotus (Ayress, 1995) Ar E
Actinocythereis microagrenon Ayress, 1995 Ak-Lw
Actinocythereis thomsoni (Hornibrook, 1952) DwRec
Acanthocythereis? reticulospinosa Ayress, 1993 Ab
Actinocythereis sp. Ayress 1993 Ab
Alataleberis paranuda Milau, 1993 Po
Anebocythereis hostizea (Hornibrook, 1952) Dh-Ld
Cletocythereis cf. bradyi Holden, 1967 Pl
Cletocythereis rastromarginata (Brady, 1880) Ak-Rec
Clinocthereis australis Ayress & Swanson, 1991
230
Ak-Rec
Cythereis contigua Hornibrook, 1952 Dm-Pl
Cythereis inlayi Hornibrook, 1952 Pl-Rec
Cythereis planalta Hornibrook, 1952 Dh-Po
Cythereis cf. brevicostata Bate, 1972 Mh
Glencoeleberis? cf. armata Jellinek & Swanson, 2003
Lwh-Po
Glencoeleberis? cf. brevicosta (Hornibrook, 1952)
Lwh-Po
Glencoeleberis? cf. incerta (McKenzie, Reyment &
Reyment, 1991) Lwh-Po
Glencoeleberis? cf. occultata Jellinek & Swanson,
2003 Lwh-Po
Glencoeleberis thomsoni (Hornibrook, 1952) Pal-Rec
Limburgina postaurora Dingle, 2009 E Mh
Marwickcythereis marwicki (Hornibrook, 1952)
Ab-Ar E
Marwickcythereis ordotormenta Whatley & Millson,
1992 Dw E
Oertliella semivera (Hornibrook, 1952) Dm-Ld
Oertliella echinata (McKenzie, Reyment &
Reyment, 1993) Ak-Lw
Philoneptunus alagracilus Whatley, Millson &
Ayress, 1992 Mh-Ab
Philoneptunus crassimurus Whatley, Millson &
Ayress, 1992 Ld-Lw
Philoneptunus eagari Whatley, Millson & Ayress,
1992 Dh
Philoneptunus eocenicus Whatley, Millson & Ayress,
1992 Dw-Dh
Philoneptunus gravizea Hornibrook, 1952 Dm-Rec
Philoneptunus hornibrooki Whatley, Millson &
Ayress, 1992 Ak-Ar
Philoneptunus paragravazea Whatley, Millson &
Ayress, 1992 Lwh-Rec
Philoneptunus paeminosus Whatley, Millson &
Ayress, 1992 Dh-Rec
Philoneptunus planaltus (Hornibrook, 1952) LwhRec
Philoneptunus praeplanaltus Whatley, Millson &
Ayress, 1992 Lwh
Philoneptunus reticulatus Whatley, Millson & Ayress,
1992 Ab-Ar
Philoneptunus swansoni Whatley, Ayress & Millson,
1992 Ab-Lwh
Philoneptunus tricostatus Whatley, Millson & Ayress,
1992 Dm-Dh
Philoneptunus sp. 1 Whatley, Millson & Ayress 1992
Lw
Philoneptunus sp. 2 Whatley, Millson & Ayress 1992
Pli-Ple
Philoneptunus sp. 3 Whatley, Millson & Ayress 1992
Ple
Philoneptunus sp. 5 Whatley, Millson & Ayress 1992
Lwh
Philoneptunus sp. 6 Whatley, Millson & Ayress 1992
Ak
Ponticocythereis praemilitaris Milau, 1993 Po
Protobuntonia hayi (Hornibrook, 1953) Ab-Ar
Rayneria? punctata Dingle, 2009 E Mh
Rugocythereis reticulata Ayress, 1993 Ab-Rec
Rugocythereis semicontigua (Hornibrook, 1953)
Ab-Lwh
Scepticocythereis cf. ornata Bate, 1972 Mh
Scepticocythereis? sp. 5044 Dingle 2009 Mh
Taracythere conjunctispina Ayress, 1995 Ak-Po
Taracythere hampdenensis (Ayress, 1993) Ab-Ak
Taracythere proterva (Hornibrook, 1953) ?Dt-Lw
Taracythere sp. Ayress 1993 Ab
Trachleberis ayressi Milau, 1993 Po
