taxonID	type	description	language	source
482787C8FFDE4B6CF6BA854BFE90E614.taxon	description	The adult tapeworms of Hymenolepis sp. 1 (the isolate numbers 18 AK 285 and 19 AK 270) were found from Ap. speciosus in Asahikawa. The following description was made based on one specimen (Fig. 3 A, B): Scolex 0.43 wide. Rostellum unarmed. Rostellar sac, 0.10 long by 0.08 wide. Suckers circular, four in number, 0.11 – 0.14 in diameter. Neck region not discernible. Mature proglottids much shorter in length than width, 0.26 long by 1.9 wide. Genital pore unilateral. Cirrus sac 0.22 – 0.29 long. Cirrus 0.03 – 0.04 long. Testes spherical, three in number in each mature proglottid, almost of equal size, 0.097 – 0.12 in diameter, arranged in straight line. Ovary lobed, 0.19 long by 0.60 wide. Vitelline gland lobed, 0.018 – 0.027 long by 0.065 – 0.10 wide. The genus Hymenolepis sensu stricto is characterized by a rudimentary unarmed rostellum and other defined morphological traits (Haukisalmi et al. 2010 a; Makarikov and Tkach 2013; Nkouawa et al. 2016). The following four species of Hymenolepis have been found from Apodemus mice in Eurasia: Hym. diminuta, Hym. Hibernia Montgomery, Montgomery, and Dunn, 1987, Hym. pseudodiminuta, and Hymenolepis apodemi Makarikov and Tkach, 2013 (Montgomery et al. 1987; Asakawa 1989; Tenora et al. 1994; Makarikov and Tkach 2013). A lengthy cirrus of mature proglottids (0.060 – 0.076 mm in length) is distinctive of Hym. apodemi (Makarikov and Tkach 2013). Although Hym. diminuta and Hym. hibernia are quite similar to each other, the latter has a lateral bulge of each proglottid extending backward over adjacent proglottids (Montgomery et al. 1987). The present phylogenetic trees of 28 S rDNA and cox 1 showed that our isolates from Hokkaido and the isolate of Hym. pseudodiminuta from Honshu (Ishih et al. 2003) might be classified as Hym. hibernia (Figs 2, 4). The values of pairwise divergence of cox 1 sequences among 14 isolates of Hymenolepis sp. 1 from Hokkaido, Hym. pseudodiminuta from Honshu, and Hym. hibernia from Korea, Europe, Turkey ranged from 0.048 to 0 (mean = 0.030). The maximum value was reduced to 0.029, when excluded the Turkish isolates.	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
482787C8FFDE4B6CF6BA854BFE90E614.taxon	description	The adult tapeworms of Hymenolepis sp. 2 (nos. JA 173, JA 175, JA 220, JA 244, and H 22 B) were found from Ap. speciosus and My. rufocanus in Otofuke. The specimens were unsuitable for morphological diagnosis. The phylogenetic trees of 28 S rDNA and cox 1 confirmed that this unknown species belongs to Hymenolepis (Figs 2, 4). The cox 1 sequences of the five isolates were completely identical to one another. Further sample collections are needed to clarify whether this unknown species is Hym. apodemi or another new cryptic species.	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
482787C8FFDC4B6DF41385AAFECEE87E.taxon	description	Members of the genus Arostrilepis are widely distribut- ed in the Holarctic region, mainly using voles as definitive hosts. Arostrilepis horrida (Linstow, 1901) was once considered to be a morphologically hypervariable and geographically widespread species (Makarikov et al. 2011; Galbreath et al. 2013; Makarikov and Hoberg 2016). However, a recent molecular phylogenetic analysis using mitochondrial cytb sequences has revealed some cryptic species within the assemblage of the so-called “ A. horrida ” (Makarikov and Kontrimavichus 2011; Makarikov et al. 2011, 2013). There are now eight species of Arostrilepis from the Palearctic region (Makarikov and Kontrimavichus 2011; Makarikov et al. 2011, 2013): A. horrida sensu stricto, A. tenuicirrosa, A. beringiensis (Kontrimavichus and Smirnova, 1991), A. microtis Gulyaev and Chechulin, 1997, A. macrocirrosa Makarikov, Gulyaev, and Kontrimavichus, 2011, A. intermedia Makarikov and