`ARC HIVES

FISHERIES AND MARINE SERVICE

Translation Series No. 4167

Results and prospects of investigation into the fauna and systematics of helminths encountered in freshwater fishes of the USSR

by O. N. Bauer, A. V. Gusev, and V. A. Roitman

Original title: Itogi i perspektivy izucheniya fauny sistematiki gel tmintov presnovodnykh ryb SSSR

From: Probl. Obshch. Gel‘mintol. 26: 125-142, 1976

Translated by the Translation Bureau (NDe) Multilingual Services Division

Department of the Secretary of State of Canada

Department of Fisheries and the Environmant Fisheries and Marine Service Pacific Biological Statiàn.

Nanaimo,.B.C.

1977

42 pages typescript

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O.N. Bauer, A.V. Gusev, V.A. Roitman TITLE IN ENGLISH TITRE ANGLAIS

Results and prospects of investigation into the fauna and syste- matics of helminths encountered in freshwater fishes of the USSR

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Itogi i perspektivy izucheniya fauny i sistematiki gel'mintov presnovodnykh ryb SSSR

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Trudy gel'mintologicheskoi laboratorii Akademii nauk SSSR (Problemy obshchei gel'mintologii)

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Trudy of the Helminthological Laboratory of the USSR Academy of Sciences (Problems of General Helminthology)

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Trudy Germintologicheskoi laboratorii Akademii nauk SSSR, vol. 26 (Problemy obshchei gel'mintologii), 1976, pp. 125-42

(125) * Results and prospects of investigation into the fauna

and systematics of helminths encountered in freshwater

fishes of the USSR •

by O.N. Bauer, A.V. Gusev, V.A. Roitman

Through the collective efforts of speci?lists of different

establishments over the past fifty years, we have accumulated ex-

tensive faunistic material which has served as a basis for detail-

ed elaboration of questions pertaining to the morphology, systema-

tics, fauna, biology and geography of all classes of p -2.rasitic

worms encountered in freshwater fishes of the USSR.

•.1. LI) The initiator of these investigations was K.I. Skryabin 0 jr• r.; 9 e: (1919). They ver3 systematically continued by his numerous stu- __.1

,

.2 O y dents and representatives of the ecoparasitologic school of cor- ‹ ry. E •

9,c

0 responding member of the USSR Academy of Sciences, V.A. Dogel'. J? .; E E

t-- • L., The collected data have served as material for monographs and .ca .2 u. 0 c Z determination tables on individual groups of helminths, faunistic D e

a-

• +The numbers in the right-hand mnrgin are the pages of the Russian text - translator

tables on the helminths encountered in fishes that spend their

entire life in a river from different regions or groups of hosts,

general theoretical publications on parasitology, as well as text

books and manuals on controlling helminthiasis in fish. The

most important of these are the 25-volume monograph of Academician

K.I. Skryabin "Trematody zhivotnykh i cheloveka" (The trematodes

dncountered in animals and human beings), "Osnovy nematodologii"

(The principles of nematodologY) (26 volumes), "Osnovy tsestodo-

logii" (The principles of cestodology) (8 volumes), "Irkantotsefaly

domashnikh i dikikh zhivotnykh" (Icanthocephali in domestic and

Wild animals) (Petrochenko, 1956),"Opredelitell paraziticheskikh

nematod" (Key to parasitic nematodes), "Opredelitell parazitov

presnovodnykh ryb SSSR" (Key to Parasites of freshwater fishes of

the USSR), "Monogeneticheskiye sosarshchiki l ikh sistema i filo-

geniya" (Monogenetic Trematoda, their system and phylogeny) (By-

khovsky, 1957), "Trematody, ikh zhiznennyye tsikly, biologiya i

evolyutsiya" (Trematodes, their life cycles, biology and evolu-

tion) (Ginetsinskaya, 1968), "Remnetsy fauny SSSR" (Ligula in

USSR fauna), "Lentochnyye chervi roda Triaenophorus - parazity

ryb" (Mapeworms of the genus Triaenophorus - parasites of fish)

(Kuperman, 1973); "Parazitofauna presnovodnykh ryb Ukr.SSR" (Pa-

rasitic fauna in freshwater fishes of the Ukrainian SSR) (Marke-

vich, 1951; "Parazity ryb Zapadnoi Sibiri" (Parasites of fishes of

Western Siberia) (Titova, 1965); "Parazity ryb vodoyomov Kazakh-

stana" (Parasites of fishesirKazakhstan) (Agapova, 1966); "Para-

zitofauna ryb ozera Baikal" (Parasitic fauna in fishes of Lake

Baikal) (Zaika, 1965); "Parazity ryb Uzbekistana" (Parasites in

fisbeseUzbekistan) (Osmanov, 1971); "Differentsiatsiya lokal'-

nykh stad nerki Oncorhynchus nerka (Walbaum)" (Differentiation

of local populations of Oncorbynchus nerka /Walbaumn (Konovalov,

1971); "Parazity ryb Amura" (Parasites in fishesethe Amur) (group

of authors, Parazitologiya, sb., XII, XXV); "Parazitofauna ryb

basseina rybovodnykh khozaistv Sibiri i Urala" (Parasitic fauna

in fishes of Siberian and lirai fisheries) (Kashkovsky and coauthors

1974); "Sravniterno-ekologicheskiy analiz parazitov ryb ozyor Ka-

relii" (Comparative ecological analysis of parasites in fishes of

Karelian 1Pkes) (Shul'man and coauthors, 1974); "Gel'minty osetro-

vykh ryb" (Hel ,ninths of sturgeons) (Skryabina, 1974); "Osnovy ger-

mintologii" (Principles of helminthology) (Skryabin, Shults, 1940);

"Kurs obshchei parazitologii (Course on general parpsitology);

"Obshchaya parazitologiya" (General parasitology); "Osnovy ger-

mintologii" (Principles of Helminthology), vol. I (Shults, Gvoz-

dev, 1970); "Stroitel'stvo gel'mintologicheskoi nauki i praktiki

SSSR"(Creation of helminthological science and practice and the

USSR) (group of authors, 1962-1972); "Osnovnyye problemy parazito-

logii ryb" (The basic problems of fish parasitology) (group of

authors, 1958); "Kurs boleznei ryb" (Course on fish diseases)

(Lyaiman, 1949); "Bolezni prudovykh ryb" (Diseases of pond fishes)

(Bauer and coauthors, 1969), etc.

Apart from these and the publications cited below, the au-

thors of the present paper have utilized the publications of many

other authors in the analysis of the data herein; the names of

these authors have unfortunately been omitted in order to make

the work more concise.

The specificity of studying each class of helminths has re-

sulted in a certain amount of nonuniformity in the presentation

of the mrterial.

The members of five classes of helminths parasitize the fresh-_

water fishes of the USSR. -

MONOGENEA.

