ChapterPDF Available

Phylum Gnathifera lesser jaw worms, rotifers, thorny-headed worms

May 2011

In book: New Zealand Inventory of Biodiversity, Volume 1. Kingdom Animalia  Radiata, Lophotrochozoa, Deuterostomia. (pp.137-158)
Chapter: Phylum Gnathifera lesser jaw worms, rotifers, thorny-headed worms
Publisher: Canterbury University Press
Editors: Dennis Gordon

Authors:

Russell Shiel

None independent consultant.

Lesley R Smales

South Australian Museum

Wolfgang Sterrer

Bermuda Natural History Museum

Ian C. Duggan

University of Waikato

Show all 6 authorsHide

The Gnathifera includes the free-living lesser jaw worms (Gnathostomulida) and parasitic thorny-headed worms (Acanthocephala) as well as the unwormlike rotifers. These groups used to be treated as separate phyla, and still are in many textbooks. In lacking a body cavity and through-gut with a permanent anus, gnathostomulids were included among flatworms prior to 1969. On the other hand, rotifers and thorny-headed worms have a body cavity, and most rotifers also have an open-ended digestive tract. Recent molecular data and new studies of morphology and anatomy have shown that the Gnathostomulida, Rotifera, and Acanthocephala are related in surprising and unsuspected ways.

Content uploaded by Russell Shiel

Content may be subject to copyright.

135

God’s ‘inordinate fondness for beetles’ is proverbial, but a fondness for

vermiform animals might equally be claimed. They may not be as

speciose as beetles (that remains to be seen), but there is certainly no

shortage of worms in this world; they occur in the ocean as much as in damp

earth and ponds, and inside plants and animals. At least a dozen major taxa

are worm-shaped. These ‘primary vermiforms’ include ‘mesozoans’ (Dicyemida

and Orthonectida), Platyhelminthes, Nemertea, Gnathifera, Gastrotricha,

Nematoda, Nematomorpha, Chaetognatha, Priapulida, Kinorhyncha, Sipuncula,

Annelida, Echiura, and Enteropneusta. Nearly all phyla contain at least some

members with vermiform bodies, though we may not think of them in this way,

but consider eels among ﬁsh, caecilians among amphibians, snakes among

reptiles. Maggots and tongue worms are vermiform arthropods, aplacophorans

are vermiform molluscs, and sea cucumbers vermiform echinoderms. It is a

lifestyle that confers particular advantages in certain environments. At the

microscopic level, the interstitial fauna that lives between grains of sand

and other particles in ponds and seaﬂoors is particularly rich in vermiforms

(Swedmark 1964); with few exceptions, such as the globular ostracods and

some ﬂattened tardigrades, practically all interstitial animals are considerably

longer than wide, including such unlikely candidates as crustaceans, cnidarians,

and ascidians. An extended cylindrical body is ideal for navigating the narrow,

three-dimensional canal system that extends between grains and particles (Riser

1984). A vermiform body also seems to be suited to the particular environment

of animal intestines, in the case of internal parasites.

The Gnathifera includes the free-living lesser jaw worms (Gnathostomulida)

and parasitic thorny-headed worms (Acanthocephala) as well as the unworm-

like rotifers. These groups used to be treated as separate phyla, and still are in

many textbooks. In lacking a body cavity and through-gut with a permanent anus,

gnathostomulids were included among ﬂatworms prior to 1969. On the other

hand, rotifers and thorny-headed worms have a body cavity, and most rotifers

also have an open-ended digestive tract. Recent molecular data and new studies

of morphology and anatomy have shown that the Gnathostomulida, Rotifera,

and Acanthocephala are related in surprising and unsuspected ways.

Ahlrichs (1995, 1997) has summarised the many common features among

these groups. A syncytial epidermis links rotifers, Seison (a marine rotifer-like

creature found on nebaliid crustaceans), and Acanthocephala, moving Ahlrichs

(1995) to propose the name Syndermata for this grouping. As revealed by

transmission (Rieger & Tyler 1995) and scanning electron microscopy (Sørensen

2003), the jaw apparatus of gnathostomulids and rotifers is remarkably similar.

That of Seison is less obviously homologous (Segers & Melone 1998), and the

EIGHT

Phylum

GNATHIFERA

lesser jaw worms, rotifers, thorny-headed worms

RUSSELL J. SHIEL, LESLEY SMALES, WOLFGANG STERRER,

IAN C. DUGGAN, SYLVIE PICHELIN, JOHN D. GREEN

ANY PHOTOS AVAILABLE?

136

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

Seisonidea may have diverged from rotifers at an early stage in their evolution.

On the other hand, Seison has similar sperm to acanthocephalans and the epi-

dermis of both groups contains bundles of ﬁlaments. On balance, morphological

and molecular evidence supports the notion that acanthocephalans may be

highly modiﬁed endoparasitic rotifers that have lost the digestive system and

jaw apparatus.

Following the suggestion of Kristensen and Funch (2000), inclusion in the

phylum Gnathifera (Ahlrichs 1997) would reduce the Gnathostomulida to one

of three classes, the others being Syndermata (comprising rotifers, Seison, and

Acanthocephala) and Micrognathozoa, the latter comprising a single genus

and species (Limnognathia maerski) from cold fresh waters in West Green-

land and the Crozet Islands (De Smet 2002). As Nielsen (2003) pointed out,

however, the name Syndermata is unnecessary if Rotifera is taken as including

Acanthocephala. Rotifera and Acanthocephala are used as class names in this

chapter.

Class Gnathostomulida – lesser jaw worms

Gnathostomulids are wholly marine. They are microscopic, free-living,

unsegmented worms whose zoological relationships are quite enigmatic.

First described as an aberrant group of turbellarian ﬂatworms (Ax 1956), the

Gnathostomulida was recognised as a distinct phylum by Riedl (1969), and

by Sterrer (1972) who proposed the currently used classiﬁcation. The phylum

comprises two orders, Filospermoidea and Bursovaginoidea, the latter divided

into two suborders, Scleroperalia and Conophoralia (Sørensen et al. 2006). The

tiniest species measure only a third of a millimetre long; giants attain three

millimetres.

Gnathostomulids differ from all known invertebrates in having an entirely

monociliated epidermis, i.e. each epidermal cell carries only a single cilium

(some gastrotrichs have part of their body monociliated). They are further

distinguished by having a bilaterally symmetrical pharynx that usually contains

complex cuticular mouthparts consisting of paired jaws and an unpaired, ventral

basal plate. The gut is straight and lacks an anus. There are paired sensory organs,

mainly in the form of bundled cilia at the anterior end, and excretory organs

in the form of paired groups of protonephridia. Respiratory and circulatory

organs are lacking.

All gnathostomulids are hermaphrodites. An unpaired, pear-shaped ovary

lies dorsally between the gut and the epidermis, extending from behind the

pharynx to about midbody region. The germinal zone is located anteriorly,

and size, maturity, and yolk content of oocytes increase posteriorly during egg

formation. A single mature egg usually takes up the hindmost half or two-

thirds of the ovary. Testes are located in the hind body; they are paired in

Bursovaginoidea and most Filospermoidea, and unpaired and dorsal to the gut

in Conophoralia. Filospermoidea, and probably Conophoralia, have a simple,

Summary of New Zealand gnathiferan diversity

Taxon Described Known Estimated Endemic Endemic

species/ undescribed/ unknown species/ genera

subspecies/ undetermined species/ etc.

varieties species/ etc. etc.

Gnathostomulida 2 10 12 4 0

Micrognathozoa 0 0 1 0 0

Rotifera 462 18 20 12 0

Acanthocephala 22 16 50 3 0

Totals 486 44 83 19 0

Taxon Marine Fresh- Terrestrial

water

Gnathostomulida 12 0 0

Rotifera 2 478 0

Acanthocephala* 30 6 2

Totals 44 484 2

* Tallies are based on the primary environment

of the host species.

Diversity by environment

137

PHYLUM GNATHIFERA

rosette- or funnel-shaped penis that is weakly muscular but richly glandular.

Located posterior to the testes, it empties into a subterminal ventral pore.

Scleroperalia are characterised by a bulbous, muscular penis that usually

surrounds a tubular penis stylet made up of eight to 10 rod-shaped cell exten-

sions. There are three types of sperm,the homology of which is uncertain:

ﬁliform sperm (with a spiral nucleus, a middle piece, and one 9+2 axoneme) in

Filospermoidea; aﬂagellate ‘dwarf’ sperm in Scleroperalia; and ‘conulus’ sperm

in Conophoralia. Filiform sperm rotate like a corkscrew, whereas dwarf sperm

and conuli are immotile.

Sperm transfer is by copulation. Filospermoidea lack a vagina and a bursa for

sperm storage; it seems that sperm are injected under the epidermis, and then

distributed throughout the body ,where they are stored prior to use in fertilis-

ation. Most Conophoralia have a permanent vagina situated dorsally behind the

ovary; this leads into a pouch-shaped bursa in which usually only one or two

sperm are stored. Scleroperalia lack a permanent vagina but are characterised by

a bursa system consisting of a caudal, rounded prebursa that connects anteriorly

to a conical bursa. The wall of the bursa is composed of ﬂattened cells that meet

laterally to form crests, and anteriorly to form a perforated mouthpiece through

which stored sperm is channeled to the mature egg.

Oviposition, at least in Scleroperalia, is by rupture of the dorsal epidermis

behind the ovary and bursa, at the spot where a vagina may be located. The

egg then becomes spherical and sticks to sand grains. Development is direct.

Cleavage, still insufﬁciently known, seems to be of the spiralian type, with

mesolecithal-epibolic gastrulation, resulting in a juvenile that lacks jaws but

has a rudimentary pharynx.

The major groups of Gnathostomulida.

CREDIT

Filospermidea

(Haplognathia)

Conophoralia

(Austrognathia)

Scleroperalia

(Gnathostomula)

basal plate

jaws

bursa

testes

conuli

penis

male

pore

ovary

testes

monociliated

epidermal cells

138

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

Gnathostomulid evolution most likely progressed from the ﬁlospermoid

condition (sperm with one 9+2 ﬂagellum, no bursa, and a simple male pore)

to the bursovaginoid condition (sperm aﬂagellate, bursa system present, and

a complex copulatory organ). In the absence of a fossil record (an afﬁnity with

conodonts was suggested by Ochietti and Cailleux (1969) but discounted by

Sterrer et al. 1985), phylogenetic relationships are still quite open. Sterrer et

al. (1985) pointed out possible relationships with Nematoda, Gastrotricha,

and Rotifera. Rieger and Tyler (1995) documented apparent homologies in jaw

ultrastructure between gnathostomulids and rotifers, and Kristensen and Funch

(2000) suggested that the Gnathostomulida be united with the Syndermata

(Rotifera plus Acanthocephala) and the new class Micrognathozoa in a phylum

Gnathifera. Molecular evidence based on 18S rRNA gene sequences supports

the taxon Gnathifera (Giribet et al. 2000, 2004).

Diversity and distribution

Although lesser jaw worms are frequently the dominant invertebrates of the

detritus-rich, oxygen-poor sands in which they generally live, knowledge of

their biology is still very scanty. It is assumed that they feed by grazing on the

microﬂora (bacteria, fungal hyphae) that coats sand grains, and that they have

extremely low oxygen requirements, in addition to mechanisms for sulﬁde

detoxiﬁcation. Found exclusively in the interstices of shallow marine sand from

the intertidal to 400 metres depth, the phylum currently comprises fewer than

100 species, of which many have a worldwide distribution (Sterrer 1998). A dozen

species are now known from New Zealand (two from shallow waters off Leigh,

North Island (Sterrer 1991), and 10 undetermined species from intertidal sand

ﬂats on the Otago Peninsula, South Island), but these probably represent less

than half of the total number expected to occur. Three of the 12 species may be

endemic. Because of the lack of resident New Zealand specialists in microscopic

‘lower’ worm taxa, it may take many more years before a more complete survey

of the regional fauna is achieved.

Class Rotifera – rotifers

Rotifers (literally, wheel bearers) are named for the beating waves of cilia,

resembling wheels in motion, on their heads. The cilia draw food particles to the

mouth and into another feature of rotifers, the jaw-like mastax that grabs and

grinds the food. Tiny creatures, rotifers are often the most abundant metazoans

of inland waters, both numerically and in terms of species numbers (Wallace

2002). Around 2,000 species are known worldwide (Segers 2002), with 480

taxa recorded to date from New Zealand and outlying islands. Australia, by

comparison, has about 710 taxa recorded, albeit with a much greater sampling

intensity over a considerably larger area. On present evidence, approximately

70% of the known New Zealand species are shared with Australia. In view of

the relatively low sampling effort to date, it is probable that these numbers

represent less than half of the rotifers likely to be found in both regions. Most

rotifers are found in fresh waters, but a small group of halophiles (‘salt lovers’)

may be found in inland salt lakes, often at very high densities, with a few

species tolerating concentrations above that of sea water. Marine rotifers are

known, but have not been well studied in the Australasian region. Cassie (1960)

reported three species of Synchaeta (Synchaetidae) in a blood-red water bloom

southwest of Kapiti Island, Cook Strait, in 1959, the colour being imparted by

the nerve ganglion and eyespot of the rotifers. Of the three, S. baltica, which is

regarded as cosmopolitan in the marine littoral and in tidal pools (Hollowday

2002), was commonest.

There are creeping, planktonic, semiplanktonic, semisessile, and sessile

forms. Several species are parasitic. Most rotifers are littoral or benthic in

Scanning electron micrograph of a rotifer

showing the paired wheel-like bands of cilia

by which the animal creates feeding currents.

CREDIT

MISSING IMAGE

139

PHYLUM GNATHIFERA

habit, and are found in the ﬂoating or submerged vegetation of lake and river

margins, ponds, weedy puddles, in damp moss, in fact any place that holds

water for more than a few days including house spoutings and bird baths. These

rotifers belong to the ‘littoral microfauna’ and are not commonly found in open

water. Nevertheless, the littoral rotifer fauna in natural habitats may provide

a signiﬁcant dietary input for macroinvertebrates, tadpoles, or juvenile ﬁsh.

Relatively few taxa are specialised for an open-water existence. The planktonic

forms tend to have a reduction of the foot, and ﬂoat with the aid of an inﬂated,

balloon-like morphology (e.g. Trochosphaera). Other species have appendages

such as bristles, ﬁns, or rudders, or produce internal oil droplets. Semiplanktonic

species living in the plant-poor or plant-free zone of lakes may retain the foot

and use an adhesive secreted from the toe-tips to attach occasionally to ﬂocculent

detritus, algal ﬁlaments, crustaceans, or other rotifers.

Most common species are small (less than a ﬁfth of a millimetre in length),

although occasionally the predatory genus Asplanchna, or colonial taxa (e.g.

Lacinularia, southeastern Australia only), may exceed 1 millimetre and be visible

to the naked eye. Rotifers have the fastest reproductive rates of any metazoan

(Nogrady et al. 1993), and can rapidly ﬁll available niches. High population

densities can thus be achieved when food is not limiting. Hence 24,000 rotifers

per litre in the Mt Bold Reservoir, South Australia (Shiel et al. 1987) is not

particularly high compared with some of the population densities (500,000 per

litre) reported from sewage ponds and aquaculture systems (Lubzens 1987).

Although such high densities are to be expected in highly eutrophic habitats,

rotifer densities in zooplankton are more commonly fewer than 1,000 per

litre, with proportionately higher densities in shallower shore areas of lakes as

habitat-increased partitioning by rooted plants increases available niches.

Diversity of New Zealand rotifers

At least 480 rotifer taxa (144 Bdelloidea/336 Monogononta) have been recorded

from New Zealand since 1859. Of these, 449 are morphospecies and 31 are

subspeciﬁc or infrasubspeciﬁc variants. Most of the named taxa are widely

distributed or cosmopolitan, with only a small proportion (less than 5%) appar-

ently restricted to New Zealand or the Australasian region.

Bdelloid rotifers (from bdella, Greek, leech; so-called because of the leech-

like gait of many species) are poorly known in the Australasian region. Largely

as a result of the efforts of J. Murray early this century (e.g. Murray 1911), the

bdelloids of New Zealand became better known than those of Australia. The

great number of bdelloid taxa from a relatively smaller area also may reﬂect

climatic and environmental differences between the two regions (Shiel &

Green 1996). In any event, the bdelloids of the wider region are badly in need

of detailed study and taxonomic revision. It is likely that more, possibly many

more, bdelloids remain to be discovered from both areas. Some 350 species

occur worldwide (Dr C. Ricci pers. comm.).