Trachyleberis brevicostata Hornibrook, 1952 Ld-Sl
Trachleberis denticulata Milau, 1993 Po
Trachyleberis hornibrooki Dingle, 2009 E Mh
Trachyleberis jilletti Ayress, 1993 Lw
Trachyleberis lytteltonsis Harding & Sylvester-
Bradley, 1953 Tt-Rec
Trachyleberis paucispinosa McKenzie, Reyment &
Reyment, 1993 Ak
Trachyleberis probesiodes Hornibrook, 1952 Sc-Wp
Trachyleberis retizea Hornibrook, 1952 Po-Pl
Trachyleberis rugibrevis (Hornibrook, 1952) Ld-Rec
Trachyleberis tridens Hornibrook, 1952 Ar-Pl
Trachyleberis zeacristata Hornibrook, 1952 Lw-Rec
XESTOLEBERIDIDAE
Microxestoleberis sp. Ayress 1993 Ld-Lw
Uroleberis minutissima (Chapman, 1926) Ak-Lw
Xestoleberis basiplana McKenzie, Reyment &
Reyment, 1993 Ak
Xestoleberis chilensis austrocontinentalis Hartmann,
1978 Ak
Xestoleberis cf. curta (Brady, 1865) Lwh-Rec
Xestoleberis paratruncata Whatley & Downing, 1983
Ak
Xestoleberis waihekeensis Milau, 1993 Po
Xestoleberis sp. 1 Ayress 1993 Lwh-Lw
Xestoleberis sp. 2 Ayress 1993 Lwh-Lw
Xestoleberis sp. Ayress 1995 Ak
INCERTAE SEDIS
Crescentocythere phoebe Ayress, 1993 Pl
Saidia limbata Colalongo & Passini, 1980 Ak
Saida torresi (Brady, 1880)*An-Rec
Saida sp. Ayress 1993 Lwh-Lw
Suborder PLATYCOPINA
CYTHERELLIDAE
Cytherella ballancei Milau, 1993 Po
Cytherella bisson Milau, 1993 Po-Pl
Cytherella chapmani Milau, 1993 Po
Cytherella elongata Swanson, 1969 Pl
Cytherella hemipunctata Swanson, 1969 Lw-Rec
Cytherella ?hemipunctata Swanson, 1969 Ak
Cytherella magna Ayress, 2006 Lw-Sc
Cytherella paranitida Whatley & Downing, 1983
Ab-Rec
Cytherella sp. Ayress, 1993 Ab-Lw
Cytherella sp. 5051 Dingle 2009 Mh
Cytherella sp. 5063 Dingle 2009 Cn
Cytherella sp. 5086 Dingle 2009 Mh
Cytherella sp. 1a Dingle 2009 Mh
Cytherelloidea paranitida Whatley & Downing, 1993
Lw
Cytherelloidea praeauricula (Chapman, 1926) AkLw
Cytherelloidea willetti Swanson, 1969* Ak-Rec E
Cytherelloidea cf. westaustraliensis Bate, 1972 Mh
Cytherelloidea n. sp. van den Bold, 1963 Rec
Cytherelloidea sp. Ayress, 1993 Lwh-Lw
Cytherelloidea sp. 1 Ayress 2006 Ld-Lw
Healdia? sp. Milau, 1993 Po
Platella sp. 5048 Dingle 2009 Mh
Platella sp. 5071 Dingle 2009 Mh
Order MYODOCOPIDA
Suborder MYODOCOPINA
SARSIELLIDAE
Sarsiella sp. Milau, 1993 Po
Class MALACOSTRACA
Subclass PHYLLOCARIDA
Order HYMENOSTRACA
HYMENOCARIDIDAE
Hymenocaris bensoni Chapman, 1934 Ord
Hymenocaris lepadoides Chapman, 1934 Ord
Order ARCHAEOSTRACA
CERATIOCARIDIDAE
Caryocaris cf. acuta Bulman, 1931 Ord
Caryocaris bulmani (Chapman, 1934) Ord
Caryocaris maccoyi (Etheridge, 1892) Ord
Caryocaris m. tumida (Chapman, 1934) Ord
Caryocaris marrii Chapman, 1934 Ord
PHYLUM ARTHROPODA
Caryocaris minima Chapman, 1934 Ord
Caryocaris wrightii Chapman, 1934 Ord
Subclass EUMALACOSTRACA
Superorder PERACARIDA
Order ISOPODA
Suborder VALVIFERA
HOLOGNATHIDAE
