Two new cestode species of Neoskrjabinolepis Spasskii, 1947 (Cyclophyllidea: Hymenolepididae) from the tundra shrew Sorex tundrensis Merriam (Mammalia: Soricidae) in Alaska and Chukotka

Two new cestode species of Neoskrjabinolepis Spasskii, 1947(Cyclophyllidea: Hymenolepididae) from the tundra shrewSorex tundrensis Merriam (Mammalia: Soricidae) in Alaskaand Chukotka

Svetlana A. Kornienko • Nikolaj E. Dokuchaev

Received: 27 March 2012 / Accepted: 23 June 2012

� Springer Science+Business Media B.V. 2012

Abstract Descriptions, illustrations and differential

diagnoses for two new species of Neoskrjabinolepis

Spassky, 1947, i.e. N. (Neoskrjabinolepis) fertilis n.

sp. and N. (Neoskrjabinolepidoides) hobergi n. sp., are

given. Neoskrjabinolepis (Neoskrjabinolepis) fertilis

was found in the shrew Sorex tundrensis Merriam on

the Seward Peninsula (Alaska, USA) and from the

mouth of the Anadyr River (Chukotka, Russia), and

N. (Neoskrjabinolepidoides) hobergi was found in

Sorex tundrensis on the Seward Peninsula (Alaska,

USA). Neoskrjabinolepis fertilis n. sp. is characterised

by having: rostellar hooks 38–42 lm long and

provided with a small epiphyseal thickening of the

handle; a long cirrus (85–100 lm), consisting of a

basal region with claw-shaped spines and a parabasal

region with small, thin needle-shaped spines; a cirrus-

sac extending well into the median field; and 55–70

eggs per gravid uterus. Neoskrjabinolepis (Neoskrja-

binolepidoides) hobergi n. sp. is characterised by

having: rostellar hooks 63–65 lm long and provided

with a large epiphyseal thickening of the handle; a

short cirrus (45–50 lm), consisting of a basal region

with small claw-shaped spines and a parabasal region

with thin, needle-shaped spines; cirrus-sac reaching

slightly into the median field; and 36–45 eggs per

gravid uterus.

Introduction

The last checklist of the helminth parasites of shrews

from North America by Kinsella & Tkach (2009)

included 34 species of cestodes (including larvae).

However, only 18 cestode species of 11 genera are

currently known for shrews of the genus Sorex L. in

Alaska (Rausch & Kuns, 1950; Voge & Rausch, 1955;

Dunagan, 1956; Olsen, 1969; Gulyaev, 2003; Melnik-

ova et al., 2003; Dokuchaev & Gulyaev, 2007;

Gulyaev et al., 2007, 2009; Kinsella, 2007; Kinsella

et al., 2008; Kinsella & Tkach, 2009). More recently,

the cestode Spasskylepis rauschi Gulyaev, Dokuchaev

& Lykova, 2010 has been found in Alaskan specimens

of Sorex cinereus Kerr (Gulyaev et al., 2010). Thus, 10

of the 12 recorded genera are common for Eurasia and

North America and only two, i.e. Lockerraushia

Yamaguti, 1959 and Vogelepis Vaucher in Czaplinski

& Vaucher, 1994, are North-American endemics. The

latter is morphologically similar to Neoskrjabinol-

epis Spassky, 1947, but Neoskrjabinolepis spp. have

recently been considered to parasitise only Palaearctic

shrews and were thought not to occur in the Nearctic

S. A. Kornienko (&)

Institute of Systematics and Ecology of Animals, Siberian

Branch, Russian Academy of Sciences, 11 Frunze Street,

Novosibirsk 630091, Russia

e-mail: [email protected]

N. E. Dokuchaev

Institute of Biological Problems of the North,

Far-East Branch, Russian Academy of Sciences,

18 Portovaja Street, Magadan 685000, Russia

e-mail: [email protected]

123

Syst Parasitol (2012) 83:179–188

DOI 10.1007/s11230-012-9383-6

(Gulyaev et al., 2009). However, in helminthological

samples from shrews in Alaska, cestodes correspond-

ing morphologically to the genus Neoskrjabinolepis

have been found (Kornienko & Gulyaev, 2008).