A. Nordmann, Professor of the Odessa University, initiated

the study of freshwater Monogenea in Russia. He described the

types of three genera, DactvloRvrus, Gyrodactvlus and Diplozoon,

which comprise a significant part of the monogenetic flukes of Eur-

asia. Prior to 1917, only fragmentary data on not more than 10

species were available. In the years that followed, the works of

K.I. Skryabin's helminthological school extended our knowledge

about this group, though they listed only the names of

a number of species, the fishes they infested and some data on

their morphology and synonymy (Levashov, 192)-4-; Skryabin, 1924,

etc.). The interest in Monogenea rapidly increased in V.A. Do-

gel's school of ecological parasitology. In the 1930's, B.Ye.

Bykhovsky described approximately 30 new species and two genera

of Monogenetic flukes. A study of the morphology, taxonomy, onto-

genesis and phylogenetic relationships enabled him to outline a

new system of Platyhelminths and Monogenea which he singled out

in an independent class (1937).

Prior to the Patriotic War and immediately following it,

a ne e fauna of parasites was discovered in fishes of the Far East;

it presently consists of over 500 species, including some 200 spe-

cies of Monogenea (Akhmerov, 1952; Gusev, 1955; Gusev, Strelkov,

1960; Strelkov, 1962, etc.). The data on monogenetic flukes of

the Palearctic were also considerably augmented (Bogolepova, 1950;

Bykhovsky, Gusev, 1950; Gvozdev, 1950; Osmanov, 1958; Bykhovsky,

Nagibina, 1959; Spassky, Roitman, 1960; Gavrilova, 1964; Gusev,

1966; Allamuratov, Koval, 1966; Dzhalilov, 1970; Dzhalilov, Ashu-

rova, 1971; Mikailov, 1974; etc.). Descriptions of new genera

(Dogielius, Falociunguism, Markewitschiana, Bvchowskvella, Para-

gvrodactvlus, Pseudacolpenteron, Paradiclvbothrium), subfamilies

and families (Ancylodiscoidinae, Diclybothriidae, Dactylogyrinae,

Ancyrocephalinae, etc.) appeared1. Most of the orders of Mono-

genea were established by Bykhovsky (1937, 1957). The taxonomic

criteria and the hosts of many of the grouPs of Monogenea were

verified or established (Bykhovsky, Nagibina, 1959; Strelkov,

1962; Gusev, 1967, etc.). This was made possible through the im-

provement of investigation techniques, higher observation require-

ments and better processing of mnterials (description and sketch-

in g).

Of great biological interest was the discovery that the mor-

phological characters of Monogenea varied depending on the size

and age of their hosts (Gusev, 1966; Gusev, Kulemina, 1971, etc.).

A similar relationship is also observed in other groups of para-

sites (Trichodina, crustaceans), which is apparently of great

adaptive importance for both components of the parasite—host

system.

. The analysis of all the material accumulated on the morpho-

logy, taxonomy, biology, frequency of occurrence and distribution

of freshwater Monogenea permits us to express a number of ideas

as to the evolutionary paths and factors of their fauna and its

genetic relationships between the different zoogeographic regions.

'The description of Gussevianus, inus, Scriabinonchus and Neo- dactylogyrinae has not been sùfficiently substantiated.

Family, sub-

family, genu's q 731 Family, sub-

eM to) family, genus r.à3 o

11 11

112 110

2

2

1 45 45

The first thing that draws our attention when analyzing the

helminthofauna is the extraordinary diversity of Monogenea. Mono-

genetic flukes are parasites with a simple cycle of development.

In large river basins, they constitute not less tban 55% of the

helminthofauna, and not less than 70%

the other groups of parasites having

when taken together with 4rartst•i) (127) a directXcycie of deve-

lopment (Protozoa, Entomostraca, Hirudinea). Monogenea occupy a

leading place -both in species composition, and in the extensity

and intensity of infestation1 . The predominant species among

them are those of the genus Dactvlogvrus (including several spe-

cies of related genera of the subfamily Dactylogyrinae), which

constitute more than 50% of all the freshwater Monogenea of the

USSR (approximately 60% in the Amur basin), about 40% of all the

helminths and more than 16% of all the freshwater species of fish

parasites. The relative significance of Ancylodiscoidinae is also

great, with 25 species of 3 genera infesting 6 species of fish.

Table 1

Monogenea encountered in freshwater fishes of the USSR

7 7

92 90 2

2

35 35

6 6

30 30

2

e/N/M

1•1•M

M11■•

11 11

Dactylogyr trine Dart ylo gyr inae

Dactylogyrus Pel lueldhaptor Al arkeyttschiana I alciungu is »twirling AI olpengeron Pseuciacolpenteron Ancylod iscott trzae Ant ybot 1 i :cadet Stlurod tirades Bychowskyella A ncyroce phaltnae

APP 2.35 112 127 224

3 ? 3 — 1

— I I 2

— 1 I 1 3 23 26

3 21 24 1 1

14 — 3 9 11

120 133 Tel r aonchidae

Tetraonchut Cyrodactylidae

Gyroclartylus Para gyrod act ylu s

Dtclyttothrttdae Dtelybothrturn Paradiclybothrturn

Dtscocotylidae Discocotyltnas Dtscocotyle Octoma(rurn Diplosooninse Dtplosoon

4

TOX• Ancyrocephalus

ncyroce photo t s. 1. 2 9 11 s

A — 3 I Total I 265 211

• Ancyrocephalus

1 The Correlation may diffe'r in favour of"helminths with a com-.plex cycle in water bodies lacking or poor in Cyprinidae.

The species abundance of parasites with a direct cycle of

development is due to their great evolutionary flexibility as com-

pared with the evolutionally more"unwieldy" helminths with a com-

plex cycle, as the first must pass through 1 -2 "adaptive barriers",

while the second must pass through several, not less than threqe of

these "barriers", depending on the number of stages or phases in

the cycle (Gusev, 1967; Strelkov, Shul'man, 1971).

The very great species diversity of the genus Dactylogyrus

(table 1) is probably due to the great flexibility of its species,

their weak specialization and their confinement to the Cyprinidae

which are abundant in our water bodies. The next most abundant

species belong to the Ancylodiscoidinae found in catfishes and

to the genus Diplozoon found in the Cyprinidae. All these are

comparatively young and primordially freshwater groups. The genus

Gyrodactylus also consists of numerous species, butlhis genus is

encountered everywhere, parasitizing even marine fishes, and has

(123; not yet been sufficiently studied. The highly specialized and,

apparently, ancient groups of the Diclybothriidae, Nitzschia and

Tetraonchus are distinguished by poor species composition and are

probably of marine origin; in any case, the fishes closest to them

are found in the sea.