Debodea mellita Hiller, 1999 (not Cirolanidae)
UCret E
Suborder CYMOTHOOIDA
CIROLANIDAE
Cirolana makikihi Feldmann, Schweitzer, Maxwell
& Kelley, 2008 Wo E
Palaega kakatahi Feldmann & Rust, 2006 Wo-Wp E
INCERTAE SEDIS
URDIDAE
Urda zelandica Buckeridge & Johns, 1996 UJur E
Superorder EUCARIDA
Order DECAPODA
Suborder PLEOCYEMATA
Infraorder GLYPHEIDEA
ERYMIDAE
Gen. et sp. indet. Mp-Mh
GLYPHEIDAE
Glyphea christeyi Feldmann & Maxwell, 1999 Ab E
Glyphea stilwelli Feldmann, 1993 Dt E
Glypheopsis antipodum Glaessner 1960 Hu E
MECOCHIRIDAE
Mecochirus marwicki Glaessner, 1960 Kh
Mecochirus? sp. Bw, Kh-Op
Infraorder ASTACIDEA
NEPHROPIDAE
Hoploparia sp. Mp
Metanephrops motunauensis Jenkins, 1972 Sw-Tt E
PARASTACIDAE
Paranephrops fordycei Feldmann & Pole, 1994 Po-Sl
E
Infraorder AXIIDEA
CALLIANASSIDAE
Callianassa awakina Glaessner, 1960 Po E
Callianassa waikurana Glaessner, 1960 Mh E
Callianassa sp. a Mh
Callianassa sp. b Tt
Callianassa sp. Cn, Mp-Mh
Callianassa sp. Ab, Lwh-Pl, Sw-Tt
Protocallianassa sp. Mp-Mh
CTENOCHELIDAE
Ctenocheles cf. maorianus Powell, 1949 Wc
Ctenocheles sp. Wc
INCERTAE SEDIS
Gen. et sp. indet. Feldmann, Schweitzer, Maxwell
& Kelley, 2008 Wo E
Infraorder GEBIIDEA
UPOGEBIIDAE
Upogebia kowai Feldmann, Schweitzer, Maxwell &
Kelley, 2008 Wo E
Upogebia sp. Ar-Lwh
Infraorder ACHELATA
PALINURIDAE
Jasus flemingi Glaessner, 1960 Pl
Linuparus korura Feldmann & Bearlin, 1988 Ab
Linuparus sp. Mp-Mh
Linuparus? sp. Mp-Mh
Infraorder ANOMURA
AEGLIDAE
Haumuriaegla glaessneri Feldmann, 1984 Mp-Mh E
GALATHEIDAE
Galathea sp. Wp-Wn
LITHODIDAE
Paralomis debodeorum Feldmann, 1998 MMio-LMio
E
PAGURIDAE
Diacanthurus clifdenensis (Hyden & Forest, 1980)
Pl E
Pagurus sp. Tt, Wp, Wn
Infraorder BRACHYURA
ATELECYCLIDAE
Trichopeltarion greggi Dell, 1969 Sw-Tt E
Trichopeltarion merrinae Schweitzer & Salva, 2000
L Mio E
CALAPPIDAE
Calappilia maxwelli Feldmann, 1993 Po E
CANCRIDAE
Lobocarcinus pustulosus Feldmann & Fordyce, 1996
Pl E
Metacarcinus novaezelandiae (Hombron &
Jacquinot, 1846) Wo-Rec
Metacarcinus cf. novaezelandiae (Hombron &
Jacquinot, 1846) Tk, Wp
Metacarcinus sp. Ak, Ld, Wp-Wn
GONEPLACIDAE
Carcinoplax temikoensis Feldmann & Maxwell, 1990
Ak-Ar E
Carcinoplax sp. Wp-Wn
Kowaicarcinus maxwellae Feldmann, Schweitzer,
Maxwell & Kelley, 2008 Wo E
Ommatocarcinus arenicola Glaessner, 1960 Pl E
Ommatocarcinus cf. arenicola Glaessner, 1960 Pl
Ommatocarcinus cf. Neommatocarcinus huttoni
(Filhol, 1886) Wp-Wn
Ommatocarcinus sp. Pl
HOMOLODROMIIDAE
Homolodromia novaezelandica Feldmann, 1993
Mp-Mh E
Homolodromia sp. Mp-Mh
MACROPHTHALMIDAE
Macrophthalmus (Hemiplax) hirtipes (Heller, 1862)
Wq–Rec E
Hemiplax?major Glaessner, 1960 Wn E
Hemiplax cf. major Glaessner, 1960 Po, Wc
Hemiplax sp. Wn-Wc
MAJIDAE
Actinotocarcinus chidgeyorum Jenkins, 1974 Sc-Tt E