For many years, only two species of Neoskrjabi-

nolepis were recognised: N. schaldybini Spassky, 1947

and N. singularis (Cholodkowsky, 1912) Spassky,

1954 (Spassky, 1947, 1954; Czaplinski & Vaucher

1994). Sometimes, N. schaldybini has been considered

a junior synonym of N. singularis (Kobuley, 1953;

Zarnowski, 1955; Kisielewska, 1958; Prokopic, 1956,

1958; Pojmanska, 1957; Rybicka, 1959), but its

validity was supported by Spassky (1954), Vaucher

(1971) and Genov (1984). Recent studies have been

undertaken on cestodes from Sorex spp. belonging to

Neoskrjabinolepis originating from various parts of the

Palaearctic Region. These studies have resulted in

redescriptions of the two known forms and the

description of a further 10 species (Kornienko et al.,

2006, 2007, 2008, 2010). Neoskrjabinolepis now

includes two subgenera whose species differ in the

pattern of their strobilar development: Neoskrjabinol-

epis (Neoskrjabinolepis) Spasskii, 1947, with gradual

strobilar maturation, and Neoskrjabinolepis (Neoskrja-

binolepidoides) Kornienko, Gulyaev & Mel’nikova,

2006, with strobila consisting of a series of proglottides

at the same developmental stage (Kornienko et al.,

2006). Both types of strobilar organisation are recorded

in cestodes found in Sorex tundrensis Merriam from

Alaska. The aim of the present article is to describe two

new species, one from each of these two subgenera.

Materials and methods

Cestodes were collected from the intestine of shrews

Sorex tundrensis caught on the Seward Peninsula

(Alaska, USA) and in the mouth of Anadyr River

(Chukotka, Russia) during the summers of 2001 and

2002. Host specimens were dissected immediately

after their death. Live cestodes were isolated, washed,

relaxed in water and fixed in 70% ethanol. They were

stained in Ehrlich’s haematoxylin, differentiated in a

3% aqueous solution of ferric ammonium-sulphate-

12-hydrate, dehydrated in an ethanol series, cleared in

clove oil and mounted in Canada balsam. Some

specimens were mounted in Berlese’s medium to

facilitate examination of the rostellar hooks and

copulatory apparatus. Type-specimens are deposited

in the collection of Zoology Museum of the Institute of

Systematics and Ecology of Animals, Novosibirsk,

Russia (ISEA) and in the Natural History Museum of

Geneva, Switzerland (MHNG).

In order to establish the stage of proglottid matu-

ration, the terminology of Mas-Coma & Galan-

Puchades (1991) is used. Measurements are given in

micrometres except where otherwise indicated. They

are presented as the range followed by the mean and

the number of the measurements (n) in parentheses.

Neoskrjabinolepis (Neoskrjabinolepis) fertilis n. sp.

Type-host: Sorex tundrensis Merriam (Mammalia:

Soricidae), the tundra shrew.

Type-locality: Seward Peninsula, Alaska, USA;

65�519.120N, 164�4145.360W.

Other locality: Vicinity of Dionisija Mountain, Chuk-

otka, Russia; 64�344.490N, 177�1848.120E; vicinity of

Shachterskii Village, Chukotka, Russia, 64�4845.420N,

177�3254.310E.

Type-material: Holotype: ISEA 18.11.6.1, collected on

10 August 2001. Paratypes: 14 slides, ISEA

18.11.6.2–18.11.6.5, 18.11.6.7–18.11.13, Seward Pen-

insula, Alaska, USA, collected on 10 August. 2001;

ISEA 18.11.6.14, vicinity of Dionisija Mountain,

Chukotka, Russia, collected on 29 July 2002; ISEA

18.11.6.15, vicinity of the Shachterskii Village, Chuk-

otka, Russia, 3 August 2002; MHNG INVE 82149,

Seward Peninsula, Alaska, USA, collected on 10

August 2001.

