REVIEW ARTICLE / ARTÍCULO DE REVISIÓN

CHROMOSOMES AND CYTOGENETICS OF HELMINTHS (TURBELLARIA,

TREMATODA, CESTODA, NEMATODA AND ACANTHOCEPHALA)

CROMOSOMAS Y CITOGENÉTICA DE HELMINTOS (TURBELLARIA, TREMATODA,

CESTODA, NEMATODA Y ACANTHOCEPHALA)

1 1 1 2 3

Tanveer A. Sofi , Fayaz Ahmad , Bashir A. Sheikh , Omer Mohi Ud Din Sofi & Khalid M. Fazili

1Post Graduate Department of Zoology, University of Kashmir, Srinagar, Kashmir, 1900 06, India.

2SK University of Agricultural Sciences and Technology, Shuhama, Aluestang Srinagar, 1900 06, India.

3Post Graduate Department of Bitechnology, University of Kashmir, Srinagar, Kashmir, 1900 06 Ph. No. India.

09797127214. stanveer96@gmail.com

Neotropical Helminthology, 2015, 9(1), jan-jun: 113-162.

ABSTRACT

Keywords: chromosomes – cytogenetics - Acanthocephala- Cestoda- - Nematoda- Trematoda- Turbellaria.

We review the literature from 1886 to 2014 and the current status of knowledge of the

chromosomes and cytogenetics of all species of Turbellaria, Trematoda, Cestoda, Nematoda, and

Acanthocephala. Karyological data are discussed and tabulated for 614 species: 115 species of

Turbellaria, 278 species of Trematoda, 117 species of Cestoda, 85 species of Nematoda and 19

species of Acanthocephala. Turbellarians are not parasitic except for a few possible exceptions

and they show a gradual reduction of the basic number of chromosomes. Trematodes are

numerous which points towards the continued efforts in this field of research. Data on

chromosomes are lacking for acetabulate cestodes of the orders: Litobothriidea,

Lecanicephalidea, Cathetocephalidea, Rhinebothriidea and Tetrabothriidea.

113

RESUMEN

Palabras clave: Acanthocephala- Cestoda- citogenética - cromosomas - Nematoda- Trematoda- Turbellaria.

En este artículo revisamos la literatura desde 1886 hasta 2014 y el estado actual del conocimiento

de los cromosomas y la citogenética de todas las especies de las familias de turbellaria,

trematoda, cestoda, nematoda y acanthocephala. Datos cariológicos son analizados y tabulados

para 614 especies: 115 especies de turbellaria, 278 especies de trematoda, 117 especies de

cestoda, 85 especies de nematoda y 19 especies de acantocephala. Los Turbelarios no son

parásitos a excepción de unas pocas posibles excepciones y muestran una reducción gradual del

número básico de cromosomas. Trematodes son numerosos requiriendo apuntar hacia los

esfuerzos continuos en este campo de investigación. Los datos sobre los cromosomas se carecen

para cestodos acetabulados de las órdenes: Litobothriidea, Lecanicephalidea,

Cathetocephalidea, Rhinebothriidea y Tetrabothriidea.

ISSN Versión impresa 2218-6425 ISSN Versión Electrónica 1995-1043

INTRODUCTION

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015

As with molecular data, cytogenetic

information can reveal differences and

similarities that may not be obvious at the

morphological level. White (1978) estimated

that more than 90% of all speciation events are

accompanied by karyotypic change. If this is

correct, then chromosomal studies should be

widely applicable to the problems of sorting

groups of morphologically similar (sibling)

species. Chromosomes are studied as a

morphological manifestation of the genome in

terms of their microscopically visible size,

shape and number, and karyology represents a

qualitative approach to phylogeny. To what

extent, though, can karyotypic features be

suitable for phylogenetic inference? Patterns

of chromosomal divergence within a group

may not necessarily parallel those of

morphological features (Gold, 1980; Baker &

Bickham, 1980), but most often species related

from a morphological point of view show

karyological affinities. If karyotypic features

are plotted over a phylogenetic tree based on

molecular or morphological data, the

processes involving chromosome evolution

might be clarified.

A karyotype is the number and appearance of

chromosomes in the nucleus of eukaryotes

(Stebbins, 1950; White, 1973). The term is also

used for the compliment set of chromosomes

in a species or an individual organism.

Karyotype describes the number of

chromosomes and what they look like under a

light microscope. Attention is paid to their

length, the position of chromosomes, any

differences between the sex chromosomes and

any other physical characteristics (King et al.,

2006). The preparation and study of

karyotypes is a part of cytogenetic. The study

of whole sets of chromosomes is known as

karyology. The chromosomes are depicted (by

rearranging a microphotograph) in a standard

format known as karyogram or ideogram: in

pairs, ordered by size and position of

centromeres for chromosomes of the same

size. The basic number of chromosomes in the

somatic cells of an individual or a species is

somatic number and is designated as 2n. In

normal diploid organisms, autosomal

chromosomes are present in two copies.

Polyploid cells have multiple copies of

chromosomes and haploid cells have single

copies. Karyotypes can be used for many

purposes such as to study chromosomal

aberrations and taxonomic relationships and to

gather information about past evolutionary

events.

Most karyotype studies use cells that are near

the end of prophase or in early metaphase of

mitosis because the chromosomes are compact

and densely staining and have a characteristic

size and shape. The chromosome size, the

location of the centromere, and patterns of

light and dark staining that occur when

chromosomes are treated with different

chemical dyes are collectively referred as

chromosome morphology and the number and

morphology of an individual's chromosomes is

called that individuals karyotype. The analysis

of karyotypes of different organisms proves

quite useful in species description and

identification (Stebbins, 1950; White, 1973).

Taxonomic identification of the helminth

parasites which causes diseases is absolutely

essential for effective treatment. Taxonomy

plays very important role in the management

and prophylaxis of diseases by biological

means. Anticipating a problem is always more

time and cost effective than responding to a

crisis, no matter how effective the response.

Systematic biology provides and integrates the

knowledge that is crucial for any effort to be

proactive in the arena of emerging parasitic

and infectious diseases. The aim is to complete

the global inventory of parasitic species, an

absolute necessity if we are to assess risk of

parasitic diseases. Taxonomy is important in

the field of Biodiversity and Conservation,

Sofi et al.

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015

Research and Studies, Agriculture and Pest

management, Quarantine, National defence,

Fisheries and Aquaculture, Parasitology and

Veterinary Science, Conservation of Plants,

Animals and Microbes.

The purpose of this study is to: (1) Find out the

chromosome number of helminths from

different vertebrates; (2) Find out the

Karyotype characteristics of helminthes; (3)

Differentiate helminths on the basis of the

karyological characteristics; (4) Role of these

studies in cytotaxonomy, and (5) Find out the

general aspects such as trends of karyotypic

evolution and sex mechanism of trematodes.

The present methodology is in accordance

with the all articles find about chromosomes of

Turbellaria, Trematoda, Cestoda, Nematoda

and Acantocephala in Google Scholar,

Elesiever, Jstor and Springer. For karyotyping,

chromosomes descriptions were on the basis of

size and centromere position (Petkeviciute &

Leshko, 1991). Relative lengths of

chromosomes were calculated by the division

of the individual chromosome length by the

total haploid length and centromeric indices

(ci) were determined by division of the length.

The terminology relating to centromere

position follows that of Levan et al. (1964). A

chromosome is metacentric (m) if the ci falls in

the range of 37.5–50.0, submetacentric (sm) if

25.0–37.5, subtelocentric (st) if 12.5–25.0 and

acrocentric (a) if < 12.5. When the centromere

position was on the borderline between two

categories, both are listed.

Chromosomes of Turbellaria

Among the turbellarians, records of

MATERIALS AND METHODS

Chromosomes and cytogenetics of helminths

chromosome numbers among the primitive

Acoela are few, but three species of the

Convolutidae have been studied and all seem

to indicate that the somatic number of very

small chromosomes is somewhat variable but

high-ranging from 20 to 30. Records for the

other orders of this basically hermaphroditic

and occasionally parthenogenetic class are

numerous and the results have been

corroborated by various researchers. Among

the Alloeocoela, members of the

Prorhynchidae, the Plagiostomidae, and the

Bothrioplanidae usually show a basic haploid

number of 10 (reduced to 5 in one species), but

the Monocelididae have the haploid number of

3, while the Pseudostomidae have either 4-5 or

8-10. Since polyploidy is common among the

turbellarians, the basic haploid chromosome

number for the alloeocoels may be 5, although

Monocelis fusca [Ruebush (1938)]; (n = 3)

does not readily fit into the schema. Among the

Tricladida there seems to be a much greater

degree of variation in the basic haploid number

of chromosomes. Among the Planariidae the

number ranges from 6 to 24 (6, 8, 12, 16, 18,

and 24 have been recorded). Curtisia and some

polycelids have n = 6. In other polycelids

polyploidy is believed to give rise to the

species having n = 12, 18, or 24. Lepori (1953a,

b) believes that in some cases among the

polycelids true fertilization does not occur, and

that the multiples of the basic chromosome

number may perhaps be due to failure to

undergo reduction divisions (a condition

Lepori terms gynogenesis). In the genus

Planaria most of the species have the basic

chromosome number of 8. In the genus

Dugesia, the number varies from 8 to 16 or to

24, again an example of possible polyploidy,

but the appearance of 12 chromosomes in

Dugesia alpina [Rappeport (1915)]; may be an

indication that the basic number is 4 rather than

8, as stated by Benazzi-Lentati, 1949. Among

the Dendrocoelidae we again find considerable

variation in the basic haploid number.

Dendrocoelum has species with 8 and others

with 16 chromosomes (possible examples of

RESULTS AND DISCUSSION

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

polyploidy). Procotyla and one variety of

Dendrocoelum lacteum [Pennypacker (1954)]

(both with 7 haploid chromosomes) and

Bdellocephala with 10 chromosomes are

variants (possible aneuploidy). Other families

such as the marine Bdellouridae and the

Procerodidae are represented by too few

species in which the basic chromosome

number has been determined (Bdelloura,

where n = 6 and fragmentation may raise the

number as high as 12; and Procerodes, with n =

6) to be indicative of their evolutionary

position, although the presence of

fragmentation may point toward a possible

compound nature of the gametic

chromosomes. Turning to the Polycladida we

again note variability in the haploid

chromosome number which ranges from 6 to 8

and occasionally to 9 or 10. The Euryleptidae,

the Leptoplanidae, and the Prosthiostomidae

seem to have the basic number of 8 while the

Planoceridae possess 10 chromosomes, but no

such consistency appears among the genera of

the Stylochidae where the basic numbers are

either 9 or 10. The single pseudocerid

examined has 9 chromosomes. It might be

suggested that in these cases possible fusion of

some of the chromosomes may have occurred,

thus reducing the number from 10 to 9, or even

to 8, and thus showing relationship to the basic

10 pattern set by the Planoceridae as a whole.

An alternative possibility is that a doubling of

one or more chromosomes has caused the

deviation from a basic number of 8, as

represented by the Euryleptidae. On purely

morphological grounds the planocerids are

regarded as the more primitive and the

euryleptids as more specialized among the

polyclads. Among the higher Rhabdocoela, we

again find considerable variation in numbers,

but the basic count is low, ranging from 2 to 6,

and it is suggested that fragmentation of one or

more chromosomes plus some group

duplication of the basic number may be

involved. Among the supposedly more

primitive rhabdocoels (Catenulidae) the basic

number runs much higher, ranging from 15 to

20 and the somatic numbers may reach to 40

[Stenostomum, Rhynchoscolex, Catenula and

Fuhrmannia = Suomina are representative

genera]. According to Pennypacker- (1954) -

such variation is of evolutionary importance in

that as a general rule a larger number of smaller

chromosomes are indicative of ancestral

conditions and a smaller number points toward

a more recently emerged species. Among the

Typhloplanidae it may be noted that

Opistomum, Krumbachia, Solenopharynx,

Amphibolella, some protoplanellids, and

Phaenocora, as well as the North American

form of Rhynchomesostoma rostratum show a

basic number of 2. (The European race of R.

rostratum has 3 chromosomes, as do

Trigonostomum, some protoplanellids, and

Co-strata.) Papi- (1950) - indicates that the

basic number for the Mesostominae is 4, but

through fusion, fragmentation, or ploidy, some

species show n = 2, 3, 5, or 8. The Dalyelliidae

all show n = 2. Jones, 1944, has indicated that

among the Macrostomidae, the basic number is

probably 3 and that a few species show

multiples in the form of 6 and 9 as the reduced

number of chromosomes (some species

however, show 2 and 8). The Graffillidae also

show at least in the genus Paravortex that the

basic number is 2, doubled in one species to 4.

The Provorticidae have a basic number of 3.

The Kalyptorhynchidae vary from 2 to 3 to 8

(Table 1).The single temnocephalid examined

has 8 chromosomes. As yet there seems to be

no complete agreement between chromosome

number and the phylogenetic position of these

forms as determined by other criteria, but on

the other hand no absolute contradictions have

come to light. Although the turbellarians are

not parasitic except for a few possible

exceptions, they have been introduced into this

discussion because they do show the gradual

reduction of the basic number of chromosomes

from a large number of small chromosomes to

a smaller number of larger units, not only

within the group as a whole, but in some cases

within races of the same species, and may

possibly give hints as to the derivation of the

117

Trematoda which do show a much greater

conformity between the chromosome number

and the taxonomic position as based on other

criteria. There seems to be considerable

evidence already available pointing to the

trematodes as evolving from a rhabdocoel

stock, and possibly from the dalyelliid group.

Cytological evidence seems to corroborate this

interpretation of a rhabdocoel ancestry, but if

such development came through the

dalyelliids, it must have been from a stock

much less specialized than the present day

forms which are regarded by many as being

among the more advanced of the rhabdocoels.

In this same connection it may be of interest to

point out that the acoelids and the alloeocelids

are regarded as the most primitive of the

turbellarian line of development and most

consistently possess the highest basic number

of chromosomes (10 in most of the species

examined). Through some form such as

Bothrioplana (n = 10) could have come the

triclads (some of the dendrocoelids show this

basic number of 10). Polyclads and the

rhabdocoels lack such close cytological

connection with a possible alloeocoelid

ancestry although some of the polyclad

planocerids and some of the rhabdocoelid

macrostomids do show practically the same

basic chromosome numbers.

