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Arostrilepis beringiensis (Kontrimavichus & Smirnova, 1991) Gulyaev & Chechulin, 1997 (Fig. 8) Syn.: Hymenolepis beringiensis Kontrimavichus & Smirnova, 1991. Hosts in Nearctic: Lemmus trimucronatus (Richardson) and Synaptomys borealis (Richardson). Localities in Nearctic: Alaska- Barrow, Arctic Coastal plain; Brooks Range; eastern Beringia, Yukon Charley Rivers National Preserve. Material examined: Vouchers include 4 specimens in Synaptomys borealis: MSB Para 1227, 1228, 1229 (field numbers AF 49480 / cyt- b sequence and 49480 A in Berlese’s; 49481 / cyt- b sequence , 2 specimens, multiple slides) by H. Henttonen et al., from the Yukon Charley Rivers National Preserve, Alaska across the Yukon River from Glen Creek Cabin (65 o 18 ’ 9 ”N, 142 o02’ 14 ”W), 8 August 2001; MSB 1233, 1234 (AF 61601 B, 61601 C, 2 specimens) by H. Henttonen et al., from Gates of the Arctic National Preserve, Brooks Range, Alaska on unnamed lake off Koyukuk River (67 o 21 ’ 11 ”N, 150 o 51 ’00”W), 23 July 2002. Palearctic specimens: MSB 1235 (LEM 229 a and 229 b, 2 specimens) ex Lemmus sibiricus by V. Fedorov and G. Jarrell on the northwestern Taymyr Peninsula, Russia (ca., 75 o 34 ’N, 94 o 29 ’E), 24–26 June 1994; the specimen from this series (designated as LEM 117 / cyt- b sequence ) from the northwest Taymyr was not available as a voucher. See Appendix 1 for listing of additional identified voucher specimens from the Nearctic and Palearctic. Description: Based on 5 specimens. Fully developed strobila 145–196 mm long, with maximum width at pregravid or gravid proglottides, 1.5–1.8 mm. Scolex slightly compressed dorso-ventrally, 255–285 (270, n = 2) wide, clearly wider than neck. Suckers ovoid in surface view, 160–250 × 110–188 (198 × 149, n = 6), with thick walls, prominent (Fig. 8 A). Neck relatively long and narrow, 160–175 (170, n = 2) wide. Dorsal osmoregulatory canals thin, 1–3 (2.1, n = 8) wide, situated predominantly in same sagittal plane as ventral canals. Ventral osmoregulatory canals 40–65 (54, n = 8) wide. Genital ducts may pass dorsally or between longitudinal osmoregulatory canals within same strobila; intersegmental variation without regularity. Development of proglottides gradual, protandrous. Mature proglottides 160–200 × 825–1120 (174 × 980, n = 12), transversely elongate, trapeziform (Fig. 8 C). Testes usually 3 in number, almost of equal size, 140–195 × 100–140 (168 × 116, n = 15), round or oval, commonly situated in triangle with flat angle or rarely, testes form one row; poral testis separated from 2 antiporal testes by female gonads. Cirrus sac relatively short, 108–127 × 23–32 (117 × 27, n = 10), antiporal part of cirrus-sac commonly does not reach ventral longitudinal canal (Fig. 8 C). Genital atrium simple, cup-shaped, deep, opens laterally about middle of lateral proglottis margin, or slightly more anteriorly. Cirrus small, 36–47 × 9–12 (41 × 10, n = 10), cylindrical; armed along entire length with relatively small (up to 2.5–2.7 long) rosethorn-shaped spines (Fig. 8 B). Internal seminal vesicle, ovoid, 50–73 × 18–26 (63 × 21, n = 10), shorter than half of cirrus sac length (Fig. 8 C). External seminal vesicle 90–135 × 45–65 (111 × 52, n = 8), with size approximately equal to that of seminal receptacle. Ovary 300–420 (365, n = 12) wide, median, fan-shaped, irregularly lobed, slightly overlapping testes (Fig. 8 C). Vitellarium 62–92 × 152–225 (73 × 176, n = 10), postovarian, median, scarcely lobed. Vagina tubular, clearly distinct from seminal receptacle; ventral to cirrus sac. Seminal receptacle relatively small, 87–115 × 30–40 (101 × 36, n = 5). Gravid proglottides 320–430 × 1300–1800 (366 × 1547, n = 8). Fully developed uterus labyrinthine, occupying entire median field and extending bilaterally beyond longitudinal osmoregulatory canals. Eggs 33–37 × 56–62, elliptical, with thin outer coat; oncosphere 12–15 × 18–21 (Fig. 8 D). Embryophore fusiform, 14–19 × 40–46, with straight polar processes. Embryonic hooks small, 7–8 long. Remarks: Results of the present morphological analysis of specimens of A. beringiensis do not indicate a high degree of differentiation between cestodes from the Nearctic and Palearctic currently isolated across the Bering Strait (Makarikov & Kontrimavichus 2011). We observed that in specimens of A. beringiensis from the Nearctic the dimensions of the eggs, embryophores and oncospheres are larger relative to those from the Palearctic. We assume that the differences in egg measurements of the specimens are associated with the methods of specimen preparation rather than with geographical divergence of two populations. Eggs of A. beringiensis from the Palearctic were mostly compressed because these specimens, stained in Ehrlich’s haematoxylin, had been mounted permanently in Canada balsam. In contrast, gravid proglottides of specimens from the Nearctic were mounted in Berlese’s medium and as a consequence the eggs were not influenced by compression. Otherwise specimens we examined representing populations of A. beringiensis in bog lemmings and Nearctic brown lemmings from North America and Siberian brown lemmings from Eurasia do not appear to be differentiated morphologically relative to host species or specific geographic localities (Makarikov & Kontrimavichus 2011). These are the first confirmed records of A. beringiensis in eastern Beringia and the Nearctic. Additionally, cestodes reported as H. horrida in L. trimucronatus (Nearctic brown lemming) from Okpilak River, Alaska (70 o08’N, 143 o 38 ’W) by Haukisalmi and Henttonen (2001) (USNPC 89245) and those from Barrow, Alaska by R.L. Rausch (RLR 29831) are referable to this species. At Siberian localities including Lopatka Peninsula, Taymyr Peninsula, New Siberian Islands, and Wrangel Island, we confirm the presence of A. beringiensis, and redetermine records and specimens (USNPC 89243 and 89244) attributed to H. horrida by Haukisalmi and Henttonen (2001) at these sites. In Eurasia most records refer to cestodes in L. sibiricus, although other species including L. amurensis Vinogradov from eastern Siberia between the Lena and Kolyma Rivers and on the New Siberian Archipelago and L. portenkoi Tchernyavsky on Wrangel Island may be represented as hosts of this species (see Musser & Carleton 2005). Arostrilepis beringiensis appears to be geographically widespread in lemmings (both Synaptomys and Lemmus) at high latitudes in North America and in lemmings of Eurasia, although an association with species of Dicrostonyx Gloger requires confirmation (E.P. Hoberg, K.E. Galbreath, unpublished data; Haukisalmi & Henttonen 2001; Makarikov & Kontrimavichus 2011). Specimens attributed to Hymenolepis horrida have been reported as common parasites in Nearctic collared lemmings (Dicrostonyx groenlandicus (Traill)) from near Barrow, Alaska (Kuns & Rausch 1950; Schiller 1952) and are also known in the Palearctic collared lemming (D. torquatus Pallas) from northern Eurasia (e.g. Ryzhikov et al. 1978); in either case, these specimens were not available for study. Interestingly, Dicrostonyx (tribe Dicrostonychini) is not considered phylogenetically close to the true lemmings (Lemmini) (reviewed in Musser & Carleton 2005), and these arvicolines may be derived from the earliest radiation of the subfamily linking Eurasia and North America and extending to 5–6 MYA (e.g. Conroy & Cook 1999). Such suggests that there is no particular basis to predict that the host distribution for A. beringiensis may extend to species of Dicrostonyx, unless it is present due to host colonization.