Analysis of the composition, distribution and morphological

peculiarities of the components of the Monogenea has shown they

that in the USSR are divided into two faunas which were, appar-

ently, formed independently of each other. One is the Palearctic

fauna which is characteristic of Siberian water bodies and the

Mediterranean and upland . Asiae subregions. The other is

characteristic of the Amur basin. The sanie type of division is

+translated literally (Nagorno-aziatskaya podoblast)-. translator

observed in all the other groups of helminths, Protozoa and crus-

taceans. The elements common in both faunas (observed in the Amur)

constitute a comparatively small addition to the main Sino-Indian

nucleus of the Far Eastern fauna.

The majority of monogenetic flukes are characterized by very

narrow host'Specificity (usually one definite species). This fre-

quently determines the parallelism of their and the hosts' evolu-

tion. A vivid example of this are the Dactvlogvrus from Pacific

gudgeon.

The abundance of monogenetic flukes in carps and catfishes

is . such that each of the majority of fishes is parasitized by se-

veral (up to 18) species of one or several genera (Dactvlogyrus,

Ancvlodiscoides, Gyrodactylus, Diplozoon). With the rare except-

ion, each specimen of the indicated species of fish is parasitized

by specific monogenetic flukes. Narrow localization on the gills ;

(Dactylor,vrus skrjabini on the "filter", other species on the gill

filaments in silver carps) has been established onlyfor certain

species; however, the majority of related species are encountered

alternately. When analyzing the fauna of Amur Monogenea, it was

assumed that the basic factors of their rich speciation may be

the specificity and the topological and ecogeographic different-

iation of populations, as well as the alternation of transgressions

and regressions of the sea.

Specificity, on the one hand, limits the evolutionary possi- I

bilities of the parasites, confining them to a certnin group of

hosts; on the other hand, it opens new paths for speciation, con-

tributing to the assimilation of narrower ecologic niches by dif-

ferent populations. With the breakdown of a formerly single basin _

into several rivers, the species of Monogenea in these rivers may

also undergo changes and diverge. Am example of this is the pre-

sence in the Amur catfish in Japan and on the continent of several

different species of monogenetic flukes besides the ones common

for both of these areas; Dactvlogvrus haplogonus Bychowsky, 1933

is encountered in vimba in the Caspian basin, whereas D. haplo-

gonoides Gussev, 1966 is found in the same fish in Black Sea rivers

If rivers that have been separated by the sea come together in the

future, we shall observe the merging of fish populations and the

exchange of the species of monogenetic flukes that had developed

in then during their isolation. It is in this way that the trans-

gressions and regressions of the sea could have influenced specia-

tion.

The study of monogenetic flukes has enabled us to alter our

previous opinion as to the correlation of speciation rates of the

parasites and hosts. According to Rubtsov (1940), the belief had

been that with the evolutionary parallelism of parasites and hosts,'

the first lagged behind the second in the rate of evolution. How-

ever, the monogenetic flukes indicate another correlation o whereby

the - first outstrip the divergence of the second:. There are also

examples of the rates of evolution coinciding. Therefore, "Rub-

tsov's law" is but an exceptional case in this type of correlation.

The method of determining the types of faunas or faunistic

complexes was first applied to the fauna of freshwater Monogenea

with the purpose of determining their origin. It was later suc-

cessfully developed and applied to other groups of helminths,

(129) Protozoa and crustaceans (Roitman, 1963a, b; Trofimenko, 1969;

Strelkov, Shultman, 1971, etc.). Each complex is characterized

by certain morphobiological features: the morphological peculia-

rities of the adhesive organ, the occurrence of this or that com-

plex in fishes, etc. These characteristics, uniting species that

are even genetically remote from each other, are an indication of

the lengthy existence and development of these complexes of spe-

cies in the conditions of a particular geographic zone, and this

is what has determined the identity of their form and biological

specificity. At least five faunistic complexes can be distinguish-

ed in the Amur fauna of monogenetic flukes: Arctic-boreal, Boreal

lowland, amphiboreal, Chinese lowland and Indian freshwater. The

Palearctic fauna is correspondingly divided into Hither Asian,

upland --- Asian, Arctic-boreal, Boreal lowland, amphiboreal and

Pontocaspian faunistic complexes.

The "nomenclature" stage in the study of freshwater Monogenea

of the USSR has basically been completed, though some interesting

new data on the Mono7onea of the upland Asian subregion have ap-

peared in recent years. Tiowever, they have not caused any signi-

ficant changes in the general concept of the fauna and systematics

of this group of helminths. The fauna of the monogenetic flukes

encountered in the rivers of Central and Eastern Siteria remains

unstudied, and helminthologists have not studied the numerous en-

demic species of fish (the Armenian and Turkish khramulya /V. sie-

boldi and V. tinca/, the Crimean and Tera barbels /j. tauricus and

B. ciscaucnsicus/ and numerous Central Asican endemic species).

It is very important to fill in these gaps.

-11-

CESTODA

Data on the cestoids encountered in freshwater fishes of the

USSR were first obtained back in the 18th century: Caryophyl-

laeus laticeps and Cvathocephalus truncatus were described in the

well-known monograph of Academician Pallas (1781). A number of

cestoids from sturgeons and other fishes were mentioned in the

12th century. The accumulation of data on the cestode fauna of

fishes in the USSR progressed rapidly leginning with the 1920 1 s.

These data have been generalized by M.N. Dubinina in the approp-

riate section of the "Key to the Parasites of Freshwater Fishes

of the USSR" (1962). Some of the widely distributed groups of

cestaids parasitic on freshwater fishes have been subjected to

thorough investigation: Proteocephalas(Freze, 1965), Ligula (Du-

binina, 1966), Triaenophorus (Kuperman, 1973). 0.P. Kulakovskaya

published a series of works on Caryophyllaceus. Work on Amphilina

(Dubinina) and some groups of Pseudophyllidea (Protasova, Kuper-

man) is underway.

Special mention should be made of the works dealing with the

cestoids parasitic on fishes of the Amur region (Zmeyev, 1936;

Akhmerov, 1960; Roitman, 1963 a-d; Roitman, Freze, 1964; Dubinina,

1971), which have indicated the distinctive features of the tape-

worms of this region. It can presently be said that the fauna of

the Cestoidea parasitic on freshwater fishes of the USSR has been

studied sufficiently, if not exhaustively.