The three orders of Monogononta (having a single ovary/germinal structure)

– Collothecacea, Flosculariacea, and Ploimida – have received considerably

more attention than have bdelloids, and as a result are somewhat better known

taxonomically and ecologically. At present, 336 monogonont taxa are known

from New Zealand (cf. 607 from Australia). As for the bdelloids, in view of

the relatively poor coverage of both regions by collectors, it is likely that many

monogononts are yet to be recorded.

Historical overview of studies

Rotifers have been studied extensively in the northern hemisphere for 300 years,

but it was not until the global collecting of Schmarda (1859) that the ﬁrst record

of New Zealand rotifers (three taxa) appeared. There were only three additional

reports by the turn of the century: Hamilton (1879, one species), Stock (1892, one

species), and Hilgendorf (1898, 16 species). A further 26 species were listed by

Synchaeta pectinata.

From Edmondson 1959

Ascomorpha ecaulis.

From Edmondson 1959

Asplanchna priodonta.

From Edmondson 1959

140

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

Hilgendorf (1902), who also compiled an updated checklist of 42 taxa for Hutton

(1904). The most signiﬁcant early contribution resulted from the Shackleton

Antarctic Expedition of 1907–1909 when Murray (1911) identiﬁed ﬁrst 67 taxa

then a further 10 (Murray 1913). A list of 22 taxa from Oamaru was given by

Morris (1912), who then collated some of the earlier records into a species list

(Morris 1913). An incidental mention appeared in Brehm (1928).

There was then an apparent hiatus until Russell, in 1944, began what was

to become a comprehensive series of species lists and methodologies (Russell

1944–1962), including several descriptions of endemic taxa. This series brought

the study of New Zealand’s rotifers to the taxonomic standards of the time,

although re-examination of some of Russell’s taxonomy (Segers 1995) indicates

that a modern reappraisal is necessary. In particular, the emerging acceptance

of non-cosmopolitanism suggests that the degree of endemicity is likely to be

higher than Russell recognised (Koste & Shiel 1987; Shiel 1995).

Further new records were included by Parr (1949), who studied rotifer com-

munities in some Otago ponds for a Master’s thesis. This represented pioneer

work on rotifer ecology in New Zealand and the only signiﬁcant research on

rotifers, other than Russell’s, between 1930 and 1960. The study was in many

ways well ahead of its time in attempting both to describe the structure and

seasonality of the rotifer communities, and to understand the causes of the

patterns observed. Unfortunately, it has remained unpublished.

Bdelloid rotifers were studied by Haigh, whose ﬁve publications (1963–1970)

added 49 new records and ﬁve new species. New Zealand bdelloids have since

been completely neglected. A brief introductory guide to some rotifers of the

Auckland region was included in Green (1969). Then followed almost 25 years

with little systematic reference to rotifers, although ecological studies mentioning

them include Jolly (1952, 1977), Stout (1969, 1978, 1981, 1984, 1991), Chapman

(1973), Green (1974, 1976), Burns and Mitchell (1980), Forsyth and McCallum

(1980), and Forsyth and James (1991). The value of most of these studies was

hampered by inadequate sampling methods (e.g. too coarse a mesh size in

nets) and poor taxonomic resolution. The most detailed study of the 1980s, an

investigation of the rotifers of the Mangere oxidation ponds in Auckland (Tolich

1988), is unpublished.

Between 1988 and 1991, L. Sanoamuang, using modern methods, made

a detailed study of rotifer ecology in Lake Grassmere and surveyed rotifer

distribution in 35 South Island lakes. Thirty new records were added to the

New Zealand rotifer fauna (Sanoamuang & Stout 1993). An elementary guide

to planktonic rotifers of New Zealand lakes was produced by Parr (1992). Shiel

and Sanoamuang (1993) described a new species of Filinia from Lake Okaro,

and Sanoamuang (1993) detailed the taxonomic features of the New Zealand

Filinia species based on scanning electron microscopic examination of jaw

elements (trophi). Most recently, Ian Duggan and colleagues detailed rotifer

communities associated with macrophytes in Lake Rotomanuka (Duggan et

al. 1998, 2001a), surveyed planktonic rotifer assemblages more widely on the

North Island (Duggan et al. 2001b, 2002a,b), and related rotifer resting-egg

densities to lake trophic state (Duggan et al. 2002c). Each study documented

new records.

Considerable knowledge of the composition of the New Zealand rotifer

fauna has thus accumulated since studies began 145 years ago. The availability

and value of this information to users such as ecologists have been very poor,

however, for a number of reasons. Until very recently, almost all taxonomic

workers failed to give comprehensive ﬁgures and keys to species, many of the

journals containing the original descriptions are obscure and not readily available

to most New Zealand workers, and the New Zealand literature is itself rather

scattered and often not easily obtained.

Gastropus hyptopus.

From Hudson & Gosse 1886

Front and side views of Testudinella patina.

From Edmondson 1959

141

PHYLUM GNATHIFERA

Endemism and taxonomic features of New Zealand’s rotifer fauna

The apparent endemicity of New Zealand’s rotifer fauna is low (less than 5%).

Some 12–15% of Australia’s rotifers are endemic on present evidence. It is likely

that endemism will be greater in both regions when taxa presently placed in

cosmopolitan taxa are examined more closely. Many of the early records were

identiﬁed by northern hemisphere systematists at a time when cosmopolitan-

ism was widely accepted. Thus, taxa only superﬁcially resembling nominate

species became lumped with them or, even if recognised by a describer as

distinct, may have been synonymised with a northern hemisphere ‘lookalike’

by a subsequent reviser.

Based on current knowledge of New Zealand taxa, there appears to be some

afﬁnity with the Australian fauna; in particular, Keratella australis, K. slacki, Lecane

eylesi, and L. herzigi are conﬁned to these two countries. Lacinularia species,

however, are notably absent in New Zealand, although eight have been recorded

in Australia and some are common components of the plankton of reservoirs

and billabongs there (Shiel & Green 1996). These species also are absent from

Tasmania. Other afﬁnities of the New Zealand fauna with that of Tasmania

include a paucity of brachionids, particularly the absence of the diverse (typically

cold-water) Notholca species group widely reported in the northern hemisphere.

Only two species of Notholca have been reported reliably from Tasmania, plus

an additional halophile Notholca from continental Australia (Shiel, unpubl.).

Notholca apparently is absent from lakes in the North Island (Duggan 2002a),

and just four species have been recorded from the South Island, indicating

possible latitudinal gradients of species in New Zealand. Other similarities

include the absence of Sinantherina and a high diversity of lecanids (Shiel &

Green 1996). Neither of the endemic monogonont species has been recorded

from the North Island.

Ecology and distribution

Seasonal changes in abundance of rotifer species in Lakes Taupo (Forsyth &

McCallum 1980) and Okaro (Forsyth & James 1991) suggested that the marked

seasonality imposed on rotifers in the northern hemisphere by the annual

temperature cycle was less distinct in New Zealand. Forsyth and James (1991)

therefore postulated that, similar to those reported for New Zealand crustacean

zooplankton, seasonal changes in the abundance and composition of food may

be more important than temperature in limiting seasonal distributions. The

results of these studies suggested that the richness of rotifers in New Zealand

lakes was low compared with that in northern temperate lakes, with typically

fewer than 10 species recorded over a year (Chapman & Green 1987).

In an investigation of the spatial and temporal dynamics of rotifers associ-

ated with rooted plants in Lake Rotomanuka, North Island, Duggan et al.

(1998, 2001b) reported 59 species over a 10-month period, a diversity higher

than that previously recorded from limnetic communities in New Zealand (e.g.

Forsyth & McCallum 1980; Forsyth & James 1991). Duggan et al. (1998) found

the changes in rotifer composition to be associated with seasonal changes in

water level and temperature, and a change from heterogeneous to homogeneous

physical and chemical conditions across the littoral from shallow to deep water.

Further, the abundances of many rotifer species differ signiﬁcantly between

different macrophyte species, except in more planktonic forms that appeared to

show little preference (Duggan et al. 2001b). These publications were globally

signiﬁcant, as few studies had previously examined the ecology of periphytic

rotifers (Duggan 2001).

The geographical distribution of planktonic rotifers in 33 North Island lakes

was reported by Duggan et al. (2001b, 2002a). Rotifer diversity within lakes,

previously thought to be low in New Zealand, was found to be comparable with

that of northern temperate lakes. Larger, deeper lakes (e.g. Taupo, Waikare-

moana) were found to have the least taxon richness, and artiﬁcial reservoirs

Euchlanis dilatata.

From Edmondson 1959

Floscularia ringens in a tube of its

own construction.

From Hudson & Gosse 1886

142

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

(e.g. Karapiro, Maraetai) had the most. Rotifer distribution patterns were most

strongly associated with gradients in trophic state when using multivariate tech-

niques. For example, Ascomorpha ovalis, Conochilus unicornis, and C. dossuarius

were associated with oligotrophic conditions, and Brachionus budapestinensis,

B. calyciﬂorus, and Keratella tropica with more eutrophic conditions. Based on

the relationship, Duggan et al. (2001b) developed a bioindicator index using

rotifers for inferring trophic state. High inorganic turbidity in some smaller

New Zealand lakes also appeared to be an important determinant of species

composition. Some species (e.g. Keratella australis and Conochilus exiguus) are

restricted in their distribution (Duggan et al. 2002a). They may be recent arrivals

into North Island habitats or are limited here by poor dispersal abilities.

In their investigation of rotifer resting-egg abundances from oligo-mesotro-

phic Lake Tikitapu and eutrophic Lake Okaro, Duggan et al. (2002c) reported

signiﬁcantly higher densities of resting eggs from the Okaro sediments. Incubat-

ing lake sediments at different temperatures did not result in the hatching of

all of the species recorded from the plankton of these lakes (e.g. Duggan et al.

2002a).

Gaps in knowledge and scope for future research

Collections of rotifers in New Zealand to date have been predominantly from

the open water of lakes and ponds. Sampling of rotifers from vegetated habitats

has been spatially and temporally restricted. For a greater understanding of the

taxonomic composition and biogeographical associations of the New Zealand

rotifer fauna, other habitats need to be given attention, including wetlands, ﬂow-

ing waters, marine and saline habitats, soil, and meiobenthic (interstitial).

Limnologists collecting rotifers on New Zealand’s subantarctic islands

should be on the look-out for Limnognathia maerski, which is superﬁcially

rotifer-like. This sole known species of the new class Micrognathozoa was ﬁrst

discovered in a Greenland spring (Kristensen & Funch 2000) but has since

been found in moss and psammon (sandy sediment) at the subantarctic Crozet

Islands in the Indian Ocean. De Smet (2002) suspects that the species may be

widespread, perhaps transported as cysts or resting eggs by waterfowl.

Class Acanthocephala – thorny-headed worms

‘Few zoologists and still fewer veterinarians and physicians ever encounter a

thorny-headed worm’ is the opening statement in a chapter on the Acantho-

cephala in Roberts and Janovy’s 1996 textbook, Foundations of Parasitology. This

adequately reﬂects both past and present awareness of these animals and their

place in the general scheme of things. Acanthocephalans are obligate internal

parasites of vertebrates. The most distinguishing feature of the group is a

proboscis at the anterior end of the body that bears hooks for attaching to the

gut of their hosts. The name Acanthocephala (‘thorny-headed’) refers to this

feature. These worms are generally thought to be rare or overdispersed but are

known to occur in all classes of vertebrates. In their monograph Biology of the

Acanthocephala, Crompton and Nickol (1985) quote Justus Mueller as saying in

his review of the parasitic helminths, ‘We have neglected the Acanthocephala

from our discussion. They are a monotonous group and few people seem inclined

to work with them.’ These few people, however, have contributed not just to

our understanding of the morphology, physiology, ecology, and biochemistry

of the group itself, but also more generally to our understanding of parasites

and parasitism.

The ﬁrst description recognisably pertaining to an acanthocephalan was

given in 1684 by Redi, who found specimens in European eels. The existence

of distinct genera and species was not recognised until the thirteenth edition

of Systema Naturae, edited by Gmelin between 1788 and 1793 (see Amin 1985).

Trichotria tetractis.

From Edmondson 1959

Scanning electron micrograph of a cystacanth

of Profilicollis antarcticus from the mud crab

Helice crassa.

Annette Brockerhoff

CAN’T FIND THIS PIC

143

PHYLUM GNATHIFERA

This relatively late appreciation of acanthocephalans is surprising given that

many of them are quite large. Macracanthorhynchus hirudinaceus, for example,

can reach up to 15 centimetres long and is distributed worldwide in free-ranging

and domestic pigs. It was ﬁrst described as Taenia hirudinacea in the belief it

was a tapeworm (Pallas 1781). The name was not stabilised into its current form

until more than a century later by Travassos (1917).

The ﬁrst generic names proposed for these worms were Acanthocephalus

by Koelreuther in 1741 and Echinorhynchus by Zoega and Mueller in 1776).

The group was given its present name, Acanthocephala, by Rudolphi, who, in

his writings of 1802, 1808, and 1809, gave it ordinal rank, with a single genus

Echinorhynchus (see Amin 1985). As work on the group continued, more spe-

cies were described and the taxonomy was revised. Signiﬁcant contributions

to classiﬁcation were made by Meyer (1932, 1933), who included all species

described to that date, as did Petrochenko (1956, 1958 – translated into English

in 1971), Yamaguti (1963), and Amin (1985).

Van Cleave (1941) argued to establish the group as a separate phylum, and

four classes comprising nine orders came to be recognised (Amin 1987).

The afﬁnities of the Acanthocephala with other phyla have been difﬁcult to

establish because their body plan has extensive adaptations to an exclusively

parasitic way of life.

Evidence for relationships with tapeworms (Cestoda) was presented by Van

Cleave (1941). The presence of a ‘cuticle’ had been suggested as a link to the

nematodes, but the structure of tegument, cuticle, and body wall is quite dif-

ferent in each group. Golvan (1958) decided that acanthocephalans were most

closely related to the Priapulida (penis worms) and therefore might be included

with the Aschelminthes. Conway-Morris and Crompton (1982) supported this

view by postulating similarities between a fossil priapulid from the Burgess

Shale deposits of the Middle Cambrian, Ancalagon, and a hypothetical proto-

acanthocephalan. This analysis relied on morphological resemblances between

the acanthocephalan proboscis and the priapulid introvert, but priapulids

have a mouth and gut, which are lacking in acanthocephalans. At present, the

aschelminth worms are widely considered to be a polyphyletic grouping (see

e.g. Winnepennickx et al. 1995; Wallace et al. 1996; D’Hondt 1997). Current

evidence using a variety of morphological, developmental, ultrastructural, and

molecular analyses treats acanthocephalans as aberrant endoparasitic rotifers

close to the Bdelloidea (Whiteﬁeld 1971; Clément 1985; Lorenzen 1985; Garey

et al. 1996; Wallace et al. 1996; Albrecht et al. 1997; Zrzavy et al. 1998; Ferraguti

& Melone 1999). Although these afﬁnities are accepted in the six-kingdom

classiﬁcation scheme of Cavalier-Smith (1998), other authorities, including

Barnes et al. (1998), Margulis and Schwartz (1998), and Monks (2001), continue

to support the Acanthocephala as a phylum, albeit linked with the Rotifera and

separate from the Aschelminthes.

Morphology

All acanthocephalans are pseudocoelomate parasites that live in the intestines

of vertebrates as adults. Sexes are separate and there may be notable sexual

dimorphism, especially in overall body size. Acanthocephalans can vary greatly

between species, from tiny (less than 2 mm long) Octospiniferoides chandleri

Bullock, 1957 to metre-long Oligacanthorhynchus longissimus (Golvan 1962).

Living acanthocephalans are often white to yellow in colour, but can be orange

or pink.

The body itself is divided into two regions. The anterior praesoma has

a retractable proboscis with rows of hooks that attach to the intestinal wall

of the host, a smooth unspined neck, a muscular sac, a proboscis receptacle

onto which proboscis invertor muscles are attached, and paired non-muscular

structures – the lemnisci. Contraction of the proboscis invertor muscles causes

the proboscis to invaginate into the proboscis receptacle. Contraction of the

Acanthocephalus galaxii from adult

Galaxias maculatus (whitebait).

From Hine 1977

144

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

proboscis receptacle forces the proboscis to evaginate. A nerve ganglion (often

referred to as the brain) is located within the receptacle. The position of the

ganglion can be taxonomically important, as is whether the proboscis is single- or

double-walled. Neck retractor muscles attach an infolding of the anterior body

wall to the inner surface of the trunk. When both the proboscis retractor muscles

and the neck retractor muscles contract, the entire anterior end is withdrawn

into the trunk. Since the size and shape of the proboscis and the size, shape,

and arrangement of the proboscis hooks are important diagnostic characters,

material that has been ﬁxed with the proboscis contracted is almost useless for

identiﬁcation purposes.