Arostrilepis cooki sp. n. (Figs. 4, 5) Type host: Myodes gapperi (Vigors) (Rodentia: Cricetidae: Arvicolinae). Other hosts: Currently unknown. Type locality: Near Meziadin Junction, British Columbia, Canada (56 o 21 ��� 55 ���N, 129 o 16 ��� 28 ���W). Other localities: Taft Creek, British Columbia (56 o 29 ��� 46 ���N, 129 o 25 ��� 31 ���W); near Bell II, British Columbia, south side Deltaic Creek (ca., 56 o 32 ��� 38 ���N, 129 o 32 ��� 39 ���W); Pattee Canyon, Missoula Co., Montana, USA (46 o 48 ���N, 113 o 57 ���W). Type material: Holotype MSB Para 1244 (field number IF 6750 / cyt- b sequence ) from type host and locality by A.M. Runck et al., 9 June 2003. Paratypes from type host species: MSB 1245 (IF 6751 / cyt- b sequence ) by A.M. Runck et al., at type locality, 9 June 2003; MSB 1249���1252 (IF 6830 C 2; 6830 C 3; 6830 C 4; 6830 C 6 / cyt- b sequence ) by A.M. Runck et al., at Taft Creek, British Columbia, 14 June 2003; MSB 1217 (JMK 02- 10), collected by J. M. Kinsella at Pattee Canyon, Montana, 10 October 2002; MSB 1246���1248 (IF 6827 C 1; 6827 C 2; 6827 C 3) by A. Runck et al., at Bell II, British Columbia, 14 June 2003. Symbiotype: Myodes gapperi (IF 6750) at type locality, skull and skeleton in MSB Mammalogy Division. Description: Based on 8 specimens. Fully developed strobila up to 150 mm long, with maximum width at postmature or pregravid gravid proglottides, 1.25���1.4 mm. Strobila flat, consisting of about 850 craspedote proglottides. Scolex slightly compressed dorso-ventrally, 280���372 (324, n = 4) wide, clearly wider than neck. Suckers unarmed, ovoid in surface view, 162���238 �� 124���195 (190 �� 161, n = 8), with thin walls, prominent (Fig. 4 A, B). Rhynchus and rostellar apparatus absent. Neck relatively long and narrow, 160���200 (178, n = 8) wide. Two pairs of osmoregulatory canals, without transverse anastomoses. Dorsal osmoregulatory canals thin, 0.5���2 (1.2, n = 10) wide, situated predominantly in same sagittal plane as ventral canals. Ventral osmoregulatory canals 30���65 (44, n = 15) wide. Position of dorsal osmoregulatory canals not always constant; loops may be situated laterally to ventral canals. Genital pores unilateral, dextral. Genital ducts usually pass dorsally to longitudinal osmoregulatory canals, position of genital ducts between osmoregulatory canals within single strobila appears rarely (Fig. 4 C, D). Development of proglottides gradual, protandrous. Strobilar part containing juvenile proglottides without external segmentation; proglottides become externally distinct at level of premature part of strobila. Mature proglottides 120���275 �� 680���1040 (198 �� 785, n = 12), transversely elongate, trapeziform (Fig. 4 C, D). Testes relatively large, usually three in number, almost of equal size, 115���175 �� 85���125 (135 �� 96, n = 25), round or oval, commonly situated in triangle; poral testis separated from two antiporal testes by female gonads. Arrangement of testes may vary (from triangle with flat angle to linear). Cirrus-sac relatively short, 190���218 �� 35���48 (205 �� 42, n = 15), with well-developed external muscular layers, commonly extends across ventral longitudinal canal (Figs. 4 D, 5 C). Genital atrium simple, infundibular, deep, opens laterally about middle or slightly anterior of lateral proglottis margin. Cirrus 88���109 (98, n = 16) long, conical, with relatively wide basal region, 19���24 (21, n = 16) in diameter, and narrow distal region, 8���14 (11, n = 16) in diameter; armed along entire length with relatively large (up to 3.3���4 long) rosethorn-shaped spines (Fig. 5 A). Internal seminal vesicle with circular musculature, ovoid, 85���120 �� 28���40 (95 �� 33, n = 15), shorter than half of cirrus-sac length (Figs. 4 D, 5 C). External seminal vesicle transversely elongate, 117���171 �� 65���103 (151 �� 78, n = 15), clearly outlined from vas deferens, slightly larger than seminal receptacle. Ovary 305���410 (329, n = 20) wide, median, fan-shaped, irregularly lobed, ventral to male genital organs, occupying substantial part of median field, slightly overlapping testes (Fig. 4 D). Vitellarium 60���117 �� 115���172 (91 �� 153, n = 20), postovarian, median, scarcely lobed. Vagina tubular, clearly distinct from seminal receptacle; ventral to cirrus-sac. Distal part of vagina 92���111 �� 9���19 (100 �� 13, n = 10), thick-walled, covered externally by dense layer of intensely stained cells; proximal part of vagina infundibular (Fig. 5 C). Conductive part of vagina 110���132 �� 10���27 (122 �� 17, n = 10), thin-walled, vastly varying in diameter depending on degree of distention with sperm. Seminal receptacle relatively small, transversely elongate, 80���122 �� 40���77 (100 �� 51, n = 14). Uterus appears as complex of fine-walled anastomosing tubes of varying length and diameter, positioned ventrally to other organs. With development of proglottides, tubular structures increase in width and uterus becomes labyrinthine. Testes remain in postmature and pregravid proglottides; cirrus-sac and vagina persist in gravid proglottides. Gravid proglottides transversely elongate, 185���360 �� 810���1330 (261 �� 1107, n = 10). Fully developed uterus labyrinthine, occupying entire median field, extending bilaterally beyond longitudinal osmoregulatory canals (Fig. 5 D). Uterus contains numerous (up to 1100) eggs. Eggs 31���38 �� 58���72, elliptic, with thin outer coat (Fig. 5 B); oncosphere 13���17 �� 18���23. Embryophore fusiform, 17���20 �� 40���48, with straight polar processes. Embryonic hooks small, 7.5���8.3 long. Etymology: This species has been named in honor of Joseph A. Cook in recognition of contributions in understanding rodent systematics and biogeography, and innovative explorations of host-parasite associations among arvicoline rodents. Remarks: Arostrilepis cooki sp. n. is distinguished from congeners by the length and shape of the cirrus (Table 2). In specimens of A. cooki the cirrus is longer in comparison to A. beringiensis, A. microtis, A. tenuicirrosa, A. gulyaevi, A. mariettavogeae and A. schilleri. The cirrus is armed with relatively large rosethornshaped spines and has a conical form; these features distinguish A. cooki from A. beringiensis, A. intermedia, A. janickii and A. schilleri, which have cylindrical cirri, and from A. microtis, A. tenuicirrosa and A. gulyaevi, in which the cirri have a conical basal region and cylindrical distal region. Compared to A. cooki, the cirrus of A. tenuicirrosa is armed with relatively small needle-shaped spines. The form and the length of the cirrus of A. cooki are most similar to those of A. macrocirrosa, but in specimens of the former the cirrus is narrower in the basal region (Figs. 11, 12). Arostrilepis cooki is characterized by a relatively narrow strobila. The cirrus-sac is shorter than that in A. horrida and A. microtis but larger than in A. beringiensis, A. janickii, A. mariettavogeae and A. schilleri. The ovary is narrower relative to those in A. horrida, A. microtis and A. gulyaevi. The scolex and the suckers are larger in comparison to A. janickii. The egg and oncosphere are large relative to those in A. horrida, A. microtis and A. janickii (see Table 2). The proximal end of the cirrus-sac in hermaphroditic mature proglottides overlaps the ventral longitudinal osmoregulatory canal. This species can be distinguished from A. beringiensis as its testes are arranged in a triangle; in the latter species, the testes form a flat angle or are situated in one row. Similarly, the testes form one row in A. microtis and A. mariettavogeae. Furthermore, the external seminal vesicle is larger than the seminal receptacle, the gravid proglottides are transversely elongate, and the polar processes of the embryophore are straight in A. cooki. This species is a specific parasite of red-backed voles (Myodes) from North America., Published as part of Makarikov, Arseny A., Galbreath, Kurt E. & Hoberg, Eric P., 2013, Parasite diversity at the Holarctic nexus: species of Arostrilepis (Eucestoda: Hymenolepididae) in voles and lemmings (Cricetidae: Arvicolinae) from greater Beringia, pp. 401-439 in Zootaxa 3608 (6) on pages 412-414, DOI: 10.11646/zootaxa.3608.6.1, http://zenodo.org/record/216055