According to more accurate.data, the fauna of helminths para-

sitic on freshwater fishes currently includes 78 species, though

the independence of some of them requires confirmation. This

number does not include the numerous, primarily larval forms which

No. of species

Subclasses, orders

philina 2

Caryophylieut 3 Parararyorhyllaeus 1 Alonobothrin m 2 Clnridacrts 2 Biarriabu I ti m 1 Brachytirus Car yo nh yllaeides 1 K hatrla 6 Brevisctler 1

Triaeno ',horn s 5 Ettbothriiim 4 JIirthjbo1hri.srn 1 1 issurobothriunt 1 Cyathocephalus 1 Bothrimonsis 1 Schysorntyle 1 Bothr tore rhalsi s 4 Polyonchobothriuni 1 Pi ,'hqllobolhris. m 3 Ligu la 3 Digram in a 2 Srhistrepha 1 us 3

Protrorerhalus 17 ngrsia 4

Silts rotarn ia 1 Para rrotroi e rim Iti s 1

N inpotarnia Parad ri t 1 biter's Cr.orbqncIa,.s 2

N I rind fern idra Cyclo phyll idea

Prviroceplialidar

Pseudo phyllidra

r Genera Families

Cestodar to Amhilintdea

Cestoda Caryophyllaeidea

Amphilin tdae

Caryophyllaeidae

Triarnorhcridar m philotylidar

Cyatheeephalidae

Bof hrioer trine Ptychnbothriidar

ilobothrtida; rigulidar •

Pram rphalidae

Nt nrotartiiidae Dar rididae

are brought into freshwater reservoirs by anadromous fishes. The

cestoids belong to two subclasses: Cestodaria (some specialists

regard it as an independent class) and Cestoda. The distribution

of species according to taxons is given in table 2.

Table 2

Cestoidea of freshwater fishes of the USSR

Total 1 78

The main background of the fauna under discussion is formed

by the Caryophyllaeidea-.(18 species), Pseudophyllidea (30 species)

and Proteocephalidae (23 species). Cyclophyllidea, parasitic (130)

mainly on higher vertebrates, are encountered in fishes only in

the larval stages in the form of cysticerci. The Nippotaeniidea,

the independence of which has been questioned by numerous cestodo-

logists, is represented by 2-3 species in the world fauna; one of

these species is known in the USSR. The taxonomically small sub-

class Cestodaria consists of two species in the USSR.

As in many of the other groups of helminths, the distribution

of cestoids according to the zoogeographic regiohs is characterized

by a noticeable difference between the fauna of the Holarctic

(on USSR territory) and the Amur transition zone. This difference

is observed even at the level of the orders, but is particular- (131)

ly obvious at the levels of genus and species. Of the 30 genera

and 78 species known in the USSR, 7 genera and 16 species have

been encountered in the Amur region alone.

The difference in the cestode fauna is less distinct between

the subregions that make up the Holarctic, primarily the Mediter-

ranean and Circumpolar subregions. The first is characterized by

an abundance of Caryophyllaeidea (not less than 12 species, i.e.

2/3 of all the USSR fauna of this order). The Pseudophyllidea

encountered in this subregion include a recently described spe-

cies of the genus Triaenorborus, as well as the genera Bathvboth-

rium (mountain rivers of the Carpathians, the Caucasus and Cent-

ral Asia) and Bothrimonus (parasitic on sturgeons), one species

of the genus Eubothrium (also encountered in sturgeons) and one

species of the genus Bothrioceohalus (in eel). The Proteocepha-

lidae of this subregion include not less than four species of

the genus Proteocephalus (from eel, sturgeons, gobies and Cat-

fishes) and the genus Silurotaenia. The cestode fauna is consider-

ably poorer in the eastern parts of the subregion (Aral district

of the Pontocaspian-Aral province and the Turkestan province).

Only four species of the Caryophyllaeidea and Proteocephalidae

are known in this area. There are no endemic species in these re-

gions either.

The Circumpolar subregion is characterized by an extreme

scarcity of Caryophyllaeidea. Only Carvophyllaeides fennica and

Khawia rossitensis are widely distributed in the water bodies of

this subregion. Glaridacris catostomi, which is widely distribut-

ed in America, has been encountered in the Far East, in the red-

side sucker of the Kolyma River. Cai-yophyllaceus laticeps para-

sitizes bream in the European district. Two species of Triaeno-

phorus, three species of Eubothrium and Cvathocephalus, as well

plerocercoids of three species of Diphyllobothrium and several

Ligula are widely distributed in this subregion. The Proteroce-

phalidae are widely represented by the genus Proteocephalus (not

less than 10 species).

The cestode fauna encountered in fish2sof the Baikal subreg-

ion resembles that of the Circumpolar subregion, except that it

is considerably poorer. We do not know of any species that are

endemic in this subregion (Zaika, 1965).

An extremely poor cestode fauna also characterizes the insuf-

ficiently studied upland Asian region. As a rule, the fishes of

this subregion are parasitized by the plerocercoids of Ligula .

which are brought into the water bodies of the subregion by fish-

eating birds, and even then very rarely. U. Dzhalikov and M. Ashu-

rova (1971) described Proteocephalus pamirensis from the Tibetan

and Kara-kul stone loaches (Nemachilus stoliczkai and N. lacus-

-nigri) of Tibetan water bodies; this species should for the

time being be considered as an endemic one for this subregion.

. The cestode fauna of the Amur region is poor on the whole,

with a mixture of Holarctic and Sino-Indian elements. A. total of

26 freshw4ter species have been indicated for this region. The

Holarctic element is represented by the genera Paracaryophyllaeus

( one species), Triaenophorus (2 species), Cyathocephalus, Euboth-

rium, Digramma (one species), Proteocephalus (3 species), Grvpo-

rhynchus and Paredilepis (one species). The Sino-Indian element

is represented by the genera Khawia (3 species), Schyzocotyle (one

species), Bothriocephalus (2 species), Polyonchobothrium (one spe-

cies), Gangesia 04 species), Pareproteocephalus (one species).

The position of two species belonging to the genera Nippotaenia

and Breviscolex remains unclear. The first genus is known only

from water bodies of Japan and the Amur basin, while the second

belongs to the subfamily Capingetinae which is extensively repre-

sented in the catostomids and sheatfishes of North America. As

we can see, the Sino-Indian element in the Amur region is far less

distinct In Cestoids than in the other groups te helminths.

-

'; • ,."

Extensive investigations on the acclimatization of fishes

that have been carried out in the USSR have resulted in the wide - distribution of the Sino-Indian species Bothriocephalus gowkongen-

_Liz and Khawia sinensis in water bodies of the Holarctic. Gange- (132)

sia Parasiluri of the Proteocephalidae has found its way into the

tributary system of Lake Baikal with the Amur catfish which was

acclimatized there (Cherepanov, 1962).

Analysis of the distribution of cestoids according to hosts

has shown that this group of helminths manifests a distinct ten-

dency to parasitize specific fish families. Sturgeons are asso-

ciated with the Amphilinidae (onè genus, two species), as well as

with Bothrimonus (one species) and Eubothrium (one species). Fish-

es of the salmon family are extensively parasitized by Eubotbrium

(2 species), Cyathocephalus, Diphyllobothrium (2 species) and Pro-

teocechalus (4 species) . . The genus Triaenophorus (5 species),

which attains maturity in the intestine of the species of only

one family, is characteristic of pikes. The Cyprinidae, which is

a widely represented family in the USSR, is noted for the presenc:

of a large number of Caryophyllaeidae, whereas in North America

and Africa this group of helminths parasitizes mainly catostomids

and catfishes, and not carps. The Cyprinidae of the USSR are also

associated with the genera Bathybothrium, Ligula and Digramma, as

well as several species of the Eenus Proteocephalus. The most

typical parasites of catfishes are Proteocephalidae with suckers

bearing spines and hooks (Gannesia - 4 species, Silurotaenia -

one species), and with a highly inflated scolex bearing numerous

folds (Parnproteocephnlus pnrasiluri).