The metasoma, or trunk, encloses the body cavity, is lined by internal muscle

layers, and contains either a male or female reproductive system attached to one

or two ligament sacs that extend from the end of the proboscis receptacle to near

the genital pore. The worm is usually ﬂattened in life but if placed in a hypotonic

solution becomes turgid, resulting in a rounded trunk and evaginated proboscis.

Such treatment prior to ﬁxation facilitates examination of internal structures and

ensures that the features of the proboscis and hooks are visible. The presence

or absence, size, type, and distribution of spines on the trunk is important

taxonomically at genus and family level within the class Palaeacanthocephala

as well as in distinguishing species in other groups. If present, they are acquired

during development in the intermediate host. Hooks and spines both originate

from connective tissue lying between the tegument and muscle layers.

The tegument, a metabolically active syncytium, consists of several micro-

scopic layers, including: a surface coat, the glycocalyx, rich in polysaccharides; a

striped zone with surface pores opening into crypts which increase the surface

area of the tegument; a felt layer with mitochondria, lipid, and glycogen droplets,

golgi complexes, and lysosomes; and a radial layer. Nuclei are found in the

basal region of the tegument and may be numerous and widely distributed or

large and few in number. The number of these nuclei is always constant in a

species and thus taxonomically informative, as is their distribution along the

trunk and whether they are whole or fragmented.

Fluid-ﬁlled channels (lacunar canals) are arranged in two independent

systems in the body wall, one for the praesoma and one for the metasoma; each

is thought to function independently. The praesomal system is continuous, with

a central channel in each of the lemnisci. The metasomal system consists of main

longitudinal canals connected to a complex network of interconnecting channels,

the location and arrangement of which have diagnostic importance.

The musculature of the body wall consists of an inner longitudinal layer

surrounded by an outer circular layer of hollow, ﬂuid-ﬁlled, tubular muscles.

These are involved, through numerous anastomosing interconnections, in

the circulation of lacunar ﬂuids to and from the lacunar channels, effectively

providing for their own nutrition and waste disposal. A rete (net) system of

modiﬁed muscle cells, highly branched anastomosing tubules, lies between the

longitudinal and circular muscles. The body wall therefore appears to provide

an alternative to a circulatory system and may also function as a hydrostatic

skeleton. The arrangement of the muscles of the proboscis is used in higher-

order classiﬁcation.

An excretory system, comprising protonephridia with ﬂame cells, is associ-

ated with the reproductive system in some members of the order Gigantorhyn-

chida. It is assumed that in other acanthocephalans the tegument is involved

in elimination of excretory wastes.

The female reproductive system consists of an organ, fragmenting into ovar-

ian balls prior to maturity, that ﬂoats freely in a ligament sac attached posteriorly

to the uterine bell. The number and persistence of ligament sacs are considered

ordinal characters. The muscular uterine bell has been interpreted as an egg-

sorting device, allowing only mature eggs to be released into the host intestine.

The female gonopore is usually subterminal.

Generalised diagrams of male (A) and female (B)

acanthocephalans.

Key: b = brain; bc = bursal cap; bu = bursa; c = cement

gland; g = genital ligament; l = lemniscus; p = proboscis;

pr = proboscis receptacle; r = retractor muscles;

s = Saefftigen’s pouch; t= testis; ub = uterine bell;

u= uterus; v = vagina. Scale bar = 1 mm.

145

PHYLUM GNATHIFERA

The male reproductive system includes two testes, sperm ducts, a penis,

cement glands, and a copulatory bursa. A Saefftigen’s pouch, cement reservoir,

and seminal vesicle may also be present. The testes are suspended within the

ligament sac by a ligament strand. The cement gland produces a substance

used to cover the external posterior end of the trunk, including the genital

pore of the female after mating has occurred. This is thought to prevent the

female from mating with other males once she has been fertilised. Three types

of cement glands are recognised – a single syncytial gland with giant nuclei

and an accessory gland in the Eoacanthocephala; two to eight elongate tubular

or pyriform glands each with a single nucleus in the Archiacanthocephala and

Polyacanthocephala; and two to eight syncytial glands with nuclear fragments

in the Palaeacanthocephala. The eoacanthocephalan arrangement is thought

to be the most primitive, but this is still being debated.

A summary of the diagnostic characters of the four main groups is given in

the table below based on Bullock (1969) and Amin (1987).

Life cycles

All acanthocephalans have a two-host life cycle, including a vertebrate and an

arthropod. The adult always occurs in the intestine of a vertebrate. The Archia-

canthocephala have a terrestrial life cycle involving birds or mammals as the

deﬁnitive hosts with insects or myriapods as the intermediate hosts. Life cycles

may also include paratenic hosts ( i.e. in which the parasite can encyst and survive

without further development until eaten by the appropriate vertebrate host).

The other three orders have aquatic life cycles with tiny crustaceans such

as amphipods as intermediate hosts. In some cases, paratenic hosts bridge the

trophic gap between arthropods (intermediate host) and vertebrates (deﬁnitive

host). This is particularly relevant where the deﬁnitive host is a top carnivore

and therefore unlikely to consume small crustaceans. Mature, embryonated

eggs are voided to the outside via the host faeces. The intermediate host eats

the eggs containing the developing larvae or acanthors and these hatch in the

midgut of the arthropod host and either penetrate into the haemocoel or attach

to the gut wall as acanthellae. During the acanthella stage, adult organs mature

Diagnostic characters of acanthocephalans

Character Palaeacanthocephala Archiacanthocephala Eoacanthocephala Polyacanthocephala

Body size Variable Mostly large Small–medium Medium–large

Host habitat Mostly aquatic Terrestrial Aquatic Aquatic

Lacunar system – main Usually lateral Dorsal and ventralor Dorsal and ventral at Dorsal and ventral

longitudinal vessels dorsal only anterior end

Cement glands 2–8 multinucleate Usually 8 uninucleate One, syncytial with 4–8 giant nuclei

few giant nuclei

Trunk spines Present or absent Absent Present or absent Present

Subcuticular nuclei Numerous amitotic Few elongated or Very few, giant Many small

fragments or few branched, remaining

highly branched fragments close

Proboscis receptacle Closed sac with Single muscle layer, Closed sac, single Closed sac, single

2 muscle layers often with ventral cleft muscle layer muscle layer

or accessory muscles

Ligament sacs Single, rupturedin Dorsal and ventral, Dorsal and ventral, Dorsal and ventral,

mature worms persistent not persistent persistent in females

Nephridia Absent Present or absent Absent Absent

Intermediate hosts Crustaceans Insects, millipedes Crustaceans Crustaceans

Deﬁnitive hosts All vertebrate classes Birds and mammals Fish, occasionallyreptiles, Fish, crocodiles

amphibians

146

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

and the worm becomes infective, as the cystacanth stage, to the deﬁnitive host.

Cystacanths may be able to be identiﬁed to genus or species level using the

morphology of the praesoma and sometimes trunk spination. In the deﬁnitive

host, the cystacanth excysts, attaches to the host’s gut wall and matures into

either an adult male or female.

Diversity of New Zealand acanthocephalans

Worldwide, there are more than 1,100 known species of Acanthocephala in

some 21 families. The New Zealand species – some 38 in 13 genera – are

relatively few. Of these, only 22 have been identiﬁed to actual species level, and

some identiﬁed only to genus may include more than one species. This fauna

represents less than 2% of estimated global acanthocephalan biodiversity.

Petrochenko (1958) reported that no acanthocephalans had been noted

in New Zealand birds or terrestrial mammals to that point in time, although

two species – Sphaerirostris physocoracis and Mediorhynchus zosteropis – had

been recorded from birds in the Polynesian region. Analysis of known species

from bird hosts was indicative of few endemic genera. Some biogeographical

regions with no endemic genera did, however, have relatively large numbers of

endemic species. This pattern is a reﬂection of the ecology of the hosts, many of

which are migratory, thus precluding sharp regional boundaries (Petrochenko

1958). A similar analysis of terrestrial mammalian hosts showed a low level

of endemism even at species level. The fact that many species are common to

hosts across several regions, however, may result from lack of understanding

of the group, with more information needed. Petrochenko (1958) noted that

the neotropical region had the largest numbers of species, the largest number

of endemic genera and species, and the largest number of primitive mammal

hosts. He therefore suggested that the neotropical region may have been the

centre of origin of acanthocephalans in mammals.

Two genera are known from marine mammals – Bolbosoma Porta, 1908 in

whales, and Corynosoma Lühe, 1904 in seals and otters. Of these genera, four

species had been recorded from Antarctic waters and eight species from the

Paciﬁc Ocean (Petrochenko 1958). Three species were common to both the

Atlantic and Paciﬁc Oceans. Again, lack of research effort had an inﬂuence

on the robustness of information available. The former Soviet Union had an

extensive research programme on marine mammals at that time, which was

Generalised cycle of Corynosoma species. Male and female worms attach to the wall of the

small intestine of whales, dolphins, seals and marine fish-eating birds. Eggs developing

into acanthors are passed out in host faeces and are eaten by crustacean intermediate

hosts. Acanthors develop through the acanthella stage to cystacanths that infect

crustacean-eating fish. Fish act as paratenic hosts, where cystacanths wait without further

development until eaten by the definitive vertebrate host.

cystacanth

adult female

egg

free-living

acanthor

acanthella

147

PHYLUM GNATHIFERA

reﬂected in 40% of known species from marine mammals being recorded from

waters surrounding the USSR coast (Petrochenko 1958).

Since 1958, some work has been done on the New Zealand fauna, with

one cosmopolitan species recorded from terrestrial mammals, at least seven

species from marine mammals, and seven species (three possibly endemic)

from birds. New species of Proﬁlicollis and Plagiorhynchus have been found in

New Zealand oystercatchers but not yet Australian ones (Smales 2001; Smales

& Brockerhoff 2002), and an acanthocephalan was described from a kiwi in

1900 as Echinorhynchus sp., a genus now reserved for species from ﬁsh.

A Polish group conducted extensive surveys in the 1980s and 1990s in

subantarctic and Antarctic waters around the South Shetland and South Orkney

Islands and South Georgia, extending our knowledge of acanthocephalans in

southern waters. Other nations with bases in the Antarctic have similarly carried

out parasitological studies in recent times.

Zdzitowiecki (1984) noted that, with the exception of uncertain records (Cor-

ynosoma bullosum (Linstow, 1892) in the northern hemisphere and Corynosoma

sermeme (Forssell, 1904) in the Southern), there is a clear distinction between

acanthocephalan parasites of Antarctic and Arctic marine-mammal and bird

hosts. He also suggested differences between acanthocephalans of Antarctic

and subantarctic hosts. The distributions of most Corynosoma species found

in hosts from New Zealand waters would tend to support his hypothesis.

Corynosoma australe Johnson & Best, 1937, for example, is found in Australian,

New Zealand, and South American waters but not in Antarctic latitudes. By

contrast, C. hannae, previously known only from leopard seals in the Antarctic,

has been found in fur seal pups and yellow-eyed penguins from St Clair Beach,

Dunedin, and the Otago Peninsula.

All presently recognisable acanthocephalan species from New Zealand

birds and mammals are classiﬁed in two of the seven families that include all

the known representatives from birds and mammals.

Of the four main groups of acanthocephalans, all but the Archiacantho-

cephala have representatives using ﬁsh as their deﬁnitive hosts (Amin 1987).

Only eight fully identiﬁed species of acanthocephalans from 20 New Zealand

ﬁsh species are known (see checklist of species). There are still several unidenti-

ﬁed acanthocephalans collected from New Zealand ﬁsh that may represent

new host records. The eight identiﬁed species are from four different parasite

families spread across nine host families. Two of these acanthocephalans –

Acanthocephalus galaxii Hine, 1977 and Aspersentis peltorhamphi (Baylis, 1944) –

are known only from New Zealand (Pichelin 2002). Micracanthorhynchina

hemi-rhamphi (Baylis 1944), Neoechinorhynchus aldrichettae Edmonds, 1971, and

Australorhynchus tetramorphacanthus Lebedev, 1967 are also present in Austra-

lian ﬁsh. The original description of Raorhynchus terebra was from Katsu-

wonuis pelamis but it is not known where the fish was caught; R. terebra

was later recorded from Hawai‘i (Dollfus, 1969). The other species, Neoechino-

rhynchus chilkaensis Poddar, 1937, was ﬁrst described from a mugilid (mullet)

from India and has only been recorded from other mullets in India and New

Zealand, and from Arripis trutta in New Zealand. Thus it appears that the few

known species of acanthocephalans present in New Zealand ﬁsh have no

common link other than that they are parasites of ﬁsh, and even then some

are freshwater and others are marine.

It may well be, as is the case in many parasite species, that the acanthocepha-

lan ﬁsh fauna of New Zealand is not depauperate but that more ﬁshes need to

be examined for parasites and/or the species are very much overdispersed.

There are no major repositories of acanthocephalans in New Zealand.

Material appears to have been deposited in an ad hoc fashion in museums in

New Zealand, Australia, and London (UK), or held as working collections at

NIWA (Wellington), California State University (USA), and Massey and Otago

Universities (New Zealand).

Corynosoma semerme from Phocarctos hookeri

(Hookers sea lion).

From Johnston & Edmonds 1953

Corynosoma clavatum from Leucocarbo colensoi

(Auckland Island shag).

From Johnston & Edmonds 1953

148

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

Health aspects – human and wildlife

If medical parasitology textbooks include a section on the Acanthocephala, it

is usually very brief and perhaps under a heading of unusual ﬁndings, since

human infections are all thought to be accidental. Not many people eat raw

insects, small crustaceans, lizards, toads, or other potential intermediate hosts.

Macracanthorhynchus hirudinaceus has occasionally been recognised as a parasite

of humans (six reports of numerous cases between 1859 and 1983). Moniliformis

moniliformis, usually found in rats, has repeatedly been found in humans.

Corynosoma strumosum, a common seal parasite (via ﬁsh paratenic hosts), has

also been found (see Schmidt 1971). Bolbosoma sp., normally found in whales,

has been reported in humans from Japan (Myazaki 1991). Not surprisingly, in

each of these cases there is either a potentially close relationship between normal

hosts (e.g. pig and rat) and humans, or eating raw ﬁsh has been implicated.

There have been no records of thorny-headed worms in New Zealanders.

Macracanthorhynchus hirudinaceus, Bolbosoma sp., and Corynosoma spp. occur in

the New Zealand fauna, so there is a remote possibility of zoonotic infections.

Not enough is known about the life cycles and potential pathogenicity of

acanthocephalans occurring in New Zealand vertebrates to assess their potential

to cause disease or death. Ranum and Wharton (1996) considered the general

threat of parasitism in the rare yellow-eyed penguin (Megadyptes antipodes) but

did not consider acanthocephalans. This was in the mistaken belief that there are

few instances of the worms occurring in ﬁsh-eating birds. An acanthocephalan

proboscis has the potential to cause serious damage to the intestinal wall of a

host. In most cases, the proboscis penetrates the outer layer of the intestinal

wall, affecting mucosa, submucosa, and the circular layer of the muscularis

externa. Necrosis, however, is slight and any inﬂammatory response limited

(Schmidt 1969). Some species of acanthocephalan stimulate the production of

dense connective-tissue nodules. In other cases, complete perforation of the

gut occurs, with the host species succumbing to peritonitis (Schmidt 1969).

Acanthocephalans have been implicated in epizootics of birds (see below). It

seems prudent to assume that these parasites do have potential to affect penguin

survival, and management strategies should be modiﬁed accordingly.

What is the impact of larval stages on intermediate hosts? Studies have

been carried out on some Otago coast crabs to address this question. Proﬁlicollis

antarcticus and P. novaezelandensis cystacanth larvae infect the hairy-handed

crab Hemigrapsus crenulatus, common rock crab H. edwardsi, and stalk-eyed

mud crab Macrophthalmus hirtipes. Worm eggs are accidentally ingested by

the crabs when they feed and encyst in the crabs’ body cavities. In general,

worm larvae do not affect crab fecundity. On the other hand, their presence

seems to increase the susceptibility of crabs to be caught by shorebirds, in part

by affecting crab behaviour and body colour (Haye & Ojeta 1998; Latham &

Poulin 2001, 2002).

Alien species

All parasites brought into New Zealand with introduced hosts could be desig-

nated alien, even when, in the case of Macracanthorhynchus hirudinaceus from

the domestic pig (Sus scrofa), the parasite is known to be cosmopolitan. The

only other known mammalian parasite is a single unidentiﬁed acanthocephalan,

still attached to the preserved intestinal wall of a ferret (Mustela furo). This speci-

men is presently housed in the Institute of Veterinary, Animal, and Biomedical

Sciences teaching collection at Massey University, Palmerston North.