Makarikov, Arseny A., Kinsella, John M. (2013): Morphological and molecular differentiation of Staphylocystis clydesengeri n. sp. (Cestoda, Hymenolepididae) from the vagrant shrew, Sorex vagrans (Soricomorpha, Soricidae), in North America. Zootaxa 3691 (3): 389-400, DOI: http://dx.doi.org/10.11646/zootaxa.3691.3.7

Staphylocystis clydesengeri n. sp. (Figures 1–3) Site in the host: small intestine. Type host: Sorex vagrans (Baird, 1857) (Soricomorpha: Soricidae). Type locality: Pattee Canyon, Missoula County, Montana, USA (46 °, 49 ” N, 113 °, 56 ” W). Other localities: vicinity of Spokane, Washington (exact coordinates not known). Material deposited: Holotype: USNPC 106801 (labelled Sorex vagrans, Pattee Canyon, Missoula County, MT, 14 July 2011, coll. J.M. Kinsella), paratypes: USNPC 106802-106808 (labelled Sorex vagrans, Pattee Canyon, Missoula County, MT, coll. J.M. Kinsella, different dates between May of 1998 to August 2009). Voucher specimens: USNPC 106809 (labelled Sorex vagrans, Pattee Canyon, Missoula County, MT, coll. J.M. Kinsella, 13 August 2011), USNPC 106810 (labelled: vicinity of Spokane, WA, 27 July 2009, coll. V. Tkach). Etymology: This species is named in honour of Clyde Senger who was the first to note the morphological differences between Staphylocystis schilleri and the form described here as a new species. Description: Based on 11 stained mounted specimens and 7 scoleces cleared in Berlese’s medium. Strobila 17.5 mm (14.4–24; 19.5; 6) mm long, with maximum width 1320 (1320–1600; 1497; 4) mm at level of gravid proglottides. Strobila consisting of 190 (138–190; 169; 6) craspedote proglottides. Scolex wider than long, 166 × 360 (166–190 × 352–430; 175 × 382; 6), clearly distinct from neck. Scolex with protracted rostellum conical, scolex with retracted rostellum slightly flattened antero-posteriorly. Suckers very muscular, thick-walled, unarmed, rounded or slightly oval, 100 × 85 (100–122 × 85–106; 109 × 97; 7). Rostellar pouch 140 × 209 (130–160 × 209– 242; 143 × 232; 5), with muscular walls, its bottom reaches approximately middle of suckers. Rostellum very muscular, apex not invaginable 92 × 99 (88–96 × 99–118; 93 × 111; 7). Rostellum armed by a single crown of 37 – 42 (average 40; 7) hooks with strongly developed dorso-ventrally flattened guard. Hook number in holotype could not be counted precisely due to overlap of several hooks in lateral regions of crown. Hook length 42 (38–44; 41.5; 13), hook blade length 21 (19–22; 21.4; 12), hook handle length 21 (19.5–23; 21.2; 12) and hook guard length 17 (16–18; 17.3; 12). FIGURE. 1. Staphylocystis clydesengeri n. sp. A. Holotype, dorso-ventral view of scolex. B. Paratype, rostellar hooks in profile and view from posterior surface showing enlarged hook guard. C. Holotype, male mature proglottis. D. Holotype, hermaphroditic mature proglottis. E. Hermaphroditic mature proglottis of Staphylocystis schilleri from Sorex palustris. (scale bars: A = 100 µm; B = 20 µm; C–E = 250 µm). FIGURE. 2. Staphylocystis clydesengeri n. sp. A. Paratype, genital ducts. B. Holotype, pregravid proglottis, showing uterus development. C. Paratype, gravid proglottis. D. Paratype, egg. E. Paratype, embryonic hooks (from left to right: median, antero-lateral, postero-lateral) (scale bars: A = 100; B, C = 250 µm; D = 20 µm; E = 10 µm). FIGURE. 3. Rostellar hooks of S. schilleri and S. clydesengeri n. sp. A. Staphylocystis schilleri from Sorex palustris collected near Missoula, Montana. B, C. Paratype of S. clydesengeri n. sp. Note that all photographs are taken at the same magnification. (scale bars: A–C = 20 µm). Osmoregulatory canals penetrate through rostellar pouch wall. Neck narrower than scolex. Ventral osmoregulatory canals 30 (28–40; 32; 7) at level of hermaphroditic proglottides, transverse anastomoses not observed. Dorsal osmoregulatory canals thin, 12 (9–12; 11.5; 7) at level of hermaphroditic proglottides, usually situated directly above ventral canals. Genital pores unilateral, dextral, genital ducts pass dorsally to both ventral and dorsal longitudinal osmoregulatory canals. Development of proglottides gradual, male and female gonads developing at approximately same rate. All segment in the strobila transversely elongated, but the length:width ratio increases from young to gravid proglottides. Mature proglottides 110 × 782 (87–130 × 782–911; 110 × 846; 5), trapezoid (Fig. 3 D). Testes relatively small, three, spherical or somewhat elongated, situated in a relatively tight triangle, one poral and two antiporal. Poral testis 51 × 57 (50–58 × 57–102; 54 × 90; 5), middle testis 51 × 67 (50–58 × 67–103; 54 × 92; 5), aporal testis 48 × 57 (48–66 × 57 – 110; 56 × 89; 5). Cirrus pouch 148 × 38 (135–168 × 37–47; 151 × 41; 9) in hermaphroditic proglottides, normally only reaching or somewhat overlapping poral osmoregulatory canals, but not crossing them. Genital atrium simple, infundibular, deep, opens laterally approximately at level of border between first and second thirds of lateral proglottis margin. Evaginated cirrus not observed. Internal seminal vesicle elongate, 103 × 33 (91– 103 × 33–42; 96 × 36; 8), occupying more than half of cirrus pouch length. External seminal vesicle ovoid, 90 × 38 (90–130 × 38–66; 106 × 50). Ovary 90 – 296 (72–95 × 272–360; 83 × 318; 5) wide, median, usually consisting of three large lobes, sometimes with secondary smaller lobes, ventral to male genital organs, occupying half to two-thirds of median field, usually overlapping testes. Vitellarium irregularly shaped, sometimes lobed 47 × 82 (40–61 × 82–132; 52 × 107; 7), postovarian, median. Copulatory part of vagina not clearly distinct from seminal receptacle; ventral to cirrus pouch. Seminal receptacle elongated, 250 long (178–260 × 54–80 × 63; 5), was not filled with sperm in mature hermaphroditic proglottides of holotype, but was filled in gravid proglottides. In majority of other specimens seminal receptacle was filled in mature hermaphroditic proglottides. Uterus first appears as transversely elongated sac, not extending beyond osmoregulatory canals, situated dorsally to other organs. With proglottis development, uterus grows and forms numerous lateral pockets and dorsoventral diverticula. Testes and vitellarium persist in postmature proglottides; cirrus pouch and vagina persist in gravid proglottides. Gravid proglottides 340 × 1286 (340–420 × 1286–1490; 382 × 1395). Fully developed uterus labyrinthine, extending into both lateral fields, saccate; walls of gravid uterus usually deeply folded or invaginated sometimes creating impression of being perforated. Uterus contains numerous small eggs. Eggs 40 – 44 × 32 – 39 (42 × 35; 8), spherical or subspherical; embryophore thin, without polar filaments, close to surface of oncosphere; oncosphere 22 – 28 × 18 – 24 (25 × 22; 8). Median embryonic hooks 16, antero-lateral and postero-lateral hooks 14.5–15. Antero-lateral