Table 1. Chromosomes of Turbellaria (From 1905 Till Date).

CERCYRINAE

Cercyra hastate 2n = 14 (3sm + 3st + 1t) Galleni & Puccinelli (1982)

Balliania thetisae 2n = 22 Gourbalt (1978)

UTERIPORIDAE

Uteriporus vulgaris 2n = 16 Ball (1976)

Foviella affinis 2n = 16 Ball (1976)

PROCERODIDAE (PROCERODINAE)

Procerodes gerlachei 2n = 12 Bohmig (1908); Ruebush

(1938)

Procerodes littoralis

2n = 14

Ball (1976, 1979); Galleni & Puccinelli (1975,

1979)

Procerodes dohrni

2n = 14

Galleni &

Puccinelli (1979)

OPISTHOBURSIDAE

Opisthobursa mexicana

2n = 14

Benazzi &

Giannini (1973)

PSEUDOCERIDAE

Yungia aurantiaca

2n = 18 (4m + 2sm + 2t)

Galleni &

Puccinelli (1985)

Dugesia lugubris

2n = 8

Galleni et al. (1989)

Polycelis nigra

2n = 16; 3n = 24

Lamatsch et al. (1998)

Dugesia polychroa

2n = 8; 3n = 12 (1m + 3a)

Lamatsch et al. (1998)

Dugesia sicula Lepori

2n = 18

Fillipi et al. (1998)

Dugesia gonocephala

2n = 16; 8m

Baguna et al. (1999); Deri et al.

(1999)

Schmidtea polychroa

2n = 8

Benazzi (1957)

Schmidtea polychroa

3n = 12

Benazzi (1957)

Schmidtea polychroa

4n = 16

Benazzi (1957)

Schmidtea polychroa

5n = 20

Benazzi (1957)

Schmidtea lugubris

2n = 8

Benazzi (1957)

Schmidtea lugubris

2n = 6

Benazzi (1957)

Schmidtea mediterranea

2n = 8

Benazzi (1957)

Schmidtea mediterranea

3n = 12

Benazzi (1957)

Dugesia benazzi

2n = 16

Baguna et al. (1999)

Dugesia hepta

2n = 14

Baguna et al. (1999)

Dugesia etrusca 2n 16 Baguna et al. (1999)

Dugesia subtentaculata 2n = 16 Baguna et al. (1999)

Dugesia sicula 3n = 27 Baguna et al. (1999)

Thysanozoon brocchi 2n =18 Schockaert (1905); Ruebush (1938)

Family Species No. and Morphology of

Chromosomes Bibliographic reference

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Chromosomes and cytogenetics of helminths

DENDROCOELIDAE

Dendrocoelum lacteum 2n =14 & 16 Pennypacker

(1954)

Dendrocoelum infernale 2n =32 Aeppli

(1951); Pennypacker (1954)

Procotyla fluviatilis 2n =14 Pennypacker

(1954)

Bdellocephala brunnea 2n =20 Momma

(1953)

BDELLOURIDAE

Bdelloura candida 2n =12-24 Pennypacker

(1954)

Bdellura candida 2n = 12 Pennypacker (1938)

EURYLEPTIDAE

Cycloporus papillosus 2n =16 Francotte

(1898); Ruebush

(1938)

Oligocladus aurritus 2n =16 Francotte

(1897); Ruebush

(1938)

Prostheceraeus vittatus

2n =12

Gerard

(1901); Ruebush

(1938)

LEPTOPLANIDAE

Leptoplana tremellaris

2n =16

Francotte (1898); Ruebush

(1938)

PROTHIOSTOMIDAE

Prosthiostomum siphunculus

2n =16

Francotte (1898); Ruebush (1938)

STYLOCHIDAE

Stylochus pilidium

2n =18

Gerard

(1901); Ruebush

(1938)

Eustylochus ellipticus

2n =20

Von Name

(1899); Ruebush (1938)

PLANOCERIDAE

Planocera inquilina

2n =20

Patterson & Wieman

(1912); Ruebush (1938)

Planocera nebulosa

2n =20

Von Name

(1899); Ruebush (1938)

CATENULIDAE

Fuhrmannia sp? (= Suomina sp?)

2n =32

Ruebush (1938)

Catenula virginiana

2n =40

Ruebush (1938)

Rhyncoscole simplex 2n =40 Ruebush (1938)

Stenostomum grandi 2n =40 Ruebush (1938)

Stenostomum sp?2n =40 Ruebush (1938)

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

RHYNCHODEMIDAE

Dolichoplana carvalhoi

2n = 14 (5m + 1sm + 1st);

3n = 21 (1a,4c,5b,6,7a,7b (m) +

1b,2a,(sm) + 3c,3d (st);

Alvarez &

Almeida (2007)

Pentacoelum hispaniense Sluys, 1989

2n = 14; 2m + 3t/st + 2sm

Vila-Farre et al. (2008)

CONVOLUTIDAE

Aphanostoma diversicolor

2n =40-60

Ruebush

(1938)

Convoluta sp?

2n =40-60

Ruebush

(1938)

Polychoerus caudatus

2n =62

Gardiner

(1898)

Polychoerus carmelensis

2n = 34

Costello (1970)

BOTHRIOPLANIDAE

Bothrioplana semperi

2n =20-30

Pennypacker

(1954)

PLAGIOSTOMIDAE

Hydrolimax grisea

2n =20

Pennypacker (1954)

Plagiostomum stetlatum

2n =10

Ruebush (1938); Pennypacker(1954)

PRORHYNCHIDAE

Geocentrophora applanatus

2n =20

Ruebush

(1938); Pennypacker (1954)

Prorhynchus stagnalis

2n =20

Ruebush

(1938); Pennypacker (1954)

PSEUDOSTOMIDAE

Pseudostomum caecum

2n =16-20

Jones (1943); Pennypacker (1954)

Pseudostomum sp?

2n =8-10

Ruebush

(1938); Pennypacker (1954)

MONOCELIDIDAE

Monocelis fusca

2n =6

Ruebush

(1938); Pennypacker (1954)

PLANARIIDAE

Curtisia foremanni

2n =12

Pennypacker

(1954)

Polycelis tenuis 2n =12 Melander (1950); Lepori (1953)

Polycelis nigra 2n =36 Lepori (1953)

Dugesia benazzi 2n =16 & 32 Benazzi-Lentati and Nardi (1950)

Dugesia gonocephala 2n =16 & 32 Schleip (1907); Benazzi- Lentati (1949)

Dugesia alpina 2n =24 Rappeport (1915); Benazzi- Lentati (1949)

Dugesia sp?2n =48 Ruebush (1938)

Planaria polychroa 2n =16 Mattieson (1904); Ruebush (1938)

Planaria torva 2n =16 Mattieson (1904); Ruebush (1938)

Phagocata fawcetti 2n = 38 Ball and Gourbault (1975)

Family Species No. and Morphology of

Chromosomes Bibliographic reference

Macrostomum beaufortensis

2n =6

Ferguson (1940)

Macrostomum tuba

2n =6

Ferguson (1940)

Macrostomum rirginianum

2n =6

Ferguson (1940)

Macrostomum hustedi

2n =12

Jones

(1944)

Macrostomum bispiralis 2n =16 Ferguson (1940)

Macrostomum kepneri 2n =18 Ferguson (1940)

DALYELLIIDAE

Castrella truncata 2n =2 Ruebush (1938)

Dalyellia abursalis 2n =2 Ruebush (1938)

Dalyellia armigera 2n =2 Ruebush (1938)

Dalyellia triangulata 2n =2 Ruebush (1938)

Dalyellia virginiana 2n =2 Ruebush (1938)

Dalyellia riridis 2n =2 Ruebush (1938)

Dalyellia (4 sp'?) 2n =2 Ruebush (1938)

TEMNOCEPHALIDAE

Temnocephalus canis

2n =16

Ruebush (1938)

TYPHLOPLANIDAE

Promesostoma marmoratum

2n =12

Ruebush (1938)

Bothriomesostoma personatum

2n =10

Ruebush (1938)

Bothriomesostoma essenii

2n =10

Papi

(1950)

Byrsophlebs sp?

2n =8

Ruebush (1938)

Mesostoma rhynchotum

2n =16

Valkanov (1938)

Mesostoma ehrenbergii (Europe)

2n =10

Papi

(1950)

Mesostoma ehrenbergii wardi (U.S.A.)

2n =8

Husted

et al.

(1939)

Mesostoma benazzii

2n =8

Papi (1950)

Mesostoma lingua

2n =4 & 6

Ruebush (1938)

Costrata subsala

2n =6

Ruebush (1938)

Costrata virginiana

2n =6

Ruebush (1938)

Costrata sp?

2n =6

Ruebush (1938)

Typhloplana viridata

2n =6

Ruebush (1938)

Trigonostomum lillei

2n =6

Ruebush (1938)

Protoplanella sp?

2n =6

Ruebush (1938)

Protoplanella sp?

2n =4

Ruebush (1938)

Opistomum pallidum

2n =4

Ruebush (1938); Papi (1952, 1953)

Opistomum sp?

2n =8

Ruebush (1938)

Krumbachia minuta

2n =4

Ruebush (1938)

Krumbachia virginiana

2n =4

Ruebush (1938)

Solenopharynx sp?

2n =4

Ruebush (1938)

AmphibolelIa virginiana

2n =4

Ruebush (1938)

Phaenocora kepneri

2n =4

Ruebush (1938)

Phaenocora lutheri

2n =4

Ruebush (1938)

Phaenocora virginiana

2n =4

Ruebush (1938)

Phaenocora sp?

2n =4

Ruebush (1938)

Rhynchomesostoma rostratum

2n =4 & 6

Volkanov

(1938); Papi (1950)

KALYPTORHYNCHIDAE

Polycystis goettei

2n =16

Ruebush (1938)

Microkalyptorhynchus virginianus

2n =6

Ruebush (1938)

Acrorhynchus reprobatus

2n =4

Ruebush (1938)

Gyratrix hermaphroditicus

2n =4

Ruebush (1938)

PROVORTICIDAE

Provortex affinis 2n =6 Ruebush (1938)

Provortex sp?2n =6 Ruebush (1938)

GRAFFILLIDAE

Paravortex cardii 2n =4 Hallez (1908); Ruebush (1938)

Paravortex gemellipara 2n =8 Ball (1916); Ruebush (1938)

MACROSTOMIDAE

Macrostomum hystrix 2n =4 Ferguson (1940)

Macrostomum viride 2n =4 Ferguson (1940)

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a = acrocetric; a, b, c, d = chromosome variants; m = metacentric; sm = sub-metacentric; st = subtelocentric acrocentric.

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recorded in most species (for a review see

Petkeviciute & Staneviciute, 2008). The

chromosome complement of Cercariaeum

crassum (Wesenberg-Lund, 1934) is unusual

among digeneans due to the low number,

2n=10. The karyotype is composed of large

and exclusively bi-armed chromosomes. Such

a karyotype presumably results from a

decrease in chromosome number through

centromere–centromere Robertsonian fusions

that have affected mono-armed chromosomes

leading to the formation of large metacentric

elements. Comparative analysis of

chromosomes of related trematode species

indicated that the reduction of chromosome

numbers resulted from centromeric fusion

rather than elimination of chromosomes

(Grossman et al., 1981). Acrocentric mono-

armed chromosomes prevail in the karyotypes

of larval B. luciopercae, 2n = 14, and larval A.

isoporum sensu Wisniewski, 1959, 2n = 14

(Petkeviciute & Staneviciute, 2008). It is

notable that the mean total length of haploid

complements (TCL) of these two species does

not exceed the TCL of C. crassum, despite

different chromosome numbers.

It may be noted, however, that all of the

heterophyids (Cryptocotyle and Acetodextra),

bucephalids (Bucephalus and Rhipidocotyle),

fasciolids (Fasciola), and zoogonids

(Zoogonus) examined (7 species), as well as 1

species of a gorgoderid (Probolitrema) and 1

of a paramphistomatid (Gigantocotyle) have a

basic number of 6 [perhaps thus indicating

some relationship to the more primitive

paramphistomatids (n = 7)]. The notocotylids

(Notocotylus) have 7 chromosomes, but too

few examples have been studied to determine

the value of such counts. One species of an

allocreadid (Bunodera), 1 gorgoderid

(Gorgoderina), and 1 schistosomatid

(Schistosomatium) have 7 chromosomes, in

addition to the 2 amphistomids (Zygocotyle

and Gastrothylax). All of the Schistosoma

species studied show n = 8, as do 2 species of

g o r g o d e r i d s ( G o r g o d e r a a n d

Chromosomes of Trematoda

A number of workers on trematode cytology,

especially Jones and his co-workers, have

pointed out the possible taxonomic value of the

study of the numbers, volume, and/or size and

shape of chromosomes, and much of our

information along these lines is based upon

their investigations. Among the Monogenea

only the Polystomidae have been investigated

for the purpose of determining the

chromosome numbers. Polystoma

integerrimum (Frolich, 1791) has the haploid

number of 4 chromosomes and Gyrodactylus

elegans has 6. This could point to a dalyelliid

ancestry (n = 2 in all species studied) although

more primitive ancestors of the dalyelliids may

have had a larger basic number of

chromosomes (Table 2). Among the Digenea,

the paramphistomatids have been regarded as

the most primitive group, but from a

cytological standpoint the basic chromosome

number is quite variable and is therefore of

little guidance in determining possible

relationships. For example, Gigantocotyle

shows a basic number of 6, Gastrothylax and

Zygocotyle have 7, Cotylophoron and

Diplodiscus have 8, while Heronimus

chelydrae has 10. No one family of the

rhabdocoel group could be regarded as being

directly ancestral based on such divergent

records. As much as the exact phylogenetic

relationships of the remaining families of the

Digenea are not as yet fully determined, even

on morphological grounds, and the various

authorities disagree as to their proper

taxonomic positions, no attempt will be made

to discuss our knowledge of chromosome

numbers in any significant succession. The

diploid chromosome numbers vary among

studied digenean taxa, from 12 to 28 (Bariene,

1993); chromosome sets with 20 or 22

elements predominate. But 56 chromosomes

were found in diploid sets of Clonorchis

sinensis (Cobbold, 1875) (Park et al., 2000).