Cosmopolitan acanthocephalans that have not been introduced with their

alien hosts are Oncicola pomatostomi, with cats (Felis catus) as deﬁnitive hosts,

and Moniliformis moniliformis, with rodents (Rattus spp.) as deﬁnitive hosts.

The latter parasite is interesting because it has not been found in the black rat

(R. rattus) or the paciﬁc rat (R. exulans) in New Zealand, both of which have

been reported from the Australian and Polynesian regions as infected (Roberts

Corynosoma hannae from Megadyptes antipodes

(yellow-eyed penguin).

Simon McDonald

149

PHYLUM GNATHIFERA

1991). It is also surprising that O. pomatostomi does not appear to have been

introduced into New Zealand. It has been recorded from Australia both in feral

cats and dingoes as well as in a wide range of paratenic hosts, especially in the

tissues of the neck of more than 30 species of birds (Edmonds 1989).

Polymorphus species may have been brought into New Zealand with mallard

and grey teal ducks. Although noted by Rind (1974) as two species of acantho-

cephalan from the same genus, the specimens listed in her report on parasites of

freshwater birds were not named to genus level. Rind (1974) used Petrochenko

(1958) as a reference text for identiﬁcation, suggesting that the specimens were

Polymorphus species. The material has not yet been located. If Polymorphus species

are present, there is a possibility they could switch from the introduced hosts

to a native duck species. Heavy infections of Polymorphus have been implicated

in mortalities within duck populations (Petrochenko 1958), so the presence of

Polymorphus infections in any such population is cause for concern.

Gaps in knowledge

The information on Acanthocephala in New Zealand is fragmentary and

sparse. It is not clear whether the low number of species is truly indicative of a

depauperate fauna or of a patchy research effort. In any case, the latter has been

compounded by the lack of a single repository for voucher and other specimens

of parasitic helminths. Much of the already sparse data on which this review

has been based may have been lost because of this. There are no specialists

currently working on acanthocephalans in New Zealand to further progress in

understanding the scope of the group in a local context.

Of the known species, two are endemic, some are found elsewhere in the

Australasian region, and the others are cosmopolitan. An acanthocephalan

noted as occurring in the North Island brown kiwi may also be endemic but

has yet to be fully identiﬁed.

Checklists of parasites from birds (Rind 1974; Week 1982), mammals (Bowie

1984; McKenna 1997), and ﬁsh (Blair 1984; Hewitt & Hine 1972) have been

drawn up and a bibliography (Barker 1973) has been prepared, all including

acanthocephalans, but there appear to be no acanthocephalan records for many

potential host species. Likewise, there appear to be no published records from

amphibians or reptiles. Intensive survey work is needed to conﬁrm the presence

or absence of acanthocephalan infections in the endemic New Zealand fauna

for which data are presently not available. When such data become available,

trends in endemism and patterns of distribution of acanthocephalans in the

New Zealand region will become amenable to analysis.

Authors

Dr Ian C. Duggan Department of Biological Sciences, Waikato University,

Private Bag 3105, Hamilton, New Zealand [i.duggan@waikato.ac.nz]

Rotifers

Dr John D. Green 532 Mooloo Road, Mooloo via Gympie, QLD 4570, Australia

[verdi.oboe@bigpond.com]

Rotifers

Dr Sylvie Pichelin Department of Microbiology and Parasitology, University of

Queensland, Brisbane, QLD 4072, Australia [s.pichelin@mailbox.uq.edu.au]

Acanthocephala

Dr Russell J. Shiel Department of Environmental Biology, The University of

Adelaide, Adelaide, South Australia 5005, Australia [russell.shiel@adelaide.edu.au]

Rotifers

Professor Lesley Smales School of Biological and Environmental Sciences, Central

Queensland University, Rockhampton, QLD 4702 Australia [l.warner@cqu.edu.au]

Acanthocephala

150

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

Dr Wolfgang Sterrer Bermuda Natural History Museum, PO Box FL 145,

Flatts FLBX, Bermuda [museum.bamz@mail.ibl.bm]

Gnathostomulida

Acknowledgments

The authors gratefully thank Anthony Saville for access to the Russell collection of

rotifers at the Canterbury Museum, Christchurch, and Amanda Freeman for her help in

tracing slides and notebooks. Hendrik Segers, University of Ghent, Belgium, commented

on taxonomic afﬁnities of the New Zealand Lecanidae. For aiding the review of Acan-

thocephala, the authors wish to thank Eileen Harris (Natural History Museum, London),

John Andrews (Victoria University of Wellington), Richard Norman (Massey University,

Palmerston North), Mark Walker (Canterbury Museum, Christchurch), Erena Barker

(Otago Museum, Dunedin), John Early (Auckland Museum), Bob McDowall (NIWA,

Christchurch), Ben Diggles (DigsFish Services, Queensland), Philip Sirvid (Museum of

New Zealand Te Papa Tongarewa, Wellington), and Simon McDonald (University of

Otago, Dunedin) for helping track down locations of material and lending specimens

for examination.

ALBRECHT, H.; EHLERS, U.; TARASCHEWSKI,

H. 1997: Syncytial organisation of acanthors of

Polymorphus minutus (Palaeacanthocephala),

Neoechinorhynchus rutili (Eoacanthocephala),

and Moniliformis monilifomis (Archiacantho-

cephala) (Acanthocephala). Parasitology Research

83: 326–338.

AHLRICHS, W. H. 1995: Ultrastruktur und Phylo-

genie von Seison nebaliae (Grube 1859) und

Seison annulatus (Claus 1876) – Hypothesen zu

phylogenetischen Verwandtschaftsverhältsnis-

sen innerhalb der Bilateria. Dissertation, Georg-

August University, Göttingen. Cuvillier-Verlag,

Göttingen. 310 p.

AHLRICHS, W. H. 1997: Epidermal ultrastructure

of Seison nebaliae and Seison annulatus, and a

comparison of epidermal structures within the

Gnathifera. Zoomorphology 117: 41–48.

AMIN, O. M. 1985: Classiﬁcation. Pp. 27–72 in

Crompton, D. W. T.; Nickol, B. B. (Eds), Biology

of the Acanthocephala. Cambridge University

Press, Cambridge.

AMIN, O. M. 1987: Key to the families and sub-

families of Acanthocephala with the erection of

a new class (Polyacanthocephala) and a new

order (Polyccanthorhynchidae). Journal of

Parasitology 73: 1216–1219.

AX, P. 1956: Die Gnathostomulida, eine rätselhafte

Wurmgruppe aus dem Meeressand. Abhandlun-

gen der Akademie der Wissenschaften und Literatur

Mainz, mathematisch-naturwissenschaftliche

Klasse 8: 1–32.

BARKER, A. S. 1973: Bibliography of the helminth

parasites of New Zealand (1879–1971). Common-

wealth Institute of Helminthology Technical

Communication 44: 1–79.

BARNES, R. S. K. 1998: The Diversity of Living

Organisms. Blackwell Science, Oxford. 345 p.

BULLOCK, W. L. 1969: Morphological features

as tools and as pitfalls in acanthocephalan

systematics. Pp. 4–43 in: Schmidt, G. D. (ed.),

Problems in Systematics of Parasites. University

Park Press, Baltimore.

BERZINS, B. 1973: Zwei neue Euchlanis-Arten,

Rotatoria. Zoologischer Anzeiger 190: 125–127.

BLAIR, D. 1984: A checklist and bibliography of

parasites of New Zealand freshwater ﬁshes.

Mauri Ora 11: 5–50.

BOWIE, J. Y. 1984: Parasites from an Atlantic

bottle-nose dolphin (Tursiops truncatus), and a

revised checklist of parasites of this host. New

Zealand Journal of Zoology 11: 395–398.

BREHM, V. 1928: Fresh-water fauna of New

Zealand, contributions to a knowledge of.

Transactions and Proceedings of the New Zealand

Institute 59: 779–812.

BROCKERFOFF, A. M.; SMALES, L. R. 2002: Pro-

ﬁlicollis novaezelandensis n. sp. (Polymorphidae)

and two other acanthocephalan parasites from

shore birds (Haematopodidae) and Scolopa-

cidae) in New Zealand, with records of two

species in intertidal crabs (Decapoda; Grapsidae

and Ocypodidae). Systematic Parasitology 52:

55–65.

BULLOCK, W. L. 1969: Morphological features

as tools and as pitfalls in acanthocephalan

systematics. Pp. 4-43 in: Schmidt, G. D. (ed.),

Problems in Systematics of Parasites. University

Park Press, Baltimore.

BURNS, C. W.; MITCHELL, S. F. 1980: Seasonal

succession and vertical distribution of zoo-

plankton in Lake Hayes and Lake Johnson.

New Zealand Journal of Marine and Freshwater

Research 14: 189–204.

CASSIE, V. 1960: A red water bloom of rotifers in

Cook Strait, New Zealand. Nature 186: 328-329.

CAVALIER-SMITH, T. 1998: A revised six-

kingdom system of life. Biological Reviews 73:

203–266.

CHAPMAN, M. A. 1973: Calamoecia lucasi

(Copepoda, Calanoida) and other zooplankters

in two Rotorua, New Zealand, lakes. Internatio-

nale Revue der gesamten Hydrobiologie 58: 79–104.

CHAPMAN, M. A.; GREEN. J. D. 1987: Zoo-

plankton ecology. In: Viner, A. B. (ed.), Inland

Waters of New Zealand. Department of Scientiﬁc &

Industrial Research Bulletin 241: 225–263.

CLÉMENT, P. 1985: The relationships of the

rotifers. Pp. 224–247 in: Conway Morris, S.;

George, J. D.; Gibson, R.; Platt, H. M. (eds), The

Origins and Relationships of Lower Invertebrates.

Clarenden Press, Oxford.

CONWAY-MORRIS, S.; CROMPTON, D. E. T.

1982: The origins and evolution of the Acantho-

cephala. Biological Reviews 57: 85–115.

CORDES, D. O.; O’HARA, P. J. 1979: Diseases

of captive marine mammals. New Zealand

Veterinary Journal 27:147-150.

CROMPTON, D. W. T.; NICKOL, B. B. 1985:

Biology of the Acanthocephala. Cambridge

University Press, Cambridge. 519 p.

DAILEY, M. D.; VOGELBEIN, W. K. 1991: Parasite

fauna of three species of Antarctic whales with

reference to their use as potential stock indicators.

Fishery Bulletin 89:355–365.

DE SMET, W. H. 2002: A new record of Limno-

gnathia maerski Kristensen & Funch, 2000

(Micrognathozoa) from the subantarctic Crozet

Islands, with redescription of the trophi. Journal

of Zoology 258: 381–393.

DOLLFUS, R. P. 1969: De Raorhynchus terebra

(Rudolphi) (Palaeacanthocephala, Rhadi-

norhynchidae). Bulletin du Muséum national

d’Histoire naturelle 41: 755–758.

DONNER, J. 1962: Neue und wenig bekannte

Bdelloidea (Rotatoria) des Bodens. Acta

Zoologica Academiae Scientiarum Hungaricae 8:

303–333.

DUGGAN, I. C. 2001: The ecology of periphytic

rotifers. Hydrobiologia 446/447: 139–148.

DUGGAN, I. C.; GREEN, J. D.; THOMPSON, K.;

SHIEL, R. J. 1998: Rotifers in relation to littoral

ecotone structure in Lake Rotomanuka, North

Island, New Zealand. Hydrobiologia 387/388:

179–197.

DUGGAN, I. C.; GREEN, J. D.; THOMPSON, K.;

SHIEL, R. J. 2001a: The inﬂuence of macro-

phytes on the spatial distribution of littoral

rotifers. Freshwater Biology 46: 777–786.

DUGGAN, I. C.; GREEN, J. D.; SHIEL, R. J. 2001b:

Distribution of rotifers in North Island, New

Zealand, and their potential use as bioindicators

of lake trophic state. Hydrobiologia 446/447:

155–164.

DUGGAN, I. C.; GREEN, J. D.; SHIEL, R. J. 2002a:

Distribution of rotifer assemblages in North

Island, New Zealand, lakes: relationships to

environmental and historical factors. Freshwater

Biology 47: 195–206.

DUGGAN, I. C.; GREEN, J. D.; THOMASSON, K.

2002b: Do rotifers have potential as bioindicators

of lake trophic state? Verhandlungen der

internationalen Vereinigung für theoretische und

angewandte Limnologie 27: 3497–3502.

DUGGAN, I. C.; GREEN, J. D.; SHIEL, R. J.

2002c: Rotifer resting egg densities in lakes of

different trophic state, and their assessment

using emergence and egg counts. Archiv für

Hydrobiologie 153: 409–420.

EDMONDS, S. J. 1989: A list of Australian

Acanthocephala and their hosts. Records of the

South Australian Museum 23: 127–133.

FERRAGUTI, M.; MELONE, G. 1999: Spermio-

genesis in Seison nebaliae (Rotifera, Seisonidea):

further evidence of a rotifer-acanthocephalan

relationship. Tissue and Cell 31: 428–440.

References

151

PHYLUM GNATHIFERA

FORSYTH, D. J.; JAMES, M. R. 1991: Seasonal

abundance and diel vertical dynamics of zoo-

plankton in Lake Okaro. Archiv für Hydrobiologie

121: 419–429.

FORSYTH, D. J.; LEWIS, M. H. 1987: The inver-

tebrates. In: Viner, A. B. (ed. ), Inland Waters of

New Zealand. Department of Scientiﬁc & Industrial

Research Bulletin 241: 265–290.

FORSYTH, D. J.; McCALLUM, I. D. 1980: Zoo-

plankton of Lake Taupo. New Zealand Journal of

Marine and Freshwater Research 14: 65–69.

GAREY, J. R.; NEAR, T. J.; NONNEMACHER, M. R.;

NADLER, S. A. 1996: Molecular evidence for

Acanthocephala as a subtaxon of Rotifera.

Journal of Molecular Evolution 43: 287–292.

GIRIBET, G.; DISTEL, D. L.; POLZ, M.; STERRER,

W.; WHEELER, W. C. 2000: Triploblastic

relationships with emphasis on the acoelo-

mates and the position of Gnathostomulida,

Cycliophora, Plathelminthes, and Chaetognatha:

a combined approach of 18S rDNA sequences

and morphology. Systematic Biology 49: 539–562.

GIRIBET, G.; SØRENSEN, M. V.; FUNCH, P.;

KRISTENSEN, R. M.; STERRER, W. 2004:

Investigations into the phylogenetic position

of Micrognathozoa using four molecular loci.

Cladistics 20: 1–13.

GOLVAN, Y. J. 1958: Le phylum des Acantho-

cephala. Première note. Sa place dans l’échelle

zoologique. Annales de Parasitologie humaine et

comparée 33 : 538–602.

GREEN, J. D. 1969: Common freshwater planktonic

animals. Tane (Journal of the Auckland University

Field Club) 15: 13–24.

GREEN, J. D. 1974: The limnology of a New

Zealand reservoir with particular reference to

the life histories of the copepods Boeckella

propinqua Sars and Mesocyclops leuckarti Claus.

Internationale Revue der gesamten Hydrobiologie

59: 441–487.

GREEN, J. D. 1976: Plankton of Lake Ototoa,

a sand dune lake in northern New Zealand.

New Zealand Journal of Marine and Freshwater

Research 10: 43–59.

HAIGH, S. B. 1963: The bdelloid rotifers of New

Zealand. Journal of the Queckett Microscopical

Club, ser. 4, 29: 161–70.

HAIGH, S. B. 1965: The bdelloid rotifers of

New Zealand. Part II. Journal of the Queckett

Microscopical Club, ser. 4, 30: 7–13, 36–41.

HAIGH, S. B. 1966: The bdelloid rotifers of

New Zealand. Part III. Journal of the Queckett

Microscopical Club, ser. 4, 30: 193–201, (1967)

223–226.

HAIGH, S. B. 1968: The bdelloid rotifers of

New Zealand. Part IV. Journal of the Queckett

Microscopical Club, ser. 4, 31: 33–41.

HAIGH, S. B. 1970: The bdelloid rotifers of New

Zealand. Part V. Microscopy 31: 326–330; (1971)

32: 17–23.

HAMILTON, A. 1879: On Melicerta ringens and

Plumatella repens. Transactions and proceedings of

the New Zealand Institute 12: 301–303.

HARRING, H. K. 1913: Synopsis of the Rotatoria.