Kontrimavichus, 2011, A. janickii Makarikov and Kontrimavichus, 2011, and A. gulyaevi Makarikov, Galbreath, and Hoberg, 2013. Morphological differential points of these species are limited to the form and size of the cirrus, and other characters may vary and overlap among all species (Makarikov et al. 2011). All the specimens of Arostrilepis obtained in this study were subjected to DNA sequencing of mitochondrial cox 1 and cytb. The values of pairwise divergence ranged at very low levels from 0.009 to 0.003 in both cox 1 and cytb, confirming the involvement of a single species. A phylogenetic tree of cytb showed that our samples should be classified as A. tenuicirrosa (Fig. 5). Voles of Myodes spp. widely distributed from Europe to Russia serve as the definitive host for A. tenuicirrosa (Galbreath et al. 2013). The present study expands the distribution range to Hokkaido. There are also records of Arostrilepis horrida sensu lato from Eothenomys smithii (Thomas, 1905) and Eothenomys andersoni (Thomas, 1905) in Honshu (Asakawa et al. 2002). Further specimens of Arostrilepis from Honshu and their DNA sequences are required to better understand the phylogeography and taxonomy of this group in Japan.	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
482787C8FFDD4B62F6BA8760FDADE7B9.taxon	description	All the six isolates of Microsomacanthus sp. were subjected to DNA sequencing of cox 1. The resultant sequences were highly homogeneous (mean pairwise divergence = 0.004). A BLAST homology search could not detect any sequences similar to them. In the case of 28 S rDNA, the unknown species showed 99.6 % similarity (1360 out of 1365 nucleotides identical) to Microsomacanthus crenatus (Goeze, 1782) from Apodemus sylvaticus (Linnaeus, 1758) in Croatia (database accession no. GU 166246). Haukisalmi et al. (2010 a) treated this isolate (M. crenatus in DNA databases) as “ Hymenolepis ” muris-sylvatici. A 28 S rDNA-based phylogenetic tree supports that the unknown species belongs to Microsomacanthus (see the isolate JA 313 in Fig. 2). It is highly probable that “ Hymenolepis ” muris-sylvatici or Microsomacanthus murissylvatici (Rudolphi, 1819) is identical to M. crenatus (Czaplinski and Vaucher 1994; Tenora 2004). Most members of Microsomacanthus exclusively use birds as definitive hosts (Yamaguti 1959; Czaplinski and Vaucher 1994). However, M. crenatus have been recorded exceptionally from Apodemus mice in Europe (Prokopič 1967; Behnke et al. 1999; Tenora 2004; Klimpel et al. 2007). Such a distant host-switching suggests a possibility that the rodent-related species belong to another genus. The confirmation of this hypothesis is challenging, because the present DNA databases lack the 28 S rDNA and cox 1 sequences of Microsomacanthus from birds. Our specimen is morphologically similar to M. murissilvatici, particularly in the sizes of strobila, scolex and suckers, when compared with the data of previous reports (Baer 1931; Prokopič 1967). A preliminary phylogeny of 28 S rDNA (Haukisalmi et al. 2010 a) suggests that “ Hymenolepis ” muris-sylvatici is sister to Rodentolepis evaginata (Barker and Andrews, 1915). Molecular phylogenetic assessments and subsequent taxonomic revisions are required for the rodent-related species of Microsomacanthus.	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
