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



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.
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
lesser jaw worms, rotifers, thorny-headed worms
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
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
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
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
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.
basal plate
epidermal cells
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.
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
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
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
(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.
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
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.
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
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)
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.
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
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.
adult female
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
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
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.
Dr Ian C. Duggan Department of Biological Sciences, Waikato University,
Private Bag 3105, Hamilton, New Zealand []
Dr John D. Green 532 Mooloo Road, Mooloo via Gympie, QLD 4570, Australia
Dr Sylvie Pichelin Department of Microbiology and Parasitology, University of
Queensland, Brisbane, QLD 4072, Australia []
Dr Russell J. Shiel Department of Environmental Biology, The University of
Adelaide, Adelaide, South Australia 5005, Australia []
Professor Lesley Smales School of Biological and Environmental Sciences, Central
Queensland University, Rockhampton, QLD 4702 Australia []
Dr Wolfgang Sterrer Bermuda Natural History Museum, PO Box FL 145,
Flatts FLBX, Bermuda []
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.
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Habrotrocha collaris (Ehrenberg, 1832)
Habrotrocha constricta (Dujardin, 1841)
Habrotrocha crenata Murray, 1905
Habrotrocha crenata var. sphagnicola Pavlovski,
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
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,
Macrotrachela quadricornifera var. Haigh, 1963 E
Macrotrachela quadricornifera Bryce, no date,
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:
A.; TIETZ, D. 1998: Phylogeny of the Metazoa
based on morphological and ribosomal DNA
evidence. Cladistics 14: 249–285.
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)
Cupelopagis vorax (Leidy, 1857)
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)
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
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)
Hexarthra brandorffi Koste, 1977
Hexarthra fennica (Levander, 1892)
Hexarthra intermedia (Wiszniewski, 1929)
Hexarthra mira (Hudson, 1871)
Hexarthra propinqua (Bartos, 1948)
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)
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)
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)
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,
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)
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)
Epiphanes brachionus (Ehrenberg, 1837)
Epiphanes macrourus (Barrois & Daday, 1894)
Epiphanes senta (Müller, 1773)
Proalides tentaculatus De Beauchamp, 1907
Rhinoglena frontalis Ehrenberg, 1853
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)
Ascomorpha ecaudis (Perty, 1850)
Ascomorpha ovalis (Bergendal, 1892)
Ascomorpha saltans Bartsch, 1870
Gastropus hyptopus (Ehrenberg, 1838)
Gastropus minor (Rousselet, 1892)
Itura aurita Wulfert, 1935
Itura myersi Wulfert, 1935
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)
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)
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)
Lindia euchromatica Edmondson, 1938
Lindia fulva Harring & Myers, 1922
Lindia pallida Harring & Myers, 1922
Lindia torulosa Dujardin, 1841
Lindia truncata (Jennings, 1894)
Microcodon clavus Ehrenberg, 1830
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)
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
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
Scaridium longicaudum (Müller, 1786)
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
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)
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)
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
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
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
M = mammal; B = bird; F = fish; Ma = marine;
Es = estuarine; Fw = freshwater; T = terrestrial.
Hosts are named in square brackets after the
parasite name.
Macracanthorhynchus hirudinaceus (Pallas, 1781)
M T [Sus scrofa, Mustela furo]
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.]