Bulletin of the American Museum of Natural

History 81: 1–226.

HAYE, P. A.; OJETA, F. P. 1998: Metabolic and

behavioural alterations in the crab Hemigrapsus

crenulatus (Milne Edwards 1837) induced by its

acanthocephalan parasite Proﬁlicollis antarcticus

Zdzitowiecki 1985). Journal of Experimental

Marine Biology and Ecology 228: 73–82.

HEWITT, G. C.; HINE, P. M. 1972: Checklist of

parasites of New Zealand ﬁshes and their hosts.

New Zealand Journal of Marine and Freshwater

Research 6: 69–114.

HILGENDORF, F. W. 1898: A contribution to the

study of the Rotifera of New Zealand. Trans-

actions and Proceedings of the New Zealand Institute

31: 107–134.

HILGENDORF, F. W. 1902: The Rotifera of New

Zealand: a revised and expanded list. Trans-

actions and Proceedings of the New Zealand Institute

35: 267–271.

HOLLOWDAY, E. D. 2002: Family Synchaetidae

Hudson & Gosse, 1886. In: Nogrady, T.; Segers,

H. (eds), Guides to the Identiﬁcation of the Micro-

invertebrates of the Continental Waters of the

World 18 (Rotifera 6): 87-211.

HUDSON, C. T.; GOSSE, P. H. 1886: The Rotifera;

or Wheel Animalicules both British and Foreign. 2

vols. Longman, Green, & Co. , London.

HUSSEY, C. G. 1981: A checklist and bibliography

of records of Rotifera (Rotatoria) in Britain.

Microﬁche. British Museum (Natural History),

London.

HUTTON, F. W. 1904: Index Faunae Novae

Zealandiae. Dulau & Co. , London. [Rotifera pp.

300–302.]

JOLLY, V. H. 1952: A preliminary study of the

limnology of Lake Hayes. Australian Journal of

Marine and Freshwater Research 3: 74–91.

JOLLY, V. H. 1977: The comparative limnology of

some New Zealand lakes. 2. Plankton. New

Zealand Journal of marine and Freshwater Research

3: 74–91.

KOSTE, W. 1978: Die Rädertiere Mitteleuropas.

Uberordnung Monogononta. 2 vols. Borntraeger,

Stuttgart.

KOSTE, W.; SHIEL, R. J. 1980: New Rotifera from

Australia. Transactions of the Royal Society of

South Australia 104: 133–144.

KOSTE, W.; SHIEL, R. J. 1987: Rotifera from

Australian inland waters. II: Brachionidae

and Epiphanidae (Rotifera: Monogononta).

Invertebrate Taxonomy 1: 949–1021.

KOSTE, W.; SHIEL, R. J. 1991: Rotifera from

Australian inland waters. VII: Notommatidae

(Rotifera: Monogononta). Transactions of the

Royal Society of South Australia 115: 111–159.

KRISTENSEN, R. M.; FUNCH, P. 2000: Micro-

gnathozoa: a new class with complicated jaws

like those of Rotifera and Gnathostomulida.

Journal of Morphology 246: 1–49.

LATHAM, A. D. M.; POULIN, R. 2001: Effect of

acanthocephalan parasites on the behaviour

and coloration of the mud crab Macrophthalmus

hirtipes (Brachyura: Ocypodidae). Marine Biology

139: 1147–1154.

LATHAM, A. D. M.; POULIN, R. 2002: Field

evidence of the impact of two acanthocephalan

parasites on the mortality of three species of

New Zealand shore crabs. Marine Biology 141:

1131–1139.

JONES, J. B. 1991: Movements of albacore tuna

(Thunnus aldunga) in the South Paciﬁc: evidence

from parasites. Marine Biology 111: 1–9.

LORENZEN, S. 1985: Phylogenetic aspects of

pseudocoelomate evolution Pp. 210–223 in:

Conway Morris, S.; George, J. D.; Gibson, R.;

Platt, H. M. (eds), The Origins and Relationships

of Lower Invertebrates. Clarenden Press, Oxford.

LOWE, D. J.; GREEN, J. D. 1987: Origins and

development of the lakes. In: Viner, A. B. (ed.),

Inland Waters of New Zealand. Department of

Scientiﬁc & Industrial Research Bulletin 241: 1–64.

McDOWALL, R. H. 1987: The native ﬁsh. In:

Viner, A. B. (ed.), Inland Waters of New Zealand.

Department of Scientiﬁc & Industrial Research

Bulletin 241: 291–306.

McKENNA, P. B. 1997: Checklist of helminth

parasites of terrestrial mammals in New

Zealand. New Zealand Journal of Zoology 24:

277–290.

MARGULIS, L.; SCHWARTZ, K. V. 1998: Five

Kingdoms – An Illustrated Guide to the Phyla of

Life on Earth. 2nd edn. Freeman & Co, New York.

520 p.

MEYER, A. 1932: Acanthocephala. In: Bronn, H. G.

(ed.), Klassen und Ordnungen des Tierreich 4:

1–332.

MEYER, A. 1933: Acanthocephala. In: Bronn, H. G.

(ed.). Klassen und Ordnungen des Tierreich 4:

333–522.

MONKS, S. 2001: Phylogeny of the Acanthocephala

based on morphological characters. Systematic

Parasitology 48: 81–116.

MORRIS, C. B. 1912: Some notes on Rotifera not

previously recorded as occurring in New

Zealand. Transactions and Proceedings of the New

Zealand Institute 45: 163–167.

MORRIS, C. B. 1913: Classiﬁed list of the New

Zealand Rotatoria, including twelve species not

previously recorded. Transactions and Proceed-

ings of the New Zealand Institute 46: 213–219.

MURRAY, J. 1911: Rotifera of New Zealand:

collected by the Shackleton Antarctic Expedition,

1907–1909. Journal of the Royal Microscopical

Society 31: 573–583.

MURRAY, J. 1913: Australasian Rotifera. Journal of

the Royal Microscopical Society 33: 455–461.

MYAZAKI, I. 1991: Acanthocephalans. Pp. 466–471

in: An Illustrated Book of Helminth Zoonoses.

Shukosa Printing Co., Fukuoka.

NIELSEN, C. 2003: Deﬁning phyla: morphological

and molecular clues to metazoan evolution.

Evolution and Development 5: 386–393.

NOGRADY, T.; POURRIOT, R.; SEGERS, H. 1995:

Rotifera Vol. 3: Notommatidae, Scaridiidae.

Guides to the Identiﬁcation of the Microinvertebrates

of the Continental Waters of the World 8: 1–248.

OCHIETTI, S.; CAILLEUX, A. 1969: Comparaison

des Conodontes et des mâchoires de Gnatho-

stomulides. Comptes rendus de l’Académie des

Sciences 268: 2664–2666.

PARR, P. J. 1949: Observations on Rotatoria in

Otago. MSc thesis, Zoology Department,

University of Otago. 27 p.

PARR, P. J. 1992: Planktonic rotifers of New

Zealand lakes. Produced by author, unpublished,

Levin.

PETROCHENKO, V. I. 1956: Acanthocephala of

Domestic and Wild Animals Volume 1. Izdatel’stvo

Akademii Nauk SSSR, Moscow. 435 p.

PETROCHENKO, V. I. 1958: Acanthocephala of

Domestic and Wild Animals Volume 2. Izdatel’stvo

Akademii Nauk SSSR, Moscow. 477 p.

PICHELIN, S.; SMALES, L.; BRAY, R. A. 2002:

A discussion on the Heteracanthocephalidae

Petrochecko, 1956 (Acanthocephala: Palaea-

canthocephala). Systematic Parasitology 52:

145–152.

PODDAR, T. N. 1937: On a new species of

Neoechinorhynchus parasite in Mugil cephalus

from Chilka Lake. Records of the Indian Museum

39: 129–131.

RANUM, A.; WHARTON, D. A. 1996: Helminth

and protozoan parasites of the alimentary

tract of the yellow-eyed penguin (Megadyptes

antipodes). New Zealand Journal of Zoology 23:

83–87.

RIEDL, R. J. 1969: Gnathostomulida from America

– this is the ﬁrst record of the new phylum from

North America. Science 163: 445–442.

RIEGER, R. M.; TYLER, S. 1995: Sister-group

relationship of Gnathostomulida and Rotifera-

Acanthocephala. Invertebrate Biology 114:

186–188.

152

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

RIND, S. 1974: Some helminth parasites of fresh-

water birds from the South Island, New Zealand,

with particular reference to trematodes of ducks.

Mauri Ora 2: 139–146.

RISER, N. W. 1984: General observations on the

intertidal interstitial fauna of New Zealand.

Tane (Journal of the Auckland University Field

Club) 30: 239–250.

ROBERTS, L. S.; JANOVY, J. 1996: Phylum

Acanthocephala: Thorny headed worms. Pp.

469–482 in: Foundations of Parasitology. William

C. Brown, Dubuque.

ROBERTS, M. 1991: The parasites of the Polynesian

rat within and beyond New Zealand. International

Journal for Parasitology 21: 777–783.

RUSSELL, C. R. 1944: A new rotifer from New

Zealand. Journal of the Royal Microscopical Society

64: 121–123.

RUSSELL, C. R. 1945: A reference list of the

Rotatoria of New Zealand, with ecological notes.

Transactions of the Royal Society of New Zealand

75: 102–123.

RUSSELL, C. R. 1947: Additions to the Rotatoria

of New Zealand, Part I. Transactions of the Royal

Society of New Zealand 76: 403–408.

RUSSELL, C. R. 1949: Additions to the Rotatoria

of New Zealand, Part II. Transactions of the Royal

Society of New Zealand 77: 351–354.

RUSSELL, C. R. 1950a: Additions to the Rotatoria

of New Zealand, Part III. Transactions of the

Royal Society of New Zealand 78: 161–166.

RUSSELL, C. R. 1950b: The design of a photo-

micrographic camera for general laboratory use.

Australian Journal of Instrument Technology 6:

107–113.

RUSSELL, C. R. 1950c: An improved form of

plankton pump apparatus. Australian Journal of

Instrument Technology 6: 179–185.

RUSSELL, C. R. 1951a: The Rotatoria of the Upper

Stillwater Swamp. Records of the Canterbury

Museum 5: 245–251.

RUSSELL, C. R. 1951b: An index of the rotifers in

the C. B. Morris collection of microscope slides

at the Cawthron Institute, Nelson. Transactions

of the Royal Society of New Zealand 79: 52–54.

RUSSELL, C. R. 1952: Additions to the Rotatoria of

New Zealand. Part IV. Transactions of the Royal

Society of New Zealand 80: 59–62.

RUSSELL, C. R. 1953a: Additions to the Rotatoria

of New Zealand Part V. Transactions of the Royal

Society of New Zealand 81: 73–78.

RUSSELL, C. R. 1953b: Some Rotatoria of the

Chatham Islands. Records of the Canterbury

Museum 6: 237–244.

RUSSELL, C. R. 1954: Additions to the Rotatoria of

New Zealand Part VI. Transactions of the Royal

Society of New Zealand 82: 461–463.

RUSSELL, C. R. 1956a: Some additions to the

Rotatoria of the Chatham Islands. Records of the

Canterbury Museum 7: 51–53.

RUSSELL, C. R. 1956b: Some rotifers from the

Fiordland district. Records of the Canterbury

Museum 7: 55–59.

RUSSELL, C. R. 1956c: A centrifuge for the separa-

tion of small aquatic animals from debris. The

Microscope 10: [1–3].

RUSSELL, C. R. 1957: Additions to the Rotatoria of

New Zealand Part VII. Transactions of the Royal

Society of New Zealand 84: 939–940.

RUSSELL, C. R. 1958a: Some rotifers from Campbell

Island. Records of the Dominion Museum 3:

137–140.

RUSSELL, C. R. 1958b: Some rotifers from Malaya.

Transactions of the Royal Society of New Zealand

85: 433–437.

RUSSELL, C. R. 1959: Additions to the Rotatoria

of New Zealand. VIII. Transactions of the Royal

Society of New Zealand 87: 69–73.

RUSSELL, C. R. 1960: An index to the Rotatoria

of New Zealand and outlying islands from

1859–1959. Transactions of the Royal Society of

New Zealand 88: 443–461.

RUSSELL, C. R. 1961a: The Rotatoria of Queens-

land, Australia. Transactions of the Royal Society

of New Zealand, n.s., 1: 235–239.

RUSSELL, C. R. 1961b: A simple method of per-

manently mounting rotifer trophi. Journal of the

Queckett Microscopical Club, ser. 4, 5: 377–378.

RUSSELL, C. R. 1961c: The mounting of rotifers

or other small animals in glycerine jelly. Journal

of the Queckett Microscopical Club, ser. 4, 5:

384–386.

RUSSELL, C. R. 1962: Additions to the Rotatoria of

New Zealand, Part IX. Transactions of the Royal

Society of New Zealand, n.s., 1: 337–341.

SANOAMUANG, L. 1993: Comparative studies

on scanning electron microscopy of trophi of

the genus Filinia Bory de St Vincent (Rotifera).

Hydrobiologia 264: 115–128.

SANOAMUANG, L.; STOUT, V. 1993: New

records of rotifers from the South Island lakes,

New Zealand. Hydrobiologia 255/256: 481–490.

SCHMARDA, L. K. 1859: Neue wirbellose Thiere

beobachtet und gesammelt auf einer Reise um die

Erde, 1853 bis 1857 . . . Band 1. Turbellarien,

Rotatorien und Anneliden. Wilhelm Engelmann,

Leipzig. xix + 66 p., 15 pls. [Rotatoria pp. 47–66.]

SCHMIDT, G. D. 1969: Acanthocephala as agents

of disease in wild mammals. Wildlife Diseases 5:

1–10.

SCHMIDT, G. D. 1971: Acanthocephalan infections

of man, with two new records. Journal of Parasit-

ology 67: 582–584.

SEGERS, H. 1995: Rotifera Vol. 2: Lecanidae

(Monogononta). Guides to the Identiﬁcation of the

Microinvertebrates of the Continental Waters of the

World 6: 1–223.

SEGERS, H. 2002: The nomenclature of the

Rotifera: annotated checklist of valid family- and

genus-group names. Journal of Naural History 36:

631–640.

SEGERS, H.; MELONE, G. 2002: A comparative

study of trophi morphology in Seisonidea

(Rotifera). Journal of Zoology 244: 201–207.

SHIEL, R. J. 1995: A guide to identiﬁcation of

rotifers, cladocerans and copepods from

Australian inland waters. Cooperative Research

Centre for Freshwater Ecology Identiﬁcation Guide

3: 1–144.

SHIEL, R. J.; GREEN, J. D. 1996. Rotifera recorded

from New Zealand, 1859–1995, with comments

on zoogeography. New Zealand Journal of

Zoology 23: 193–209.

SHIEL, R. J.; KOSTE, W. 1979: Rotifera recorded

from Australia. Transactions of the Royal Society

of South Australia 103: 57–68.

SHIEL, R. J.; KOSTE, W. 1986: Australian Rotifera:

Ecology and biogeography. Pp. 142–150 in: De

Deckker, P.; Williams, W. D. (eds), Limnology

in Australia. CSIRO/Dr W. Junk, Melbourne &

Dordrecht. 671 p.

SHIEL, R. J.; KOSTE, W. 1993: Rotifera from

Australian inland waters. IX. Gastropodidae,

Synchaetidae, Asplanchnidae (Rotifera:

Monogononta). Transactions of the Royal Society

of South Australia 117: 111–139.

SHIEL, R. J.; SANOAMUANG, L. 1993: Trans-

Tasman variations in Australasian Filinia

populations. Hydrobiologia 255/256: 455–462.

SMALES, L. R. 2002: Plagiorhynchidae Meyer, 1931

(Acanthocephala) from Australasian birds and

mammals with descriptions of Plagiorhynchus

(Plagiorhynchus) menurae Johnston, 1912 and

P. (P. ) allisonae n. sp. Systematic Parasitology 51:

207-216.

SØRENSEN, M. V. 2003: Further structures in

the jaw apparatus of Limnognathia maerski

(Micrognathozoa), with notes on the phylogeny

of the Gnathifera. Journal of Morphology 255:

131–145.

SØRENSEN, M. V.; STERRER, W.; GIRIBET, G.

2006: Gnathostomulid phylogeny inferred from

a combined approach of four molecular loci and

morphology. Cladistics 22: 32–58.

STERRER, W. 1972: Systematics and evolution

within the Gnathostomulida. Systematic Zoology

21: 151–173.

STERRER, W. 1991: Gnathostomulida from Fiji,

Tonga and New Zealand. Zoologica Scripta 20:

107–128.