TREMATODA

The first reliable data on the trematodes encountered in

fishes of our country probably belong to the renowned zoologist,

Karl Ernest Ber (1795-1876), Academician of the Russian Academy

of Sciences. He has described numerous species of trematodes,

including species of the genera Buceohalus and Aspidogaster (1827).

Five years later, A.D. Nordmann dealt extensively with the para-

sites encountered in the eyes of animals and human beings. He

has described five Diplostomum species that parasitize fishes.

During the second half of the 19th century and at the beginning

of the 20th century, fragmentary data on trematodes parasitic on

fish were found in a number of works. Intensive investigation of

the trematode fauna encountered in fishes of our country began in

the 1920's.

A great contribution to the study of trematodes parasitic on

the fishes of USSR waters has been made by K.I. Skryabin, V.A.

Dogel, V.P. Koval, I.Ye. Bykhovskaya-Pavlovskaya, T.A. Ginetsin-

skaya, E.M. Lyayman, S.V. Pigulevsky, A.A. Shigin, D.A. Razmashkin.

V.Ye. Sudarikov and many other researchers. In their works, they

have described new species, elaborated the systems of certain tax-

ons, deciphered life cycles, determined geographic distribution

and studied the trematode fauna parasitic on fish.

Researchers are presently continuing their profound study of

the ontogeneses of trematodes, their interrelationships with dif-

ferent categories of hosts, are searching for new methods of

systematizing the imago and larval forms of trematodes, and are

studying the population dynamics and the factors influencing the

variability of trematodes at all the levels of their ontogenesis.

Researchers are also beginning to deal with questions per-

taining to the biochemistry and physiology of trematodes parasi-

tic on fish.

Unfortunately, the systematics of the large taxons of trema-

todes (orders, superorders, subclasses) has not yet been developed

sufficiently. According to the classifications accepted among So-

viet helmintholgists (Shults, Gvozdev, 1970), the trematodes en-

countered in freshwater fishes of the USSR belong to three sub-

classes: Aspidogastridea, Bucephaldea and Protostomatidea. A

total of 73 species of imagoes and approximately 60 larval forms

of trematodes (the species of which have been determined) have

been recorded in the different taxonomic and ecologic groups of

the different zoogeographic areas of the country. The distribu-

tion of species according to taxons is given in table 3. (133

Sixty-seven species of the subclass Prosostomidea that be-

long to the order Fasciolida and six of its suborders dominate

in the fauna of trematode imagoes parasitic on fishes that spend

their entire life in a river. The other two subclasses are by far

not as well represented.

In the suborders of Fasciolida, there are genera with an

abundant species composition (Allocreadium, Asymphilodora l Phyllo-

distomum), but most of the generic groups are represented by 1-2

species.

The zoogeographic distribution of the trematodes encountered

in freshwater fishes is highly irregular.

11.121e_3.

Trematodes encountered in freshwater fishes of the

USSR (imagoes)

etïbblass, 1No. of order, sub- Family Genus 'specie order Asiddopsfritlm elspidot:asfrid a :Is pidogastridat. :ls !thing,: si el- 2 Bury phalid,n human, Urfa Duceldmhdot. En rrphaln s 9

RhiPlibefdgie 9 ■

Prosttsiontidra 1 nscielida lns. Wahl Cord oder idat. Phyllodislt,mn nt 17 .11Incrt ad La i , •llorreadiidar ..11 locrend in In 12

AuWidumns 1 ffiingdera 1 Crvffidoshnimm -

3Mrwrdidir, AsYmiddyhdMra 8 Milnuorthis 3

Oprcodidm• Nico/M 1 Pia e io porils 1;

Psendosilltnrrostantn 1 S !timer° slonnt

Ori, illmnadiidar Orkrilg,nadium 9 -

Azygifita • l:yeiidar A.P/gi'i 4 liens turnia I so parorch (dur 1 sopurarchis 1 PttrantrItiennwMtn PtitIodiscidnr AnturvIrtnut 1 Sangtankolata Sanoinieohda, .irisguittii , uhl 5

• Total 73

The trematode fauna encountered in fishes in the provinces

(Pontocaspian-Aral and Baltic) of the Mediterranean subregion is

the most abundant as to the number of species (36). Three species

are con.i:on to both of these provinces, eleven species are distribut-

ed only in the Pontocaspian-Azov area and two are found only in the

Baltic province. We observe a distinct peculiarity in the trematode

fauna encountered in fishes of the Amur region for which imagoes of

29 species of trematodes have been indicated; 21 of these species

have never been encountered in the other zoogeographic regions.

The trematode fauna•encountered in fishes of the European

and Siberian districts of the Arctic Sea province consists of al-

most an equal numler of species (25 and 24 respectively). Nine-

(134) teen species are common to both districts, and each district

has a species that is encountered in other regions.

The trematode fauna parasitizing fishes of the Pacific pro-

vince, Circumpolar subregion (6 species) and upland Asian sub-

region (14 species) is extremely poor. However, these territories

have not yet been sufficiently studied, and so the figures given

here are preliminary ones.

The most widely distributed trematodes in the European dist-

rict of the Circumpolar subregion are Allocreadium isoporum, A.

transversale, Bucephalus polvmorphus, Bunodera luciopercae, Crepi-

dostomum farionis, C. metoecus, 8 species of the genus Phvllodis-

tomum and Azvgia lucii. The overwhelming majority of these spec-

ies is encountered in fishes of the other regions of the Palearc-

tic, except for Phyllodistomum dogelii. In the Siberian district,

the trematode fauna parasitic on fish consists of almost the same

species. The number of Phyllodistomum species is greater (9),

but the species composition of this group is somewhat different

than in the European district: Ph. elongatum is not encountered,

but Ph. pigulevskii and Ph. phoxini are. Azygia robusta, two spe-

cies of Crepidostomum and two species of Allocreadium are widely

distributed. Most of the species are encountered in other dist-

ricts of the Palearctic. The trematode fauna of the Pacific pro-

vince consists mainly of two species of Crepidostomum, one species

of the genus Allocreadium (A. transversale) and one of the genus

Phyllodistomum (Ph. conostomum). Ail these species have been ob-

-21- ,

served in fishes of contiguous territories.