Actinotocarcinus maclauchlani Feldmann, 1993
Sw-Tt E
Jacquinotia edwardsii (Jacquinot, 1853) Wp-Rec E
Leptomithrax atavus Glaessner, 1960 Tk E
Leptomithrax elongatus McLay, Feldmann &
MacKinnon, 1995 Sw E
CRUSTACEA
Leptomithrax garthi McLay, Feldmann &
MacKinnon, 1995 Sw-Tt E
Leptomithrax griffini Feldmann & Maxwell, 1990
Ab-Ar E
Leptomithrax irirangi Glaessner, 1960 Wo E
Leptomithrax aff. irirangi Glaessner, 1960 Sw
Leptomithrax uruti Glaessner, 1960 E Tt
Leptomithrax cf. uruti Glaessner, 1960 Tt
Leptomithrax sp. Tt
Micromithrax? minisculus Feldmann & Wilson, 1988
Dm-Dh
Notomithrax allani Feldmann & Maxwell, 1990
Ak-Ar E
Notomithrax minor (Filhol, 1885) Wc – Rec
Notomithrax sp. Wc
MENNIPIDAE
Galene proavita Glaessner, 1960 Pl-Sc E
Galene sp. Wp-Wn
Menippe sp. Pl
Pseudocarcinus sp. Tk
PORTUNIDAE
Ovalipes cf. catharus (White, 1843) Wn-Wc
Ovalipes sp. A Wp
Ovalipes sp. Wn-Wc
Pororaria eocenica Glaessner, 1980 Ak-Ar E
Portunus sp. Lwh, Lw
Rhachiosoma granuliferum (Glaessner, 1960) Dp-Ar
E
Gen. et sp. indet. Dm-Dh, Ab-Ak
PSEUDOZIIDAE
Tongapapaka motunauensis Feldmann, Schweitzer,
Maxwell & Kelley, 2008 Wo E
RANINIDAE
Hemioon novozelandicum Glaessner, 1980 Cn E
Laeviranina keyesi Feldmann & Maxwell, 1990
Ak-Ar E
Laeviranina perarmata Glaessner, 1960 Ab E
Laeviranina pororariensis (Glaessner, 1980) Ak-Ar E
Lyreidus bennetti Feldmann & Maxwell, 1990 Ak-Ar
E
Lyreidus elegans Glaessner, 1960 Po-Pl E
Lyreidus waitakiensis Glaessner, 1980 Ab E
Lyreidus sp. Sw
Gen. et sp. indet. Ab
TORYNOMMIDAE
Eodorripe spedeni Glaessner, 1980 Mp-Mh E
Torynomma flemingi Glaessner, 1980 Mp-Mh E
Torynomma planata Feldmann, 1993 Mp-Mh E
TUMIDOCARCINIDAE
Tumidocarcinus dentatus Glaessner, 1960 Lwh-Ld E
Tumidocarcinus cf. dentatus (Glaessner, 1960) Lwh
Tumidocarcinus giganteus Glaessner, 1960 Pl-Tt E
Tumidocarcinus cf. giganteus Glaessner, 1960 Lw-Po,
Sw-Tk
Tumidocarcinus tumidus (Woodward, 1876) Ab-Ld E
Tumidocarcinus cf. tumidus (Woodward, 1876)
Lwh-Ld
Tumidocarcinus? sp. Ak-Ld, Po-Sc
VARUNIDAE
Austrohelice manneringi Feldmann, Schweitzer,
Maxwell & Kelley, 2008 Wo E
Miograpsus papaka Fleming, 1981 Tt E
231
NEW ZEALAND INVENTORY OF BIODIVERSITY
Developmental stages of New Zealand Decapoda
Compiled by W. R. Webber
Following are the larvae and/or pre- or post-larvae described to date, of species listed in the
decapod species list above. Species named below are those with one or more developmental
stages described in the literature. Names and dates in brackets indicate publications in which
larvae are described, not species authorities. However, Jaxea novaezealandiae (Gebiidea)
was described in the same paper as the adult and two polychelid species were described
from the larvae, thus authors in brackets after these names are also the original authorities.