STERRER, W. 1998: Gnathostomulida from the

(sub)tropical northwestern Atlantic. Studies on

the Natural History of the Caribbean Region 74:

1–178.

STERRER, W.; MAINITZ, M.; RIEGER, R. M.

1985: Gnathostomulida: enigmatic as ever. Pp.

183–199 in: Conway-Morris, S.; George, J. D.;

Gibson, R.; Platt, H. M. (eds.), The Origins and

Relationships of Lower Invertebrates. Clarendon

Press, Oxford. 394 p.

STOCK, A. 1892: On a New Zealand variety of

Floscularia coronetta Cubitt. Transactions of the

New Zealand Institute 25: 193.

STOUT, V. M. 1969: Lakes in the mountain region

of Canterbury, New Zealand. Verhandlungen

internationale Vereinigung für theoretische und

angewandte Limnologie 17: 404–413.

STOUT, V. M. 1978: Effects of different silt loads

and of hydro-electric developments on four

large lakes. Verhandlungen internationale Vereini-

gung für theoretische und angewandte Limnologie

20: 1182–1185.

STOUT, V. M. 1981: Some year to year ﬂuctuations

in a natural and in an artiﬁcial lake, South Island,

New Zealand. Verhandlungen internationale

Vereinigung für theoretische und angewandte

Limnologie 21: 699–702.

STOUT, V. M. 1984: Seasonal plankton cycles in

two adjacent lakes in the South Island, New

Zealand. Verhandlungen internationale Vereini-

gung für theoretische und angewandte Limnologie

22: 1226–1230.

STOUT, V. M. 1991: Year to year ﬂuctuations in

zooplankton in a small mountain lake. Verhan-

dlungen internationale Vereinigung für theoretische

und angewandte Limnologie 24: 1145–1148.

SUDZUKI, M. 1959: A new proalid rotifer,

Liliferotrocha urawensis n. g. n. sp. , from a

marsh in Urawa, Japan. Scientiﬁc Report, Tokyo

Kyoiku Daigaku, Section B9: 9–33.

SWEDMARK, B. 1964: The interstitial fauna of

marine sand. Biological Reviews 39: 1–42.

TOLICH, G. 1988: The population dynamics of

rotifers in the Mangere oxidation ponds.

MSc thesis, University of Auckland.

VAN CLEAVE, H. J. 1941: Relationships of the

Acanthocephala. American Naturalist 75: 31–47.

WALLACE, R. L. 2002: Rotifers: exquisite metazoans.

Integrative and Comparative Biology 42: 660–667.

WALLACE, R. L.; RICCI, C.; MELONE, G. 1996:

A cladistic analysis of pseudocoelomate

(aschelminth) morphology. Invertebrate Biology

115: 104–112.

WALLACE, R. L.; SNELL, T. W. 1993: Rotifera.

Pp. 187–248 in: Thorp, J. H.; Covich, A. P. (eds),

Ecology and Classiﬁcation of North American

Freshwater Invertebrates. Academic Press, New

York. 911 p.

153

PHYLUM GNATHIFERA

WEEKES, P. J. 1982: Checklist of helminth

parasites of birds in New Zealand. New Zealand

Journal of Zoology 9: 451–460.

WHITFIELD, P. J. 1971: Phylogenetic afﬁnities of

Acanthocephala: an assessment of ultrastruc-

tural evidence. Parasitology 63: 49–58.

WINNEPENNICKX, B.; BACKELJAU, T.; MACKEY,

L. Y.; BROOKS, J. M.; DE WACHTER, R.;

KUMAR, S.; GAREY. J. R. 1995: 18S rRNA data

indicate that the Aschelminthes are polyphyletic

in origin and consist of at least three distinct

clades. Molecular Biology and Evolution 12:

1132–1137.

YAMAGUTI, S. 1963: Acanthocephala. Systema

Helminthum Volume 5. Wiley Interscience, New

York. 423 p.

ZDZITOWIECKI, K. 1984: Some Antarctic

acanthocephalans of the genus Corynosoma

Checklist of New Zealand Gnathifera

All the Gnathostomulida are marine; their classiﬁcation follows Sterrer (1972). Clas-

siﬁcation of the rotifers follows Sefers (2002).

E = endemic species; M= marine.

Class GNATHOSTOMULIDA

Order FILOSPERMOIDEA

HAPLOGNATHIIDAE

Haplognathia sp. indet. 1 Sterrer unpubl.

Haplognathia sp. indet. 2 Sterrer unpubl.

Haplognathia sp. indet. 3 Sterrer unpubl.

Haplognathia sp. indet. 4 Sterrer unpubl.

PTEROGNATHIIDAE

Pterognathia sp. indet. 1 Sterrer unpubl.

Pterognathia sp. indet. 2 Sterrer unpubl.

Pterognathia sp. indet. 3 Sterrer unpubl.

Pterognathia sp. indet. 4 Sterrer unpubl.

Order BURSOVAGINOIDEA

Suborder SCLEROPERALIA

GNATHOSTOMULIDAE

Gnathostomula maorica Sterrer, 1991

Gnathostomula sp. indet. Sterrer unpubl.

Suborder CONOPHORALIA

AUSTROGNATHIIDAE

Austrognathia novaezelandiae Sterrer, 1991

Austrognatharia sp. indet. Sterrer unpubl.

Class ROTIFERA

Subclass BDELLOIDEA

Order PHILODINAVIDA

PHILODINAVIDAE

Philodinavus paradoxus (Murray, 1905)

Order PHILODINIDA

HABROTROCHIDAE

Habrotrocha angusticollis (Murray, 1905)

Habrotrocha angusticollis var. attenuata (Murray,

1906)

Habrotrocha angusticollis var. monteti de Koning,

1947

Habrotrocha angusticollis var. reversa Bartos, 1952

Habrotrocha aspera (Bryce, 1892)

Habrotrocha bidens Gosse, 1851

Habrotrocha bidens var. Haigh, 1967 E

Habrotrocha collaris (Ehrenberg, 1832)

Habrotrocha constricta (Dujardin, 1841)

Habrotrocha crenata Murray, 1905

Habrotrocha crenata var. sphagnicola Pavlovski,

1938

Habrotrocha elegans (Milne, 1886)

Habrotrocha elusa Milne, 1916

Habrotrocha eremita Bryce, 1894

Habrotrocha ﬂava Bryce, 1915

Habrotrocha ﬂava var. Bryce, 1915

Habrotrocha ﬂaviformis de Koning, 1947

Habrotrocha gibbosa de Koning, 1947

Habrotrocha gracilis var. Montet, 1915

Habrotrocha insignis var. Bryce, 1915

Habrotrocha lata (Bryce, 1892)

Habrotrocha leitgebii (Zelinka, 1886)

Habrotrocha ligula Bryce, 1913

Habrotrocha ligula var. aligula Burger, 1948

Habrotrocha munda Bryce, 1913

Habrotrocha perforata (Murray, 1906)

Habrotrocha proxima Donner 1953

Habrotrocha puella var. excedens Donner, 1962

Habrotrocha pulchra (Murray, 1905)

Habrotrocha pusilla (Bryce, 1893)

Habrotrocha pusilla nuda Donner, 1950

Habrotrocha pusilla textrix f. bicornis Haigh, 1966 E

Habrotrocha pusilla textrix f. brevilabris Donner, 1950

Habrotrocha pusilla textris f. longilabris Donner, 1950

Habrotrocha reclusa Milne, 1886

Habrotrocha rosa Donner, 1949

Habrotrocha scepanotrochoides de Koning, 1947

Habrotrocha serpens Donner, 1949

Habrotrocha spicula Bryce, 1913

Habrotrocha tridens Milne, 1886

Habrotrocha tridens globigera Donner, 1950

Habrotrocha tripus (Murray, 1907)

Otostephanus auriculatus (Murray, 1911)

Otostephanus auriculatus var. bilobatus Hauer, 1939

Otostephanus donneri Bartos, 1959

Otostephanus torquatus (Bryce, 1913)

Scepanotrocha corniculata Bryce, 1910

Scepanotrocha rubra Bryce, 1910

Scepanotrocha setifera Haigh, 1963 E

Scepanotrocha simplex var. de Koning, 1947

PHILODINIDAE

Ceratotrocha cornigera (Bryce, 1893)

Ceratotrocha delicata (Donner, 1949)

Didymodactylos carnosus Milne, 1916

Dissotrocha aculeata Ehrenberg, 1832

Dissotrocha aculeata f. dukesi Russell, 1959 E

Dissotrocha macrostyla (Ehrenberg, 1838)

Macrotrachela brevilabris de Koning, 1947

Macrotrachela brevilabris var. aliena Donner, 1964

Macrotrachela concinna (Bryce, 1912)

Macrotrachela decora Bryce, 1912

Macrotrachela ehrenbergi (Janson, 1893)

Macrotrachela extensa Haigh, 1966 E

Macrotrachela festinans Donner, 1949

Macrotrachela habita (Bryce, 1894)

Macrotrachela insolita de Koning, 1943

Macrotrachela ligulata Haigh, 1966 E

Macrotrachela microcornis (Murray, 1911)

Macrotrachela multispinosa Thompson, 1892

Macrotrachela multispinosa var. brevispinosa

(Murray, 1908)

Macrotrachela musculosa (Milne, 1886)

Macrotrachela nana (Bryce, 1912)

Macrotrachela nixa Donner, 1962

Macrotrachela obtusa Haigh, 1967 E

Macrotrachela papillosa Thompson, 1892

Macrotrachela plicata (Bryce, 1892)

Macrotrachela punctata (Murray, 1911)

Macrotrachela quadricornifera Milne, 1886

Macrotrachela quadricornifera var. ligulata Berzins,

1950

Macrotrachela quadricornifera var. Haigh, 1963 E

Macrotrachela quadricornifera Bryce, no date,

n.nud.?

Macrotrachela timida Milne, 1916

Macrotrachela tuberilabris de Koning, 1947

Macrotrachela zickendrahti Richters, 1902

Mniobia bredensis de Koning, 1947

Mniobia burgeri Bartos, 1951

Mniobia dentata Haigh, 1963 E

Mniobia dentata var. Haigh, 1967 E

Mniobia incrassata Murray, 1905

Mniobia lobata Haigh, 1967 E

Mniobia magna (Plate, 1889)

Mniobia obtusicornis Murray, 1911

Mniobia obtusicornis var. Murray, 1911

Mniobia placida Haigh, 1963 E

Mniobia punctata Haigh, 1963 E

Mniobia russeola (Zelinka, 1891)

Mniobia scabrosa Murray, 1911

Mniobia scarlatina (Ehrenberg, 1853)

Mniobia symbiotica (Zelinka, 1886)

Mniobia tarda Donner, 1949

Mniobia tetraodon (Ehrenberg, 1848)

Mniobia tetraodon var. Haigh, 1971 E

Philodina acuticornis Murray, 1902

Philodina brevipes Murray, 1902

Philodina citrina Ehrenberg, 1832

Philodina erythophthalma Ehrenberg, 1830

Philodina ﬂaviceps Bryce, 1906

Philodina grandis Milne, 1916

Philodina megalotrocha Ehrenberg, 1832

Philodina nemoralis Bryce, 1903

Philodina nitida Milne, 1916

Philodina plena Bryce, 1894

Philodina proterva Milne, 1916

Philodina roseola Ehrenberg, 1832

Philodina rugosa Bryce, 1903

Philodina rugosa var. Haigh, 1966 E

Philodina squamosa Murray, 1906

Philodina tranquila Wulfert, 1942

Philodina vorax (Janson, 1893)

Pleuretra alpium (Ehrenberg, 1853)

Pleuretra brycei (Weber, 1898)

Pleuretra humerosa (Murray, 1905)

Pleuretra lineata Donner, 1962

Rotaria curtipes (Murray, 1911)

Rotaria exoculis de Koning, 1947

Rotaria macroceros (Gosse, 1851)

Rotaria macrura (Ehrenberg, 1832)

Rotaria montana (Murray, 1911)

Rotaria neptunia (Ehrenberg, 1832)

Rotaria rotatoria (Pallas, 1766)

Rotaria sordida (Western, 1893)

Rotaria sordida var. ﬁmbriata Murray, 1906

Rotaria tardigrada (Ehrenberg, 1832)

parasitising Pinnipedia, with descriptions of

three new species. Parasitologia Polonica 29:

359–377.

ZRZAVY, J.; MIHULKA, S.; KEPKA, P.; BEZDEK,

A.; TIETZ, D. 1998: Phylogeny of the Metazoa

based on morphological and ribosomal DNA

evidence. Cladistics 14: 249–285.

154

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

Order ADINETIDA

ADINETIDAE

Adineta acuticornis Haigh, 1967 E

Adineta barbata Janson, 1893

Adineta cuneata var. gracilis Haigh, 1963

Adineta elongata Rodewald, 1935

Adineta gracilis Janson, 1893

Adineta longicornis Murray, 1906

Adineta steineri Bartos, 1951

Adineta tuberculosa Janson, 1893

Adineta vaga (Davis, 1873)

Adineta vaga var. minor Bryce, 1893

Bradyscela clauda (Bryce, 1893)

Subclass MONOGONONTA

Order COLLOTHECACEA

ATROCHIDAE

Cupelopagis vorax (Leidy, 1857)

COLLOTHECIDAE

Collotheca ambigua (Hudson, 1883)

Collotheca coronetta (Cubitt, 1869)

Collotheca libera (Zacharias, 1894)

Collotheca mutabilis (Hudson, 1885)

Collotheca ornata (Ehrenberg, 1832)

Collotheca ornata var. cornuta (Dobie, 1849)

Collotheca pelagica Rousselet, 1893

Collotheca spinata (Hood, 1893)

Order FLOSCULARIACEA

CONOCHILIDAE

Conochilopsis causeyae (Vidrine, McLaughlin &

Willis, 1985)

Conochilus coenobasis (Skorikov, 1914)

Conochilus dossuarius Hudson, 1875

Conochilus exiguus Ahlstrom, 1938

Conochilus hippocrepis (Schrank, 1830)

Conochilus natans (Seligo, 1900)

Conochilus unicornis Rousselet, 1892

FLOSCULARIIDAE

Floscularia conifera (Hudson, 1886)

Floscularia ringens (Linnaeus, 1758)

Limnias ceratophylli Schrank, 1803

Ptygura crystallina (Ehrenberg, 1834)

Ptygura melicerta (Ehrenberg, 1832)

Ptygura melicerta var. socialis (Weber, 1888)

Ptygura velata (Gosse, 1851)

HEXARTHRIDAE

Hexarthra brandorfﬁ Koste, 1977

Hexarthra fennica (Levander, 1892)

Hexarthra intermedia (Wiszniewski, 1929)

Hexarthra mira (Hudson, 1871)

Hexarthra propinqua (Bartos, 1948)

TESTUDINELLIDAE

Pompholyx complanata Gosse, 1851

Pompholyx sulcata Hudson, 1885

Testudinella incisa (Ternetz, 1892)

Testudinella incisa emarginula (Stenroos, 1898)

Testudinella mucronata (Gosse, 1886)

Testudinella parva (Ternetz, 1892)

Testudinella patina (Hermann, 1783)

Testudinella patina f. aspis (Carlin, 1939)

Testudinella reﬂexa (Gosse, 1887)

Testudinella striata (Murray, 1913)

TROCHOSPHAERIDAE

Filinia brachiata (Rousselet, 1901)

Filinia cornuta (Weisse, 1847)

Filinia longiseta (Ehrenberg, 1834)

Filinia longiseta limnetica (Zacharias, 1893)

Filinia novaezealandiae Shiel & Sanoamuang, 1993

Filinia opoliensis (Zacharias, 1898)

Filinia passa (Müller, 1786)

Filinia cf. pejleri Hutchinson, 1964

Filinia terminalis (Plate, 1886)

Order PLOIMIDA

ASPLANCHNIDAE

Asplanchna amphora Hudson & Gosse, 1889

Asplanchna brightwelli Gosse, 1850

Asplanchna intermedia Hudson in Hudson &

Gosse, 1886

Asplanchna priodonta Gosse, 1850

Asplanchna sieboldi (Leydig, 1854)

Asplanchna silvestris Daday, 1902

Asplanchnopus multiceps Schrank, 1793)

BRACHIONIDAE

Anuraeopsis ﬁssa (Gosse, 1851)

Anuraeopsis navicula Rousselet, 1910

Brachionus angularis Gosse, 1851

Brachionus angularis bidens Plate, 1886

Brachionus bidentatus Anderson, 1889

Brachionus budapestinensis (Daday, 1885)

Brachionus calyciﬂorus Pallas, 1766

Brachionus caudatus Barrois & Daday, 1894

Brachionus caudatus var. aculeatus (Hauer, 1937)