Allocreadiid species possess comparatively

large chromosomes, up to 13–14 mm, but low

haploid numbers of six, seven or eight were

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Chromosomes and cytogenetics of helminths

Phyllodistomum), 1 troglotrematid

(Paragonimus), 3 species of allocreadids

(B u n o d e r a , C re p i d o s t o m u m and

Allocreadium), 1 species of a rhopaliid

(Rhopalias), and 2 species of the reniferids

(Staphylodora and Telorchis), in addition to

the 2 species of paramphistomatids

(Cotylophoron and Diplodiscus). Two species

of the azygiids (Azygia and Proterometra), 2

species of the plagiorchids (Eustomas and

Glypthelmins), 1 species of a spirorchid

(S p i r o r c h i s ) , 1 p r o n o c e p h a l i d

(Macrovestibulum), 1 lecithodendriid

(Brandesia), 1 monorchid (Asymphylodora), 1

hemiurid (Halepegus), and 1 reniferid

(Auridistomum), in addition to 1 race of the

paramphistomatid Diplodiscus (temperatus)

have 9 chromosomes. Those having a basic

number of 10 chromosomes are 1 species of a

cyclocoelid (Cyclocoelum), 2 species of

dicrocoelids (B rachy coelium and

D i c ro c o e l i u m ) , 1 c l i n o s t o m a t i d

(C l i n o s t o m u m ) , a n d 1 h e m iur i d

(Isoparorchis). Only Heronimus of the

paramphistomatids falls in this cate-gory. Four

species of plagiorchiids (3 Pneumonoeces and

1 Plagitura), 1 echinostomid (Parorchis), 2

lecithodendriids (Acanthatrium and

Loxogenes), and 15 species of reniferids (1

Dasymetra, 1 Lechriorchis, 1 Natriodora, 6

Neorenifer, 2 Pneumatophilus, 1 Renifer, and

3 Telorchis) have 11 chromosomes. In these

cases again no direct relationship to the

paramphistomatids can be noted in terms of

chromosome number, although in terms of the

presence of an increased number of

chromosomes as indicating a possible

primitive condition, these forms might be

regarded as less specialized than others of the

Digenea. It should be pointed out, however,

that in many cases it is not only in the matter of

actual number of chromosomes that

similarities (relationships) may be indicated:

total volume of chromatic material, shapes and

sizes of the chromosomes, point of spindle

attachment to individual chromosomes, and

behavior during division may afford evidence

of equal importance. It is also definite that

sufficient differentiation occurs to facilitate

identification of species. One species of

Cephalogonimus has 14 chromosomes which

may be a case of doubling of the usual number

of 7 in this genus. If one follows the belief of

Ciordia (1949), who doubts the existence of

polyploidy among the Trematodes, this would

be an example of extreme aneuploidy-

duplication of individual chromosomes and

not of the set as a unit. The matter of the

presence or absence of the so-called

heterochromosome ("sex chromosome") has

not been determined in most of the trematode

species examined, but in a few cases the

recognition of two types of sex cells differing

from each other in terms of the number, size,

shape, volume or behavior of the chromosomal

elements would seem to indicate that such

sexual differentiation does occur. As examples

we may mention the studies on Schistosoma

and Schistosomatium. The earlier observations

on Schistosoma haematobium, S. mansoni, and

S. japonicum (Katsurada, 1904); seemed to

indicate that two types of sperm could be

identified and that adult males possessed 15

and adult females possessed 16 somatic

chromosomes. This would seem to mean that

an X-O condition obtained in these forms.

Other studies reported the numbers as 14 and

16, respectively, and were interpreted as

showing the presence of a 2X + 12 and a 4X +

12 chromosome complex. Niyamasena (1940)

in his studies on S. mansoni found the somatic

number of chromosomes to be 16 in each sex,

and could not find any evidence of the presence

of recognizable sex chromosomes although the

possibility of an X-Y condition could not be

ruled out. Most recent studies support the

finding of 16 chromosomes as the diploid

number in all adults of all three of the above

Schistosoma species and the inability to

recognize sex chromosomes as being present,

16 chromosomes are also present in each sex of

S. mansoni carcariae (Bilharz, 1852). Recent

studies on Schistosomatium douthitti (Bilharz,

1852); have presented 2 different

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

interpretations of what is undoubtedly an

example of the presence of distinguishable sex

chromosomes. One study (Woodhead, 1957)

indicates the presence of a single "X"

chromosome in the male, while the other

(Short, 1957) presents evidence of the

heterogametic condition as prevailing in the

female. Studies in this laboratory seem to

substantiate this second interpretation. The

somatic (diploid) number of chromosomes in

each sex is 14, with the male showing a pair of

large V-shaped chromosomes that are not

matched in the female. In the latter case there is

a single large V-shaped chromosome

apparently paired with a single rod-shaped

body. This rod-shaped chromosome does not

appear in any of the male cells. It is interpreted

as indicative of a ZZAA condition in the male

and a ZWAA condition in the female. Short &

Menzel (1957) report a similar condition in the

cercariae of Ornithobilharzia canaliculata,

where 2n = 16. No other records of the

presence of recognizable sex chromosomes

among the trematodes seem to have been

substantiated by recent investigations. No

records of the "diminution" phenomenon have

been brought to our attention. Le Roux (1958)

in a study of mammalian blood flukes, has

suggested a division of the genus

Schistosonmai into several groups:

Schistosoma (S. haematobium, type),

Afrobilharzia (A. mansoni, type),

Sinobilharzia (S. japonicum type),

Rhodobilharzia (R. margrebovici, type), and

Eurobilharzia (E. bomfordi, type). These

genera are in addition to the already recognized

genera of Bivitellobilharzia, Heterobilharzia

and Schistosomatium as members of this group

of flukes. Restudy of the chromosomes of

these species might help to evaluate such a

separation. The same technique might also

help solve the complexities of the taxonomy of

avian Schistosomes.

Table 2. Chromosome number of Trematoda from 1902 Till Date.

Family Species No. and Morphology of Chromosomes Reference

SCHISTOSOMIDAE

Schistosum japonicum Katsurada, 1904 2n = 16 (6t +2at) Short & Menzal (1960)

Austrobilharzia variglandis 2n=16 Short & Menzal (1960)

Gigantobilharzia huronensis 2n=16(+B) Short & Menzal (1960); LoVerde & Kuntz (1981)

Heterobilharzia americana 2n=20(ZW); ZZ/ZW, m/st;

6m+4sm+2sm-st+ 8st

Short & Grossman (1986)

Texas (Male) 2n=20(ZZ); ZZ, m; 4m + 2m-s m+4sm-

st+2sm+2st-1+6st

Short et al. (1987)

Louisiana (Female)

2n=20(ZWA); WA, m; 3m+2m -sm

+4sm-st+2sm+2st-t+6st

Short et al. (1987); Britt (1947)

Ornithobilharzia huronensis

2n=16

(XY)

Short & Menzel (1960)

Schistosoma bovis, Schistosoma haematobium,

Schistosoma intercalatum, Schistosoma mattheei

2n=16

Short & Menzel (1960); Grossman et al. (1981a)

Schistosoma japinicum

2n=16; ZZ/ZW, m/sm

Grossman et al. (1981b)

Schistosoma mekongi

2n=16; ZZ/ZW, sm/sm; 4sm+8 a + 4t

Grossman et al. (1981b)

Schistosoma mansoni

2n=16 (ZW No. 1)

Short & Menzel (1960); Short et al. (1979);

Grossman et al. (1980)

Schistosoma mansoni

2n=16; ZZ/ZW; st/st-sm; 2m + 4 m-sm

+ 4 s t-sm + 4st + 2t

Atkinson (1980); Short & Grossman et al. (1981a)

2n=16 (ZW No. 2); m/st; 4m + 2sm-m +

2m-sm + 2 s m-st+ 2st + 2t-st

Atkinson (1980); Grossman et al. (1981); Grossman

(1981)

Schistosomatium douthitti

2n=14 (ZW No. 1);

m/st; 4m + 2sm-m + 2m-sm + 2 s m-st+

2st + 2t-st

Short & Menzel (1960); Short & Grossman (1981);

Puente &

Short (1985); Short

(1957); Short & Menzel

(1959)

Trichobilharzia physellae,

2n=16

Short &

Menzel (1960); Short (1983)

Trichobilharzia stagnicola

2n=16

Short &

Menzel

(1960); Short (1983)

Trichobilharzia szidati

2n=16 (5m+2sm+1sm-st)

Spakulova et al. (1996)

Trichobilharzia regent Horak, Kolarova et Dvorak, 1998 2n=16 ((5m+1sm+1sm-m+1sm-

m(Z)+1m(W)+1sm (Supernumerary B) Spakulova et al. (2001)

Schistosoma rodhaini 2n=16; ZZ/ZW, s t /st; 2m+2sm-m+4sm-

st +4st +2t-st Atkinson (1980); Grossman

et al. (1981a); Short & Grossman (1981)

Schistosoma haematobium 2n=16; ZZ/ZW, s t/st;8m +8st Short (1983)

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Family Species No. and Morphology of Chromosomes Reference

Megalodiscus temperatus 2n=18; Van der Woude (1954); Saksena (1969)

Diplodiscus amphichrus magnus 2n=18; 6m + 6sm+6a Saksena (1962)

Schistosoma bovis

2n=16;

LoVerde &

Kuntz (1981); Short (1983)

Schistosoma matthei

2n=16;

LoVerde &

Kuntz (1981); Short (1983)

Schistosoma intercalatum

2n=16;

Short

(1983)

Schistosoma margrebowiei

2n=16;

Grossman et al.

(1981a); Short (1983)

Schistosoma japonicum

2n=16; ZZ/ZW, sm/sm

Gao Longsheng et al.

(1985)

Schistosomatium sp.

2n=14; ZZ/ZW, m/a; 12m+2m-sm

Barsiene et al.

(1989)

Bilharziella polonica

2n=16; ZZ/ZW, st-sm (Male); 4m 4sm-

m+2m-sm+4sm+2st

Barsiene

Stanyavichyute

(1993)

Ornithobilharzia caniculata

2n=16; ZZ/ZW

Short

(1983)

Austrobilharzia variglandis

2n=16; ZZ/ZW, sm-m/a; 12m+2sm +2a

Barsiene et al.

(1989)

Triehoilharzia physellae

2n=16;

Triehoilharzia szidati

2n=16; 6m+2sm-m+6sm+ 2st-sm

Barsiene &

Stanyavichyute (1993)

Trichobilharzia

sp. 1

2n=18; 14m +4sm-m

Barsiene et al.

(1989)

Trichobilharzia sp. 2

2n=16; 12m +2sm-m+2sm

Britt

(1947)

Gigantobilharzia huronesis

2n=16;

Britt

(1947)

PRONOCEPHALIDAE

Macrovestibulum kepneri

2n = 20

Jones et al. (1945)

PARAMPHISTOMIDAE

Ceylonocotyle dicranocoelium

2n = 18

Subramanyam &

Venkat Reddy (1977)

Cotylophoron cotylophorum

2n=16

Subramanyam &

Venkat Reddy (1977)

Cotylophoron sp.

2n=16

Subramanyam &

Venkat Reddy (1977)

Fischoederius elongates

2n=16

Subramanyam &

Venkat Reddy (1977)

Gastrothylax crumenifer

2n=18; 4m +6sm+2a+6st

Romanenko (1974); Subramanyam & Venkat Reddy

(1977)

Gigantocotyle explanatum

2n=18

Subramanyam &

Venkat Reddy(1977)

Liorchis scotiae

2n=18

Romanenko (1974)

Megalodiscus (Diplodiscus) temperatus

2n=20

Grossman &

Cain (1981)

Paramphistomum cervi

2n=14

Venkat Reddy &

Subramanyam (1975)

Paramphistomum epiclitum

2n=18

Subramanyam &

Venkat Reddy (1977)

Paramphistomum ichikawal

2n=18

Romanenko (1974)

Paramphistomum microbothrium

2n=18

Mutafova (1983a)

Stichorchis subtriquetrus

2n=18

Romanenko (1974)

Diplodiscus temporatus

2n = 16

Cray (1909)

2n=14

Sey

(1971)

Paramphistomum microbothrium

2n=18, 2 s m + 10m + 6st

Mutafova

(1983a)

Paramphistomum explanatum

2n=18, 2sm+16a

Sharma & Lal

(1984)

Paramphistomum hiberniae

2n=12

Willmott

(1950)

Paramphistomum ichikawai

2n=18; 2 m + 4 s m + 12st

Romanenko

(1974)

Paramphistomum epiclitиm

2n=18; 2 m + 14sm + 2st

Subramanyam &

Venkat Reddy (1977)

Paramphistomum crassum

2n= 14;

Srivastava &

Iha

(1964a)

2n= 14; 4m +l0sm

Subramanyam &Venkat Reddy

(1977)Paramphistomum cervi

2n=18; 6m + 4sm + 8 st

Rhee et al. (1987a)

Paramphistomum elongatum

2n=16; 6m+6sm +4st

Dhingra

(1955a)

Paramphistomum sp.( Planorbarius corneus) 2n=18; 2m + 2m-sm+6sm + 8st Barsiene (1991)

Paramphistomum sp. (Planorbis planorbis)

2n=18; 2m + 10sm + 6st

Barsiene

(1991b)

Gigantocotyle bothycotyle

2n= 12;

Willmott

(1950)

Gigantocotyle explanatum

2n=18; 4m+l0sm + 4a

Venkat Reddy &

Subramanyam

(1975b);

Subramanyam &

Vekat Reddy

(1977)

2n=18; 4m + 14a

Romanenko

(1972)

Liorchis scotiae

2n=18; 2m + 8 sm + 8a

Romanenko

(1974)

2n=18; m + sm Britt (1947)Gastrothylax cruminifer

2n=14; Dhingra (1955a)

Fischoederium cobboldi 2n=18; 8 m + l0sm Rhee et al. (1988)

Zygocotyle lunata 2n=14; Willey & Godman (1951)

Cotylophoron elongatum 2n=16; Dhingra (1955b)

Cyclonocotyle dicranocoelium 2n=18; 4m + 10sm + 4st Britt (1947)

Cyclonocotyle orthocoelium 2n=18; Sharma et al. (1968)

Cyclonocotyle dawesi 2n=20; Britt (1947)

Cyclonocotyle scoliocoelium 2n=22; Britt (1947)