Etymology: The species name is derived from the

Latin fertilis (=fertile) and refers to the great number

of eggs in the gravid proglottides of this species.

Description (Figs. 1, 2)

Gravid specimens 8.0–10.0 mm (8.9 mm, n = 10)

long; strobila consists of 190–220 (200, n = 10)

proglottides. Strobilation gradual. Premature and

mature proglottides with developing uterus wider than

long; pregravid and gravid proglottides almost equal

in width and length. Internal formation of proglottides

distinct before external segmentation is apparent;

external segmentation appears at level of postmature

proglottides.

Scolex 250–290 wide (260, n = 7), distinctly wider

than neck (Fig. 1A). Suckers round, 100–130 100–110

180 Syst Parasitol (2012) 83:179–188

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Fig. 1 Neoskrjabinolepis (Neoskrjabinolepis) fertilis n. sp. Holotype ISEA, 18.11.6.1: A. Scolex; B. Rostellar hook of cestodes from

Alaska; C. Rostellar hook of cestodes from Chukotka; D. Mature proglottides, ventral view; E. Mature proglottides, dorsal view;

F. Gravid proglottides. Scale-bars: A, D–F, 100 lm; B, C, 20 lm

Syst Parasitol (2012) 83:179–188 181

123

(95 9 90, n = 7), with well-developed musculature.

Rostellar apparatus complex. Rhynchus short, 62–65

long, 78–87 wide. Rostellum sac-like, 125–140 9

100–110 (121 9 72, n = 7); its walls consist of

external layer of longitudinal muscular fibres and

internal layer of circular muscle bundles. Rostellar

hooks 10 in number, arranged in single row, 38–42

(41, n = 15) long, with characteristic pincer-like

shape; claw-like blade longer than guard and axis of

blade almost parallel to axis of guard; blade with

curved middle region; handle provided with small

epiphyseal thickening (Fig. 1B, C). When rostellar

apparatus retracted, rostellar hooks with blades

directed anteriorly. Rostellar pouch voluminous,

140–145 9 110–140 (144 9 129, n = 7), reaches

beyond level of posterior margins of suckers; its wall

consists of longitudinal muscular fibres and circular

muscular bundles.

Proglottides acraspedote. Mature proglottides

15–17 9 230–260 (15 9 246, n = 10) [as external

segmentation appears only at level of postmature

proglottides, length of mature proglottides is measured

as distance between genital atria of adjacent proglot-

tides] (Fig. 1D), with lateral fields 37–38 wide; gonads

densely situated in median field, with Neoskrjabino-

lepidoides-type organisation (as defined by Gulyaev &

Fig. 2 Neoskrjabinolepis (Neoskrjabinolepis) fertilis n. sp. Holotype, ISEA 18.11.6.1; paratypes, ISEA 18.11.6.10 & 18.11.6.14:

A. Cirrus-sac; B. Vagina; C. Cirrus of cestodes from Alaska; D. Cirrus of cestodes from Chukotka. Scale-bars: 50 lm

182 Syst Parasitol (2012) 83:179–188

123

Kornienko, 2009). Gravid proglottides 180–210 9

280–300 (200 9 290, n = 10). Osmoregulatory canals

arranged in pairs, without transverse anastomoses;

ventral canals 4–6 wide; dorsal canals 1–2 wide.

Genital pores unilateral, dextral. Genital atrium sim-

ple, 5–6 deep, 5–6 in diameter.

Testes 3, of almost equal size, 15–23 9 34–50

(20 9 45, n = 12), situated in line; poral testis

separated from 2 antiporal testes by external seminal

vesicle (Fig. 1E). Diameter of testes greater than

proglottis length and, consequently, dense dorsal

testicular field formed in mature region of strobila.