embryonic hooks much more robust than slender postero-lateral and median hooks. Remarks: Morphological differentiation. The only other Staphylocystis species previously reported in North America is S. schilleri originally described from Sorex cinereus in Wisconsin (Rausch & Kuns 1950). The two species share general morphological characteristics such as the hook shape, overall scolex and strobila anatomy including triangular arrangement of testes and gravid uterus expanding into lateral fields of proglottides. But S. clydesengeri n. sp. differs from S. schilleri in several significant morphological features. The two species can be most easily distinguished based on the size and number of their rostellar hooks. Staphylocystis clydesengeri has 37 – 42 (average 40) hooks 39 – 44 µm (average 40.7 µm) long while S. schilleri has 22 hooks 27 – 30 µm long according to the original description. In our material of S. schilleri from Montana the hook number was higher (28 – 30, average 29). We could not count hooks in the holotype of S. schilleri very precisely due to their dense arrangement in the crown, but it looked like the number 22 is correct or at least very close to the real situation. In our experience, counting rostellar hooks in the laterally positioned crowns of Staphylocystis on total mounts usually results in underestimations of their numbers comparing with the crowns of the same species from the same sample cleared and apically oriented (or squished) in Berlese’s medium. The cirrus pouch in mature (hermaphroditic) proglottides of S. schilleri distinctly crosses the poral ventral osmoregulatory canal. In S. clydesengeri the cirrus pouch only barely reaches or, rarely, slightly crosses the poral ventral osmoregulatory canal. Gonads in S. clydesengeri occupy the central part of the middle field of the proglottis and do not reach the canals on either side. In fully mature proglottides of S. schilleri gonads normally fill the middle field of the proglottis entirely and reach or even overlap osmoregulatory canals on either side of the proglottis. Mature proglottides in heat-killed relaxed specimens of the two species distinctly differ in shape. In the new species proglottides are relatively shorter than in S. schilleri (Fig. 1 D, E). Finally, the two species differ in the early development of the uterus and its extent in gravid proglottides. In the new species the early uterus has numerous pockets and chambers from the very beginning while in S. schilleri the uterus at is initially formed as a two-winged saccular structure. The fully developed uterus in the new species extends well into the lateral fields while in S. schilleri it is confined to the median field of the proglottides and may only somewhat overlap the osmoregulatory canals on either side. Staphylocystis clydesengeri clearly differs from all Palearctic species parasitic in Sorex. While there are multiple characters separating these species, the rostellar hook number and size are discriminative enough, therefore we are not providing other differentiating features. The new species has 37 – 42 hooks 39 – 44 µm long while S. furcata has 23 – 30 hooks 23 – 27 µm long (Vaucher 1971, Genov 1984), S. sibirica has 28 – 30 hooks 32 – 35 µm long (Morozov 1957, Gulyaev & Shakhmatova 1990) and S. amurensis has 34 hooks 19 – 22 µm long (Karpenko 1984, 2004). Vampirolepis novosibirskiensis Sawada & Kobayashi, 1994 described from Sorex araneus, Sorex caecutiens and Sorex minutus in the Novosibirsk region and Altai Krai, Russian Federation (Sawada & Kobayashi 1994), clearly belongs to Staphylocystis. By the combination of morphological characteristics such as rostellar hook shape, size and number, organization of mature segments and other features, it is morphologically indistinguishable from Staphylocystis furcata. Therefore, we consider V. novosibirskiensis a junior synonym of S. furcata. We do not provide here differentiation of the new species from Staphylocystis spp. parasitic in crocidurine shrews because of the very high level of specificity among shrew hymenolepidids to the host genus (see the detailed discussion below). These two groups of shrews do not share their cestodes even when they occur in the same habitat, let alone on different continents. In the absence of any crocidurine shrews in North America we are certain that our specimens described herein as a new species cannot belong to one of the numerous Staphylocystis species known from Crocidura and related genera. In addition, Staphylocystis from the two groups of shrews deminstrate a high level of genetic divergence (see the Molecular differentiation below). Molecular differentiation. The aligned trimmed sequenced rRNA fragments from 5 specimens of S. clydesengeri (sequence length 2856 base pairs), one S. schilleri (2866 base pairs) and one S. furcata (2862 base pairs) comprised a short portion of the 18 S gene, complete ITS 1 spacer, complete 5.8 S gene, complete ITS 2 spacer, and partial 28 S gene. Due to introduced gaps the length of pairwise alignments between the three species varied from 2862 to 2872 base pairs. No intraspecific variability was detected among 4 replicates of S. clydesengeri. The two North American species S. clydesengeri and S. schilleri differed in only 3 bases while both these species differed from the Palaearctic S. furcata in 7 bases. Sequences of mitochondrial genes nad 1 and 16 S provided further convincing evidence of the status of S. clydesengeri as a new species. The nad 1 alignment of 4 sequences of S. clydesengeri (2 from Montana and 2 from Washington), one S. schilleri and one S. furcata was 663 bases long and had no gaps. Among 4 sequences of the new species only one specimen from Montana had a single base substitution. Results of pairwise comparisons are presented in Table 1. Staphylocystis clydesengeri and S. schilleri differed in 33 bases while the two North American species differed from S. furcata in 45 – 49 positions (Table 1). The 16 S alignment of 3 sequences of S. clydesengeri (2 from Montana and one from Washington), one S. schilleri and one S. furcata was 452 bases long and had no gaps. No intraspecific variability was detected in 16 S sequences of 4 sequenced specimens of S. clydesengeri. Results of pairwise comparisons are presented in Table 1. Staphylocystis clydesengeri and S. schilleri differed in 11 bases while the two North American species differed from S. furcata in 11 – 16 positions (Table 1). Thus, molecular data strongly support the status of S. clydesengeri as a new species. We do not provide a detailed molecular differentiation between S. clydesengeri and members of the genus known from crocidurine shrews because the new species shows much greater levels of interspecific sequence divergence from Staphylocystis parasitic in crocidurine shrews than from Staphylocystis parasitic in Sorex. For instance, the approximately 1400 base pair long sequence of the nuclear ribosomal 28 S gene of S. clydesengeri obtained in the present study was identical to that of S. furcata collected in the Ukraine. In the same DNA region, S. clydesengeri had 41 nucleotide differences from Staphylocystis brusatae (Vaucher, 1971) collected from lesser white-toothed shrews Crocidura suaveolens Pallas in the Ukraine (GenBank JQ 260805) and in 29 nucleotides from Staphylocystis sp. collected from Asian house shrews Suncus murinus Linnaeus in Thailand (V. Tkach, unpublished data). This provides additional evidence of the significant differences between Staphylocystis from soricine and crocidurine shrews.