Stuncardia dilymphosa 2n=18; Sharma & Nakhasi (1974)

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

Family Species No. and Morphology of Chromosomes Reference

Diplodiscus subclavatum

2n=20; 2sm -m + 8sm + 2st-sm + 6 s t +

Petkeviciute

et al.(1989b)

2n=20; 2 m + 12 + sm + 4st + 2a

Britt (1947)

2n=21-30;

Britt (1947)

Notocotylus noyeri Joyeux, 1922

2n=20; 2sm-m+12sm+4st+2a

Barsiene &

Grabda-Kazubska (1991b)

Gastrodiscoides hominis

2n=20;

Romenenko

(1974)

Stichorchis subtriguentrus

2n=20;

Britt (1947)

Heronimus chelydrae

2n=18; 4sm 14a

Guilford

(1955)

ZOOGONIDAE

2n = 10

Goldschmidt (1905)

Zoogonus mirus

2n = 12

Schreiner (1908); Gregoire (1909); Wassermann

(1913)

DICROCOELLIDAE

Dicrocoelium lanceolatum

2n = 20

Goldschmidt (1908); Dingler (1910)

Dicrocoelium lanceolatum

2n=24

Romanenko (1979)

Eurytremum pancreuticum

2n=26

Romanenko (1979)

Paradistomoides orientalis

2n=28

Scharma &

Nakahasi (1974)

Dicrocoelium lanceatum

2n=24; 22m + 2sm

Sharma &

Nakhasi

(1974)

Eurytrema coelomaticum

2n=26; 1 0 m + 2 sm+12st + 2t

Moriyama

(1982a, 1982b)

Eurytrema pancreaticum

2n=26; 10m+4sm + 8st + 4t

Britt (1947)

Paradistomoides orientalis

2n=28; 1 4 s t + 10st + 4t

Dhar &Sharma

(1984)

BRACHYCOELIDAE

Brachycoelium salamandrae

2n = 20

Von Kemnitz (1913)

HETEROPHYIDAE

Cryptocotyle lingua

2n = 12

Cable (1931); Britt (1947)

Apophallus miiehlingii

2n = 14 ( 3m+4sm)

Barsiene et al. (1995)

ECHINOSTOMIDAE

Parorchis acanthus

2n = 22

Rees (1939)

Episthmium bursicola

2n=18; 12m+4sm+2st

Barsiene &Kiseliene

(1990a)

Echinochasmus baleocephallus

2n=14; 6m+2m-sm+4sm+2a

Britt (1947)

Parorchis acanthus

2n=22;

Rees

(1939)

2n=22;

Churchill (1950)Echinostoma reuolutum

2n=22; 2m +20st

Mutafova & Kanev (1986)

Echinostoma revolutum (L. stagnalis)

2n=22; 2m+4sm+2 st-sm +10st+4a

Barsiene & Kiseliene (1991)

2n=22; 2m 2sm-m+4sm+14st

Britt (1947)Echinostoma revolutum (L. ovata)

2n=24; 2 В

Britt (1947)

Echinostoma jurini

2n=22; 6m+8sm+4st+4a

Britt (1947)

Echinostoma miyagawai

2n=22; 2m+2sm+2st-sm+16st

Barsiene & Kiseliene, 1991

2n=22; 2m + 20st

Mutafova & Kanev (1986)

2n=22; 2m+2sm-m+2sm+12st+4a

Barsiene &

Kiseliene

(1991)

Echinostoma echinatum

2n=22; 2m + 2sm-m +2sm+16st

Britt (1947)

Echinostoma barbosai;

Echinostoma echinatum 2n=22; 2sm + 20a Mutafova & Kanev (1983)

Echinostoma hortense

2n=20; 2m+2m-sm+8sm-st + 8st-t

Terasaki et al. (1982)

Echinostoma tinetorchis

2n=22; 2m + 2sm-st+12st+4st + 2t

Britt (1947)

Echinostoma caproni

2n=22; 4sm-st, t

Richard &

Voltz (1987)

Neoacanthoparyphium echinatoides 2n=20; 2m + 4 s t + 1 4a Barsiene & Kiseliene (1990b)

Moliniella anceps 2n=20; 2m + 8 s t + 1 0a Barsiene et al. (1990b)

Isthmiophora metis (из Lymnaea stagnalis) 2n=20; 4m+4sm+2st+10asm Britt (1947)

Pegosomum asperиm 2n=20; Aleksandrova & Podgornova (1978)

Lymnaea saginatum 2n=20; Britt (1947)

Echinoparyphium recurvatum 2n=20; 2m + 2sm + 16t Mutafova et al. (1987)

Echinoparyphium recurvatum А (Lymnaea auricularia,

Lymnaea ovata) 2n=20; 2m-+4sm + 14st Barsiene (1991а)

Echinoparyphium recurvatum В (Lymnaea corиus,

Lymnaea palustris) 2n=20; 4m + 6sm+10a Britt (1947)

Echinoparyphium pseudorecurvatum A (PI. planorbis 2n=20; 2sm -m+4 sm+4st +10a Britt (1947)

Echinoparyphium pseudorecurvatum В(A. acronicus) 2n=20; 2sm-m + 2st-sm + 8st + 8a Britt (1947)

Echinoparyphium bacutus (Vatvata piscinatis) 2n=20; 2 m+12 s t+2a-st+4a Britt (1947)

2n=20; 2m-sm + 2sm + 16t Mutafova et al.(1987); Mutafova & Kanev (1984)Echinoparyphium aconiatum

2n=20; 2m + 2st-sm+ 4st+4st-a +8a Barsiene & Kiseliene (1990b)

2n=20; 4 m + 1 6 s t Mutafova et al.,1986Hypoderaeum conoideum 2n=20; 2 m + 4 s m + 12st Barsiene & Kiseliene, 1990b

Cathamaesia hians (Lymnaea stagnalis ПНР) 2n=20; 8m 4sm-m +4sm+4st Barsiene (1990)

Cathamaesia hians (Planorbis planorbis. ПНР) 2n=20; 4m + 4sm-m+8sm +2st-s m + 2 s

tBarsiene (1991b)

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Family Species No. and Morphology of Chromosomes Reference

BUCEPHALIDAE

Bucephalus elagans

2n = 12

Woodhead (1931)

Bucephalus pusillus

2n = 12

Woodhead (1931); Britt

(1947)

Rhipidocotyle papillosum

2n=12

Ciordia

(1956)

TROGLOTREMATIDAE (PARAGONIMIDAE)

Paragonimus kellicotti

2n = 16

Chen (1937)

Paragonimus kellicotti

2n = 22 (2a + 9sm)

Loverde (1979)

Paragonimus ohirai

2n = 22 (2a + 9sm)

Loverde (1979)

Paragonimus miyazakii

2n=22

Sakaguchi &

Tada (1975, 1976); Terasaki (1977);

Hirai et al.

(1985)

Paragonimus ohirai; 2 geographical races (Paragonimus

iloktsuemensis; Paragonimus sadoensis)

2n=22

Sakaguchi &

Tada (1975, 1976); Terasaki (1977);

Hirai et al.

(1985)

Paragonimus westermani

2n=22

LoVerde (1979); Hirai et al.

(1985); Sugiyama et al.

(1985)

Paragonimus westermani, (Paragonimus pulmonalis)

2n=22

Sakaguchi &

Tada (1976b); Terasaki (1977);

Agatsuma &

Habe (1985); Hirai et al. (1985)

Paragonimus westermani

2n=22

Blair (2000)

Paragonimus heterotremus

2n=22 (1m+4st+3m/sm+3sm/st)

Komalamisra (2005)

Paragonimus kellicotti

2n=16

Benazzi &Benazzi Lentati (1976)

2n=22;

Britt (1947)

2n=22; 4m + 18sm

Britt (1947)

Paragonimus ohirai

2n=22; 2 m + 1 0 sm + 10st

Hirai еt аl.

(1985)

2n=22; 4 m + 10sm + 8st

Sakaguchi &

Tada

(1976а); Terasaki (1977)

3 n = 33;

Sakaguchi &

Tada

(1976a); Miyazaki (1978)

2n=22; 6m+8sm+8st

Pengpeng et al.

(1986)

2n=33; 2m + 8 st + 12m, sm, st

He Lian-Yin et al.

(1982)

Paragonimus westermani

2n=22; m + 6 st + 2 sm + 12m, sm, st

Britt (1947)

Paragonimus westermani Shoawu,

Fujian

2n=22; 2m + 4st + 4sm + 12m, sm, st

Britt (1947)

Paragonimus westermani filipinus

2n=22; 2m + 6m-sm + 6sm-st + 8st

Terasaki (1983)

Paragonimus westermani westermani

2n=22; 8m + 6sm-st + 8st

Britt

(1947)

Paragonimus skrjabini

2n=22; 6m+6sm-m + 2sm +8st

Li and Zheng

(1983)

3n=33; 2m+6m- sm+6sm-st+ 8st

Terasaki

(1980); Sakaguchi &

Tada (1976b)

2n=22; 2m + 6m-sm + 6sm- st + 8st

Terasaki

(1977)

Paragonimus pulmonalis

3n=33; 3 m + 12sm + 6 st +12a

Hirai еt аl.

(1985)

2n=22; 2m + 6m-sm+6sm-st + 8st

Terasaki

(1977); Sakaguchi &

Tada (1980)Paragonimus iloktsuenensis

2n=22; 2m + 10sm+10st Hirai еt аl. (1985)

2n=22; Terasaki (1977); Sakaguchi & Tada (1980)Paragonimus sadoensis 2n=22; 2m + 10sm+ l0st Hirai et al. (1985)

Paragonimus peruvian us 2n=22; 2m+6m-sm + 6sm-

st + 8st Terasaki (1978)

Paragonimus hueitungensis 2n=22; 2m + 8 st + 12m, sm, st He Lian-Yin et al. (1982)

Euparagonimus cenocopiosus 2n=22; 6 m + 8 s m + 8 s t Lei Changqui et al. (1985)

AZYGIIDAE

Proterometra macrostoma 2n =18 Anderson (1935)

Azygia acuminata 2n = 18 Britt (1947)

Azygia lucii 2n=20; 10m + 6 a + 4 st Barsiene (1991b)

ALLORCEADIIDAE

Allocreadium isoporum 2n =16 Britt (1947)

Crepidostomum serpentinum 2n = 16 Britt (1947)

Bunodera saculata 2n = 16 Britt (1947)

Bunodera luciopercae 2n = 14 Britt (1947)

Cercariaeum crassum Wesenberg-Lund, 1934 2n=10 (1m+1sm+sm-m+1m-sm) Petkeviciute et al. (2011)

Allocreadium fasciatusi 3n=21; 3 m + 12sm + 6st Ramanjaneyulu &Madhavi (1983)

Bunodera sacculata 3n=23; Cannon (1971)

Allocreadium fasciatusi 3n=21 Ramanjaneyulu & Madhavi (1984)

Allocreadium handiai 2n=14 Ramanjaneyulu & Madhavi (1984)

Bunodera luciopercae 2n=14 (2m+1sm/st+4a) Petkeviciute & Staneviciute (2008)

Allocredium isoporum 2n=14 (2m+5a) Petkeviciute & Staneviciute (2008)

Crepidostomum serpentinum 2n=14 (1m+5a) Petkeviciute & Staneviciute (2008)

Cercariaeum crassum Wesenberg-Lund, 1934 2n=10 (1m+1sm+2sm-m+1m-sm) Petkeviciute et al. (2011)

CLINOSTOMIDAE

Clinostomum marginatum 2n = 20 Britt (1947)

LECITHODENDRIIDAE

Loxogenes bicolor 2n = 22 Britt (1947)

Acanthatrium pipistrella 2n = 22 Britt (1947)

Ganeo kumaonensis 2n=20; Saksena (1969)

Acanthatrium pipistrella 2n=22; Britt (1947)

Mahroarchis ranarum 2n=22; Saksena (1969)

Pleurogenoides medians 2n=22; 16m + 4st + 2a Barsiene, Grabda-Kazubska (1991c)

Pleurogens claviger

2n=22; 1 2 m + 6 s t + 4a

Barsiene, Grabda-Kazubska (1991c)

Pleurogonidum orientalis

2n=18;

Saksena

(1969)

Prosotocus kashabia

2n=12;

Britt (1947)

Ganeo tigrinum

2n=22

Subramanyam &

Venkat Reddy (1977)

CEPHALOGONIMIDAE

Cephalogonimus americanus

2n = 28

Britt (1947)

GORGODERIDAE

Probolitrema californiense

2n = 12

Markell (1943)

Gorgoderina attenuate

2n = 14

Britt (1947)

Gorgodera amplicava

2n = 16

Britt (1947)

Phyllodistomum folium

2n=18

Petkeviciute et al. (2003)

Gorgodera amplicava

2n=16;

Britt (1947)

Gorgodera pagenstecheri

2n=18; 2m +2sm +2st-a+12а

Barsiene

(1991)

Gorgoderina attenuata

2n=14;

Willey & Koulish

(1950)

Probilotrema californiense

2n=12;

Britt (1947)

Phyllodistomum spatula

2n=16;

Dhingra (1954a)

Phyllodistomum pungitti

2n=18; 2 m + 12st +4a

Britt (1947)

PLAGIORCHIDAE

Eustomas chelydrae

2n = 18

Britt (1947)

Glypthelmins quieta

2n = 18

Britt (1947)

Plagitura salamandra

2n = 22

Britt (1947)

Pneumobites breviplexus

2n = 22

Britt (1947)

Pleumocoeces medioplexus

2n = 22

Pennypacker (1936)

Pleumocoeces similiplexus

2n = 22

Pennypacker (1940)

Pleumocoeces similiplexus

2n = 22

Britt (1947)

Trematorchis ranarum

2n=18

Subramanyam &

Venkat Reddy (1977)

Glypthelmins guieta

2n=18;

Britt (1947)

Plagitura salamandrae

2n=22;

Britt (1947)

Haematoloechus mediplexus

2n=22;

Burton

(1960)

Haematoloechus parviplexus 2n=22; Pennypacker (1936)

Haematoloechus semiplexus 2n=22; Britt (1947)

Haematoloechus similis 2n=22; 12m+6sm + 2sm-m+2st Barsiene, Grabda-Kazubska (1988b)

Haematoloechus asper 2n=22; 14m+4sm + 2sm-m +2st Britt (1947)