The considerable qualitative diversity of the trematode fauna

encountered in fishes of the provinces of the Central Asian sub-

region is due to the predominance of species of the genera San-

guinicola (3), Allocreadium (5), Asymphvlodora (5), Sphaerostoma

(2) and Phyllodistomum (7). Species of Aspidogaster, Rhipidico-

tvle, Orientocreadium, Plagioporus and Nicolla are widely encoun-

tered in the fauna of trematodes. The ranges of the species of

Rhipidicotvle (1), Sanguinicola (1), Allocreadium (2), Paleorchis

(1), Pseudosphaerostoma (1), Phyllodistomum (3) and Orienthocrea-

dium (1) do not exceed the boundaries of the Pontocaspian-Aral

province. The trematode fauna encountered in fishes of the Baltic

province is of a mixed nature. About one-half of the species com-

prising it are also common to the European district of the Arctic

province, and the majority of the remaining species are common to

the Pontocaspian-Aral province. The distribution of Plagioporus

occidentalis outside the Baltic province has not been verified.

Lke the fauna of the other groups of helminths, the trema-

tode fauna encountered in fishes of the Amur basin is quite dis-

tinctive. Almost one-third of the species parasitize mainly carps

belonging to the genus Allocreadium. It is interesting to note

the appearance in the fauna of species belonging to the genera Iso-

Parorchis and Amurotrema which are not encountered anywhere- else

in the USSR except in the Amur basin, the disappearance from the

fauna of numerous trematode species that are widely distributed

in the Holarctic (A. lucii, Bucephalus polvmorphus, Rhipidodoco-

tvle jalense, Bunodera D_uciopercae, etc.) and the appearance of

species of this group that parasitize fishes of the Sino-Indian

freshwater type of fauna. The presence of species similar in

their morphology to species widely distributed in the districts

and provinces of the Holarctic should be noted. These include

Aspidogaster conchocola, Bucephalus skriabini, Sanguinicola skrja-

bini, some species of Allocreadium and Phvllodistomum, etc.

Apart from the trematodes for which fishes serve as defini-

tive hosts, freshwater representatives of the ichthyofauna are

frequently infested by the metacercariae of trematodes, the ima-

goes of which parasitize amphibians, reptiles, birds and mammals,

as well as human beings. We know of approximately 60 species of

larval forms parasitic on fish. Mn st of them belong to the order

Strigeidida (41 species, 16 genera, 5 families). These include (l35)'

imagoes of pathogenic fish parasities (the genera Diplostomum,

Posthodiplostomum, Ichthyocotvlurus). The remaining species of

larvae are distributed in the following family taxons: Climosto-

matidae (2 species of 2 genera), Heterophyidae (5 species of 6

genera), Opisthorchidae (5 species of 4 genera), Nanophyetidae

(1 species of 1 genus) and Echinostimatidae (2 species of 2 genera

Therefore, we presently know of more than 130 species of tre-

matodes belonging to 49 genera and 22 families that parasitize

freshwater fishes of the USSR.

ACANTHOCEPHALA

Prior to the Great October,Socialist Revolutioh, the data

available on the acanthocephalans encountered in freshwater fish-

es of the USSR were of an extremely fragmentary nature. A great

contribution to the science of this group of helminths was made

by N.N. Kostylyov who in the 1920'5-1930's described a number of

genera and species that were new to the fauna of the USSR. The

first to generalize the distribution of acanthocephalans parasitic

on freshwater fishes of the USSR was I.Ye. Bykhovskaya (1936).

Extensive material on this group of helminths is presented in the

first volume of a compendium on acanthocephalans by V.I. Petro-

chenko (1956). The first substantial data on the acanthocephalans

encountered in fishes of the Amur region are presented in the works

of A.Kh. Akhmerov, O.S. Dombrovskaya-Akhmerova (1941) and A.Kh.

Akhmerov (1959). These dat have been augmented by I.L. Sokolov-

skaya.

Acanthocephalans place last in the helminthofauna of fresh-

water fishes of the USSR as to the number of species. There are

37 species of acanthocephalans in all, including several species

of marine origin; the latter are brought into fresh waters by ana-

dromous fishes. They belong to two subclasses, Neoechinorhynchi-

nea and Echinorhynchinea, the latter being distinctly predominant.

The distribution of species according to taxons is given in table 14

The distribution of acanthocephalans according to zoogeogra-

phic regions is characterized by a significant difference between

the fauna of the Holarctic (within the USSR) and the Amur transi-

tion zone. The widely distributed acanthocephalans of the Holarc-

tic include species of the genera Pseudechinorhynchus ( 1 species)

Metechinorhynchus (2 species), Acanthocephalus (2 species), Pom-

phorhynchus (1 species), as well as Neoechinorhynchus rutili (a

total of 7 species). They comprise the nucleus of the fauna in

the majority of basins. This nucleus is augmented by Leptorhyn-

choides plagiplephalus in the Mediterranean subregion, and by the

endemic species Echinorhynchus baeri (parasitic on trout) and

Pomphorhynchus kostylewi, as well as the Hither Asian element

Quadrigyrus cholodkowskii, on the very outskirts of the subregion

in the Black Sea district (Lake Sevan). The fauna of acanthoce-

phalans is extremely poor in the Aral district of the Pontocaspian-

Aral province and in the Turkestan province of the above-indicated

subregion (Osmanov, 1971). Only N. rutill and 0. cholodkowskii

have been observed here. The latter also inhabits the Balkhash

province of the upland Asian region where it is the sole repre-

sentative of the Acanthocephala.

The main nucleus of the acanthocephalan fauna of the Holarc-

tic is somewhat poorer in the Circumpolar subregion: the genus

Acanthocephalus is encountered only in the European district, and

Pomohorhynchus laevis only at the southernmost boundaries of the

Siberian district. At the same time, the number of Neoechinorhyn-

chus species increases to four (Kamchatka) from west to east, and

Paracanthocephalus tenuirostris (Konovalov, 1971), which was first

described from the Amur basin, is encountered in the Lena basin

and the inland water bodies of Kamchatka.

The acanthocephalan fauna parasitic on fishes is far more

abundant in the Amur transition zone than in the Holarctic. The

Neoechinorhynchinea are represented here by five species of Neo-

echinorhynchus, two species of the family Dendronucleatidae that

is s, far known only from the Amur, and by one species of the fa-

mily Hebesomidae that was described earlier from China, as well a

by one species of the family Acanthogyridae. The Amur region is

in many ways similar to the Neoarctic as to the abundance of Neo-

echinorhynchinea (Hoffman, 1967).

Sever hi 111.1- 111111 ,

,N7 h t ii • rb tin eh idg

Arewi/weyr q

/...e lit lenr h gm- h

Pnlyti.rphifM

tuThinorlonchid fif.

11 ehesnmid«

111.m.lronuclentirine

. n loop-War

(shmdrigeidar

Erlem nr hem r lj idae.