Echinorhynchus gadi Zoega in Mueller, 1776 F Ma
[Macruronus novaezelandiae]
Echinorhynchus sp. F Ma
[Nemadactylus macropterus, Hoplostethus
Aspersentis peltorhamphi (Baylis, 1944) F Ma
[Peltorhamphus novaezeelandiae, Rhombosolea
leporina, R. plebeia]
Australorhynchus tetramorphacanthus Lebedev, 1967
F Ma
[Pagrus auratus, Seriola lalandi, Trachurus
Micracanthorhynchina hemirhamphi (Baylis, 1944)
F Ma
[Hyporhamphus ihi]
Raorhynchus terebra (Rudolphi, 1819) F Ma
[Katsuwonus pelamis]
Rhadinorhynchus sp. F Ma
[Thunnus alalunga]
Plagiorhynchus allisonae Smales, 2002 B Ma
[Haematopus finschi, H. unicolor]
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]
Neoechinorhynchus aldrichettae Edmonds, 1971 F
[Aldrichetta forsteri, Anguilla australis]
Neoechinorhynchus chilkaensis Podder, 1937 F Ma
[Aldrichetta forsteri, Arripis trutta, Contusus
Neoechinorhynchus sp. F Ma
[Latridopsis ciliaris]
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]
... Clearly, NZ sole is not their definitive host. As a matter of fact, adult specimens have been recovered from the large intestine of Hydrurga leptonyx (leopard seal), Arctophoca forsteri (long-nosed fur seal) and Phocarctos hookeri (NZ sea lion) (Zdzitowiecki, 1984;Shiel et al., 2009;Hernández-Orts et al., 2017) from Antarctica and NZ, while immature specimens have been recovered from the fisheating birds Leucocarbo chalconotus (Stewart Island shag), Phalacrocorax punctatus (spotted shag) and Megadyptes antipodes (yellow-eyed penguins) (Shiel et al., 2009;Hernández-Orts et al., 2017) from NZ. Pleuronectid fish are considered to be paratenic hosts for this parasite and cystacanths have been reported from our sampling of NZ sole and Colistium guntheri (NZ brill) (this study, but see Hernández-Orts et al., 2017). It is believed that the life cycle for C. hannae might involve amphipods, with teleosts being necessary for concentrating the cystacanths for transmission up the food chain to pinnipeds, but in which no parasite development occurs (Zdzitowiecki & Presler, 2001). ...
... Clearly, NZ sole is not their definitive host. As a matter of fact, adult specimens have been recovered from the large intestine of Hydrurga leptonyx (leopard seal), Arctophoca forsteri (long-nosed fur seal) and Phocarctos hookeri (NZ sea lion) (Zdzitowiecki, 1984;Shiel et al., 2009;Hernández-Orts et al., 2017) from Antarctica and NZ, while immature specimens have been recovered from the fisheating birds Leucocarbo chalconotus (Stewart Island shag), Phalacrocorax punctatus (spotted shag) and Megadyptes antipodes (yellow-eyed penguins) (Shiel et al., 2009;Hernández-Orts et al., 2017) from NZ. Pleuronectid fish are considered to be paratenic hosts for this parasite and cystacanths have been reported from our sampling of NZ sole and Colistium guntheri (NZ brill) (this study, but see Hernández-Orts et al., 2017). It is believed that the life cycle for C. hannae might involve amphipods, with teleosts being necessary for concentrating the cystacanths for transmission up the food chain to pinnipeds, but in which no parasite development occurs (Zdzitowiecki & Presler, 2001). ...
... The other acanthocephalan species, A. peltorhamphi, has been reported from the NZ sole previously and is included in the checklist of Hine et al. (2000). The NZ sole is indeed one of the definitive hosts for this parasite as it has also been reported from the pleuronectids Rhombosolea leporina T. Anglade and H.S. Randhawa (yellowbelly flounder) and R. plebeia (NZ flounder) (Shiel et al., 2009). The A. peltorhamphi cystacanths are likely acquired by the teleost fish feeding on infected amphipods (Zdzitowiecki & Presler, 2001), although this has yet to be determined empirically for NZ pleuronectid fish. ...
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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.
... (Hector) and pink cusk-eel, Genypterus blacodes (Forster) from the South Island of New Zealand and the Auckland and Campbell Islands. Shiel et al. [20] reported specimens of Corynosoma hannae in the long-nosed fur seal, Arctophoca forsteri (Lesson) (as Arctocephalus forsteri) and yellow-eyed penguin, Megadyptes antipodes (Hombron & Jacquinot) from St Clair Beach, Dunedin, and the Otago Peninsula, respectively. More recently, García-Varela et al. [21] provided molecular data apparently for C. australe from New Zealand sea lion, Phocarctos hookeri (Gray) from Enderby Island. ...
