Keywords: Capillariidae, Paracapillaria, Ophidiocapillaria, taxonomy, synonymy, snakes, Paracapillaria sonsinoi, P. mingazzinii, P. colubra, P. vipe rae, P. ptyasi ...
On the validity of the species considered synonyms of Paracapillaria sonsinoi (Parona, 1897) (Nematoda: Capillariidae)
1
Vulko Y . B l S E R K O V , Ferenc M É S Z Á R O S
institute
2
and Nesho C H I P E V
1
of Parasitology, Bulgarian Academy of Sciences, Acad. G. Bonchev Str., Bl. 25, 1113 Sofia, Bulgaria,
Zoological Department, Hungarian Natural History Museum, 1088, Budapest, Baross u. 13. Hungary (Received 15 June, 1994)
Abstract: The revision proposed by Moravec (1986) of the species in the genus Paracapillaria parasitizing snakes is discussed. On the basis of morpho logical and morphometric differences, host range, localization in the host and geographical distribution it is concluded that there are insufficient grounds for accepting the synonymy suggested by this author. The nematodes Para capillaria sonsinoi, P. mingazzinii, P. colubra, P. viperae, P. ptyasi, P. xochimilcensis and P. heterodontis are considered to be distinct species within the genus Paracapillaria at present. K e y w o r d s : Capillariidae, Paracapillaria, Ophidiocapillaria, taxonomy, synonymy, snakes, Paracapillaria sonsinoi, P. mingazzinii, P. colubra, P. vipe rae, P. ptyasi, P. xochimilcensis, P. heterodontis. INTRODUCTION Ten nematode species have been described within the genus Paracapillaria, parasi tizing mainly the digestive tract of poikilotherm vertebrates (Moravec, 1986). Moravec (1986) revised the genus Paracapillaria and proposed a new subgenus Ophidiocapillaria for species parasitizing snakes. He considered Paracapillaria (Ophidiocapillaria) sonsinoi (Parona, 1897) to be the type species of the new subgenus. Moravec (1986) regarded the following species as synonyms of P. sonsinoi: Trichosoma sonsinoi Parona, 1897, Trichosoma mingazzinii Rizzo, 1902, Capillaria heterodontis Harwood, 1932, C. xochimilcensis Cabalero et Cercero, 1943, Capillaria colubra Pence, 1970, Capillaria ptyasi Wang, 1982 and Paracapillaria viperae Biserkov, Georgievet Genov, 1985. In his monographical work Baker (1987) included T. sonsinoi, T. mingazzinii, C. heterodontis, C. xoehimücensis, C. colubra, C. ptyasi and P. viperae in the genus Capillaria as separate species.
Table 1. Comparison of the species of the genus Paracapillaria Mendonca, 1963
Species
P. mingazzinii P. colubra [Pen- P. viperae [Bi- P. ptyasi [Wong P. xochimilcensis P. heterodon
P. sonsinoi
P. sonsinoi
P. sonsinoi
[Skrjabin et
[Moravec
[Moravec
[Skrjabin et.
al. (1957)]
(1986)]
(1986)]
al. (1957)]
Locality Italy Italy France USA Host species C. viridiflavus N . rhombifera N . natrix N . maura Location intestine urin. bladder intestine rectum Female Body length 11.5-13.0 49.71-56.71 24.96-29.10 21.0-24.0 in mm Length of ant. 4.5 6.53-7.40 6.0 8.91-10.20 part in mm not elevated elevated not elevated elevated Vulvar lips 70-84 75-81 52-58 72-81 Egg length in n 26-30 Egg width in /j, 56 33-36 33-36 Ratio ant. : 1:1.8 1:4.5-1:4.6 1:3 1:2.8-1:2.9 post.part Male Body length 10.0-11.0 18.14-19.49 32.64-39.78 14.0-16.0 in mm Length of ant. 6.0 6.6 6.58-9.18 4.5 part in mm Spicule length 1.0 2.27-2.45 4.23-4.40 1.5-1.8 in mm about Ratio spicule: 1:11 1:7.9 1:7.7-1:9 1:9 whole length Spicular she short, smooth long, furrowed long, furrowed long, furro ath wed Ratio 1:0.66 1:1.7-1:1.9 1:3.3-1:3.9 1:2.5 ant.:post.part
ce (1970)]
serkov et al.
(1982)]
(1985)]
[Skrjabin et all.