Cirrus-sac cigar-shaped, 100–130 9 12–15 (120 9 13,

n = 12), slightly winding, passes deeply into median

field but does not reach median line of proglottis

(Fig. 2A). Cirrus long, 85–100 (93, n = 8), cylindri-

cal, armed with small, thin, needle-shaped spines

along its entire length; its basal region armed with

small, claw-shaped spines (Fig. 2C, D). Ductus ejac-

ulatorius forms several coils. Internal seminal vesicle

small, 25–35 9 10–12 (28 9 10, n = 12), occupies

less than quarter of cirrus-sac length even when filled.

External seminal vesicle 32–35 9 15–17 (33 9 17,

n = 10), connected to cirrus-sac by long, narrow

duct.

Vitellarium subspherical, 25–28 9 15–17 (26 9

17, n = 11), situated antiporally to ovary. Ovary

transversely elongate, compact, 10–15 9 90–110

(14 9 103, n = 11), situated in poral half of median

field, overlaps cirrus-sac, external seminal vesicle and

testes ventrally (Fig. 1D). Vagina thin-walled, passes

ventrally to cirrus-sac. Conductive part of vagina

gradually enlarges and passes into small, sac-like

seminal receptacle (Fig. 2B).

Uterus not extending into lateral fields. Uterine wall

thin, membranous throughout morphogenesis of uterus.

Number of eggs in uterus 55–70 (Fig. 1F).

Neoskrjabinolepis (Neoskrjabinolepidoides) hobergi

n. sp

Type-host: Sorex tundrensis Merriam (Mammalia:

Soricidae), the tundra shrew.

Type-locality: Seward Peninsula, Alaska, USA;

65�519.120N, 164�4145.360W.

Type-material: Holotype: ISEA 18.11.19.1, collected

on 7 August 2001. The holotype and another specimen

belonging to the same species are mounted on one

slide. The specimen marked N 1 was chosen as the

holotype. Paratypes 4: MHNG INVE 82150, ISEA

18.11.19.3–18.11.19.5; collected on 7 August 2001.

Etymology: This species is named for Dr Eric P.

Hoberg in recognition of his contributions to

helminthology.

Description (Figs. 3, 4)

Gravid specimens 8.0–10.0 mm (8.6 mm, n = 4)

long; strobila consists of 480–520 (510, n = 4)

proglottides. Strobilation serial; pregravid or gravid

strobila usually consisting of 4 series of proglottides,

each containing proglottides at same developmental

stage (first series of juvenile or premature proglottides;

second section of mature proglottides; third series of

postmature proglottides; fourth series of pregravid or

gravid proglottides); each series consists of c.110–130

proglottides. Strobilar regions containing juvenile,

premature or mature proglottides without external

segmentation; proglottides externally distinct at level

of postmature region of strobila.

Scolex 300–320 wide (316, n = 4), clearly wider

than neck (Fig. 3A). Suckers round, 100–110 9

100–110 (105 9 100, n = 4), with well-developed

musculature. Rostellar apparatus complex. Rhynchus

short, 65–66 long, 100–110 wide. Rostellum sac-like,

120–130 9 100–110 (121 9 72, n = 4); its walls

consist of external layer of longitudinal muscular fibres

and internal layer of circular muscular bundles.

Rostellar hooks 10 in number, arranged in single row,

63–66 (64, n = 5) long, with characteristic pincer-like

shape; blade twice length of handle and guard

(Fig. 3B), curved medially; handle provided with

large epiphyseal thickening. When rostellar appara-

tus retracted, blades of rostellar hooks are directed

anteriorly. Rostellar pouch voluminous, 180–190 9

170–180 (186 9 173, n = 5), reaches beyond level of

posterior margins of suckers; its wall consists of

longitudinal muscular fibres and circular muscular

bundles. Neck 160–180 (173, n = 4) wide.