Literature sources for the species list below are cited in the References section, above.
PHYLUM CRUSTACEA
Class MALACOSTRACA
Order DECAPODA
Suborder DENDROBRANCHIATA
SERGESTIDAE
Sergestes arcticus [Gurney & Lebour 1940; Wear 1985]
SOLENOCERIDAE
Solenocera comata [Gurney 1924; Wear 1985]
Suborder PLEOCYEMATA
Infraorder STENOPODIDEA
STENOPODIDAE
Stenopus hispidus [Gurney 1936, 1942]
Infraorder CARIDEA
ALPHEIDAE
Alpheus euphrosyne richardsoni [Packer 1983, 1985]
Alpheus socialis [Packer 1983, 1985]
Alpheopsis garricki [Packer 1983, 1985]
Betaeopsis aequimanus [Packer 1983, 1985]
ATYIDAE
Paratya curvirostris [Ch’ng 1973; Wear 1985]
CAMPYLONOTIDAE
Campylonotus rathbunae [Pike & Williamson 1966;
Wear 1985]
CRANGONIDAE
Aegaeon lacazei [De Simón 1979; Packer 1983, 1985]
Philocheras australis [Thomson & Anderton 1921;
Packer 1983, 1985]
Philocheras chiltoni [Packer 1983, 1985]
Philocheras hamiltoni [Packer 1983, 1985]
Philocheras pilosoides [Packer 1983, 1985]
HIPPOLYTIDAE
Alope spinifrons [Lebour 1955; Packer 1983, 1985]
Hippolyte bifidrostris [Packer 1983, 1985]
Hippolyte multicolorata [Packer 1983, 1985]
Nauticaris marionis [Packer 1983, 1985]
Tozeuma novaezealandiae [Packer 1983, 1985]
OGYRIDIDAE
Ogyrides delli [Packer 1983, 1985]
PALAEMONIDAE
Palaemon affinis [Lebour 1955; Packer 1983, 1985]
Periclimenes yaldwyni [Packer 1983, 1985]
Periclimenes (Periclimenes) sp. [Packer 1983, 1985]
Infraorder ASTACIDEA
NEPHROPIDAE
Metanephrops challengeri [Wear 1976]
PARASTACIDAE
Paranephrops planifrons [Hopkins 1967]
Infraorder AXIIDEA
CALLIANASSIDAE
Callianassa filholi [Gurney 1924; Lebour 1955;
Wear 1965a]
Infraorder GEBIIDEA
LAOMEDIIDAE
Jaxea novaezealandiae [Wear & Yaldwyn 1966]
UPOGEBIIDAE
Acutigebia danai [Gurney 1924]
232
Infraorder PALINURA
PALINURIDAE
Jasus edwardsii [Batham 1967; Lesser 1974]
Sagmariasus verreauxi [Lesser 1974; Kittaka et al.