Arostrilepis gulyaevi sp. n. (Figs. 2, 3) Type host: Myodes rufocanus (Sundevall) (Rodentia: Cricetidae: Arvicolinae). Other hosts: Rarely Microtus oeconomus Pallas. Type locality: Bol'shekhekhtsirskiy Nature Reserve, Khabarovskiy Kray, Russia (ca., 48 �� 16 ���N, 134 �� 45 ���E). Other localities: Baikal Nature Reserve, Republic of Buryatia, Russia (ca., 51 o 21 ���N, 105 o 17 ���E); Anadyr, Chukotka Autonomous Okrug, Russia, 1 km SE of confluence of Markova and Anadyr Rivers (64 o 40 ��� 57 ���N, 170 o 26 ��� 26 ���E); Buyunda River, Magadanskaya Oblast���, Russia (62 o 20 ���N, 153 o 21 ���E); Chaun Biological Field Station, Chaun River, Chukotka Autonomous Okrug, Russia (69 o00���N, 170 o 50 ���E). Type material: Holotype ISEA No. 18.28. 8.1 from type host and locality by Y. Melnikova, 18 July 2003. Paratypes from type locality and type host species: MSB Para 1262 (ISEA No.18.28.8.4) by Y. Melnikova, 17 July 2003; ISEA No. 18.28. 8.7 by Y. Melnikova, 18 July 2003; No. 18.28. 8.8 by Y. Melnikova, 21 July 2003; and No. 18.28. 8.10 by Y. Melnikova, 21 July 2003. Paratypes from other localities: MSB Para 1263 (field number RLR 43095) in type host species by L. Smirnova at Chaun Biological Field Station, 16 July 1975; MSB 1259, 1261 (IF 5657 / cyt- b sequence ; 5657 C 1) in type host species by A. Lahzuhtkin and K.E. Galbreath at Buyunda River, Russia, 4 July 2002; MSB 1253, 1254 (IF 5079 C 1 / cyt- b sequence ; 5079 C 2) in type host species, by N.E. Dokuchaev at Anadyr, Russia, 23 July 2002; and MSB 1256 (IF 5094 / cyt- b sequence ) in type host species by N.E. Dokuchaev at Anadyr, Russia, 24 July 2002. See Appendix 1 for additional identified vouchers in M. rufocanus and Microtus oeconomus from the Baikal Reserve and Magadanskaya Oblast���. Symbiotype: Type host specimen not deposited in a museum archive. Description: Based on 10 specimens. Fully developed strobila 180���300 mm long, with maximum width at postmature or pregravid gravid proglottides, 1.9���3.4 mm. Strobila flat, consisting of 900���1500 craspedote proglottides. Scolex slightly compressed dorso-ventrally, 250���350 (292, n = 6) wide, clearly wider than neck. Suckers unarmed, ovoid in surface view, 177���253 �� 135���185 (221 �� 153, n = 12), with thin walls, prominent (Fig. 2 A, B). Rhynchus and rostellar apparatus absent. Neck relatively long and narrow, 137���185 (154, n = 8) wide. Two pairs of osmoregulatory canals, without transverse anastomoses. Dorsal osmoregulatory canals thin, 0.5���1.5 (0.9, n = 15) wide, situated predominantly in same sagittal plane as ventral canals. Ventral osmoregulatory canals 60���170 (117, n = 15) wide. Position of dorsal osmoregulatory canals not always constant; loops may be situated laterally to ventral canals. Genital pores unilateral, dextral. Genital ducts may pass dorsally or between longitudinal osmoregulatory canals within single strobila; intersegmental variation not showing regularity (Fig. 2 C, D). Development of proglottides gradual, protandrous. Strobilar part containing juvenile proglottides without external segmentation; proglottides become externally distinct at level of premature part of strobila. Mature proglottides 215���300 �� 1350���2030 (248 �� 1619, n = 10), transversely elongate, trapeziform (Fig. 2 C, D). Testes relatively large, usually three in number, almost of equal size, 173���375 �� 95���216 (238 �� 144, n = 20), oval or pear-shaped, commonly situated in triangle; poral testis separated from two antiporal testes by female gonads. Arrangement of testes may vary (from triangle with flat angle to linear). Cirrus-sac relatively short, 192���235 �� 32���45 (206 �� 39, n = 20), with well-developed external muscular layers (Figs. 2 D, 3 C). Proximal part of cirrus-sac commonly not reaching or rarely overlapping ventral longitudinal canal or rarely overlapping. Genital atrium simple, infundibular, deep, opens laterally about middle or slightly anterior of lateral proglottis margin. Cirrus 53���72 (64, n = 28) long, conical, with relatively wide basal region, 16���24 (19, n = 28) in diameter, and narrow distal region, 9���15 (12, n = 28) wide; armed along entire length with relatively large (up to 4���4.5 long) rosethorn-shaped spines (Fig. 3 A). Internal seminal vesicle with circular musculature, ovoid, 85���120 �� 27���35 (103 �� 31, n = 18), shorter than half of cirrus-sac length (Figs. 2 D, 3 C). External seminal vesicle transversely elongate, 195���390 �� 55���84 (266 �� 69, n = 16), clearly outlined from vas deferens, slightly larger than seminal receptacle or with size approximately equal to that of seminal receptacle. Ovary 575���715 (633, n = 17) wide, median, fan-shaped, irregularly lobed, ventral to male genital organs, occupying a substantial part of median field, slightly overlapping testes (Fig. 2 D). Vitellarium 100���142 �� 215���386 (111 �� 283, n = 17), postovarian, median, scarcely lobed. Vagina tubular, not clearly distinct from seminal receptacle; ventral to cirrus-sac. Distal part of vagina 62���92 �� 7���16 (73 �� 10, n = 15), thick-walled, covered externally by dense layer of intensely stained cells; proximal part of vagina infundibular (Fig. 3 C). Conductive part of vagina 255���320 �� 9���35 (290 �� 19, n = 10), thin-walled, vastly varying in diameter depending on degree of distention with sperm. Seminal receptacle relatively small, transversely elongate, 150���340 �� 30���80 (198 �� 60, n = 14). Uterus appears as complex of fine-walled anastomosing tubes of varying length and diameter, positioned ventrally to other organs (Fig. 3 D). With development of proglottides, tubular structures increase in width and uterus becomes labyrinthine. Uterus may pass dorsally or between longitudinal osmoregulatory canals within same strobila; intersegmental variation does not show any regularity. Testes remain in postmature and pregravid proglottides; cirrus-sac and vagina persist in the gravid proglottides. Gravid proglottides transversely elongate, 340���540 �� 1700���3410 (466 �� 2343, n = 10). Fully developed uterus labyrinthine, occupying entire median field, extending bilaterally beyond longitudinal osmoregulatory canals (Fig. 3 E). Uterus contains numerous (up to 3000) eggs. Eggs 30���37 �� 58���68, elliptic, with thin outer coat (Fig. 3 B); oncosphere 13���15 �� 16���19. Embryophore fusiform, 16���20 �� 40���49, with straight polar processes. Embryonic hooks small, 7.2���8 long. Etymology: The name of this species has been dedicated to the memory of Vladimir D. Gulyaev, in recognition of his critical studies on hymenolepidid cestodes of rodents and faunistic explorations in central Siberia. Remarks: Arostrilepis gulyaevi sp. n. is distinguished from 9 recognized congeners by the length and shape of the cirrus (Table 2). In specimens of A. gulyaevi the cirrus is shorter relative to those in A. horrida, A. macrocirrosa, A. microtis and A. intermedia, but longer in comparison to A. beringiensis and A. mariettavogeae. In A. gulyaevi the length and form of the cirrus is similar to those in A. tenuicirrosa and A. microtis: the cirrus with conical basal region and cylindrical distal region, this species can be distinguished from A. tenuicirrosa as its cirrus is twice as wide as that of A. tenuicirrosa and armed with relatively large rosethorn-shaped spines; in the latter species, the cirrus is armed with relatively small needle-shaped spines (Figs. 11, 12). Arostrilepis gulyaevi is characterized by a relatively wide strobila and ovary (Table 2). The proximal end of the cirrus-sac in hermaphroditic mature proglottides commonly does not reach the ventral longitudinal canal or rarely overlaps it. This species can additionally be distinguished from A. beringiensis as its testes are arranged in a triangle; in the latter species, the testes form a flat angle or are situated in one row. Similarly, the testes form one row in A. microtis and A. mariettavogeae. Furthermore, the gravid proglottides are transversely elongate and the polar processes of the embryophore are straight in A. gulyaevi. This species is a specific parasite of red-backed voles (Myodes) from the eastern Palearctic region., Published as part of Makarikov, Arseny A., Galbreath, Kurt E. & Hoberg, Eric P., 2013, Parasite diversity at the Holarctic nexus: species of Arostrilepis (Eucestoda: Hymenolepididae) in voles and lemmings (Cricetidae: Arvicolinae) from greater Beringia, pp. 401-439 in Zootaxa 3608 (6) on pages 409-412, DOI: 10.11646/zootaxa.3608.6.1, http://zenodo.org/record/216055