Skrjabinoeces sp. 2n=22; 10m + 2sm-m + 4sm +2sm-st +

4st Petkeviciute et al. (1990)

Monodistomum salamandra 2n=20; Britt (1947)

Encylometra colubrimurorum 2n=12; Saksena (1969)

Staphylodora bascaniensis 2n=16; Britt (1947)

2n=20; Sanderson (1959)Haplometra cylindracea

2n=22; 4m + 8 s m-m + 4 sm +6st Barsiene, Grabda-Kazubska (1988a)

Plagiorchis sp. (L. stagnalis,

ПНР) 2n=22; 2m + 8sm-m + 4sm +

+ 8st Barsiene, Grabda-Kazubska (1988b)

Opisthioglyphe ranae 2n=22; 2m + 6sm-m+6sm+ 4st + 4a Barsiene, Grabda-Kazubska (1988a)

Opisthioglyphe ranae (L. stagnalis) 2n=22; 6m +6sm + 2sm-st

+2st +4a-st+2a Petkeviciute et al. (1990)

Leptophallus nigrouenosus 2n=20; 12m + 2sm-m + 2sm +2st + 2a Barsiene, Grabda-Kaka (1988а)

Paralepoderma progeneticum 2n=20; 14m + 2sm + 2st-a + 2a Barsiene, Grabda-Kaka (1991a)

Paralepoderma brumpti 2n=20; 10m+2m-sm + 2sm+ 2st + 4a Petkeviciute et al. (1990)

Omphalometra flexuosum 2n=20; 4sm-m + 4 sm + 4st +8a Barsiene, Grabda-Kazubska (1991a)

RENIFERIDAE

Natriodera verlatum 2n = 22 Britt (1947)

Dasymetra villicoeca 2n = 22 Britt (1947)

Pneumatophilus leidyi 2n = 22 Britt (1947)

Pneumatophilus variabilis 2n = 22 Britt (1947)

Lechriorchis abduscens 2n = 22 Britt (1947)

Renifer ellipticus 2n = 22 Britt (1947)

Neorenifer wardi 2n = 22 Britt (1947)

Neorenifer georgianus 2n = 22 Britt (1947)

Neorenifer aniarum 2n = 22 Britt (1947)

Neorenifer orula 2n = 22 Britt (1947)

Neorenifer drymarchon 2n = 22 Britt (1947)

Neorenifer elongates 2n = 22 Britt (1947)

Staphylodora bascaniensis 2n = 16 Britt (1947)

Auridistomum chelydrae 2n = 18 Britt (1947)

Telorchis robustus 2n = 16 Britt (1947)

Telorchis lobusus 2n = 22 Britt (1947)

Telorchis medius 2n = 22 Britt (1947)

Telorchis corti 2n = 22 Britt (1947)

RHOPALIADIDAE

Rhopalias macracanthus Chandler, 1932 2n = 16 Ciordia (1949)

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DICLYBOTHRIIDAE

Diclybothrium hamulatum (Simer, 1929) Price, 1942

2n = 12

Pickle &Jones (1967)

SPIRORCHIIDAE

Spirorchis magnitestis

2n = 18 (4a/t + 1a + 3sm)

Teehan &

Short (1989)

Spirorchis parvus

2n = 18 (4a/t + 1a + 3sm)

Teehan &

Short (1989)

Spirorchis magnitestis

2n=18; 2m + 16а

Jones &

Mayer

(1953)

Spirorchis parvus

2n=18; 2m + 16st

Grossman

еt al.

(1981b)

Spirorchis sp.

2n=18; 2 sm + 6 st-sm +8t-st + 2t

Teehan &

Short

(1989)

CONVOLUTIDAE

Convolute convolute

2n = 16 (7m-sm + 1st)

Birstein (1990)

Baltalimania agile

2n = 14

Birstein (1990)

MIROPHALLIDAE

Microphallus pygmaeus

2n = 18

Birstein & Mikhailova (1990)

Microphallus piriformis

2n = 18; 8m + 2sm + 4st + 4?

Birstein & Mikhailova (1990)

Microphallus triangulatus

2n = 18

Birstein & Mikhailova (1990)

Microphallus pygmaeus

2n=18; 8m + 2sm + 4st +4?

Britt (1947)

Microphallus triangulatus

2n=18; 8m + 6 s m + 4?

Britt

(1947)

MONORCHIIDAE

2n=18; Dhingra (1955a, 1955b)

2n=20; 6m+2m -sm + 2sm+ 4st + 6a Dhingra (1955a)

2n=20; 12m + 6sm-m + 2 s t-sm Dhingra (1955a)

Asymphylodora sp.

2n=22; 8m + 8sm-m+2sm-

st + 2st Dhingra (1955a)

Palaeorchis sp.2n=14; 2m + 2sm-m+6sm+ 2st + 2a Dhingra (1955a)

Asymphylodora spp.2n = 20 (5m+4sm+1st) Barsiene et al. (1995)

NOTOCOTYLIDAE

Notocotylus attenuates, Notocotylus imbricatus, 2n=20 Petkeviciute & Barsiene (1988)

Notocotylus ephemera 2n=20,21 Petkeviciute & Barsiene (1988)

Notocotylus filamentis 2n=14; Ciordia (1950)

2n=20; 2m + 6sm-m+ 4sm+ 2st + 6a Petkeviciute & Barsiene (1988)

2n=21; Britt (1947)

Notocotylus ephemera

2n=22; Britt (1947)

2n=22; 4m + 4sm-m+4sm+ 8a Britt (1947)Notocotylus attenuatus 2n=20; 4sm + ? Rao & Venkat Reddy (1982)

Notocotylus imbricatus 2n=20; 2 m + 10sm +2st+6a Petkeviciute & Barsiene (1988)

2n=20; 2m + 2m-sm + 4sm+ 4st +8a Petkeviciute et al. (1989а)

2n=21; Britt (1947)

Notocotylus noyeri

2n=20; 4m + 4sm +4st +8t Barsiene & Grabda-Kazubska (1991b)

2n=20; 2m +6sm+4 st+8a Barsiene et al. (1990)

2n=21; Britt (1947)

Notocotylus sp. (Anisus acronicus,)

2n=21-30; Britt (1947)

CYCLOCOELIDAE

Cyclocoelium oculeum 2n=20;4m + 6sm + 8st + 2a Taft & LeGrande (1979)

Cyclocoelum bivesiculatum 2n=20; Dhingra (1954a)

FASCIOLIDAE

Fasciola gigantica 2n=20 (2sm+1t+7st) Romanenko & Pleshanova (1975); Subramanyam &

Venkat Reddy (1977)

Fasciola gigantica 2n=20 (6sm+1m-sm+3st) Venkat Reddy & Subramanyam (1973);

Subramanyam & Venkat Reddy (1977)

Fasciola hepatica 2n=20 (1sm-m+5st+2sm+1m+1sm) Romanenko & Pleshanova (1975)

Fasciola hepatica 2n=20 (5sm+4st+1t) Li et al. (1988)

Fasciola hepatica 2n=20 (1sm-m+4st+5sm) Spakulova & Kralova (1991)

Fasciola hepatica 2n=20 (1m+5sm+4st) Reblanova et al. (2011)

Fasciola sp.2n=20 Sakaguchi & Wakako (1976); Sakaguchi (1980);

Moriyama et al. (1979); Rhee et al. (1987); Yin & Ye

(1990)

Fasciola sp. 3n=30 (8sm+2st) Sakaguchi and Nakayama (1975); Sakaguchi &

Wakako (1976);

Fasciola gigantica 2n=20 Sakaguchi (1980)

Fasciola hepatica 2n=12

2n=22 (1sm-m+1sm+9st) Srimuzipo et al. (2000);

Henneguy (1902); Schubmann (1905); Schellenburg

(1911)

Fascioloides magna 2n=22 (9st+1sm-m+1sm) Reblanova et al. (2010)

Fasciolopsis buski 2n = 14 (6m+1t) Gao (1985)

Fasciolopsis buski 2n = 14 (4m+2sm+1t) Dai (1990)

Parafasciolopsis fasciolaemorpha 2n=20 (1m+1t+6st+2sm) Barsiene (1990)

3n=30 Britt (1947)

2n=20/30; Britt (1947)

2n=20; 2m + 10sm + 8st Rhee et al. (1987b)

2n=20/30; Sakaguchi, Yoneda (1976)

2n=20; Britt (1947)

Fasciola sp.

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3n=30; 3m + 12sm+ 15st Rhee et al. (1987b)

2n=20 Sanderson (1953, 1959)Fasciola hepatica 2n=20; 6 m + 2 sm + 1 2 st Li et al. (1988)

2n=20; 4sm+4sm-s t + 12st

Moriyama et al.

(1979)

2n=20; 2m + 12sm + 6st

Subramanyam &

Venkat Reddy (1977)

3n=30;

Sakaguchi

(1980)

Fasciola giganticа

2n=20/30;

Mariyama et al.

(1979)

PHILOPHTHALMIDAE

Philophthalmus gralli

2n=20

Grossman & Cain (1981)

Philophthalmus sp.

2n=20

Venkat Reddy &

Subramanyam (1971)

Philophthalmus sp. (Georgia, USSR; Bulgaria)

2n=20

Mutafova et al. (1986)

Philophthalmus megalurus

2n=20;

Kahlil & Cable

(1968)

Philophthalmus indieus 2n=20; 2sm + 2 a + 1 6t Subramanyam & Venkat Reddy (1977)

Philophthalmus hegeneri 2n=20; Fried (1975)

Philophthalmus sp.2n=20; 2sm + 18a Mutafova (1983b)

Philophthalmus gralli 2n=20; 8sm + 12a LoVerde (1978)

HEMIURIDAE

Isoparorchis euritremum; Isoparorchis hypselobargi 2n=20 (XY) Chattopadhyay & Manna (1987)

Halipegus occidualis 2n=18; Jones (1956); Guilford1(1961)

Halipegus eccentricus 2n=22; Guilford (1961)

Isoparorchis eurytreum 2n=18; Srivastava & Iha (1964b)

2n=20; 10sm + 8a + 2XY

( X = sm; Y = a) Chattopadhyay & Manna (1987); Dhingra (1954b)Isoparorchis hypselobagri

2n=18; 4m + 14a Srivastava & Iha (1964b); Iha (1975)

DIPLOSTOMATIDAE

Diplostomum indisticum; Diplostomum mergi;

Diplostomum pseudospathaceum; Diplostomum

spathaceum

2n=20 Romanenko & Shigin (1977); Mutafova &

Niewiadomska (1988)

Tylodelphys clavata 2n=20 Romanenko & Shigin (1977)

Diplostomum sp. 1 2n=20; 6m + 4sm-m + 2st-

sm + 8 s t Barsiene & Staneviciiite (1991)

Diplostomum sp. 2 2n=20; 6m+2sm-m + 4sm

+ 4st + 2st- a + 2a Britt (1947)

Diplostomum baeri 2n=20; 2m + 2sm-m + 6sm

+ 6st + 4a Barsiene еt аl. (1990а); Barsiene & Staneviciute

(1991)

Diplostomum mergi 2n=20; 10m + l0t Romenenko & Shigin (1977)

Diplostomum pseudospathaceum 2n=20; 6m + 4sm + 6st + 4a Barsiene & Staneviciute (1991) Barsienеet al. (1991)

Diplostomum paracaudum 2n=20; 6m+6sm +8st Barsienе et al. (1990a)

Barsiene & Staneviciute (1991)

2n=20; 8m + 2m-sm + 8st + 2? Romenenko &Shigin (1977)Tylodelphys clavata

2n=20; 6m + 2sm-m + 4st+ 2st-a + 6 a Barsiene (1991c)

Proalorioides tropidonotis 2n=16; Saksena (1969)

Posthodiplostomum cuticola 2n=20; 4 sm + 6 st + l0t Barsiene (1991c)

STRIGEIDAE

Gogatea serpentium indica 2n=16 Subramanyam & Venkat Reddy (1977)

Ichthyocotylurus erraticus (Rudolphi, 1809) 2n=20 (4m+2sm+1sm-st+3st-a) Bell et al. (1998)

Ichthyocotylurus variegates (Creplin, 1825) 2n=20 (4m+1sm+1m-sm+4st) Bell et al. (1998)

Apatemon gracilis (Rudolphi, 1819) 2n=20 (3m+1m-sm+3sm-

st+1sm+1a+1st-a) Bell et al. (1998)

Apatemon gracilis 2n=20 (3m+3sm-st+1sm+2a+1st-a) Petkeviciute & Staneviciute (1999)

Cotylurus cornutus (Lymnaea zazuriensis) 2n=20; 2m + 6st + 2a-s t + 1 0a Barsiene et al. (1990)

Apatemon gracilis (Lymnaea ovata) 2n=20; 6m + 4sm + 4sm-st +2st + 4a Petkeviciute (1991)

2n=20; 2 m + 6 s m + 4st + 2a-st + 6a Barsiene (1992)Apatemon minor (Planorbarius planorbis)

2n=21;

Apatemon fuligulae 2n=20; 4 m + 8 s m + 4 s t + 6a Barsiene et al. (1990)

OPISTHORCHIIDAE

Opisthorchis felineus 2n=14 Romanenko (1973)

Opisthorchis felineus 2n=14 (4sm+3m) Polyakov et al. (2010)

Clonorchis sinensis 2n=56 (3m+1m-sm+16sm+8st) – Korea

(2m+2m/sm+16sm+8st) – China Park et al. (2000)

Clonorchis sinensis 2n=56 Park & Young (2003)

Opisthorchis felineus (Rivolta, 1884) 2n=14 (2m/sm+5) Zadesenets et al. (2012)

Opisthorchis viverrini (Poirier, 1886) 2n= 12 (2sm+1sm+1sm/st+1st/a +1a) Zadesenets et al. (2012)

Metorchis xanthosomus (Creplin, 1846) 2n=14 Zadesenets et al. (2012)

Metorchis billis (Braun, 1893) 2n=14 Zadesenets et al. (2012)

Clonorchis sinensis (Cobbold, 1875) 2n=14, 2n=56 Zadesenets et al. (2012)

Opisthorchis viverrini 2n=12 (4m+1sm+1a) Kaewkong et al. (2012)

TRANSVERSOTREMATIDAE

Transversotrema patialense 2n=20 Madhavi & Ramanjaneyulu (1986)