1997]
POLYCHELIDAE
Gen. et sp. indet. (as Eryonicus fagei) [Bernard
1953]
Gen. et sp. indet. (as Eryonicus scharffi) [Selbie 1914]
SCYLLARIDAE
Ibacus alticrenatus [Atkinson & Boustead 1982]
Scyllarus sp. Z [Webber & Booth 2001]
Infraorder ANOMURA
CHIROSTYLIDAE
Gastropyychus novaezelandiae [Pike & Wear 1969]
Uroptychus n. sp. [Pike & Wear 1969]
GALATHEIDAE
Munida gregaria [Roberts 1973]
PAGURIDAE
Pagurixus hectori [Roberts 1971; Wear 1985]
Pagurus novizealandiae [Greenwood 1966; Wear 1985]
Pagurus traversi [Thomson & Anderton 1921; Wear
1985]
Porcellanopagurus edwardsi [Roberts 1972; Wear 1985]
PARAPAGURIDAE
Sympagurus dimorphus [Lemaitre & McLaughlin
1992]
PORCELLANIDAE
Petrocheles spinosus [Wear 1965b, 1966]
Petrolisthes elongatus [Greenwood 1956; Wear
1964b, 1965c]
Petrolisthes novaezelandiae [Greenwood 1956; Wear
1964a, 1965d]
Infraorder BRACHYURA
ATELECYCLIDAE
Trichopeltarion fantasticum [Wear & Fielder 1985]
BELLIIDAE
Heterozius rotundifrons [Wear & Fielder 1985]
CANCRIDAE
Metacarcinus novaezelandiae [Wear & Fielder 1985]
CYMONOMIDAE
Cymonomus bathamae [Wear & Fielder 1985
DROMIIDAE
Metadromia wilsoni [Wear & Fielder 1985]
GONEPLACIDAE
Neommatocarcinus huttoni Wear & Fielder 1985
GRAPSIDAE
Leptograpsus variegatus [Wear & Fielder 1985]
Planes major [Wear & Fielder 1985]
Planes marinus [Wear & Fielder 1985]
HOMOLIDAE
Dagnaudus petterdi [Williamson 1965; Wear &
Fielder 1985]
Homola orientalis [Wear & Fielder 1985]
HYMENOSOMATIDAE
Amarinus lacustris [Wear & Fielder 1985]
Elamena longirostris [Wear & Fielder 1985]
Elamena momona [Wear & Fielder 1985]
Elamena producta [Wear & Fielder 1985]
Halicarcinus cookii [Wear & Fielder 1985]
Halicarcinus innominatus [Wear & Fielder 1985]
Halicarcinus planatus [Wear & Fielder 1985]
Halicarcinus varius [Horn &Harms 1988]
Halicarcinus whitei [Wear & Fielder 1985]
Hymenosoma depressum [Wear & Fielder 1985]
Neohymenicus pubescens [Wear & Fielder 1985]
INACHIDAE
Achaeus curvirostris [Wear & Fielder 1985]
Cyrtomaia lamellata [Wear & Fielder 1985]
INACHOIDIDAE
Pyromaia tuberculata [Webber & Wear 1981; Wear
& Fielder 1985]
LATREILLIIDAE
Eplumula australiensis (Wear &Fielder 1985)
LEUCOSIIDAE
Bellidilia cheesmani [Wear & Fielder 1985]
MACROPHTHALMIDAE
Macrophthalmus (Hemiplax) hirtipes [Wear & Fielder
1985]
MAJIDAE
Eurynolambrus australis [Webber & Wear 1981;
Wear & Fielder 1985]
Jacquinotia edwardsi [Webber & Wear 1981; Wear &
Fielder 1985]
Leptomithrax longimanus [Webber & Wear 1981;
Wear & Fielder 1985]
Leptomithrax longipes [Webber & Wear 1981; Wear
& Fielder 1985]
Leptomithrax tuberculatus mortenseni [Wear &
Fielder 1985]
Notomithrax minor [Webber & Wear 1981; Wear &
Fielder 1985]
Notomithrax peronii [Webber & Wear 1981; Wear &
Fielder 1985]
Notomithrax ursus [Webber & Wear 1981; Wear &
Fielder 1985]
OZIIDAE
Ozius truncatus (Wear & Fielder 1985)
PILUMNIDAE
Pilumnopeus serratifrons [Wear & Fielder 1985]
Pilumnus lumpinus [Wear & Fielder 1985]
Pilumnus novaezelandiae [Wear & Fielder 1985]
PINNOTHERIDAE
Nepinnotheres novaezelandiae [Wear & Fielder 1985]
PLAGUSIIDAE
Plagusia chabrus [Wear & Fielder 1985]
PORTUNIDAE
Liocarcinus corrugatus [Wear & Fielder 1985]
Nectocarcinus antarcticus [Wear & Fielder 1985]
Ovalipes catharus [Wear & Fielder 1985]
Portunus pelagicus [Wear & Fielder 1985]
Scylla serrata [Wear & Fielder 1985]
RANINIDAE
Lyreidus tridentatus [Wear & Fielder 1985]
VARUNIDAE
Austrohelice crassa [Wear & Fielder 1985]
Cyclograpsus insularum [Wear & Fielder 1985]
Cyclograpsus lavauxi [Wear & Fielder 1985]
Hemigrapsus crenulatus [Wear & Fielder 1985
Hemigrapsus sexdentatus [Wear & Fielder 1985]