Arostrilepis macrocirrosa Makarikov, Gulyaev & Kontrimavichus, 2011 (Fig. 9) Hosts in Nearctic: Myodes rutilus (Pallas); rarely Microtus xanthognathus (Leach), Microtus pennsylvanicus (Ord) and Tamiasciurus hudsonicus (Erxleben). Localities in Nearctic: Alaska - N Central region, SE region; Seward Peninsula (Bering Landbridge National Preserve); Brooks Range (Kobuk Valley National Park, Noatak National Preserve); Wrangell-St Elias National Preserve; Yukon Charley Rivers National Preserve; and Lake Clark National Park, northern Alaska Peninsula. Material examined: Vouchers include 4 specimens in Myodes rutilus: MSB 1306 (field number- AF 55147, 2 slides/ cyt- b sequence ) by E.P. Hoberg et al., at Braye Lakes in the Wrangell-St. Elias National Preserve, Alaska (62 o01��� 52 ���N, 141 o07��� 46 ���W), 26 July 2001; MSB 1285 (AF 37468, 2 slides) by H. Henttonen near Bonanza Creek Research Station, Alaska (62 o 42 ���N, 148 o 16 ���W), 1 August 2000; MSB 1302, 1303 (AF 49374, 1A and 1 B, 3 slides/ cyt- b sequence ) by H. Henttonen et al., from Yukon Charley Rivers National Preserve, Alaska, NW of Mt. Kathryn on Yukon River, S of Woodchopper (65 o 12 ��� 16 ���N, 143 o 34 ��� 22 ���W), 3 August 2001; MSB 1307 (AF 55196, 2 slides) by E.P. Hoberg et al., at Rex Creek, northwestern slope of Mt. Holmes in the Wrangell-St. Elias National Preserve, Alaska (61 o 18 ��� 37 ���N, 142 o 30 ��� 56 ���W), 30 July 2001; MSB 1304 (AF 49977) ex Tamiasciurus hudsonicus by H. Henttonen et al., from the Yukon Charley Rivers National Preserve, Alaska at Glenn Creek Cabin (65 o 18 ���07���N., 142 o05��� 23 ���W), 12 August 2001; MSB 1351 (RLR 15515, 2 slides) ex T. hudsonicus by R.L. Rausch near Chitina, Alaska (31.6 miles west on highway), 8 October 1955. Palearctic specimens: MSB 1221 (H- 15 / cyt- b sequence ) ex M. rutilus, by A. Lavikainen, on the Tunguska River, Irkutsk Oblast���, Russia (59 o06��� 36 ���N, 108 o 23 ��� 54 ���E), 8 August 2003. See Appendix 1 for listing of additional identified voucher specimens from the Nearctic and Palearctic. Description: Based on 4 specimens. Fully developed strobila 145���196 mm long, with maximum width at pregravid or gravid proglottides, 0.9���1.3 mm. Scolex slightly compressed dorso-ventrally, 275���385 (321, n = 3) wide, clearly wider than neck. Suckers ovoid in surface view, 162���210 �� 130���191 (182 �� 162, n = 8), with thick walls, prominent (Fig. 9 A). Neck relatively long and narrow, 130���185 (157, n = 3) wide. Dorsal osmoregulatory canals thin, 1���2.5 (1.7, n = 10) wide, situated predominantly in same sagittal plane as ventral canals. Ventral osmoregulatory canals 35���70 (53, n = 10) wide. Genital ducts may pass dorsally or between longitudinal osmoregulatory canals within same strobila; intersegmental variation without regularity. Development of proglottides gradual, protandrous. Mature proglottides 167���240 �� 670���980 (205 �� 825, n = 15), transversely elongate, trapeziform (Fig. 9 C). Testes usually 3 in number, almost of equal size, 125���210 �� 90���132 (161 �� 107, n = 15), round or oval, commonly situated in triangle; poral testis separated from 2 antiporal testes by female gonads. Cirrus sac relatively large, 208���245 �� 39���48 (229 �� 43, n = 15), commonly overlapping ventral longitudinal canal (Fig. 9 C). Genital atrium simple, cup-shaped, deep, opens laterally about middle of lateral proglottis margin. Cirrus large, 95���123 (106, n = 20) long, conical, with relatively wide basal region, 27���34 (30, n = 20) in diameter, and narrow distal region, 17���24 (19, n = 20) wide; armed along entire length with relatively small (up to 4 long) rosethorn-shaped spines (Fig. 9 B). Internal seminal vesicle, ovoid, 86���113 �� 30���40 (101 �� 36, n = 15), shorter than half of cirrus sac length (Fig. 9 C). External seminal vesicle 88���145 �� 35���72 (115 �� 53, n = 8), with size approximately equal to that of seminal receptacle. Ovary 270���395 (342, n = 10) wide, median, fan-shaped, irregularly lobed, slightly overlapping testes (Fig. 9 C). Vitellarium 70���115 �� 140���195 (92 �� 167, n = 8), postovarian, median, scarcely lobed. Vagina tubular, clearly distinct from seminal receptacle; ventral to cirrus sac. Distal part of vagina 105���112 �� 15���25 (107 �� 19, n = 8), thick-walled. Seminal receptacle relatively small, 135���155 �� 18���33 (147 �� 72, n = 8). Gravid proglottides 490���611 �� 815���1240 (552 �� 987, n = 10). Fully developed uterus labyrinthine, occupying entire median field and extending bilaterally beyond longitudinal osmoregulatory canals. Eggs 23���26 �� 38���43, elliptical, with thin outer coat; oncosphere 11���13 �� 14���15 (Fig. 9 D). Embryophore fusiform, 12���15 �� 23���26, with straight polar processes. Embryonic hooks small, 7.5���8.2 long. Remarks: Results of the present morphological analysis of specimens of A. macrocirrosa do not indicate a high degree of differentiation between cestode populations from the Nearctic and the Palearctic now isolated by the Bering Strait. Similar to observations in A. beringiensis, we noted differences in egg dimensions between conspecific cestodes in the Nearctic and Palearctic (Makarikov et al. 2011). We attribute these minor differences to methods of specimen preparation and mounting medium. Otherwise, specimens we examined representing populations of A. macrocirrosa in red backed voles and limited numbers of cestodes in red squirrels across the Holarctic do not appear to be differentiated morphologically relative to specific geographic localities (Makarikov et al. 2011). These constitute the first confirmed geographic records for A. macrocirrosa from eastern Beringia and the Nearctic. Arostrilepis macrocirrosa is a dominant cestode of Myodes voles at high latitudes across the Holarctic. Occurrence in other arvicolines and sciurids appear to represent incidental infections; A. macrocirrosa in red squirrels from Alaska is known based on multiple hosts and specimens collected at the Yukon Charley Rivers National Preserve, Lake Clark National Park and near Chitina (Appendix 1). Records in Myodes voles are currently limited to M. rutilus from North America, and further collections will be necessary to determine if there is a broader geographic range extending to temperate latitudes in the Nearctic. It is not known if A. macrocirrosa and A. cooki are parapatric and restricted to different species of Myodes, or whether populations of these cestodes occur in sympatry., Published as part of Makarikov, Arseny A., Galbreath, Kurt E. & Hoberg, Eric P., 2013, Parasite diversity at the Holarctic nexus: species of Arostrilepis (Eucestoda: Hymenolepididae) in voles and lemmings (Cricetidae: Arvicolinae) from greater Beringia, pp. 401-439 in Zootaxa 3608 (6) on pages 422-423, DOI: 10.11646/zootaxa.3608.6.1, http://zenodo.org/record/216055