Brachionus caudatus f. vulgatus Ahlstrom, 1940

Brachionus leydigii (Cohn, 1862)

Brachionus lyratus Shephard, 1911

Brachionus novaezealandiae (Morris, 1912)

Brachionus plicatilis Müller, 1786

Brachionus quadridentatus Hermann, 1783

Brachionus rubens Ehrenberg, 1838

Brachionus urceolaris Müller, 1773

Brachionus zahniseri Ahlstrom, 1934

Keratella ahlstromi Russell, 1951 E

Keratella australis Berzins, 1963*

Keratella cochlearis (Gosse, 1851)

Keratella cochlearis f. micracantha (Lauterborn,

1900)

Keratella crassa Ahlstrom, 1943

Keratella hispida Lauterborn, 1898

Keratella javana Hauer, 1937

Keratella procurva Thorpe, 1891

Keratella quadrata (Müller, 1786)

Keratella quadrata var. edmondsoni Ahlstrom, 1943

Keratella sancta Russell, 1944

Keratella slacki (Berzins, 1963)*

Keratella tecta (Gosse, 1851)

Keratella tropica (Apstein, 1907)

Keratella valga (Ehrenberg, 1834)

Notholca foliacea (Ehrenberg, 1838)

Notholca paciﬁca (Russell, 1962) E

Notholca squamula (Müller, 1786)

Notholca striata (Müller, 1786)

Plationus patulus (Müller, 1786)

Platyias quadricornis (Ehrenberg, 1832)

DICRANOPHORIDAE

Aspelta angusta Harring & Myers, 1928

Apelta aper (Harring, 1913)

Dicranophoroides caudatus (Ehrenberg, 1834)

Dicranophorus dolerus Harring & Myers, 1928

Dicranophorus edestes Harring & Myers, 1928

Dicranophorus epicharis Harring & Myers, 1928

Dicranophorus forcipatus (Müller, 1786)

Dicranophorus grandis (Ehrenberg, 1832)

Dicranophorus luetkeni (Bergendal, 1892)

Encentrum insolitum Myers, 1936

Encentrum marinum (Dujardin, 1841)

Encentrum putorius eurycephalum Wulfert, 1936

Encentrum saundersiae (Hudson, 1885)

Paradicranophorus hudsoni (Glasscott, 1893)

EPIPHANIDAE

Epiphanes brachionus (Ehrenberg, 1837)

Epiphanes macrourus (Barrois & Daday, 1894)

Epiphanes senta (Müller, 1773)

Proalides tentaculatus De Beauchamp, 1907

Rhinoglena frontalis Ehrenberg, 1853

EUCHLANIDAE

Diplois daviesiae Gosse, 1886

Euchlanis alata Voronkov, 1912

Euchlanis calpidia (Myers, 1930)

Euchlanis deﬂexa (Gosse, 1851)

Euchlanis dilatata Ehrenberg, 1832

Euchlanis dilatata var. crassa Myers, 1938

Euchlanis dilatata var. lucksiana Hauer, 1930

Euchlanis forcipata Russell, 1962

Euchlanis lyra Hudson, 1886

Euchlanis meneta Myers, 1930

Euchlanis oropha Gosse, 1887

Euchlanis parameneta Berzins, 1973

Euchlanis parva Rousselet, 1892

Euchlanis phryne Myers, 1930

Euchlanis pyriformis Gosse, 1851

Euchlanis triquetra Ehrenberg, 1838

Manfredium eudactylotum (Gosse, 1886)

GASTROPODIDAE

Ascomorpha ecaudis (Perty, 1850)

Ascomorpha ovalis (Bergendal, 1892)

Ascomorpha saltans Bartsch, 1870

Gastropus hyptopus (Ehrenberg, 1838)

Gastropus minor (Rousselet, 1892)

ITURIDAE

Itura aurita Wulfert, 1935

Itura myersi Wulfert, 1935

LECANIDAE

Lecane arcula Harring, 1914

Lecane arcuata (Bryce, 1891)

Lecane aspasia Myers, 1917

Lecane bulla (Gosse, 1851)

Lecane closterocerca (Schmarda, 1859)

Lecane cornuta (Muller, 1786)

Lecane decipiens (Daday, 1913)

Lecane elasma Harring & Myers, 1926

Lecane eylesi Russell, 1953*

Lecane ﬂexilis (Gosse, 1886)

Lecane furcata (Murray, 1913)

Lecane galeata (Bryce, 1892)

Lecane gissensis (Gosse, 1886)

Lecane glypta Harring & Myers, 1926

Lecane hamata (Stokes, 1896)

Lecane herzigi Koste, Shiel & Tan, 1988*

Lecane hornemanni (Ehrenberg, 1834)

Lecane latissima Yamamoto, 1955

Lecane leontina (Turner, 1892)

Lecane ludwigii (Eckstein, 1883)

Lecane luna (Müller, 1776)

Lecane luna var. presumpta Ahlstrom 1938

Lecane lunaris (Ehrenberg, 1832)

Lecane lunaris constricta (Murray, 1913)

Lecane lunaris perplexa (Ahlstrom, 1938)

Lecane nana (Murray, 1913)

Lecane ploenensis Voigt, 1902

Lecane pyriformis (Daday, 1905)

Lecane rhacois Harring & Myers, 1926

Lecane cf. rotundata (Olofsson, 1918)

Lecane cf. rhytidaHarring & Myers, 1926

Lecane similis Russell, 1959 E

Lecane stichaea Harring, 1913

Lecane subtilis Harring & Myers, 1926

Lecane tethis (Harring & Myers, 1926)

Lecane cf. ungulata (Gosse, 1887)

LEPADELLIDAE

Colurella adriatica Ehrenberg, 1831

Colurella colurus (Ehrenberg, 1830)

Colurella colurus f. compressa Lucks, 1912

Colurella gracilis (Hilgendorf, 1898)

Colurella hindenbergi Steinecke, 1917

Colurella obtusa (Gosse, 1886)

Colurella salina Althaus, 1957

Colurella uncinata (Müller, 1773)

Colurella uncinata f. bicuspidata (Ehrenberg, 1832)

Colurella uncinata f. deﬂexa (Ehrenberg, 1834)

Lepadella (Heterolepadella) ehrenbergi (Perty, 1850)

Lepadella (Lepadella) acuminata (Ehrenberg, 1834)

Lepadella biloba Hauer, 1958

Lepadella dactyliseta (Stenroos, 1898)

Lepadella latusinus (Hilgendorf, 1898)

Lepadella latusinus var. americana Myers, 1934

Lepadella oblonga (Ehrenberg, 1834)

155

PHYLUM GNATHIFERA

Lepadella ovalis (Müller, 1786)

Lepadella patella (Müller, 1786)

Lepadella quadricarinata (Stenroos, 1898)

Lepadella rhomboides (Gosse, 1886)

Lepadella triptera (Ehrenberg, 1830)

Lepadella vitrea (Shephard, 1911)*

Lepadella whitfordi Ahlstrom, 1938

Squatinella geleii Varga, 1933

Squatinella mutica (Ehrenberg, 1832)

Squatinella rostrum (Schmarda, 1846)

LINDIIDAE

Lindia euchromatica Edmondson, 1938

Lindia fulva Harring & Myers, 1922

Lindia pallida Harring & Myers, 1922

Lindia torulosa Dujardin, 1841

Lindia truncata (Jennings, 1894)

MICROCODONIDAE

Microcodon clavus Ehrenberg, 1830

MYTILINIDAE

Lophocharis salpina (Ehrenberg, 1834)

Mytilina bisulcata (Lucks, 1912)

Mytilina mucronata (Müller, 1773)

Mytilina trigona (Gosse, 1851)

Mytilina ventralis (Ehrenberg, 1832)

Mytilina v. macracantha (Gosse, 1886)

NOTOMMATIDAE

Cephalodella apocolea Myers, 1924

Cephalodella auriculata (Müller, 1773)

Cephalodella biungulata Wulfert, 1937

Cephalodella catellina (Müller, 1786)

Cephalodella cf. delicata Wulfert, 1937

Cephalodella eva (Gosse, 1887)

Cephalodella exigua (Gosse, 1886)

Cephalodella forﬁcata (Ehrenberg, 1832)

Cephalodella forﬁcula (Ehrenberg, 1832)

Cephalodella gibba (Ehrenberg, 1832)

Cephalodella globata (Gosse, 1887)

Cephalodella gracilis (Ehrenberg, 1832)

Cephalodella intuta Myers, 1924

Cephalodella lipara Myers, 1924

Cephalodella megalocephala (Glasscott, 1893)

Cephalodella mucronata Myers, 1924

Cephalodella panarista Myers, 1924

Cephalodella pheloma Myers, 1924

Cephalodella physalis Myers, 1924

Cephalodella plicata Myers, 1924

Cephalodella sterea (Gosse, 1887)

Cephalodella strepta Myers, 1924

Cephalodella tantilla Myers, 1924

Cephalodella tenuior (Gosse, 1886)

Cephalodella ventripes (Dixon-Nuttall, 1901)

Eosphora anthadis Harring & Myers, 1922

Eosphora ehrenbergi Weber, 1918

Eosphora thoides Wulfert, 1935

Eothinia elongata (Ehrenberg, 1832)

Monommata astia Myers, 1930

Monommata appendiculata Stenroos, 1898

Monommata caeca Myers, 1930

Monommata longiseta (Müller, 1786)

Monommata maculata Harring & Myers, 1924

Notommata allantois Wulfert, 1935

Notommata aurita (Müller, 1786)

Notommata caudata Collins, 1872

Notommata cyrtopus Gosse, 1886

Notommata falcinella Harring & Myers, 1921

Notommata saccigera Ehrenberg, 1832

Notommata tripus Ehrenberg, 1838

Pleurotrocha petromyzon (Ehrenberg, 1830)

Taphrocampa annulosa Gosse, 1851

Taphrocampa selenura Gosse, 1887

PROALIDAE

Bryceella tenella (Bryce, 1897)

Bryceella voigti (Rodewald, 1935)

Proales decipiens (Ehrenberg, 1832)

Proales doliaris (Rousselet, 1895)

Proales gigantea (Glasscott, 1893)

Proales longidactyla Edmondson, 1948

Proales parasita (Ehrenberg, 1838)

Proales sordida Gosse in Hudson & Gosse, 1886

Proales theodora (Gosse, 1887)

Proalinopsis staurus Harring & Myers, 1924

SCARIDIIDAE

Scaridium longicaudum (Müller, 1786)

SYNCHAETIDAE

Polyarthra dolichoptera Idelson, 1925

Polyarthra major Burckhardt, 1900

Polyarthra minor Voigt, 1904

Polyarthra remata Skorikov, 1896

Polyarthra vulgaris Carlin, 1943

Synchaeta baltica Ehrenberg, 1834 M

Synchaeta cecilia Rousselet, 1902

Synchaeta cf. curvata Lie-Petersen, 1905 M

Synchaeta cf. fennica Rousselet, 1909

Synchaeta longipes (Gosse, 1887)

Synchaeta monopus Plate, 1889

Synchaeta neapolitana Rousselet, 1902

Synchaeta oblonga Ehrenberg, 1832

Synchaeta pectinata Ehrenberg, 1832

Synchaeta stylata Wierzejski, 1893

Synchaeta tremula (Müller, 1786)

Synchaeta triophthalma Lauterborn, 1894

TRICHOCERCIDAE

Ascomorphella volvocicola (Plate, 1886)

Elosa worralli Lord, 1891

Trichocerca agnatha Wulfert, 1939

Trichocerca bidens (Lucks, 1912)

Trichocerca birostris (Minkiewicz, 1900)

Trichocerca brachyura (Gosse, 1851)

Trichocerca cavia (Gosse, 1886)

Trichocerca chattoni (Beauchamp, 1907)

Trichocerca collaris (Rousselet, 1896)

Trichocerca cristata Harring, 1913

Trichocerca cylindrica (Imhof, 1891)

Trichocerca cf. dixonnuttalli (Jennings, 1903)

Trichocerca elongata (Gosse, 1886)

Trichocerca cf. gracilis (Tessen, 1890)

Trichocerca helminthodes (Gosse, 1886)

Trichocerca iernis (Gosse, 1887)

Trichocerca inermis (Linder, 1904)

Trichocerca insignis (Herrick, 1885)

Trichocerca insulana Hauer, 1937/38)

Trichocerca intermedia (Stenroos, 1898)

Trichocerca lata (Jennings, 1894)

Trichocerca longiseta (Schrank, 1802)

Trichocerca lophoessa (Gosse, 1886)

Trichocerca marina (Daday, 1890)

Trichocerca mus Hauer, 1938

Trichocerca myersi (Hauer, 1931)

Trichocerca orca (Murray, 1913)

Trichocerca plaka Myers, 1938

Trichocerca porcellus (Gosse, 1851)

Trichocerca pusilla (Jennings, 1903)

Trichocerca rattus (Müller, 1776)

Trichocerca relicta Donner, 1950

Trichocerca ripli Berzins, 1973

Trichocerca rosea (Stenroos, 1898)

Trichocerca rousseleti (Voigt, 1902)

Trichocerca ruttneri (Donner, 1953)

Trichocerca scipio (Gosse, 1886)

Trichocerca similis (Wierzejski, 1893)

Trichocerca stylata (Gosse, 1851)

Trichocerca sulcata (Jennings, 1894)

Trichocerca tenuior (Gosse, 1886)

Trichocerca tigris (Müller, 1786)

Trichocerca tigris f. monostyla Russell, 1951 E

Trichocerca tortuosa (Myers, 1936)

Trichocerca uncinata (Voigt, 1902)

Trichocerca vernalis (Hauer, 1936)

Trichocerca weberi (Jennings, 1903)

TRICHOTRIIDAE

Macrochaetus altamirai (Arevalo, 1918)

Macrochaetus collinsi (Gosse, 1867)

Trichotria pocillum (Müller, 1776)

Trichotria tetractis (Ehrenberg, 1830)

T. tetractis similis (Stenroos, 1898)

Trichotria truncata (Whitelegge, 1889)

SPECIES INQUIRENDAE

Names in the New Zealand literature that cannot

be related to known species, or possibly valid

species for which the description and/or ﬁgures are

inadequate under terms of the modern Interna-

tional Code of Zoological Nomenclature.

Callidina (=Macrotrachela) bihamata Gosse in

Hudson & Gosse, 1886 (in Hilgendorf 1902)

Callidina (=Macrotrachela) quadridens Hilgendorf,

1989 (in Hilgendorf 1898)

Cathypna (=Lecane) hudsoni Lord, 1890 (in

Hilgendorf 1902)

Dicranophorus n. sp. (in Parr 1949)

Encentrum n. sp. (in Parr 1949)

Epiphanes n.sp. (in Parr 1949)

Keratella serrulata (Ehrenberg, 1838) (in Russell

1961). Possibly a misinterpretation of a comment

in Brehm (1928). ?Not known from New Zealand.

Kellicottia longispina (Kellicott, 1879) (in Duggan

et al. 2002b). Possibly a contaminant from a

Northern Hemisphere collection – the taxon is

absent from Australasia .

Notommata pentophthalma Hilgendorf, 1898

Philodina cloacata Hilgendorf, 1902

Philodina microps Gosse, 1887 (in Hilgendorf 1902)

Planoventer varicolor Hilgendforf, 1898

Proales n. sp. (in Parr 1949)

Trichocerca cimolius (Gosse, 1886) (in Hilgendorf

(1902)

The following lecanids were recorded and ﬁgured

in C. R. Russell’s laboratory notes (Canterbury

Museum). According to Hendrik Segers, who

revised the Lecanidae globally (Segers 1995) and

subsequently checked a copy of Russell’s notes, the

New Zealand taxa are not the nominate species:

Lecane lauterborni Hauer, 1924

Lecane opias (Harring & Myers, 1926)

Lecane tenuiseta Harring, 1914

Lecane rugosa (Harring, 1914)

Lecane verecunda Harring & Myers, 1926

Class ACANTHOCEPHALA

Marine Acanthocephala in the following list

pertain to the New Zealand EEZ but also include

records from subantarctic waters where the species

almost certainly range into the EEZ. Records

of larval forms from intermediate hosts are not

included.

M = mammal; B = bird; F = ﬁsh; Ma = marine;

Es = estuarine; Fw = freshwater; T = terrestrial.

Hosts are named in square brackets after the

parasite name.

