Synopsis of the Nematoda Parasitic in Amphibians and Reptiles

332 Pages·1987·59.54 MB·English

by  Michael R Baker

ial University
SYNOPSIS OF THE NEMATODA PARASITIC
IN AMPHIBIANS AND REPTILES
Michael R, Baker
No. 11 1,987
MEMORIAL UNIVERSITY OF NEWFOUNDLAND
OCCASIONAL PAPERS IN BIOLOGY
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Contents
Baker, M.R. 1987. Synopsis of the Nematoda Parasitic in Amphibians and Reptiles.
SYNOPSIS OF THE NEMATODA
PARASITIC IN AMPHIBIANS AND REPTILES
M. R e BAKER
Department of Zoology,
University of Guelph,
Guelph, Ontario, Canada N1G 1C1
March, 1987

CONTENTS
....................................................
INTRODUCTION 4
.................................................
ACKNOWLEDGEMENT 4
..............................................
THE PARASITE FAUNA 5
.......................................
LIST OF TAXONOMIC CHANGES 10
- ................................................
ORDER TYLENCHIDA 12
MYENCHOIDEA ..................................................
Myenchidae 12
- ..................................................
ORDER ENOPLIDA 12
TR ICH UROIDEA .................................................
Trichuridae ................................................ 12
Cystoopsidae 15
DIOCTOPHYMATOIDEA ............................................
Dioctophymatidae 16
.................................................
Rhabditidae .......................................... 17
Angiostomatid.a.e. ............................................. 17
Rhabdiasidae ........................................ 18
Strongyloididae 23
DIPLOGASTEROIDEA ...........................................
Cylidrocorporidae 24
..................................................
-
ORDER OXYURIDA 25
OXYUROIDEA .............................................
Pharyngodonidae 25
-
ORDER ASCARIDIDA e............. ..................... u . . 51
COSMOCERCOIDEA ...............................................
Cosmocercidae ............................................. 51
Kathlaniidae. .................................*................ 67
Atractidae 77
HETERAKOIDEA .................................................
Heterakidae ................................................ 84
Ascaridiidae 91
SEURATOIDEA ..................................................
Seuratidae ........................................ 93
Chitwoodchaba.u.d.i.id.a.e. ......................................... 94
Cucullanidae ......................................... 94
Quimperiidae 95
ASCARIDOIDEA .................................................
Ascarididae 99
- ................................................
ORDER SPIRURIDA a114
CAMALLANOIDEA ................................................
Camallanidae 114
DRACUNC ULO I DEA ...............................................
Dracunculidae 119
..............................................
Micropleuridae 120
GNATHOSTOMATOIDEA ............................................
Gnathostomatidae 121
PHYSALOPTEROIDEA .............................................
physalopteridae 124
SPIRUROIDEA ..................................................
Spiruridae 139
THELAZ IOIDEA ..............................................
Rhabdochonidae 139
HABRONEMATOIDEA ..............................................
Cystidicolid.a.e. ...............................................1 40
Hedruridae ................................................1 40
Tetrameridae 142
DIPLOTRIAENOIDEA .............................................
Diplotriaenidae 143
FILARIOIDEA ...............................................
Onchocercidae 145
...............................................
ORDER STRONGYLIDA 157
STRONGYLOIDEA ................................................
Strongylidae 157
DIAPHANOCEPHALOIDEA ..........................................
Diaphanocephalidae 159
TRICHOSTRONGYLOIDEA ............................................
Amphibiophilidae. .............................................1 69
Dictyocaulidae .........................................1 69
Herpetostrm.g.y.l.i.d.a.e. ..........................................1 70
Molineidae ................................................ 170
Nicollinidae 177
...................................
INDEX TO GENERA AND SUBGENERA 178
HOST-PARASITE .L.I.S.T. ...............................................
AMPHIBIANS. .....................................................2 15
REPTILES 229
....................................................
BIBLIOGRAPHY 258
INTRODUCTION
A modern classification of vertebrate nematode parasites to the level
of genus is available in the CIH Keys to the Nematode Parasites of
Vertebrates (1974-83). However there is a need for up to date summaries of
species since previous compendia are out of date (e.g. Yorke & Maplestone
of
1926; Yamaguti 1961; ~ssentials Nematodology series edited by ~krjabin
and colleagues). This is especially true for the nematode parasites of the
lower terrestrial vertebrates which have historically received relatively
l i t t l e attention from zoqlogists. The present study aims to f i l l this need
for the nematode parasites of amphibians and reptiles. In addition,
published records have been brought together to give summaries of host and
geographical distributions of individual genera and species.
Despite the fact that the present work is essentially a compendium, a
number of taxonomic changes have been introduced. In most cases the reasons
are technical. An example would be the discovery of a hitherto unreported
homomymy.
Each nematode order has been treated separately, with its own
index to genera, subgenera, species and subspecies. Only adult worms have
been included. Recent revisions are indicated for each nematode family and
genus. For each genus or subgenus a numbered list of species is given
starting with the type species. For each species a list of synonyms is given
which is restricted to the first generic combination in which a species
epithet was published. For example, Stronqylus bialata Molin, 1861, is
listed as a synonym of Oswaldocruzia filiformis, but Oswaldocruzia bialata
(Molin, 1861) Travassos, 1917 is not listed. A literature reference for each
synonymy is also given. This is followed by a listing of the host(s) from
which the type specimens were taken (see "ex"), geographical locality, and
any pertinent note concerning taxonomy or biology. Subsequent literature
records (hosts, locality, literature reference) £01l ow (see "Rec"). For some
-
frequently reported species (e.g. Aplectana acuminata) hosts, localities and
references are grouped together. Also some early reports are not listed if
these have been reviewed in more recent papers. For frequently reported
species, i.e. nematodes of Western mropean amphibians and reptiles, this
has saved much duplication in the listing of records.
An attempt was made to gather complete information on each species.
Original references were in many cases examined. Otherwise, the
of and
Helminthological Abstracts, Index Catalogue Medical Veterinary
Zoology, and the Zoological Record were found to be particularly useful. It
is inevitable that given the limitations of these sources, there w i l l be
some omissions. The bib1i ography covers on1y references published after
of
1900. Earlier papers are listed in the Index Catalogue Medical
Veterinary Zoology.
For the identification of hosts, only the most obvious errors have been
pointed out. Many older reports were based on obsolete classifications of
amphibians and reptiles. In addition, the taxonomy of many amphibian and
reptilian groups is still far from stable. Despite these problems
identification to the level of host family has been generally reliable.
ACKNOWLEDGEMENT
This study was supported by funds made available through the Department
of Zoology, and the Dean's Office, College of Biological Sciences,
University of Guelph.