Staphylocystis Villot, 1877, Published as part of Makarikov, Arseny A. & Kinsella, John M., 2013, Morphological and molecular differentiation of Staphylocystis clydesengeri n. sp. (Cestoda, Hymenolepididae) from the vagrant shrew, Sorex vagrans (Soricomorpha, Soricidae), in North America, pp. 389-400 in Zootaxa 3691 (3) on page 391, DOI: 10.11646/zootaxa.3691.3.7, http://zenodo.org/record/216838

Arostrilepis rauschorum sp. n. (Figs. 6, 7) Type host: Microtus oeconomus Pallas (Rodentia: Cricetidae: Arvicolinae). Other hosts: Microtus longicaudus (Merriam), M. pennsylvanicus (Ord), M. miurus Osgood and M. xanthognathus (Leach). Type locality: Adjacent toToolik Lake, Brooks Range, Alaska, near US Department of Energy research site (ca., 68 o 38 ���N, 149 o 36 ���W). Other localities: North-central interior, Alaska, near Bonanza Creek Research Station (62 o 42 ���N, 148 o 16 ���W); Gates of Arctic National Preserve, Brooks Range, Alaska, SE side of Walker Lake (67 o06���N, 154 o 16 ���W); Yukon Charley Rivers National Preserve, Alaska, across Yukon River from Glenn Creek Cabin (65 o 30 ���N, 142 o03���W); Kobuk Valley National Park, Alaska, Baird Mountains, Salmon River (67 o 36 ��� 11 ���N, 159 o 47 ��� 20 ���W); and Pattee Canyon, Missoula Co., Montana (46 o 48 ���N, 113 o 57 ���W, and 46 o 49 ���N, 113 o 58 ���W). Type material: Holotype MSB Para 1208 (field number FIN 4072484) from type host and locality by H. Henttonen and G. Batzli, 24 July 1984. Paratypes MSB1360, 1362 (AF 37462 C 1 / cyt- b sequence ; 37462 C 3) ex M. pennsylvanicus, by H. Henttonen at Bonanza Creek, 1 August 2000; MSB 1204 (FIN 2070784) from type host species and locality by H. Henttonen and G. Batzli, 7 July 1984; MSB 1211 (FIN 5220884) from type host species and locality by H. Henttonen and G. Batzli, 22 August 1984; MSB 1205, 1206 (FIN 3120884 -1, 2 slides; 3120884 - 2) from type host species and locality by H. Henttonen and G. Batzli, 12 August 1984; MSB 1363 (AF 42531 / cytb sequence ) from type host and locality by H. Henttonen, 8 August 2000; MSB 1365 (AF 42657) ex M. oeconomus by H. Henttonen et al., near Toolik Lake, 8 August 2000; MSB 1215 (JMK 02-04) ex M. longicaudus, by J.M. Kinsella at Pattee Canyon, Montana (46 o 48 ���N, 113 o 57 ���W), 8 October 2002; MSB 1369 (AF 49499 / cyt- b sequence ) ex M. longicaudus by H. Henttonen et al., from Yukon Charley, 8 August 2001; MSB 1370, 1371 (AF 59099 C3, 2 slides; 59099 C4, 2 slides) ex M. xanthognathus by A. M. Runck et al., from Gates of the Arctic National Preserve, 6 August 2002; and MSB 1216 (JMK 2009, 2 slides) ex M. pennsylvanicus, by J.M. Kinsella at Pattee Canyon, Montana (46 o 49 ���N, 113 o 58 ���W), 20 August 2009. See Appendix 1 for listing of additional identified voucher specimens. Symbiotype: Host specimen not deposited in a museum archive. Description: Based on 13 specimens. Fully developed strobila 120���185 mm long, with maximum width at pregravid or gravid proglottides, 2.4���3.8 mm. Strobila flat, consisting of 750���950 craspedote proglottides. Scolex slightly compressed dorso-ventrally, 240���300 (266, n = 7) wide, clearly wider than neck. Suckers unarmed, ovoid in surface view, relatively small, 130���190 �� 120���155 (166 �� 137, n = 15), with thin walls (Fig. 6 A, B). Rhynchus and rostellar apparatus absent. Neck relatively long and narrow, 125���190 (152, n = 10) wide. Two pairs of osmoregulatory canals, without transverse anastomoses. Dorsal osmoregulatory canals thin, 2���3.5 (2.6, n = 10) wide, situated predominantly in same sagittal plane as ventral canals. Ventral osmoregulatory canals 30���58 (42, n = 10) wide. Position of dorsal osmoregulatory canals not always constant; loops may be situated laterally to ventral canals. Genital pores unilateral, dextral. Genital ducts usually pass dorsally to longitudinal osmoregulatory canals, position of genital ducts between osmoregulatory canals within same strobila appears rarely (for no more than 10 % proglottides) (Fig. 6 C, D). Development of proglottides gradual, protandrous. Strobilar part containing juvenile proglottides without external segmentation; proglottides become externally distinct at level of premature part of strobila. Mature proglottides 170���230 �� 1330���1700 (202 �� 1485, n = 16), transversely elongate, trapeziform (Fig. 6 C, D). Testes relatively large, usually three in number, almost of equal size, 130���252 �� 110���194 (217 �� 164, n = 35), oval or pear-shaped, commonly situated in one row; poral testis separated from two antiporal testes by female gonads. Arrangement of testes may vary (triangle or triangle with flat angle). Cirrus-sac relatively large, 210���242 �� 31���42 (225 �� 37, n = 23), with well-developed external muscular layers. Antiporal part of cirrus-sac commonly not reaching or rarely overlapping ventral longitudinal canal (Figs. 6 D, 7 C). Genital atrium simple, cup-shaped, deep, opens laterally about middle or slightly anterior of lateral proglottis margin. Cirrus 77���92 (83, n = 18) long, conical, with relatively wide basal region, 18���23 (19, n = 18) in diameter, and narrow distal region, 12���15 (13, n = 18) in diameter; armed along entire length with relatively large (up to 3.5���4 long) rosethorn-shaped spines (Fig. 7 A). Internal seminal vesicle with circular musculature, ovoid, 87���122 �� 25���38 (102 �� 30, n = 23), shorter than half of cirrus-sac length (Figs. 6 D, 7 C). External seminal vesicle transversely elongate, 165���270 �� 55���87 (218 �� 70, n = 18), clearly outlined from vas deferens, with size approximately equal to seminal receptacle. Ovary 455���585 (537, n = 25) wide, median, fan-shaped, irregularly lobed, ventral to male genital organs, occupying substantial part of median field, slightly overlapping testes (Fig. 6 D). Vitellarium 75���123 �� 145���240 (96 �� 193, n = 25), postovarian, median, scarcely lobed. Vagina tubular, clearly distinct from seminal receptacle; ventral to cirrus-sac. Distal part of vagina 96���120 �� 8���18 (107 �� 13, n = 12), thick-walled, covered externally by dense layer of intensely stained cells; poral part of vagina infundibular (Fig. 7 C). Conductive part of vagina 170���230 �� 9���30 (210 �� 19, n = 10), thin-walled, vastly varying in diameter depending on degree of distention with sperm. Seminal receptacle relatively small, transversely elongate, 165���280 �� 50���92 (220 �� 68, n = 18). Uterus appears as complex of fine-walled anastomosing tubes of varying length and diameter, positioned ventrally to other organs. With development of proglottides, tubular structures increase in width and uterus becomes labyrinthine. Testes remain in postmature and pregravid proglottides; cirrus-sac and vagina persist in gravid proglottides. Gravid proglottides transversely elongate, 300���500 �� 1750���3800 (373 �� 2600, n = 16). Fully developed uterus labyrinthine, occupying entire median field and extending bilaterally beyond longitudinal osmoregulatory canals (Fig. 7 D). Uterus contains numerous (up to 2100) eggs. Eggs 22���35 �� 50���68, elliptical, with thin outer coat (up to 0.5); oncosphere 11���17 �� 15���22 (Fig. 7 B). Embryophore fusiform, 14���20 �� 38���47, with straight polar processes. Embryonic hooks small, 7.5���8.3 long. Etymology: Arostrilepis rauschorum sp. n. is named