Subclass ARCHIACANTHOCEPHALA

Order OLIGOACANTHORHYNCHIDA

OLIGOACANTHORHYNCHIDAE

Macracanthorhynchus hirudinaceus (Pallas, 1781)

M T [Sus scrofa, Mustela furo]

Subclass PALAEACANTHOCEPHALA

Order ECHINORHYNCHIDAE

ECHINORHYNCHIDAE

Acanthocephalus galaxii Hine, 1977 F Fw E

[Anguilla australis, A. dieffenbachii, Galaxias

argenteus, G. maculatus, Gobiomorphus cotidianus,

Retropinna retropinna]

Acanthocephalus sp. F Fw

[Anguillla sp., Gobiomorphus sp.]

156

THE NEW ZEALAND INVENTORY OF BIODIVERSITY

Echinorhynchus gadi Zoega in Mueller, 1776 F Ma

[Macruronus novaezelandiae]

Echinorhynchus sp. F Ma

[Nemadactylus macropterus, Hoplostethus

atlanticus]

HETERACANTHOCEPHALIDAE

Aspersentis peltorhamphi (Baylis, 1944) F Ma

[Peltorhamphus novaezeelandiae, Rhombosolea

leporina, R. plebeia]

RHADINORHYNCHIDAE

Australorhynchus tetramorphacanthus Lebedev, 1967

F Ma

[Pagrus auratus, Seriola lalandi, Trachurus

novaezelandiae]

Micracanthorhynchina hemirhamphi (Baylis, 1944)

F Ma

[Hyporhamphus ihi]

Raorhynchus terebra (Rudolphi, 1819) F Ma

[Katsuwonus pelamis]

Rhadinorhynchus sp. F Ma

[Thunnus alalunga]

Order POLYMORPHIDA

PLAGIORHYNCHIDAE

Plagiorhynchus allisonae Smales, 2002 B Ma

[Haematopus ﬁnschi, H. unicolor]

POLYMORPHIDAE

Bolbosoma balaena (Gmelin, 1790) M Ma

[Balaenoptera borealis]

Bolbosoma physeteris Gubanov, 1952 M Ma

[Physeter macrocephalus]

Bolbosoma turbinella (Diesing, 1851) M Ma

[Balaenoptera borealis]

Bolbosoma physeteris Gubanov, 1952 M Ma

[Physeter macrocephalus]

Bolbosoma vasculosum (Rudolphi, 1819) M Ma

[Tursiops truncatus]

Corynosoma australe Johnston & Best, 1937 M Ma

[Arctocephalus forsteri, Phocarctos hookeri,

Hydrurga leptonyx]

Corynosoma bullosum (Linstow, 1892) M Ma

[Physeter macrocephalus, Mirounga leonina]

Corynosoma clavatum Goss, 1941 B Ma

[Leucocarbo colensoi ]

Corynosoma semerme (Forssell, 1904) M Ma

[Phocarctos hookeri]

Corynosoma hannae Zdzitowiecki, 1984 M Ma

[Arctocephalus forsteri, Megadyptes antipodes]

Corynosoma pseudohammani Zdzitowiecki, 1984

M Ma

[Leptonychotes weddellii]

Corynosoma semerme (Forssell, 1904) M Ma

[Phocarctos hookeri]

Corynosoma shackletoni Zdzitowiecki, 1985 B Ma

[Pygoscelis papua]

Corynosoma spp. B M Ma

Corynosoma shackletoni Zdzitowiecki, 1985 B Ma

[Pygoscelis papua, Megadyptes antipodes,

Phocarctos forsteri, Hydrurga leptonyx]

Polymorphus sp. B Fw

[Anas sp., A. platyrhynchos, A. superciliosa]

Proﬁlicollis antarcticus Zdzitowiecki, 1985 B Ma

[Haematopus ﬁnschi, Limosa lapponica]

Proﬁlicollis novaezelandensis Brockerhoff & Smales,

2002 B Ma E

[Haematopus ﬁnschi, Limosa lapponica]

Subclass EOACANTHOCEPHALA

Order NEOECHINORHYNCHIDA

NEOECHINORHYNCHIDAE

Neoechinorhynchus aldrichettae Edmonds, 1971 F

Ma/Fw

[Aldrichetta forsteri, Anguilla australis]

Neoechinorhynchus chilkaensis Podder, 1937 F Ma

[Aldrichetta forsteri, Arripis trutta, Contusus

richei]

Neoechinorhynchus sp. F Ma

[Latridopsis ciliaris]

SPECIES INQUIRENDAE

Echinorhynchus [s.l.] B T E

[Apteryx mantelli]

Pomphorhynchus turbinella Leiper & Atkinson, 1915

M Ma

Megaptera novaeangliae]

Genera et spp. indet. F B M Ma Fw T

[Neochanna apoda, N. diversus, Gobiomorphus

huttoni, Retropinna retropinna, Mustela furo,

Haematopus ﬁnschii, Limosa lapponica]

Gaining insights into the ecological role of the New Zealand sole (Peltorhamphus novaezeelandiae) through parasites

Article

Full-text available

Apr 2017
J HELMINTHOL

Despite tapeworms comprising the bulk of parasite communities of sharks in marine ecosystems, little is known about their life cycles and more specifically about the potential intermediate hosts they utilise as transmission routes. In the absence of morphological features required for specific identifications of larval tapeworms from potential intermediate hosts, recent molecular advances have contributed to linking larval and adult parasites and, in some instances, uncovering unknown trophic links. Host-parasite checklists are often the first source of information consulted to assess the diversity and host specificity of parasites, and provide insights into parasite identification. However, these host-parasite checklists are only useful if encompassing the full spectrum of associations between hosts and parasites. A checklist of New Zealand fishes and their parasites has been published, but recent parasitological examinations of commercial fish species reveal that the checklist appears to be far from complete. We focused our current study on a comprehensive survey of macroparasites of a commercial species, the New Zealand sole (Peltorhamphus novaezeelandiae) off the coast of Otago, New Zealand. Specifically, we were expecting to recover marine tapeworms using sharks as their definitive hosts that are generally underreported in parasite surveys. Surprisingly, a large proportion of non-tapeworm parasites we recovered were not previoulsy reported from this fish species, including tapeworms, flukes, round worms, and thorny-headed worms. A discussion on the potential ecological roles played by this fish species in the transmission of parasites is included.

Novel morphological and molecular data for Corynosoma hannae Zdzitowiecki, 1984 (Acanthocephala: Polymorphidae) from teleosts, fish-eating birds and pinnipeds from New Zealand

Article

Oct 2016

Are zooplankton invasions in constructed waters facilitated by simple communities?

Article

Samantha Maree Parkes

The invasion of non-indigenous species is considered to be one of the leading causes of biodiversity loss globally. My research aimed to determine if constructed water bodies (e.g., water supply reservoirs, dams and ponds) were invaded by zooplankton with greater ease than natural water bodies, and whether this was due to a lower biodiversity, and therefore lower 'biotic resistance', in constructed water bodies. Sediment cores were collected from a cross-section of 46 lakes, ponds and reservoirs (23 natural and 23 constructed) throughout the North Island, New Zealand. Diapausing zooplankton eggs were separated from the sediments and hatched to assess species composition and richness. In addition, the distributions of non-indigenous zooplankton were examined to determine if they occurred more frequently in constructed water bodies than in natural ones. Species composition results showed that natural water body zooplankton communities appeared to consist mainly of a core group of truly planktonic species. However, the species assemblages of constructed water bodies were more varied, comprising of a number of littoral and benthic species, and a large number of species that were recorded from only a single water body. A canonical correspondence analysis indicated that Trophic Level Index explained a significant amount of variation in zooplankton community composition of natural waters (p = 0.002). Distance to nearest water body and number of water bodies within a 20 km radius explained significant amounts of variation in community composition of constructed water bodies (p = 0.040 and 0.038 respectively). Average species richness was slightly higher for natural water bodies than constructed water bodies (18.47 and 15.05 respectively), although overall there was a lot of variation for both natural and constructed water body datasets. A stepwise linear regression indicated that latitude and approximate maximum depth of water body were significant predictors of natural water body species richness (p = 0.002 and 0.016 respectively). However, no significant predictors of species richness were elucidated for constructed water bodies. The non-indigenous calanoid copepods Sinodiaptomus valkanovi and Boeckella minuta were only found in constructed water bodies. However, the non-indigenous cladoceran Daphnia galeata was recorded in both natural and constructed water bodies. The non-indigenous calanoid copepods are more likely to establish populations in constructed water bodies due to the absence of key species (i.e. native calanoid copepods), whose presence in natural waters seemingly provides 'biotic resistance'. The invasion success of D. galeata in constructed and natural waters may be attributed to the absence of a superior competitor, as native Daphnia populations, for example, are rare in the North Island. My results suggest that species richness may not be as important as species composition in influencing the ease with which non-indigenous species invade constructed water bodies. The core group of species found in natural water bodies are likely to be better adapted to pelagic conditions, and therefore better at resisting invaders, than the more varied constructed water body assemblages.

Life on the farm: are zooplankton communities in natural ponds and constructed dams the same?

Article

Full-text available

Jul 2020

Constructed waters (e.g. dams and retired quarries) are commonly found to have a different zooplankton composition than are natural waters, and to be more readily invaded by non-indigenous species. Constructed ponds are common on farmland, but zooplankton research in these areas is scarce. Consequently, our aims were to (1) compare zooplankton communities between natural ponds and dams in rural environments and, (2) examine environmental determinants of zooplankton community composition among rural ponds. Thirty-eight ponds on farmland in New Zealand were sampled for zooplankton in winter–spring 2018 and summer 2019. All ponds were eutrophic, and zooplankton taxa typical of such conditions were common in both pond types (e.g. Brachionus, Keratella and Polyarthra species). Zooplankton community composition differed statistically between each type, although we deemed this difference to be ecologically insignificant (one-way ANOSIM, r = 0.09, P = 0.014). Prevalence of non-indigenous species was low, with 7% of farm dams and 2% of natural ponds being invaded, indicating that farm ponds are not acting as ‘stepping-stones’ for invaders across landscapes. Macrophyte abundance and concentration of humic substances were the dominant environmental variables measured determining zooplankton distribution among ponds.

Grass carp (Ctenopharyngodon idella) translocations, including hitchhiker introductions, alter zooplankton communities in receiving ponds

Article

Full-text available

Jun 2017

In the present study we tested the effects of translocations from aquaculture facilities of grass carp, one of the most commonly used species in aquaculture globally, to constructed ponds in the Auckland region, New Zealand. Primarily, we were interested in whether zooplankton assemblages in recipient ponds are affected by the concomitant introduction of ‘hitchhikers’ with fish releases. Zooplankton community composition was quantified in 34 ponds that had been subject to grass carp release and 31 that had no grass carp introductions. A significant difference in zooplankton community composition was observed between ponds that had received grass carp translocations and those that had not. Differences in community composition between ponds with and without carp releases could be attributed to both the: (1) effects of activity of grass carp through habitat modification; and (2) establishment of hitchhiking zooplankton species originating from aquaculture ponds, including non-native species. Effective measures to curb the proliferation of non-native taxa within aquaculture facilities, and to mitigate the accidental movement of non-native taxa with translocations from these facilities, are required to reduce future zooplankton introductions.

Are zooplankton invasions in constructed waters facilitated by simple communities?

Article

Dec 2012

Aim Constructed water bodies (e.g. water supply and hydroelectricity dams, ornamental ponds) are invaded at faster rates than natural waters, but the mechanisms that lead to this pattern are uncertain. We aimed to determine whether constructed lakes have lower zooplankton species richness or differ in species composition relative to natural waters, which might allow them to be invaded more readily. Location N orth I sland, N ew Z ealand. Methods Sediment cores were collected from 23 natural and 23 constructed lakes, ponds and reservoirs. Zooplankton diapausing eggs were hatched from sediments to assess species composition and richness. Results Average species richness between natural and constructed water bodies was not significantly different. Stepwise linear regression indicated latitude and maximum depth were significant predictors of species richness in natural waters ( P = 0.002 and 0.016, respectively). No significant predictors were elucidated for constructed waters. Species in natural waters consisted mainly of planktonic species, while assemblages in constructed waters were more varied, comprising of more littoral and benthic species, and included many species recorded from only single water bodies. Canonical correspondence analysis indicated that trophic state explained the greatest proportion of variation in species composition of natural waters ( P = 0.002). Distance to nearest water body and number of water bodies within a 20 km radius (i.e. opportunity) explained the greatest proportion in constructed waters ( P = 0.040 and 0.038, respectively). Main conclusions Natural and constructed waters had similar species richness per water body, but community composition varied between them. The core group of species found in natural waters are better adapted to pelagic conditions, and may therefore be better at reducing establishment rates of new arrivals than assemblages in constructed water bodies, which had more varied assemblages. Our study suggests that manipulating new water bodies, so that they develop mature communities faster, may reduce establishment rates of non‐indigenous species.

The freshwater aquarium trade as a vector for incidental invertebrate fauna

Article

Aug 2010

Ian C. Duggan

The aquarium trade has a long history of transporting and introducing fish, plants and snails into regions where they are not native. However, other than snails, research on species carried “incidentally” rather than deliberately by this industry is lacking. I sampled invertebrates in the plankton, and from water among bottom stones, of 55 aquaria from 43 New Zealand households. I recorded 55 incidental invertebrate taxa, including copepods, ostracods, cladocerans, molluscs, mites, flatworms and nematodes. Six were known established non-indigenous species, and eight others were not previously recorded from New Zealand. Of the latter, two harpacticoid copepod species, Nitokra pietschmanni and Elaphoidella sewelli, are not native to or known from New Zealand, demonstrating the aquarium trade continues to pose an invasion risk for incidental fauna. The remaining six species were littoral/benthic rotifers with subtropical/tropical affinities; these may or may not be native, as research on this group is limited. A variety of behaviours associated with the set-up and keeping of home aquaria were recorded (e.g., fish and plants in any home were sourced from stores, wild caught, or both, and cleaning methods varied), which made prediction of “high risk” behaviours difficult. However, non-indigenous species had a greater probability of being recorded in aquaria containing aquatic plants and in those that were heated. Methods for disposal of aquarium wastes ranged from depositing washings on the lawn or garden (a low risk for invasion) to disposing of water into outdoor ponds or storm-water drains (a higher risk). It is recommended that aquarium owners be encouraged to pour aquarium wastes onto gardens or lawns—already a common method of disposal—as invasion risk will be minimised using this method.

Phylum Rotifera

Chapter

Dec 2016

The second volume in this landmark series includes an introductory chapter followed by 15 taxonomically specific chapters with identification keys to Nearctic freshwater invertebrates (protozoa through arthropods). Other than the second chapter's treatment of the diverse, multi-phyla array of protozoa (ciliates, flagellates, and amoebas), each of the remaining 14 chapters is limited to a single phylum. Chapters include the following six sections: Introduction, Limitations (to taxonomic identification of that group), Terminology and Morphology (information needed to use the keys), Material Preparation and Preservation, Identification Keys, and References. This volume complements Volume I's global coverage of “Ecology and General Biology,” and thus, only minimal information on those topics is included here. Volume II is devoted to the identification of inland water invertebrates of the Nearctic Bioregion including Canada, the continental United States, Mexico north of the Tropic of Cancer, Bermuda, and Greenland. Most keys terminate at either the level of genus or species, and generalized distribution data is included.

An immature polymorphid acanthocephalan from a European hedgehog (Erinaceidae) from New Zealand

Article

Full-text available

Jun 2010

This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

Field evidence of the impact of two acanthocephalan parasites on the mortality of three species of New Zealand shore crabs (Brachyura)

Article

Full-text available

Sep 2002

The effects of the acanthocephalan parasites Profilicollis antarcticus and P. novaezelandensis on the fecundity and mortality of three species of shore crab (Macrophthalmus hirtipes, Hemigrapsus edwardsi and H. crenulatus) are examined. The number of eggs produced by female crabs was strongly correlated (all P<0.05) with carapace width; parasite load was not a significant determinant of female fecundity. Mortality was inferred from reduction in the mean number of parasites per crab in the largest crab size-classes, indicating that heavily infected individuals are removed from the population. Mortality attributable to the parasites was observed for all three species of crabs, although the effect of parasites varied in both time and space; significant curvilinear regressions between parasite load and crab size-classes were not found in all samples. Crab mortality appears to be influenced by more than the pathological influences exerted by the parasites. Parasite-induced behavioural alterations may cause crab hosts to be more susceptible to predation by definitive hosts. We support this suggestion with three lines of evidence: the lack of parasite effects on fecundity, the weakening of the parasite effect on mortality during the time of year when birds are absent, and previous indications of parasite-mediated alterations in crab burrowing behaviour.

Seasonal abundance and diel vertical dynamics of zooplankton in Lake Okaro

Article