482787C8FFD24B63F66C87E2FC6AE533.taxon	description	The adult tapeworms of Paranoplocephala kalelai (nos. 19 AK 378, 19 AK 412, 19 AK 419, 19 AK 436, 19 AK 454 - 2, and 19 AK 454 - 3) were found from My. rufocanus in Asahikawa. This is the first record from Japan. The following description was made based on one specimen (Fig. 6 A – C): Scolex distinctly wider than neck, 0.59 in maximum width. Rostellum absent. Suckers protruding, crateriform, four in number, 0.28 – 0.29 in diameter. Mature proglottids shorter in length than width, 0.40 – 0.45 long by 0.74 – 0.78 wide. Genital pore located on posterior half of lateral margin. Testes spherical, 26 – 30 in number, forming compact group, 0.04 – 0.06 in diameter. Cirrus sac 0.13 – 0.15 long by 0.07 – 0.08 wide. Seminal receptacle pyriform or ovoid, 0.13 – 0.15 long by 0.17 – 0.19 wide. Ovary irregularly lobed, 0.40 – 0.48 in width. Vitellarium asymmetrically bilobed, 0.19 – 0.22 long by 0.07 – 0.10 wide. Gravid proglottids longer in length than width, 1.09 – 1.29 long by 0.82 – 0.93 wide. Uterus labyrinthine, occupying entire field of proglottid. A phylogenetic tree of cox 1 showed that P. kalelai, Paranoplocephala omphalodes (Hermann, 1783) sensu stricto, Paranoplocephala macrocephala (Douthitt, 1915), and Paranoplocephala jarrelli (Haukisalmi, Henttonen, and Hardman, 2006) are distinguishable from one another, and that our isolates from Hokkaido should be classified as P. kalelai (Fig. 7). The tree reconfirmed that the Fennoscandian isolates of P. kalelai could be divided into two clades named as Narvik and Kilpisjärvi (Haukisalmi et al. 2004). The Hokkaido isolates of P. kalelai showed a sister relationship to the Narvik clade. The values of pairwise divergence between the Narvik and Kilpisjärvi clades and between the Hokkaido and Narvik clades were 0.034 and 0.015, respectively. The sequencing of 28 S rDNA supports the species identification (see the isolate 19 AK 378 in Fig. 2). Members of the genus Paranoplocephala Lühe, 1910 are widely distributed in the Holarctic region (Haukisalmi et al. 2014). Currently, the species complex of P. omphalodes sensu lato has been divided into several species (Haukisalmi and Henttonen 2003; Haukisalmi et al. 2004, 2007; Vlasenko et al. 2019). Most species of Paranoplocephala parasitize Microtus voles, whereas P. kalelai is specific to Myodes voles in Fennoscandia (Tenora et al. 1985; Haukisalmi et al. 2004, 2007). Our data suggest that P. kalelai is distributed widely from Fennoscandia to the Far East, along with the geographic expansion of Myodes voles.	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
482787C8FFD14B61F439854BFAC3E40F.taxon	description	The former Hyd. taeniaeformis sensu lato is a cryptic species complex, including Hyd. taeniaeformis and Hydatigera kamiyai Iwaki, 2016 (Galimberti et al. 2012; Jia et al. 2012; Nakao et al. 2013; Lavikainen et al. 2016). Both species prevail in Hokkaido, using domestic cats and rodents as definitive and intermediate hosts, respectively (Iwaki et al. 1993; Asakawa and Fukumoto 1997; Okamoto et al. 2007). In general, the host specificity of both species is strict in selecting intermediate hosts, namely, murids (e. g., R. norvegicus) for Hyd. taeniaeformis and cricetids (e. g., My. rufocanus) for Hyd. kamiyai (Nonaka et al. 1994; Iwaki et al. 1994 a; Lavikainen et al. 2016). However, in this study, fully developed strobilocerci of Hyd. taeniaeformis were found from cricetids. In Japan, the adult tapeworms of Hyd. taeniaeformis sensu lato have been found from the following carnivores: the cat, Felis catus Linnaeus, 1758; the dog Canis lupus familiaris Linnaeus, 1758; the Tsushima leopard cat, Prionailurus bengalensis euptilurus (Elliot, 1871); the raccoon, Procyon lotor (Linnaeus, 1758) (Oishi and Kume 1973; Yagisawa 1978; Uga et al. 1983; Matoba et al. 2003; Yasuda et al. 2005). A special attention should be paid to the raccoon, an invasive species from North America, because it is now increasing in number in Hokkaido (Ikeda et al. 2004). Both Hyd. taeniaeformis and Hyd. kamiyai are also alien species probably due to the anthropogenic movement of domestic cats or commensal rodents into Japan (Lavikainen et al. 2016).	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