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OMPHALOMETRIDAE

Rubenstrema exasperatum 2n=16 (3m+4sm-m+1sm(X1)+1st(X) Mutafova &Kanev (1996)

NEODIPLOSTOMATIDAE

Neodiplostomum seoulense 2n=20 (2m+5sm/st+3t) Park et al. (1998)

ASPIDOGASTREA

Aspidogaster conchicola 2n=10 (1st+4a) Petkeviciute (2001b)

Cotylogaster occidentalis 2n=12 (2m+2sm+2a) Loverde & Fredericksen (1978)

Cotylapis insignis 2n=22 Loverde & Fredericksen (1978)

DIPLOZOIDAE

Diplozoon paradoxum 2n=8 (3m+1a) Koskova et al. (2011)

Paradiplozoon bliccae 2n=14 (7a) Koskova et al. (2011)

Paradiplozoon sapae 2n=14 (7a) Koskova et al. (2011)

Paradiplozoon nagibinae 2n=14 (7a) Koskova et al. (2011)

Eudiplozoon nipponicum 2n=7 Koroleva (1968b)

Paradiplozoon megan 2n=7 Koroleva (1968b)

Diplozoon paradoxum 2n=8, (3m+1a) Koroleva (1968a,b)

Paradiplozoon bliccae (syn. Diplozoon gussevi) 2n=14, (7a) Koroleva (1968a,b)

Paradiplozoon bliccae (syn. Diplozoon markevitchi) 2n=14, (7a) Koroleva (1968b, 1969)

Paradiplozoon sapae 2n=14, (7a) Koroleva (1969)

Paradiplozoon nagibinae 2n=14, (7a) Koroleva (1969)

Paradiplozoon pavlovskii 2n=14, (7a) Koroleva (1968a,b)

Paradiplozoon homoion 2n=14, (7a) Koroleva (1968a,b)

Diplozoidae sp. 2n=14, (7a) Bovet (1967)

Diplozoidae sp. (sp. n.) 2n=10, (2m+3a) Koroleva (1969)

Diplozoidae sp. 2n=7 Baer & Euzet (1961)

Diplozoidae sp. 2n=7 Bovet (1967) Incorrect data

according to Koroleva (1968b)

PSILOSTOMIDAE

Psilotrema sp.2n=16; 4m + 2sm-m + 2sm+ 8st Britt (1947)

Sphaeridiotrema globulus 2n=14; 4m + 4sm-m + 4s m+ 2a Britt (1947)

SANGUINICOLIDAE

Sanguinicola sp.2n=22; 1 6 m + 2 sm + 4st Britt (1947)

BRACHYLAEMIDAE

Leucochloridiomorpha constantiae 2n=16; Filippone & Fried (1974)

CYATHOCOTYLIDAE

Gogotea serpentium 2n=16; Saksena, 1969, Subramanyam, Venkat Reddy (1977)

CRYPTOGONIMIDAE

Acetodexira amiuri 2n=12; Perkins (1956)

Atrophecaecum bur minis 2n=14; 14sm Madhavi & Ramanjaneyulu (1988)

OPECOELIDAE

Sphaerostoma bramae 2n=24; Gresson (1958)

EUCOTILIDAE

Cercaria pectinata 2n=12; 6m + 2st+4m-sm Ieyama & Ozaki (1987)

Cercaria tapidis 2n=16; 8m + 2st-sm + 2st +2sm-st +2sm-

mBritt (1947)

Sm =sub-metacentric; a = acrocentric; m =metacentric. t =telocentric. X =sex chromosome. Y =sex chromosome. ZZ =sex chromosomes. ZW =sex chromosomes.

Chromosomes of Cestoda

Studies of the chromosomal patterns among

the Cestoidea have not been numerous. Most

of the recent ones have originated from one

laboratory, and there seems to have been no

broad sampling of the Class as a whole. No

records of studies on members of the

Cestodaria have come to our attention, and

attempts in this laboratory have not been

successful in obtaining reliable results. Among

the Cestoda only one of the nine commonly

recognized orders, the Cyclophyllidea, seems

to have been examined for chromosome

numbers. Among the cyclophyllideans the

family Hymenolepididae is represented by

records from four genera (8 species). These

indicate that in Diorchis (ralli and reynoldsi)

and in Protogynella (blarinae), the diploid

number of chromosomes is 10. One species of

Hymenolepis (H. fraterna) has 10

chromosomes but in 3 other species (H.

ananthocephalus (van Gundy, 1935), H.

diminuta (Rudolphi, 1819), and H. serpentulus

(Goeze, 1782) the number is 12. Twelve

chromosomes are also present in an

unidentified species of Aploparaksis. H.

serpentulus is represented by two sub-races

(sturni and turdi), each with the same number

of chromosomes (12), but the two sets of

chromosomes show such consistent and easily

129

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

recognizable differences that the races can be

readily separated on the basis of cytological

grounds alone, without reference to collection

records. The Hymenolepididae, while not

having a common basic number of

chromosomes, are cytologically quite uniform.

The same may not be said for the Dilepididae.

For example, Dipylidium caninum (Linnaeus,

1758) has the diploid number of 10

chromosomes; Liga brasiliensis has 14

chromosomes, and unnamed species of

Anonchotaenia and Choanotaenia each have

1 6 c h r o m o s o m e s . A m o n g t h e

Anoplocephalidae, Avitellina centripunctata

(Rivolta, 1874) has somatic chromosomes,

Oochoristica has 10, and Moniezia expansa

(Rudolphi, 1805) and M. planissima (Moniez,

1879) have 12. For the Taeniidae records of 2n

= 16 for Hydatigera taeniaeformis (Batsch,

1786) and 2n = 20 for Taeniarhynchus

saginatus (Goeze, 1782) and Taenia serrata=

pisiformis (Bloch, 1780) are found, although

some investigators, including the present

author, find only 16 chromosomes in the two

latter species. Among the Nematotaeniidae,

two races of Baerietta desmognathi (Douglas,

1957) have been studied, one race having 2n =

8 and the other 2n = 16 (Table 3). This

difference may be an example of polyploidy

but examination of a greater series of

specimens would be necessary before any

definite statement should be made. It would

appear from the above records that the Cestoda

do not show any completely definite

taxonomic pattern of chromosomal numbers

above the species level. Perhaps such evidence

would be forthcoming following a broader

sampling among the Cestodaria and the

Cestoda. We believe, on other grounds, that the

Cyclophyllidea are probably the most

specialized of the Cestoda, but study of

chromosome numbers, structure, or behavior,

has not as yet seemingly substantiated such a

conclusion.

Karyology represents a conspicuous gap in the

phylogenetic evaluation of the Cestoda and of

other flatworms, despite the fact that

chromosome structure and gene location are of

evolutionary relevance. Cytogenetic features,

alone or in concert with other modern

character-based approaches, might provide

information not only on phylogeny but also on

systematic interrelationships within the target

group. Unfortunately, only nine out of 16

eucestode orders and up to 2% or 115 known

species have been studied karyologically. Most

early cytogenetic studies have been

exclusively focused on the number of

chromosomes; 74 species (63.5%) have been

studied for chromosome morphology.

Table 3. Summary of chromosomes and karyotype data of Cestoda (Tapeworms) (1907–Till Date).

Order/ Family Number

2n [3n] Morphology References

I. SPATHEBOTHRIIDEA

Acrobothriidae

Cyathocephalus truncates (Pallas, 1781)

2m+2sm+5a

Petkeviciute (1996a)

II.DIPHYLLOBOTHRIIDEA

Diphyllobothriidae

12-16

Smyth (1946)

Schistocephalus solidus (Muller, 1776)

5m+4a

Petkeviciute (1996b)

Diphyllobothrium dendriticum (Nitzsch, 1824)

18 (9–18)

7m+2sm

Wikgren & Gustafsson (1965)

18 (8–22)

7m + 2sm

Wikgren &

Gustafsson(1965)Diphyllobothrium ditremum (Creplin, 1825)

(=Diphyllobothrium osmeri)

7m + 2sm

Petkeviciute (1992)

Diphyllobothrium latum (Linnaeus, 1758)

18 (15–28)

7m + 2sm

Wikgren &

Gustafsson (1965)

Diphyllobothrium ursi Rausch, 1954 18 Wolcott (1959)

Ligula intestinalis (Linnaeus, 1758) 18 6m +3sm Petkeviciute (1992)

130

2n [3n]

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Chromosomes and cytogenetics of helminths

Spirometra erinaceieuropaei

(Rudolphi, 1819) (= Diphyllobothrium erinacei)

[27]

Sasada (1978)

Spirometra mansonoides (Muller, 1935) (= S. mansoni)

[27]

Liu & He (1989)

III. CARYOPHYLLIDEA

Balanotaeniidae

Balanotaenia bancrofti Johnston, 1924

Grey (1979)

Capingentidae

Capingens singularis Hunter, 1927

Grey (1979)

Caryophyllaeidae

3m + 1sm +3a

Mackiewicz &

Jones (1969)

Hunterella nodulosa

Mackiewicz et McCrae, 1962

3m + 4a

Grey (1979)

Archigetes sp. (=appendiculatus)

Motomura (1929)

Biacetabulum biloculoides Mackiewicz et McCrae, 1965

Grey (1979)

[30]

Grey (1979)

20 [30]

10m

Petkeviciute &

Kuperman (1992)

Caryophyllaeus laticeps (Pallas, 1781)

10m

Bombarova et al. (2009)

Glaridacris laruei Lamont, 1921

3m + 1sm + 4a

Grey &Mackiewicz (1974), Grey (1979)

Glaridacris confusus Hunter, 1927

Grey (1979)

Glaridacris catostomi Cooper, 1920

20 [30]

8m + 2sm

Grey (1979), Grey & Mackiewicz (1980)

Glaridacris vogei Mackiewicz, 1976

8m + 1sm + 1a

Grey (1979)

Monobothrium hunter Mackiewicz, 1963

9m + 1a

Grey (1979)

Isoglaridacris folius Fredrickson et Ulmer, 1965

1m + 8a

Grey (1979)

Isoglaridacris jonesi

Mackiewicz, 1972

2m + 7a

Grey (1979)

Isoglaridacris bulbocirrus Mackiewicz, 1965

18 [27]

Grey (1979)

Archigetes appendiculatus

Motomura (1929)

Lytocestidae

[24]

4m + 3a+ 1 minute

Jones & Mackiewicz (1969)

Atractolytocestus huronensis Anthony, 1958

[24] 4m + 3a + 1

minute Kralova-Hromadova et al. (2010)

Caryoaustralus sprenti Mackiewicz et Blair, 1980 6 Grey (1979)

Khawia iowensis Calentine et Ulmer, 1961 16 5m + 3a Grey (1979)

Khawia rossittensis (Szidat, 1937) 16 Grey (1979)

Khawia sinensis Hsu, 1935 16 3m + 5a Petkeviciute (1998)

3m + 5a

Mutafova & Nedeva (1999)

Khawia saurogobii Xi et al., 2008

3m + 5a

Orosova et al. (2010b)

Lytocestus indicus (Moghe, 1925)

Vijayaraghavan & Subramanyam (1977)

Bombarova et al. (2009)Caryophyllaeides fennica (Schneider, 1902)

10m

Orosova et al. (2010a)

Notolytocestus minor Johnston et Muirhead, 1950

Grey (1979)

IV. TRYPANORHYNCHA

Lacistorhynchidae

Lacistorhynchus tenuis (Van Beneden, 1858)

8m or sm

Jones (1954)

V. BOTHRIOCEPHALIDEA

Triaenophoridae

Triaenophorus crassus Forel, 1868

7m + 1sm + 1a

Petkeviciute &Ieshko (1991)

Triaenophorus nodulosus (Pallas, 1781)

5m + 7sm + 1a

Petkeviciute & Ieshko (1991)

Bathybothrium rectangulum (Bloch, 1782)

8m + 1sm

Spakulova & Scholz (1999)

Eubothrium crassum (Bloch,

1779) 16 3m + 2sm + 3a Petkeviciute & Bondarenko (2001)

Eubothrium rugosum (Batsch, 1786) 16 3m + 2sm + 3a Petkeviciute & Kuperman (1991)

Eubothrium salvelini (Schrank, 1790) 16 2m + 3sm + 3a Petkeviciute & Bondarenko (2001)

Bothricephalidae

Bothriocephalus gregarious Renaud, Gabrion et Pasteur,

1983 14 4m + 3a Petkeviciute (2003)

Bothriocephalus acheilognathi Yamaguti, 1934 14 6m + 1m-sm Nedeva & Mutafova (1988)

Bothriocephalus claviceps (Goeze, 1782) 14 7m Petkeviciute (2003)

12 Bazitov (1978)Bothriocephalus scorpii Muller, 1776

Order/ Family Number

Morphology References

131

Moniezia expansa (Rudolphi, 1805)

probably

12 or 14

Child (1907)

Oochoristica sp.