Proglottides acraspedote, transversely elongate.

Mature proglottides 11–13 9 310–350 (11 9 336,

n = 4) (Fig. 3C) [as external segmentation appears

only at level of postmature proglottides, length of

mature proglottides is measured as distance between

genital atria of adjacent proglottides], with length/

width ratio 1:26–28; lateral fields 50–60 wide; gonads

densely situated in median field. Gravid proglottides

Syst Parasitol (2012) 83:179–188 183

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Fig. 3 Neoskrjabinolepis (Neoskrjabinolepidoides) hobergi n. sp. Holotype, ISEA 18.11.19.1: A. Scolex; B. Rostellar hook; C. Mature

proglottides, dorsal view; D. Mature proglottides, dorsal view; E. Transition between the series of pregravid and gravid proglottides.Scale-bars: A, C–E, 100 lm; B, 30 lm

184 Syst Parasitol (2012) 83:179–188

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20–22 9 330–340. Osmoregulatory canals 2 pairs,

without transverse anastomoses; ventral canals 4–6

wide; dorsal canals 2–3 wide. Genital pores unilateral.

Genital atrium simple, 7–8 deep, 3–4 in diameter.

Testes 3, of almost equal size, 20–30 9 40–54

(20 9 35, n = 8), situated in line; poral testis sepa-

rated from 2 antiporal testes by external seminal

vesicle (Fig. 3C). Diameter of testes larger than

proglottis length and, consequently, dense dorsal

testicular field is formed in mature strobilar region.

Cirrus-sac cigar-shaped, 72–80 9 10–14 (76 9 11,

n = 4), thin-walled, crosses poral osmoregulatory

canals and reaches slightly into median field of

proglottis (Fig. 4A). Cirrus short, cylindrical, 45–50

(47, n = 4); its basal region armed with small, claw-

shaped spines; parabasal region provided with thin,

needle-shaped spines. Length of needle-shaped spines

decreases towards apex of cirrus (Fig. 4B). Ductus

ejaculatorius forms several coils. Internal seminal

vesicle small, 17–22 9 8–10 (21 9 8, n = 4), occu-

pies not more than quarter of cirrus-sac length even

when filled. External seminal vesicle 30–32 9 20–22

(30 9 20, n = 4), connected to cirrus-sac by narrow

duct.

Vitellarium, 16–19 9 8–11 (17 9 11, n = 4), sit-

uated antiporally to ovary. Ovary transversely elon-

gate, compact, 7–10 9 80–90 (9 9 84, n = 4),

medial or may be in poral half of median field; in

latter case, ovary overlaps cirrus-sac, external seminal

vesicle and testes ventrally (Fig. 3D). Vagina thin-

walled, passes ventrally to cirrus-sac (Fig. 4A).

Uterus extends into lateral fields of both postmature

and gravid proglottides. Uterine wall thin, membra-

nous throughout morphogenesis of uterus. Number of

eggs in uterus 36–45 (Fig. 3E).

Remarks

Currently, four species are included in the subgenus

N. (Neoskrjabinolepis), which is characterised by a

gradual maturation along the strobila. These are

N. schaldybini Spassky, 1947, N. longicirrosa Kor-

nienko, Gulyaev & Melnikova, 2006, N. pilosa

Kornienko, Gulyaev & Melnikova, 2007 and N. plagis

Kornienko, Gulyaev & Melnikova, 2007.