THE PARASITE FAUNA
The nematode parasite fauna of amphibians and reptiles contains
representatives from seven orders. However, study of host distributions a t
the family and superfamily level (Table 1) reveals only four families and
one superfamily (Diaphanocephaloidea) which are unique to the amphibians and
reptiles, whereas 36 families are shared with other vertebrates (and rarely
invertebrates) or are mainly free-living forms. The only superfamilies
reported from vertebrates which are unknown (as adults) in amphibians and
reptiles are the secernentean superfamilies Subuluroidea and
Ancylostomatoidea, and the adenophorean superfamilies Dioctophymatoidea and
Muspiceoidea. This lack of uniqueness is probably the result of parasite
exchange between vertebrate groups, an apparently common theme in nematode
evolution (Anderson 1984). This does not mean that host-parasite coevolution
has not occurred. In fact a t taxonomic levels below the family, evidence for
coevolution is abundant. Chabaud (1981) has summarized an analytical method
by which correlation of parasite morphological evolution, host distribution
and evolution, and zoogeography, often reveals a probable host group in
which a nematode group evolved. This method has been applied to the nematode
parasites of amphibians and reptiles (Baker 1984) and nematode groups
reported from these hosts which were acquired by transfer from other hosts
(Table 2) have been distinguished from those which probably evolved in
amphibians and reptiles (Table 3). A few groups occur widely and their
original host groups cannot be distinguished (Table 4). Details of the
analysis were given earlier (Baker 1984).
In the evolutionary history of the four largest orders of nematode
parasites of vertebrates (Oxyurida, Strongylida, Ascaridida and Spirurida),
amphibians and reptiles have played important roles (see Baker 1984).
1) Oxyurida: Oxyuridans probably evolved in the ~rthropodab efore they were
acquired by vertebrates. Representatives in amphibians and reptiles,
however, are morphologically closer to the primitive pin worms of arthropods
than are the two families occurring in mammals (and rarely birds).
2) Strongylida: The greatest diversity of superfamilies, families and genera
and the most primitive representatives (Strongyloidea) occur mainly in
mammals. Nevertheless, amphibians and reptiles have played an important role
in the early evolution of some families of trichostrongyles
(Amphibiophil idae, Molineidae, Dictyocaul idae) and one superf amily
(Diaphanocephaloidea) has evolved in modern snakes.
3 ) Ascaridida: The ascarididans probably evolved in the amphibians and
reptiles and these vertebrates still harbour an impressive variety of
Cosmocercoidea, Seuratoidea, Heterakoidea and Ascaridoidea. The
cosmocercoids, which are morphologically the most primitive ascarididans,
are especially w e l l represented in a wide variety of amphibians and
reptiles.
4) Spirurida: Mammals and birds are the main host groups. The origin of the
Spirurida is complicated by the fact that a t least some superfamilies
apparently evolved independantly from ancestral Ascaridida. The spiruridans
are relative1y poor1y represented in the amphibians and reptiles, a1t hough
in individual superfamilies a few distinct genera, subfamilies or rarely
families have evolved in them.
*
Table 1. Systematic characteristics of the superfamilies of nematode
parasites occurring as adults in amphibians and reptiles. Numbers
in parenthesis in "Families" column refer to total number of
families in superfamil y. Similarly, numbers in parenthesis in
"Genera" column refer to tota1,number of genera in superfamily.
........................................................................................................................................................
Families
Families Genera Unique Shared
Adenophorea
Enopl ida
Trichinelloidea
Secernentea
Tylenchida
Myenchoidea
Rhabditida
Rhabditoidea
Diplogasteroidea
Oxyur ida
Oxyuroidea
Ascaridida
Cosmocercoidea
Heterakoidea
Seuratoidea
Ascaridoidea
Spirurida
Camallanoidea
Dracunculoidea
hathostomatoidea
Physalopteroidea
Spiruroidea
Thelazioidea
Habronematoidea
Diplotriaenoidea
Filarioidea
Strongylida
Strongyloidea
Diaphanocephaloidea
Trichostrongyloidea
Total
* Information on on free-living forms from following reference: Goodey, J.B.
1963. Soil and freshwater nematodes. 2nd ed., Methuen & Co., New York, 544
P