10 small

Jones (1945)

Taeniidae

18 Lukashenko et al. (1965)

18 Sakamoto et al. (1967)

18 4sm + 5a Rausch & Rausch (1981)

Echinococcus multilocularis Leuckart, 1863

18 1sm + 8sm-a Mutafova & Svilenov (1985)

18 1sm +8sm-a Smyth (1962)Echinococcus granulosus (Batsch, 1786) Mutafova & Svilenov (1985)

Echinococcus vogeli Rausch et Bernstein, 1972 18 Rausch & Rausch (1981)

Taenia macrocystis (Diesing, 1850) 18 Rausch & Rausch (1981)

Taenia (= Hydatigera) taeniaeformis Batsch, 1786 16 Jones & Ciordia (1956)

Taenia rileyi Loewen, 1929 18 Rausch & Rausch (1981)

Taenia crassiceps (Zeder, 1800) 16 2m + 6a Smith et al. (1972)

T. crassiceps (ORF strain) 14 1m + 6a Smith et al. (1972)

20 Jones & Ciordia (1956)

16 Walton (1959)

Taenia pisiformis (Bloch, 1780)

18 Romanenko & Movsessian (1988)

18 7m + 1sm + 1a Liu & He (1987b)Taenia hydatigena Pallas, 1766

5m + 1sm

Petkeviciute (2003)

Tetracampos ciliotheca Wedl, 1861 (=Polyonchobothrium

clarias (Woodland, 1925))

4m + 2sm

Badawy and Noor El-Din (1998)

Bothriocephalus acheilognathi Yamaguti, 1934

6m + 1m-sm

Sofi & Ahmad (2014)

VI. TETRAPHYLLIDEA

Phyllobothriidae

Pelichnibothrium speciosum Monticelli, 1889

1m + 1sm + 6a

Petkeviciute & Regel (1993)

VII. PROTEOCEPHALIDEA

Proteocephalidae

Acanthotaenia sp. (= A. multitesticulata)

Vijayaraghavan &

Subramanyam (1980a)

Glanitaenia

(= Proteocephalus) osculate (Goeze, 1782)

1m + 3sm + 4a

Petkeviciute (2001a)

Proteocephalus longicollis (Zeder, 1800) (=Proteocephalus

exiguus)

4m + 5sm

Hanzelova et al. (1995)

Proteocephalus macrocephalus (Creplin, 1815)

7m + 2sm

Scholz et al. (1997)

6m + 1sm + 2a

Spakulova & Hanzelova(1992)Proteocephalus percae (Muller, 1780)

2m + 7sm

Petkeviciute (1993)

VIII. NIPPOTAENIIDEA

Nippotaeniidae

7m + 7sm

Bombarova et al. (2005)

Nippotaenia mogurndae Yamaguti et Miyata, 1940

Bombarova et al. (2009)

IX. CYCLOPHYLLIDEA

Mesocestoididae

Mesocestoides vogae Etges, 1991 (=Mesocestoides corti)14 5m + 2sm Raghunathan & Voge (1974)

Nematotaeniidae Cylindrotaenia (= Baerietta)

desmognathi (Douglas, 1957) 8, 16 Douglas (1957)

Cylindrotaenia (= Baerietta) diana (Helfer, 1948) 28 Douglas (1963); diploid number assessed

Distoichometra bufonis Dickey, 1921 (=Distoichometra

kozloffi)16 Douglas (1963); diploid

number assessed

Nematotaenia dispar (Goeze, 1782) 28 Vijayaraghavan & Subramanyam (1980b)

Anoplocephalidae

Avitellina centripunctata (Rivolta, 1874) 8 Walton (1959)

Moniezia benedeni (Moniez, 1879) (=Moniezia planissima) probably

12 or 14 Child (1907)

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

Order/ Family Number

Morphology References

132

12 2m + 3sm + 1a Liu & He (1987a)

12 1m-sm + 5a Mutafova & Gergova (1994)

Microsomacanthus spasskii Tolkacheva, 1965 6 3m Petkeviciute & Regel (1994)

Microsomacanthus spiralibursata (Czaplinski, 1956) 6 2m + 1m-sm Petkeviciute & Regel (1994)

Passerilepis crenata (Goeze, 1782) (= Hymenolepis

serpentulus sturni = H. serpentulus turdi)12 4m + 2? Jones (1945)

Protogynella blarinae Jones, 1943 10 Jones (1945)

Pseudodiorchis (= Diorchis) reynoldsi (Jones, 1944) 10 1sm + 4a Jones (1945)

Retinometra giranensis (Sugimoto, 1934) 12 Petkeviciute & Regel (1994)

Rodentolepis erinacei (Gmelin, 1790) 12 6a Mutafova & Gergova (1994)

10 1m + 4a Jones (1945)Rodentolepis fraterna (Stiles, 1906)

12 6a Jones & Ciordia (1955)

12 6a Hossain & Jones (1963)Rodentolepis microstoma (Dujardin, 1845) 12 6a Proffitt & Jones (1969)

Rodentolepis myoxi (Rudolphi, 1819) 12 3m + 3sm Casanova et al. (2000)

Rausch & Rausch (1981)

Romanenko & Movsessian (1988)

18 (12-22)

Movsessian & Margarian (1991)

Walton (1959)

Taenia saginata Goeze, 1782

10a

Jones & Wyant (1957)

Paruterinidae

Anonchotaenia sp.

1m + 7

Jones (1945)

Anonchotaenia globate (Von Linstow, 1879)

2m + 1a + 3

Jones (1945)

Francobona (= Rhabdometra) similis (Ransom, 1909)

2m + 1a + 3?

Jones (1945)

Dipylidiidae

Jones (1945)

4m-sm+ 1a + 3?

Bovt (1973)

4m + 1sm + 3a

Liu & He (1988)

Dipylidium caninum

(Linnaeus, 1758)

4sm + 6

Margarian (1989)

Davaineidae

Cotugnia meggitti Yamaguti, 1935

Gupta & Greval (1971)

Davainea proglottina (Davaine, 1860)

probably 9a

Jones (1951)

Raillietina echinobothrida (Megnin, 1880)

6m +1sm + 2?

Margarian (1989)

Raillietina tetragona (Molin, 1858)

Margarian (1989)

Skrjabinia caucasica Petrochenko et Kireev, 1966

4m + 4sm

Margarian (1989)

Dilepididae

Anomotaenia bacilligera (Krabbe, 1869)

6m + 2m-sm

Petkeviciute et al. (2006)

Choanotaenia sp. 16 7sm + 1a Jones (1945)

Dilepis undula (Schrank, 1788) 18 4m + 3sm + 2a Petkeviciute et al. (2006)

Liga brasiliensis (Parona, 1901) 14 6sm + 1a Jones (1945)

Molluscotaenia crassiscolex

(von Linstow, 1890) 12 5m + 1sm Petkeviciute et al. (2006)

Hymenolepididae

Anatinella spinulosa (Dubinina, 1953) 12 Petkeviciute & Regel (1994)

Aploparaksis sp. 12 Jones (1945)

Aploparaksis brachyphalos (Krabbe, 1869) 12 Petkeviciute & Regel (1994)

Aploparaksis filiformis Spasskii, 1963 12 Petkeviciute & Regel (1994)

Aploparaksis furcigera (Rudolphi, 1819) 12 Petkeviciute & Regel (1994)

Aploparaksis occidentalis Prudhoe et Manger, 1967 12 Petkeviciute & Regel (1994)

Aploparaksis retroversa Spasskii et Gubanov, 1961 12 Petkeviciute & Regel (1994)

Cryptocotylepis (=Hymenolepis) anthocephalus (van

Gundy, 1935) 12 1sm + 1a + 4? Jones (1945)

Dicranotaenia fallax (Krabbe, 1869) 12 Petkeviciute & Regel (1994)

Diorchis ralli Jones, 1944 10 2m + 1a + 2? Jones (1945)

Fimbriaria sp. 10 Petkeviciute (2002)

Hymenolepis citelli McLeod, 1933 12 6a Ward et al. (1981)

12 Jones (1945)

12 6a Kisner (1957)

Hymenolepis diminuta (Rudolphi, 1819)

12 Douglas (1962)

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Chromosomes and cytogenetics of helminths

Order/ Family Number

Morphology References

133

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

12 1m-sm + 5a Jones & Ciordia (1955)

12 1m + 5a Mutafova & Gergova (1994)

Rodentolepis nana (Siebold, 1852)

12 6a Goldschmidt et al. (2006)

Rodentolepis straminea (Goeze, 1782) 12 2m + 2sm + 2a Spakulova & Casanova (1998)

Sobolevicanthus Mastigopraedita (Polk, 1942) 12 1m + 1a-sm +4a Petkeviciute & Regel (1999)

Sobolevicanthus spasskii Kornyushin, 1969 12 1m + 1a-sm +4a Petkeviciute & Regel (1999)

Wardium fryei Mayhew, 1925 12 6a Bondarenko & Petkeviciute (1998)

Wardium retracta (Von Linstow, 1905) 12 Petkeviciute & Regel (1994)

Dioecocestidae Gyrocoelia pagollae Cable et Meyers, 1956 12 Coil (1972)

8 Coil (1970)Shipleya inermis Fuhrmann, 1908 8 2sm + 2a Rausch & Rausch (1990)

Order/ Family Number

Morphology References

Sm = sub-metacentric; a =acrocentric; m =metacentric.

Chromosomes of Nematoda

Examination of over 87 species of nematodes,

together with later observations by other

workers gives available data on 1 genus and 1

species of the Desmadoridae, 1 genus and 4

species of the Rhabditidae, 2 genus and 8

species of the Rhabdiasidae, 3 genus and 5

species of the Strongyloididae, 9 genera and 24

species or subspecies of the Ascaridae, 1 genus

and 3 species of the Anisakidae, 1 genus and 1

species of the Kathlaniidae, 2 genera and 2

species of the Oxyuridae, 2 genera and 6

species of the Heterakidae, 3 genera and 5

species of the Strongylidae, 1 genus and 1

species of the Heligmosomidae, 2 genera and 2

species of the Trichostrongylidae, 3 genera and

4 species of the Metastrongylidae, 1 genus and

1 species of the Onchocercidae 1 genus and 1

species of the Diplogastridae 1 genus and 1

species of the Camallanidae, 1 genus and 1

species of the Rhabdochonidae, 1 genus and 1

species of the Acuariidae, 2 genera and 2

species of the Spiruridae, 3 genera and 3

species of the Physalopteridae, 1 genus and 1

species of the Setariidae, 1 genus and 1 species

of the Cosmocercidae, 1 genus and 1 species of

the Spirocercidae, 1 genus and 1 species of the

Parasitaphelenchidae, 1 genus and 1 species of

the Gordiidae, 1 genus and 1 species of the

Chordodidae, 1 genus and 4 species of the

Trichinellidae and 1 genus and 1 species of the

Trichosomoididae (Table 4). Over one-half of

these forms show a somatic number of

chromosomes greater than the number present

in the zygote or in the "stem" cells of the

embryo, and supports the postulation that the

germ-cell-number is composed of compound

chromosomes in many instances. Some

authors, however, interpret this phenomenon

as indicative of fragmentation during the

formation of the somatic chromosomes (in

Parascaris equorum (Walton 1924)) the

diploid number may be 2 or 4 or 6 depending

upon the variety being studied, and the somatic

number of chromosomes may amount to rather

high figures). “Diminution” or the loss of

chromatic material is known to occur in at least

9 species of nematodes, as well as occurring in

other phyla such as the Protozoa, the

Arthropoda, and the Chordata. This loss has

been interpreted as an attempt to stabilize the

embryonic environment of the "stem" or

"germ-track" cells during development

(Ubisch, 1943). It is known that DNA is

released from the nucleus at this time and such

more-or-less universal release of nuclear

chemicals may on occasion have a physical

manifestation (particulates). Such a release

may also be associated in some manner with

the so-called "chromosome fragmentation"

noted in some somatic cells. Lin, 1954,

suggests a comparison of this elimination of

chromatic material (heterochromatin is mainly

DNA) from the collective germ-cell

chromosomes to the macronuclear phenomena

noted in so many protozoa. Examination for

the presence or absence of recognizable "sex

chromosomes" (heterochromosome) has

afforded records indicating that 37 species of

nematodes rather definitely show the presence

134

Family Species No. and Morphology of

Chromosomes Reference

COSMOCERCIDAE

Cosmocerca kashmirensis Fotedar, 1959

2n=16

Fotedar et al. (1973)

TRICHINELLIDAE

Trichinella nelsoni

2n=6 (female)

2n=5 (male)

Mutafova & Komandarev (1976)

Trichinella spiralis

2n=6

Thomas (1965)

Trichinella spiralis

2n=6 (female)

2n=5 (male)

Mutafova et al. (1982)

Trichinella pseudospiralis

2n=6 (female)

2n=5 (male)

Mutafova et al. (1982)

Trichinella nativa 2n=6 (female)

2n= 5 (male) Mutafova et al. (1982)

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Chromosomes and cytogenetics of helminths

of an "X" chromosome-sometimes appearing

as a complex of 2 or more members in certain

species-and in 6 species the presence of a "Y"

chromosome (however in only one species,

Contracaecum incurvum, has subsequent

work substantiated the presence of such a "Y"

element). Jeffrey and Haertl's 1938 objection

to interpreting apparently unpaired and

l a g g i n g c h r o m o s o m e s a s be i n g

heterochromosomes has not held up in the light

of more investigations (Nigon & Robert, 1952;

Lin, 1954). Heterochromosomes may appear

in forms in which the sperm acts only as a

stimulating agent (true fertilization never

taking place) and also in parthenogenetic

species (Nigon & Roman, 1952, in

Strongyloides ratti). From the taxonomic

standpoint, the number and behavior of the

chromatic elements seem to have a great deal

of uniformity within species of a single genus,

and even among species of closely related

genera, and therefore may be of some

significance. In the first place the so-called

"diminution" phenomenon is found only in

members of the Ascaridata (Nematoda),

primarily among the Ascaridae. Secondly, the

number of the so-called "X"-bodies is greater

than 1 only in the Ascari data ("X" may be

composed of 1, 2, 5, 6, or 8 units). The presence

of an undoubted "Y"-body is reported only

from the Ascaridata (Contracaecum incurvum

Good-rich, 1916). Thirdly, there is a certain

consistency of total chromosome numbers in

various groups. The Strongyloididae have the

reduced number of 3; the Rhabdiasidae all

show the haploid number of 6; the Rhabditidae

as a rule have n = 7, although one form has n =

9; 11 of the 12 species of the Strongylata show

n = 6 (one has n = 8); the Heterakidae seem to

group around n = 5, and the Oxyuridae around

n = 4 (occasionally 8). Most of these groups are

quite homogeneous and the similarity of

chromosome numbers could be expected; on

the other hand, the Ascaridata are much more

heterogeneous in composition and the wide

variation in chromosome numbers should

equally be expected. These variations as found

within a closely knit group have been

interpreted as examples of fragmentation or

duplication of one or more chromosomes

(aneuploidy) rather than as cases of

duplication of sets of chromosomes

(polyploidy). The closely related

Nematomorpha show a somewhat similar

situation in that members of the genus Gordius

show n= 2 or n = 4. Paragordius, however, has

n = 7; a situation not in keeping with the

interpretation of close relationships being

indicated by the number of chromosomes

present.

When one turns to the Platyhelminthes, one

finds a somewhat similar picture, although this

phylum is far less homogeneous than the

Nematoda, with such variant groups as the

non-parasitic Turbellaria and the parasitic

Trematoda and Cestoda.

Table 4. Chromosome Number of Nematoda from 1886 Till Date.