Neoskrjabinolepis fertilis n. sp. is morphologically

close to N. longicirrosa and N. pilosa. Despite the

similar lengths of the rostellar hooks (39–42, 41–45

Fig. 4 Neoskrjabinolepis (Neoskrjabinolepidoides) hobergi n. sp. Holotype, ISEA 18.11.19.1; paratype, ISEA 18.11.19.4: A. Cirrus-

sac and vagina; B. Cirrus. Scale-bars: 25 lm

Syst Parasitol (2012) 83:179–188 185

123

and 45–49 lm, respectively), the cirrus-sac (100–130,

150–165 and 120–140 lm, respectively) and the

cirrus (85–100, 122–125 and 100–110 lm, respec-

tively), there are significant differences in hook shape

and the number of eggs. The blade of the hooks of

N. fertilis has a curvature in the middle region, in

contrast to the almost straight blade of hooks of

N. longicirrosa and N. pilosa. The blade in N. pilosus

is twice the length of the guard, whereas in N. fertilis

this ratio is much less. In N. fertilis the rostellar pouch

reaches beyond level of the posterior margins of the

suckers, but in N. pilosa it extends almost to the neck.

Furthermore, there are differences in the number of

eggs per gravid uterus: this is 16–20 in N. longicirrosa,

35–47 in N. pilosa and 55–70 in N. fertilis. Further-

more, N. pilosa has not been found outside Western

Siberia (Kornienko 2011; Kornienko et al., 2008).

This new species differs from N. plagis in the

smaller size of its rostellar hooks (39–42 and

52–55 lm, respectively), the length of the cirrus

(85–100 and 45–50 lm, respectively) and by having

more than twice the number of eggs per gravid uterus.

The lengths of the hooks of N. fertilis are similar to

those of N. schaldybini (39–42 and 38–43 lm, respec-

tively). However, the new species has a longer cirrus-

sac (100–130 vs. 73–90 lm) and longer cirrus (85–100

vs. 40–42 lm). The two species also differ from one

another on the basis of the cirral armament. The

parabasal region of the cirrus of N. schaldybini is

armed with several large, claw-shaped spines, and the

middle and distal regions of the cirrus are armed with

sparsely distributed, sabre-shaped spines; whereas the

cirrus of N. fertilis is armed with small, thin, needle-

shaped spines along its entire length.

Currently, the subgenus N. (Neoskrjabinolepido-

ides), which is characterised by strobila with a series

of proglottides at the same stage of development,

contains six species: N. (Neoskrjabinolepidoides)

singularis (Cholodkowsky, 1912) Spasskii, 1954,

N. nadtochijae Kornienko, Gulyaev & Melnikova,

2006, N. corticirrosa Kornienko, Gulyaev & Melnik-

ova, 2007, N. kedrovensis Kornienko, Gulyaev &

Melnikova, 2007, N. nuda Kornienko, Gulyaev,

Melnikova & Georgiev, 2008 and N. merkushevae

Kornienko & Binkien _e, 2008.

Neoskrjabinolepis hobergi n. sp. is most similar to

N. singularis, a species with a Euro-Siberian geo-

graphical range, which does not occur in the Far East

(Kornienko 2011; Kornienko & Binkiene, 2008;

Kornienko & Gulyaev, 2008; Kornienko et al.,

2008). Although N. hobergi and N. singularis are

similar in terms of their rostellar hooks (63–66 and

56–65 lm in length, respectively), cirrus (45–50 and

40–47 lm in length, respectively) and the number of

eggs (35–45) in the gravid uterus, there are differences

in the number of proglottides in series and in the shape

of the gravid proglottides. Their shape in N. hobergi is

much wider than long, whereas in N. singularis it is

much longer than wide, and the number of proglottides

in N. singularis is about 350 but exceeds 520 in

N. hobergi. Furthermore, N. hobergi is unique in

Neoskrjabinolepis in that its uterus crosses the longi-

tudinal osmoregulatory canals and reaches into the

lateral fields of postmature, pregravid and gravid

proglottides.