tis [Pence
(1957)]
(1970)]
C. constrictor oviducts
Bulgaria V. ammodytes intestine
China P. mucosus intestine
Mexico T angustirostris intestine
USA H . contortrix rectum
29.2-36.5
23.0-25.0
10.2-11.9
13.6-13.8
24-26
5.8-6.9 elevated 61-70 28-31
7.5-8.5 elevated 70-80 27-36
5.0-5.2 -
4.5-4.8 elevated
63-72 28-32
-
7.4-8.4 elevated 63-66 25-30
1:4
1:2.0-1:2.1
1:1-1:1.3
1:1.9-1:2
1:2.1-1:2.2
19.6-26.4
19.0-21.0
9.9-12.4
10.0-10.1
16.5-22.5
5.5-6.9
7.5-8.7
5.0-5.4
3.8-4.2
7-11.5
2.2-2.9
2.24-2.46
0.92-1.12
1.46-1.60
2.55-3.25
1:9
1:7.9-1:8.6
1:10.7-1:11
1:6.3-1:6.8
1:6.5-1:6.9
-
-
long, non fur
1:0.98-1:1.4
1:1.4-1:1.6
1:0.96-1:1.4
USA
long, furrowed long, furrowed
rowed 1:2.7
1:1.3-1:1.6
DISCUSSION Having analyzed the morphometric characters from the descriptions of the different samples of P. sonsinoi presented by Moravec (1986), we assume that in his concept P. sonsinoi is a composite species. There are several grounds for such understanding: 1. The zoogeographical distribution of hosts. Relying on the similarity in the geographical distribution of European capillariid species and the close relationship of their hosts, Moravec (1986) has concluded that there is only one nematode species and that the observed morphological and metricdifferences should be regarded as intraspecific variation. However, the geographical isolation between the type localities of T. sonsinoi and T. mingazzinii, and the localities of the newly described materials should be considered. Moravec (1986) has redescribed P. sonsinoi using materials from Southern France (Bognas-Agde, Hérault) from Natrix maura (L.) and from the USA (Lousiana) from Natrix rombifera. The original description of P. sonsinoi made by Parona (1897) was from Italy (Pisa) from Zamenis viridifiavus Lacepede (=Coluberviridiflavus), and that of Trichosoma mingazzinii was from the island of Sicily from Tropidonotus natrix (L.) (=Natrix natrix). We are not convinced that the entire Southern Europe can be regarded as a geographically homogeneous territory. Between Pisa (Italy) and Southern France lay the mountain ranges of the Northern Apennines and the Western Alps. The island of Sicily is even more isolated. As for the species from North and South America and Eastern Asia, it is obvious that they originate from different zoogeographical regions. The close relations between the host species, put forward by Moravec (1986) to support his views, are in fact reduced to their belonging to one and the same family. However, these hosts belong to different genera (Coluber and Natrix), which have basically different habitats and lifestyles. Furthermore, P. viperae is a parasite of snakes which belong to another family (Viperidae). We consider as quite improbable the existence of only one species of capillariid nematode parasitizing hosts as ecologically diverse as the snakes belonging to the families Colubridae and Viperidae inhabiting, moreover, territories a great distance apart. We believe that there are strong enough reasons to expect that parasite speciation had occurred in these hosts as the result of their geographical and ecological isolation. 2. Localization in the host Capillaria colubra could be hardly synonymized with any of the other known species of the genus because of its specific localization. Pence (1970) stressed the fact that C. colubra was found only in the oviducts of Coluber constrictor and never in the host gut. The parasite has been established only in females and never in male hosts. Collins (1973) confirmed this site specificity of C. colubra in other hosts as well (Natrixspp.). 3. Morphometric characteristics. There are considerable morphological and mor phometric differences among the 9 descriptions (Table 1) regarded as synonyms of P. sonsinoi by Moravec (1986). Combinations of the taxonomically significant characteris tics of capillariids (e.g. spicule length, length and morphology of spicule sheath, morphology of vulvar lips, egg dimensions, the relations anterior/posterior body length and spicule length/body length) allow a clear differentiation of seven species. The closest similarity was observed between the characteristics of P. xochimilcensis and
P. heterodontis. The samples from France used in the redescription of P. sonsinoi by Moravec (1986) have morphometric characteristics similar to those of P. viperae and P. mingazzinii (Table 1). However, clear differences were found in the egg surface sculpture of P. xochimilcensis (few big plates with wide furrows) and of P. viperae (many small plates with narrow furrows). Finally, none of the descriptions of specimens from one locality (population) overlaps even half of the established range of variability in the taxonomic characters. In conclusion, we regard P. sonsinoi, P. mingazzinii, P. xochimilcensis, P. heterodon tis, P. colubra, P. viperae and P. ptyasi as distinct species until a more sound concept for synonymization, based on additional data on the morphological variability of species included in the genus Paracapillaria, is presented. The synonymization suggested by Moravec (1986) is, on the one hand, not based on examination of the type materials and, on the other hand, significant differences in nematode morphology and geo graphical distribution, host species range and within host localization exist among Paracapillaria species described in the literature so far. Obviously, further studies are needed to clarify the species composition of capillariid nematodes parasites of reptiles from the Palaearctic and particularly those from Europe. Biserkov, V. I., Mészáros, F. és Chipev, N.: Paracapillaria sonsinoi (Parona, 1897) (Nematoda: Capillariidae) és szinonimjai A szerzők hét Paracapillaria taxon önállóságát bizonyítják morfometriai tulajdonságaik, a gazdában való megtelepedés helye és zoogeográfiai elterjedésük alapján. REFERENCES Baker M . R. (1987): Synopsis of the Nematoda parasitic in Amphibians and Reptiles. Memorial University of Newfoundland Occasional Papers in Biology, 11: 1-325. Collins R. E (1973): New hosts and locality records for Capillaria colubra Pence, 1970. Journal of Parasitology, 59 (6): 1020. Moravec F. (1982): Proposal of a new systematic arrangement of nematodes of the family Capillariidae. - Folia Parasitologic
(Praha), 29: 119-132.
Moravec F. (1986): Review of Capillariid nematodes (Capillariinae) parasitic in amphibians and reptiles. Part 3. Genus Paracapillaria. - Vest. Ceskoslov. Spot Zool., 50:573-589. Pence D . B. (1970): Capillaria colubra sp. n. from the oviducts of Coluber constrictor priapus. - Journal of Parasitology, 56 (2): 261-264. Skrjabin K. I., Shihobalova N . P. and Orlov I. V (1957): Trichocephalid and capillariid parasites of man and animals and the diseases caused by them (In Russian). - In: Osnovy Nematodologii (Ed. K. I. Skrjabin), Moscow, Izd. A N USSR, Volume 6, pp. 587. Wang P. Q. (1982): Studies on nematodes of the family Capillariidae from Fujian (in Chinese, Engl, summary). -Acta Zootaxonom. Sinica, 7:117-126.