135

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

Family Species No. and Morphology of

Chromosomes Reference

ASCARIDIDAE

Ascaris lumbricoides (=suis) 2n=48 Homedes (1933)

Neoascaris vitulorum 2n=18 Homedes (1933)

Ophidascaris filaria 2n=14 Walton (1959)

Parascaris equorum, var. 1 2n=18 Li (1937)

Parascaris equorum, var. 2 2n=12 Li (1934)

Parascaris equorum, var. 3 2n=6 Walton (1924)

Parascaris equorum, var. 4 2n=4 Walton (1924)

Parascaris equorum, var. 5 2n=2 Walton (1924)

Toxacara cati

2n=18 Walton (1924)

Toxacara canis 2n=36 Walton (1924)

Toxacara vulpis 2n=24 Walton (1924)

Toxoscaris leonina 2n=40 Mutafova (1995)

Baylisascaris transfuga 2n=36 Mutafova (1995)

Hexametra sp.

2n=22 Mutafova (1995)

Toxocara canis

2n=22 Mutafova (1995)

Toxocara cati

2n=22

Mutafova (1995)

Ascaridia galli

2n=10

Mutafova (1995)

Ascaridia compare

2n=10

Mutafova (1995)

Ascaridia dissimilis

2n=10

Mutafova (1995)

Ascaris suum

2n=48

Mutafova (1975)

Ascaridia galli Schrank, 1788

2n = 10 (Female)

2n=9 (Male)

Mutafova (1976)

Ascaridia dissimilis (Vigueras, 1931)

2n=10 (Female)

2n=9 (Male)

Mutafova (1976)

Ascaris megalocephala

2n=4

Merlin et al. (2003)

Ascaris lumbricoides var. suum

2n=24

Merlin et al. (2003)

SPIROCERCIDAE

Mastophorus muris

2n=8+XX/XO

Spakulova et al. (2000)

STRONGYLOIDIDAE

Strongyloides ratti

2n=5 (Male)

2n=6 (Female)

Harvey & Viney (2001)

Strongyloides ratti

2n=6

Nignon & Roman (1952)

Strongyloides papillosus

2n=6

Chang & Graham (1957)

Strongyloides stercoralis

2n=5 (Male)

2n=6 (Female)

Harvey & Viney (2001)

DIPLOGASTRIDAE

Pristionchus pacificus

2n=12

Mitreva et al. (2005)

ONCHOCERCIDAE

Brugia malayi

2n=10

Mitreva et al. (2005)

PARASITAPHELENCHIDAE

Bursaphelenchus xylophilus

2n=12

Hasengawa et al. (2006)

DESMADORIDAE

Spirina parasitifera

2n=14

Cobb (1928)

RHABDITIDAE

Rhabditis aspera

2n=14

Walton (1940)

Rhabditis monhysterii

2n=14

Nignon (1949)

Rhabditis pellio

2n=14

Walton (1940)

Rhabditis aberrans

2n=18

Kruger (1913); Walton (1940)

Caenorhabditis elagans

2n=12

Mitreva et al. (2005)

Caenorhabditis briggsae

2n=12

Mitreva et al. (2005)

Caenorhabditis remanei

2n=12

Mitreva et al. (2005)

Caenorhabditis japonica 2n=12 Mitreva et al. (2005)

RHABDIASIDAE

Rhabdias briggsiae 2n=12 Nignon & Dougherty (1949)

Rhabdias bufonis 2n=12 Schleip (1911)

Rhabdias elegans 2n=12 Nignon & Dougherty (1949)

Rhabdias filleborni 2n=12 Dreyfus (1937)

Trichinella nelsoni 2n=6 (female)

2n= 5 (male) Mutafova et al. (1982)

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Chromosomes and cytogenetics of helminths

ANISAKIDAE

Contracaecum clavatum

2n=24

Walton (1940)

Contracaecum incurvum

2n=42

Goodrich (1916)

Contracaecum spiculigerum

2n=10

Walton (1924)

KATHLANIIDAE

Cruzia tentaculata

2n=12

Walton (1924)

HETERAKIDAE

Ascaridia galli

2n=10

Walton (1940)

Heterakis dispar

2n=10

Gulick (1911); Walton (1940)

Heterakis gallinae

2n=10

Gulick (1911); Walton (1940)

Heterakis papillosa

2n=10

Gulick (1911); Walton (1940)

Heterakis sp?

2n=10

Walton (1924)

Heterakis spumosa

2n=12

Walton (1924)

Heterakis gallinarum

2n=10

Mutafova (1995)

OXYURIDAE

Passalurus ambiguus

2n=8

Meves (1920)

Syphacia obvelata

2n=16

Walton (1924)

TRICHOSOMOIDIDAE

Trichosomoides crassicauda

2n=8

Walton (1924)

HELIGMOSOMIDAE

Nematospira turgida

2n=12

Walton (1924)

STRONGYLIDAE

Cyclostomum tetracanthum

2n=12

Walton (1940)

Strongylus edentatus

2n=12

Kultz (1913); Walton (1940)

Strongylus equinum

2n=12

Kultz (1913); Walton (1940)

Strongylus vulgaris

2n=12

Kultz (1913); Walton (1940)

Stephanurus dentatus

2n=12

Tromba & Steele (1957)

METASTRONGYLIDAE

Dictyocaulus filaria

2n=12

Struckmann (1905); Walton (1940)

Dictyocaulus viviparous

2n=12

Walton (1940)

Filaroides mustelarum

2n=16

Carnoy (1886); Walton (1940)

Metastrongylus elongatus

2n=12

Gulick (1911); Walton (1940)

TRICHOSTRONGYLIDAE

Haemonchus contortus

2n=12

Bremner (1955)

Trichostrongylus tenuis

2n=12

Gulick (1911); Walton (1940)

Haemonchus contortus

2n=12

Mitreva et al.

(2005)

ACUARIIDAE

Dispharynx spiralis

2n=12

Walton (1924)

CAMALLANIDAE

Camallanus lacustris

2n=12

Walton (1940)

PHYSALOPTERIDAE

Physaloptera turgida

2n=10

Walton (1924)

Proleptus robustus

2n=16

Carnoy (1886); Walton (1940)

Physaloptera clausa

2n=10

Mutafova (1995)

RHABDOCHONIDAE

Cystidicola farionis 2n=12 Mulsow (1912)

SPIRURIDAE

Protospirura muris 2n=10 Walton (1924)

Spirura talpae 2n=16 Carnoy (1886); Walton (1940)

SETARIIDAE

Setaria equina 2n=12 Meves (1915)

GORDIIDAE

Gordius (all spp?) 2n=4, 8 SvAbenik (1909); Meyer (1913);

Muhldorf (1914); Vejdovsky (1912);

Camerano (1890)

CHORDODIDAE

Paragordius varius 2n=14 Montgomery (1904)

Family Species No. and Morphology of

Chromosomes Reference

137

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

Chromosomes of Acanthocephala

Turning to the Acanthocephala, we again find

very few records of the examination of the

chromosomal components of the various

species. Apparently only 5 species belonging

to 3 ge nera have been studied :

Macracanthorhynchus hirudinaceous (Noe,

1914); of the Archiacanthocephala, and

Echinorhynchus (acus, haeruca, and

polymorphus) (Hamann, 1891) and

Pomphorhynchus proteus (Von Voss 1910) of

t h e P a l a e a c a n t h o c e p h a l a . I n

Macracanthorynchus the diploid number is 6,

and strong evidence seems to indicate that the

male is heterogametic and that an X-Y pair of

heterochromosomes is present. In the female

the 2X condition seems to be present (Jones &

Ward, 1950). Earlier reports of other than 6

chromosomes being characteristic of this

species may be due to faulty technique or to

mi si de nt if ication of th e wo rms.

Pomphorhynchus seems to have 8 somatic

chromosomes while the 3 Echinorhynchus

species show 16 (Table 5). This definite ratio

relationship between members of the

Echinorhynchidae may or may not be of

phylogenetic importance, but definitely does

support the separation of Pomphorhynchus

from Echinorhynchus, a separation which

some authorities have questioned on purely

morphological grounds. It is interesting to note

that in the only genus in which more than one

species has been studied there is a common

chromosome number. Similar examples may

appear in other genera, with a broader

sampling. Such constancy would not be

surprising in view of the known constancy of

nuclear numbers among certain of the

Acanthocephala. No evidence of the presence

of recognizable heterochromosomes among

the Palaeacanthocephala has been presented as

far as can be determined. It would seem, on the

basis of the evidence at hand, that while

constancy in chromosome numbers may seem

to occur within a genus, far too few records are

available for making any general statement.

The finding of heteromorphic conditions in

one species of the Acanthocephala should

encourage search for similar phenomena in

other members of the phylum. As far as the

actual chromosome numbers are concerned,

one can only say that the low numbers reported

are in accord with the high specialization of the

Phylum, but do not give recognizable clues as

to possible relationships, either within groups

above the species level, or as to possible

derivation from other phyla.

Table 5. Chromosome number of Acanthocephala from 1891 Till Date.

Family Species No. and Morphology of

Chromosomes Reference

OLIGACANTHORHYNCHIDAE

Macracanthorhynchus hirudinaceus 2n=12 Noe (1914); Jones & Ward (1950)

Macracanthorhynchus hirudinaceus 2n=6 Robinson (1964)

ECHINORHYNCHIDAE

Pomphorhynchus proteus 2n=16 Von Voss (1910)

Echinorhynchus acus 2n=32 Hamann (1891)

Echinorhynchus haeruca 2n=32 Hamann (1891)

Echinorhynchus polymorphus 2n=32 Hamann (1891)

Echinorhynchus gadi Mueller, 1776 2n=16 Robinson (1965)

Echinorhynchus gadi 2n=16 Walton (1959)

Echinorhynchus truttae 2n=7/8 Parenti et al. (1965)

Acanthocephalus ranae (Schrank, 1788) 2n=16 Robinson (1965)

2n=8 John (1957)

Acanthocephalus ranae 2n=16 Walton (1959)

Acanthocephalus lucii 2n=6 (2sm-m+1sm) 1st (X)

1m (Supernumerary B) Spakulova et al. (2002)

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Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Chromosomes and cytogenetics of helminths

MONILIFORMIDAE

Moniliformis dubius

2n=8 (female)

2n=7 (male)

Robinson (1965)

POLYMORPHIDAE

Polymorphus minutes (Goeze, 1782)

2n=16

Robinson (1965)

Polymorphus minutes

2n=16

Walton (1959)

POMPHORHYNCHIDAE

Pomphorhynchus laevis Mueller, 1776

2n=8

Robinson (1965)

Pomphorhynchus laevis

2n=7/8

Mutafova &Nedeva (1988); Fontana et

al.

(1993)

Pomphorhynchus laevis

2n=6+X

Bambarova et al. (2007)

Pomphorhynchus tereticollis

2n=6+X

Bambarova et al. (2007)

RHADINORHYNCHIDAE

Leptorhyncoides plagicephalus (Westrumb, 1821) 2n=14 Fontana et al. (1993)

Leptorhyncoides thecatus 2n=5/6 Bone (1974)

Family Species No. and Morphology of

Chromosomes Reference

CONCLUSION

Although the turbellarians are not parasitic

except for a few possible exceptions, they have

been introduced into this paper because they

do show the gradual reduction of the basic

number of chromosomes from a large number

of small chromosomes to a smaller number of

larger units, not only within the group as a

whole, but in some cases within races of the

same species, and may possibly give hints as to

the derivation of the Trematoda which do show

a much greater conformity between the

chromosome number and the taxonomic

position as based on other criteria.

Studies of trematode material have been

relatively numerous, and the results point

definitely toward the desirability of continued

efforts in this field of research. The discovery

of the presence of heterochromosomes,

particularly of the fact that some forms show

the female as the heterogamic sex, is of

especial interest and importance. Some

definite taxonomic relationships are

recognizable, but since in some cases they

substantiate other types of evidence used in

establishing phylogenetic position and in other

cases the results seem to be contradictory,

further observations are in order. Perhaps some

of the apparent contradictions may be

eliminated as our knowledge increases. Such

definitely has been the case among the

Turbellaria, as mentioned in the body of this

paper. In the digenetic trematodes studied till

to date, most variations in the chromosome

numbers within a genus are seldom greater

than + 1 or 2 bivalents. Thus the mechanism for

an addition or deletion of the chromosome

must operate at a low level or inefficient level

in this group. This suggest that the differences

in the have come about by a doubling of the

whole sets of chromosome but by a gradual

addition or losses. Each change which

represents aneuploid condition becomes

stabilized. When variation in the chromosome

number exceeds 1 or 2 bivalents, it probably

represents successive aneuploid conditions,

each change followed by a period of stability in

the new chromosome number.

Study of cestode material has been quite

limited, and such few records as have been

noted do not give any appreciable taxonomic

clues, except perhaps to indicate that some

present taxa are too heterogeneous to be of real

validity. Many questions on the systematics of

basal cestode groups still remain controversial

and phylogenetic interrelationships of the

a c b

TCL, total complement length.; ?, uncertain value.; m, metacentric chromosome; sm, submetacentric chromosome; a, acrocentric chromosome (i.e. telocentric and

subtelocentric of Levan et al. (1964).

139

Neotropical Helminthology. Vol. 9, Nº1, jan-jun 2015 Sofi et al.

groups remain only partially resolved

(Kodedova et al., 2000; Mackiewicz, 1981,

1982, 1994, 2003; Olson et al., 2001, 2008;

Waeschenbach et al., 2007). Unfortunately, no

cytogenetic information is available on the two

most basal cestodarian lineages Gyrocotylidea

and Amphilinidea, as well as for

Haplobothriidea and Diphyllidea.

Study of the parasitic nematodes has given a

considerable amount of valuable taxonomic

information, and here again; the field is by no

means exhausted. The cytologist has a broad

field of endeavor in the study of parasitic

forms. With the newer techniques and the

newer optical equipment now available, new

knowledge of the physiology, morphology and

phylogenetic relationships of such parasitic

forms as those discussed is just at the threshold

of a whole vista of new concepts and new

interpretations.

Study of the Acanthocephala gives clues as to

relationships only at the species level, but such

work as has been done promises interesting

results.

The authors are highly thankful to the

Department of Zoology for providing the

Laboratory and Library facility, the first author

is also thankful to Prof. Fayaz Ahmad for

compiling the paper.

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