Polludiorh mr Witty'

Arledlim.P . Inthidde

Poi!' Inoi.

7

1

.-

1 • 1

1

1 1

1 •1

*

1 •

-25-

Table 4

Acanthocephalans encountered in freshwater fishes of the USSR

Subclasses No. of Families Genera & orders species

ro,rhinoriofierhus

lb.hreonia

Wildrormavala

Acanthorrphalorhurichnidrs

Qundriggrus

nothr,rh tin rleni firs

Erliuhrltwirffil*

114echinrwlignehuç

Fr 1 t

id)? lignelmides

.levaliptroffilluA

Sdamlimaqqpielins

Lerlor4nrimides

Pam nherhqnch is

Heinirlrldim.rhylichns

Merufbuqi.dinoriqmdtlis

11.11n.sonia

Total

Marine forms brought into fresh waters by anadromous fishes.

The Echinorhynchinea are also quite diverse, particularly

those of the order Echinorhynchida. Here we know of 8 species

belonging to the family Echinorhynchidae. The most frequehtly

encountered are two species of the genus Paracanthocephalus which

parasitize the most diverse fishes and in considerable numbers.

The Palearctic genus Acanthocephalus is represented in the Amur

basin by the aberrant species A. parallelotestis which requires

redescription, as it has been described on the basis of one male

specimen. The genera Bolborhynchoides (1 species) and Echinorhyn-

choides (1 species) have been described for the Amur basin alone,

and the genus Sachalinorhynchus (1 species) for the rivers of

Sakhalin. The Palearctic element is represented by Pseudechinorhyn-

chus borealis and the near-Palearctic species Metechinorhynchus

salmonis, as well as by M. crvophilus which is so far known only

from the Amur. The distinctive nature of the acanthocephalans

from the Amur region is augmented by the presence of the two gene-

ra Hemirhadinorhvnchus (1 species) and Pseudorhadinorhvnchus (2

species) of the order Polymorphida which is not repremented at all

in the freshwater fishes of the Palearctic.

However, even though the acanthocephalan fauna of the Amur

region has not yet been determined fully enough and further disco-

veries are possible, it already numlers 19 species; at the same

time, only 17 purely freshwater acanthocephalans are known from

all the thoroughly studied water basins of the USSR that belong to

the Holarctic and are characterized by the most diverse conditions.

The specificity of acanthocephalans parasitic on freahwatà137:

fishes of the USSR is very weakly defined. The presence of any

species is determined mainly by the nature of the food consumed by

the fishes. Feeding on small demersal-crustaceans (gammarids and

isopods) is conducive to infestation by Echinorhynchinea; Neoechi-

norhynchinea infest fishes that feed on ostracods, the intermediate

hosts of this group of acanthocephalans. Perhaps, the narrowest

specificity is characteristic only of Leptorhynchoides plagicepha-

lus which parasitizes only sturgeons. At the same time, a narrow

specificity is characteristic of acanthocephalans found within the

first intermediate hosts 'where . the parasite undergoes consider-

able metamorphosis and where the formation of the genital system

is completed. At this stage we encounter even species specificity.

For example, Echinorhvnchu salmonjs develops only in Pontoporeia

affinis.

NEMATODA

The Nematoda comprise one of the large groups of parasites

in the helminthofauna of freshwater fishes of the USSR. They at-

tack the overwhelming majority of fishes inhabiting the water bo-

dies of our country.

Parasitic roundworms have been studied quite thoroughly from

the faunistic aspect. Ninety-four species of nematodes belonging

to 31 genera, 15 families, 3 orders and 2 subclasses are presently

known in the freshwater fishes of various taxonomic and ecologic

groups on the territory of the USSR. Most of these species para-

sitize fishes in their mature form, and the other species in the

form of larvae, the imagoes of which parasitize vertebrates of dif-

ferent classes, as well as human beings. (Only the larval forms of

definite and unquestionable species have been recorded).

The subclass Adenophorea is represented by the families Capil-

lariidae and Cystoopsidae of the order Trichocephalida and by Dioc-

tophymidae of the order Dioctophymida. Eleven species of the genus

Capillaria have been recorded in the sturgeons, salmons, carps, go-

bies, perches and cods from the fresh waters of all the zoogeogra-

phic regions. The life cylces and other details of the biology

of these parasites are almost unknown. The only species of the

genus Cystoopsis (Cystoopsidae), C. acipenseris, is an obligate

parasite of sturgeons in the Pontocaspian-Azov area and the Amur R.

Unlike the Capillariidae and Cystoopsidae that parasitize in

fishes in a mature form, the Dioctophymidae (the genera Dioctophyme

and Eustrongyloides) are encountered in pikes, sturgeons, carps,

catfishes, perches, salmons and other fishes in the larval form.

The larvae of the only species of Dioctophyme are distributed in

the southern parts of the country, while the three species of Eu-

strongyloides are encountered everywhere.

The Sencernentea, which belong to different suborders of the

order Spirurida, are more numerously represented in the freshwater

fishes of the USSR. They belong primarily to four families of the

superfamily Thelazioidea and two families of the superfamily Physa-

lopteroidea. Two families of the Thelazioidea, Rhabdochonidae

Skrjabin, 1946 and Ascarophiididae Trofimenko, 1967, dominate in

the numler of species over the other groups of nematodes.

The rhabdochonids belong to 16 species of 4 genera (Rhabdo-

chona - 11 species, Filochona - 3 species, Cystidicola - 1 species,

Prosungulonema - 1 species). The Rhabdochona and Filochona para-

sitize mostly carps, less frequently sturgeons, salmons and perch-

es; the largest number of these parasites is observed in the Ponto-

caspian-Aral province. The only species of the genus Cystidicola (13E

(C. farionis) is an obligate parasite of salmons, and is distribut

ed mainly in the water bodies of the Circumpolar subregion. The

monotypic genus Prosungulonema has been confirmed for the nematode!

of the Chinese perch from the Amur River.

The Ascarophiididae are represented in fresh waters by 9 spe-

cies belonging to 4 genera (Ascarophis - 3 species, Cysticycoloi-

des - 1 species, Salvelinema - 1 species, Comephoronema - 2 speciez•

Of the three species of AscaroPhis, two parasitize sturgeons of

the Pontocaspian-Aral region, Siberia and the Amur basin, and one

species is encountered in cods and gobies of the Azov-Black Sea

basin. The systematics and biology of the Ascarophiq species re-

quire further study. On USSR territory, the genus Cvstidicoloides

is represented by one species, C. tenuissima, an obligate parasite

of salmons and graylings, and is distributed everywhere in the ri-

ver basins of the Arctic province. Salvelinema salmonicola para-

sitizes Pacific salmons and is distributed in the river basins of

the Pacific Ocean.