in honor of Robert L. Rausch and Virginia R. Rausch in recognition of their seminal and critical studies of parasites of arvicoline rodents, rodent systematics and biogeography at high latitudes of the Holarctic, and insights about the historical development of the Beringian fauna. Further, with the passing of Robert Rausch on 6 October 2012, this species and our recent studies across Beringia are dedicated to his memory and the legacy established by an extraordinary pioneer of parasitology and mammalogy in the north. Remarks: Arostrilepis rauschorum sp. n. is distinguished from congeners by the length and shape of the cirrus (Table 2). In A. rauschorum the cirrus is longer relative to those in A. beringiensis, A. tenuicirrosa, A. mariettavogeae, A. schilleri and A. gulyaevi, but smaller in comparison to A. macrocirrosa and A. cooki (Figs. 11, 12). The cirrus is armed with relatively large rosethorn-shaped spines and has a conical form; these features distinguish A. rauschorum from A. beringiensis, A. intermedia, A. janickii and A. schilleri (cylindrical cirrus), A. microtis and A. gulyaevi (cirrus with wide conical basal region and a cylindrical distal region) and A. tenuicirrosa (cirrus armed with relatively small needle-shaped spines and having a conical basal region and a very narrow cylindrical distal region). Arostrilepis rauschorum is characterized by a relatively wide strobila and ovary and a large seminal receptacle. The cirrus-sac is shorter than in A. horrida, but longer than in A. beringiensis, A. intermedia, A. janickii, A. mariettavogeae and A. schilleri. The egg and oncosphere are large relative to those in A. janickii, A. mariettavogeae and A. schilleri (see Table 2). The proximal end of the cirrus-sac in hermaphroditic mature proglottides commonly does not reach the ventral longitudinal canal or rarely overlaps it. Specimens of A. rauschorum are distinguished from A. horrida, A. macrocirrosa, A. tenuicirrosa, A. intermedia, A. schilleri, A. gulyaevi and A. cooki as its testes form one row; in the latter species, the testes are arranged in a triangle. Furthermore, the gravid proglottides are transversely elongate and the polar processes of the embryophore are straight in A. rauschorum. This species is a specific parasite of voles of the genus Microtus from North America., Published as part of Makarikov, Arseny A., Galbreath, Kurt E. & Hoberg, Eric P., 2013, Parasite diversity at the Holarctic nexus: species of Arostrilepis (Eucestoda: Hymenolepididae) in voles and lemmings (Cricetidae: Arvicolinae) from greater Beringia, pp. 401-439 in Zootaxa 3608 (6) on pages 416-419, DOI: 10.11646/zootaxa.3608.6.1, http://zenodo.org/record/216055

Family Hymenolepididae Ariola, 1899, Published as part of Makarikov, Arseny A. & Kinsella, John M., 2013, Morphological and molecular differentiation of Staphylocystis clydesengeri n. sp. (Cestoda, Hymenolepididae) from the vagrant shrew, Sorex vagrans (Soricomorpha, Soricidae), in North America, pp. 389-400 in Zootaxa 3691 (3) on page 391, DOI: 10.11646/zootaxa.3691.3.7, http://zenodo.org/record/216838

Arostrilepis cf. janickii Makarikov & Kontrimavichus, 2011 (Fig. 10) Host in Nearctic: Microtus oeconomus Pallas. Material examined: Vouchers include 3 specimens in M. oeconomus: MSB 1267 (field number AF 36025 C 2) by A.M. Runck et al. near Nome, Alaska, Skookum River (64 o 43 ��� 11 ���N, 163 o 58 ��� 35 ���W), 20 July 2000. MSB 1268, 1269 (AF 36738 C 2, C 3) by A.M. Runck et al., on Seward Peninsula, Alaska, Pilgrim Hot Spring Road (65 o05���09���N, 164 o 54 ���05���W), 1 August 2000. Sequence data for cyt- b are not currently available for these specimens from the Nearctic. See Appendix 1 for listing of additional identified voucher specimens from the Palearctic. Description: Based on 3 specimens. Fully developed strobila about 115 mm long, with maximum width at pregravid or gravid proglottides, up to 1.9 mm. Scolex slightly compressed dorso-ventrally, 273 wide, clearly wider than neck. Suckers ovoid in surface view, 170���190 �� 120���150 (184 �� 134, n = 4), with thin walls, prominent (Fig. 10 A). Neck relatively long and narrow. Genital ducts commonly pass dorsally to longitudinal osmoregulatory canals. Development of proglottides gradual, protandrous. Mature proglottides 160���190 �� 870���975 (177 �� 913, n = 5), transversely elongate, trapeziform (Fig. 10 C). Testes usually 3 in number, almost of equal size, 113���135 �� 70���90 (124 �� 78, n = 7), oval or pear-shaped, commonly situated in triangle or rarely, testes form one row; poral testis separated from 2 antiporal testes by female gonads. Cirrus sac relatively short, 157���180 �� 29���37 (167 �� 32, n = 11), commonly extending across ventral longitudinal canal (Fig. 10 C). Genital atrium simple, cup-shaped, deep, opens laterally about middle of lateral proglottis margin. Cirrus small, 62���81 �� 13���15 (69 �� 13, n = 13), cylindrical, armed along entire length with relatively small (up to 3.5 long) rosethorn-shaped spines (Fig. 10 B). Internal seminal vesicle ovoid, 70���87 �� 24���29 (80 �� 25, n = 5), shorter than half of cirrus sac length (Fig. 10 C). External seminal vesicle 104���120 �� 26���33 (114 �� 29, n = 5), with size approximately equal to that of seminal receptacle. Ovary 365���450 (405, n = 11) wide, median, fan-shaped, irregularly lobed, overlapping testes (Fig. 10 C). Vitellarium 67���90 �� 145���186 (77 �� 162, n = 7), postovarian, median, scarcely lobed. Vagina tubular, clearly distinct from seminal receptacle; ventral to cirrus sac. Seminal receptacle relatively small, 97���114 �� 27���36 (106 �� 31, n = 5). Gravid proglottides 340���390 �� 1650���1900 (369 �� 1802, n = 6). Fully developed uterus labyrinthine, occupying entire median field and extending bilaterally beyond longitudinal osmoregulatory canals. In present material outer coat of eggs very compressed, elliptical, with thin outer coat; oncosphere 10���11 �� 14���16. Embryophore fusiform, 13���14 �� 33���41, with straight polar processes. Embryonic hooks small, 6.5���7 long. Remarks: Specimens attributed to A. cf. janickii are largely consistent with the original description, particularly with respect to the structure, dimensions and spination of the cirrus (based on a partial view), dimensions of the cirrus sac and distribution of the testes (Makarikov & Kontrimavichus 2011). The few available specimens from the western Seward Peninsula, however, could not be completely evaluated, as the cirrus was not fully evaginated in these cestodes, nor was sequence data available; this does not preclude the possibility of another undescribed species being represented. Confirmation will require additional specimens from which these characters may be assessed. Assuming a correct identification, based on the current data, specimens of A. janickii had not been previously reported in arvicolines from the Nearctic and these would be the first confirmed records from eastern Beringia. A minimum geographic distribution in Microtus from the western Seward Peninsula may be established. The species has not been found to the east in the Brooks Range or to the south where A. rauschorum appears to be common., Published as part of Makarikov, Arseny A., Galbreath, Kurt E. & Hoberg, Eric P., 2013, Parasite diversity at the Holarctic nexus: species of Arostrilepis (Eucestoda: Hymenolepididae) in voles and lemmings (Cricetidae: Arvicolinae) from greater Beringia, pp. 401-439 in Zootaxa 3608 (6) on page 424, DOI: 10.11646/zootaxa.3608.6.1, http://zenodo.org/record/216055