Neoskrjabinolepis hobergi differs from all other

species in this subgenus by the size of the rostellar

hooks, cirrus-sac and cirrus, its cirral armament, the

number of proglottides and the number of eggs per

gravid uterus. Its rostellar hooks are larger (63–66 lm)

than those of N. corticirrosa (48–53 lm), N. nad-

tochijae and N. nuda (40–45 lm), N. kedrovensis

(36–38 lm) and N. merkushevae (35–37 lm). The

cirrus-sac of N. hobergi only slightly crosses the poral

excretory canals, whereas the long cirrus-sacs of

N. nuda, N. nadtochijae and N. kedrovensis extend

far into the median field of the proglottides. In terms of

the length of the cirrus-sac, that of N. hobergi

(72–80 lm) is similar to N. corticirrosa (90–95 lm)

and N. merkushevae (60–65 lm). However, the latter

species can be distinguished from each of these by

differences in armament of the cirri. The parabasal

region of each cirrus in N. corticirrosa is armed with

several small, claw-shaped spines and the remainder

with scarce, sabre-shaped spines. The middle region of

the cirrus of N. merkushevae is provided with sparse,

needle-shaped spines which decrease in number dis-

tally, and the parabasal region and distal end are

unarmed. The parabasal region of each cirrus of

N. hobergi is armed with thin, needle-shaped spines

which decrease in numbers towards the apex. More-

over, these species differ in the number of proglottides

and the number of eggs in gravid proglottides. More

than 520 proglottides are found in N. hobergi, c.420 in

N. corticirrosa and only c.200 in N. merkushevae. The

gravid uterus of N. hobergi contains 36–45 eggs,

186 Syst Parasitol (2012) 83:179–188

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whereas that of N. corticirrosa and N. merkushevae are

characterised by 20 and 12–16 eggs per gravid uterus,

respectively.

On the basis of the above comparisons, we recognise

the two species described above as new to science.

Discussion

North-American shrews (Sorex spp.) have been

recorded as hosting 20 species of cestodes belonging

to 12 genera of the families Hymenolepididae and

Dilepididae (Melnikova et al., 2003; Dokuchaev &

Gulyaev, 2007; Gulyaev et al., 2007, 2009, 2010;

Kinsella, 2007; Kinsella et al., 2008; Kinsella &

Tkach, 2009). Taxonomic composition of tapeworm

fauna parasitising Nearctic shrews appears to be

similar to that of shrews in northern Asia. In North

American shrews, there are autochthonous species

belonging to 10 cestode genera which are also known

from the Palaearctic: Staphylocystis Villot, 1877,

Skrjabinacanthus Spassky & Morosov, 1959, Ecri-

nolepis Spassky & Karpenko, 1983, Mathevolepis

Spassky, 1948, Lineolepis Spassky, 1958, Staphylo-

cystoides Yamaguti, 1959, Soricinia Spassky &

Spasskaja, 1954, Spasskylepis Schaldybin, 1964,

Monocercus Villot, 1882 and Neoskrjabinolepis Spas-

sky, 1947. There are only two genera endemic to the

Nearctic, i.e. Vogelepis Vaucher in Czaplinski &

Vaucher, 1994 and Lockerraushia Yamaguti, 1959.

Our record of species of the genus Neoskrjabinolepis

in North-American shrews refutes the recent assump-

tion that in North America species of Neoskrjabinol-

epis are replaced by species of the morphologically

related but distinct genus Vogelepis (see Gulyaev,

2003).

Only one cestode species from shrews, Lineolepis

pribilofensis (Olsen, 1969) Dokuchaev & Gulyaev,

2007, has previously been recorded as occurring at

both sides of the Bering Strait (Dokuchaev & Gulyaev,

2007). The geographical distribution of N. fertilis n.

sp. in both Alaskan and Chukotkan shrews provides

further evidence for the presence of a close faunal

relationship between the Palaearctic and the Nearctic.

Acknowledgements This work was funded by the Russian

Fund for Fundamental Research (Project Nos. 11-04-49010-a,

11-04-00870-a, 11-04-00342) and by the National Science

Foundation ‘‘Beringian Coevolution Project’’ (NSF 0196095

and 0415668).

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