Comephoronema oschmarini, a parasite of burbot, has been dis-

covered in Lake Baikal, the Yenisei River, in western Taimyr (Lake

Keta) and Lake Onega; another species of the genus, C. wereschagini ll

is known from the gobies of Lake Baikal.

Species of the genus Spinitectus (Spinitectidae) have been

encountered in gobies and cods of the Azov-Black Sea basin, in the

swallows and sea basses of the Amur river, and in eels from the

rivers of the Baltic Sea. The Spinitectus are on the whole a poor-

ly studied group of nematodes, which requires further analysis.

The haplonematids in USSR fauna are represented by the genera

Cottocomephoronema (1 species) and Haplonema (1 species). Cotto-

comephoronema problematica l the only species of the genus, is an

obligate parasite of the burbot in the European and Siberian dist-

ricts. Haplonema orthocephalum is still the only representative

of the genus; these nematodes have been recorded in salmons of the

Amur.

The larvae of Desmidocercella numidica and related species

(Desmidocercidae) are frequently encountered in perches and carps

of the Caspian, Aral and Baltic basins. The larvae of spirurids

of the group Agamospirura are even more widespread in sturgeons,

carps, perches, catfishes and swallows of various water bodes.

Three species from the superfamily Physalopteroidea are known

to parasitize fish: two belong to the genus Cvclozone (Cyclozoninae

and are obligate parasites in sturgeons of the Pontocaspian-Aral

region; the third one, Gnathostoma hispidum (Gnathostomatidae), in-

fests perches and carps of the Aral basin in the larval stage.

The ascarid nematodes known in fishes mostly belong to several

genera of the family Anisakidae. As a rule, fishes act as the se-

cond intermediate (supplementary) or reservoir host, which explains

the frequent occurrence of larvae of the Anisakidae in fishes in

various areas. The species diagnosis of Anisakidae larvae has been

insufficiently developed as to morphological characters, and so it

is as yet impossible to estimate the number of species of the generi

Anisakis, Contracaecum and Porrocaecum that are encountered in the

freshwater fishes of the USSR. We know for certain that the fresh-

water nematode fauna encountered in fishes of the USSR includes 6

species of the genus Contracaeceum (in sturgeons, carps, perches

and salmons inhabiting water bodies close to the sea), the Chinese

perch of the Amur River, two species of Goezia (in catfishes, pikes

gobies, salmons and cods) in the Pontocaspian-Aral region and one

species of the genus RaDhidascaris (obligate parasite of pikes,

encountered within the range of the host, with the exception of

Kamchatka, in the larval stage parasitizes numerous groups of fish)

The larvae of Streptocara crassicauda have been detected in carps,

perches and dragonets (in the Azov-Black Sea basin).

The nematodes of the suborder Camallanata belong to two fami-

lies. Two genera, Camallanus and Procamallanus, represent the fa-

mily Camallanidae: the first genus includes four known species

encountered in perches, carps, cods and other groups of fishes

from water bodies of the Arctic province and the Amur River; the

second genus consists of two species that parasitize catfishes (139)

and sturgeons of the Amur River and Aral Sea basin.

The family Cucullanidae is represented by two genera on USSR

territory: Cucullanus (8 species - parasites of salmons, sturgeons,

perches, carps and other groups of fish; encountered in all the zoo-

geographic areas) and Bulbodacnilis (1 species - parasite of sal-

mons of the Pacific province of the Palearctic).

A large group of nematodes of the family Dracunculidae falls

into four genera: Philometra (7 species, mainly parasites of carps

in most river basins), Philometroides (3 species, parasities of the

Cyprinidae of various regions), Philonema (1 species, obligate par a .

site of salmons in the rivers of the Arctic and Pacific oceans),

Thwaitia (3 species encountered in the carps of most of the pro-

vinces of the Palearctic, with the exception of the Pacific pro-

vince) and Clavinema (1 species, attacks catfishes of the Amur R.).

The sole representative of the family Pingidae, Pingus sinen-

sis, has been discovered in the snakeheads of the Amur basin.

The family Skrjabillanidae has in recent years been scrutiniz-

ed from different aspects. The group includes four species of ne-

matodes that parasitize the carps of the Caspian and Aral basins.

Two of these species belong to the genus Skriabillanus, and the

other two to the genera Agrachanus and Molnaria. The distribution

of the individual species requires further investigation.

This summarizes, in general, the taxonomie position of nema-

tode species and their distribution on USSR territory.

The above data indicate that from the faunistic aspect the

helminths encountered in the fresh waters of the USSR have been

studied quite thoroughly. There is not likely to be a sudden in-

crease in the quantitative and qualitative composition of the in-

dividual classes of the helminthofauna, in spite of the fact that

BON@ areas of the USSR have not even been studied.

At the same time, the systematics of numerous taxons of dif-

ferent ranks within the classes of helminths requires further pro-

found improvement. This should inevitably affect the approximate

total indices of species numbers which at present are:

Classes No. of species

Classes No. of species

Monogenoidea 450 Acanthocephaloidea 37 Cestoidea 78 Nematoidea 94 Trematoidea 130

Total 798

The present level in the development of helminthology deter-

mines the new tasks pertaining to the study of the fauna and sys-

tematics of the helminths encountered in freshwater fishes. In

general, they are as follows:

1. The development of experimental faunology with the purpose

of studying the principles governing speciation, the morphological

variability in helminths and the natural phylogenetic relationships

between the taxons of different ranks.

2. The extensive introduction of karyological, biochemical

and mathematical methods of investigation, as well as electron

and scanning microscopy, etc., in order to increase the accuracy

of the results and to determine the new tendencies in morphological

and taxonomic research.

3. The improvement of the methods (quantitative and qualita-

tive) of studying helminth populations, determination of intraspe-

cific categories and evaluation of their role in the evolution of

individual groups of helminths.

4. The continuation of studies dealing with insufficient- (140)

ly studied groups of helminths: Dactvlozvrus and Gvrodactylus of

the Monogenea; Pseudophyllidea of the Cestoda; Sanguinicola, the

Allocreadiidae and Gordoderidae of the Trematoda; the Neoechino-

rhynchidae and freshwmter Echinorhynchidae of the Acanthocephaaa;

and the Capillariidae, Cucullanidae, Spinitectus, Philometroides,

etc. of the Nematoda.

5. The study of the helminthofauna encountered in freshwater

fishes of the central and eastern parts of Siberia and the upland.

Asian subregion, as well as endemic species of fishes.

6. The generalization of faunistic data on the individual re-

gions and groups of hosts in order to solve the general theoretical

problems pertaining to the ecology and geography of freshwater hel-

minths parasitic on fish. The compilation of a key to the para-

sites of freshwater fishes of the USSR and of regional tables on

the helminthofauna of fishes.

7. The training of helminthologists familiar with the new

methods of investigation, and the Coordination of research work

with specialists in other fields.

8. The improvement of the collection aspect of the investiga-

tions.

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