... References Zdzitowiecki [12,13]; Stryukov [40]; Shiel et al. [20]; Present study. ...
... Interestingly, Zdzitowiecki [13] designated the gravid female as the holotype which is not usual taxonomic practice. Later, C. hannae was reported by Stryukov [40] from the intestine of H. leptonyx from the Balleny Islands, D'Urville Sea, Antarctica, and by Shiel et al. [20] from A. forsteri and M. antipodes from New Zealand. The specimens from fish-eating birds, fur seal and sea lion from New Zealand in this study exhibit all the morphological features of C. hannae (i.e. ...
... A total of 71 species of zooplankton were found in natural and constructed water bodies throughout the North Island, New Zealand, during this study (Table 3). All species have been previously recorded in New Zealand, except for the rotifer Cephalodella stenroosi (Shiel et al., 2009 ...
... All of these species have been previously recorded in New Zealand, except for one. To date there has been no other record of the rotifer Cephalodella stenroosi in New Zealand (Shiel et al., 2009). Cephalodella stenroosi was identified based on its toes with a dorsal lump, not tooth, the absence of an eye spot, and the trophi were as figured in Nogrady et al. (1995). ...
... It may be that this species is a new arrival to New Zealand, but it is much more likely that it has just never been documented or identified. Although studies of zooplankton in New Zealand now span over 150 years, many of the early studies are of little use to ecologists as they lacked comprehensive figures and keys to species (Shiel et al., 2009). In addition to this, most rotifer collections have been from the open water of lakes and ponds, and therefore benthic and periphytic species are likely to be very poorly described (Duggan et al., 1998; Shiel et al., 2009). ...
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.
... 13.0) was similar (t-test, P ¼ 0.356). Three species were recorded that had not previously been identified in New Zealand; these were Erignatha clastopis and Octotrocha speciosa, for which neither genera had been recorded in New Zealand previously, with Cephalodella theodora also being recorded for the first time (Table 2; Shiel et al. 2009). A single individual of E. clastopis was found in one farm dam, whereas both C. theodora and O. speciosa were recorded in the same natural pond, with only single individuals being found of each also; based on distributions elsewhere, all are likely to be native. ...
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.
... Twenty-five taxa were present in single ponds only. With the exception of Microcodides robustus, present in McLennan Pond, all taxa have been recorded previously in New Zealand (Shiel et al. 2009). M. robustus has an almost cosmopolitan distribution globally, with the exclusion of Africa to date, but it is rarely encountered (Segers 2007). ...
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.
... A total of 71 zooplankton species were recorded (Table 1). All have been previously recorded in New Zealand, except the rotifer Cephalodella stenroosi (Shiel et al., 2009), which was hatched from a constructed pond in Whangamata. The raw average numbers of species per water body were 10.43 in natural and 9.22 in constructed water bodies, while the Chao2 nonparametric estimator indicated richness of 18.47 and 15.05, respectively. ...
AimConstructed 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. LocationNorth Island, New Zealand. 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. ResultsAverage 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 conclusionsNatural 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 common Asian native distribution of N. pietschmanni and E. sewelli is consistent with the predominant global regions for aquarium fish culture facilities and exports (Cheong 1996), which indicates the likely pathway for introduction. Some or all of the rotifer species not previously recorded may potentially be native to New Zealand, as species found in this study typically have broad distributions, while little research has been conducted in New Zealand of the littoral or benthic fauna (Shiel et al. 2009). However, of the four Lecane species recorded, L. nana, L. signifera and L. aculeata are noted by Segers (1995) to be more common in ''tropical and subtropical waters'', or ''warm waters''. ...
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.
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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.