482787C8FFD14B61F42C8025FE5CE657.taxon	description	This species is widely distributed in the northern hemisphere (Deplazes et al. 2019). The larval cysticerci were already found from Ap. speciosus and Mi. montebelli in Honshu (Sato and Kamiya 1989; Uchida et al. 1990, 2001; Ihama et al. 2000). Canid carnivores serve as the definitive hosts (Freeman 1962), and the cysticerci multiply in the subcutaneous tissue and body cavity of wide-ranging mammalian intermediate hosts including humans (Delvalle 1989; Miyaji et al. 1990; Deplazes et al. 2019). In Hokkaido, the main definitive host is still unclear.	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
482787C8FFD14B67F6B58782FD47E676.taxon	description	This species is well known to cause human alveolar echinococcosis, which is one of the most important zoonoses in the Holarctic region. A small number of the human cases occur every year in Hokkaido (Takahashi et al. 2005; National Institute of Infectious Diseases 2019; Kamiyama 2020). The red fox, Vulpes vulpes Linnaeus, 1758, is the main definitive host of E. multilocularis (Romig et al. 2017; Tsukada et al. 2000). It seems that E. multilocularis has been recently introduced into Hokkaido from the Kurile islands by migrant foxes on drift ice or the anthropogenic movement of foxes and rapidly spread throughout Hokkaido (Yamashita 1956). A lower genetic divergence among the parasite population in Hokkaido supports this hypothesis (Nakao et al. 2003; Okamoto et al. 2007). Red foxes are colonizing urban areas because they can adapt to the artificial environment (Tsukada et al. 2000; Uraguchi et al. 2009). One of the present isolates was obtained from an urban park in Asahikawa. Conclusion. Our collections and literature search revealed that the cestode fauna of murid and cricetid rodents in Japan consists of at least 30 species from 6 families (Table 2). Among them, 23 species occur in Hokkaido. The species composition is strongly affected by the nearby Eurasian continent. It is most likely that the diverse cestode fauna is caused by rodent migrations over land bridges between Hokkaido and Sakhalin and between Hokkaido and Honshu during the late Pleistocene. The recent anthropogenic introduction of host animals seems to be responsible for the distribution of T. crassiceps, Hyd. taeniaeformis, Hyd. kamiyai, and E. multilocularis. This report contains the first records of A. tenuicirrosa, P. kalelai, and T. crassiceps from Hokkaido. The former two species correspond to the first records in Japan. Several cestodes are still waiting to be further investigated and placed into suitable taxonomic positions by an integrated approach of molecular phylogeny, morphology, and ecology. In this study, nuclear and mitochondrial DNA barcodes were generated for some cestodes from rodents in Hokkaido (Table 1). The DNA barcoding system has a great potential to revise existing classification systems and to accelerate the discovery of cryptic species. In order to expand its potential, it is necessary to enhance DNA databases with reliable data from cestode taxonomists across the world. Moreover, it is most important to standardize DNA markers for the taxonomy of cestodes. In previous studies on rodent cestodes, various regions of nuclear and mitochondrial DNA were used as DNA markers for phylogenetic analyses (Haukisalmi et al. 2004; Haukisalmi et al. 2010 b; Makarikov et al. 2013). The use of the markers, however, lacks consistency. We especially recommend the primer set, JB 3 and CO 1 - R trema (Miura et al. 2005), because it enables the amplification of cox 1 sequences (approximately 800 bases in length) from most platyhelminths. The sequencing of mitochondrial cox 1 and nuclear 28 S rDNA is a minimum requirement for the species and genus level identification of cestodes.	en	Sasaki, Mizuki, Anders, Jason Lee, Nakao, Minoru (2021): Cestode fauna of murid and cricetid rodents in Hokkaido, Japan, with assignment of DNA barcodes. Species Diversity 26: 255-272, DOI: 10.12782/specdiv.26.255
