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Ochotona hyperborea yesoensis Kishida, 1930 Ochotona yoshikurai Kishida, 1932 Holotype.— Botanic Garden and Museum, Field Science Center for Northern Biosphere, Hokkaido University HUNHM06803, adult male. Collected by Forestry Bureau of the Hokkaido Government Office in 11 October 1928. Type locality.— “In Prov. Kitami, a northern part of that Island ” (Kishida 1930). Notsukeushi, Kitami according to the specimen label. Description.— Pikas of the F genetic lineage; eastern acoustic race. Separation from the only spatial neighbor of this subspecies: O. h. fedoseevi ssp. n., is possible on the basis of genetic traits only. Distribution.— Sikhote-Alin Range, Sakhalin, and Hokkaido Islands. The range of this subspecies is isolated from those of other subspecies of northern pikas. Nomenclatural notes.— No available names were described from the mainland part of this subspecies’ distribution. Sakhalin pikas were described as Ochotona yoshikurai Kishida, 1932. The nominal taxon Ochotona yesoensis Kishida, 1930, was described from Hokkaido Island; that region was not included in this study (except for bioacoustics). Nevertheless, Kartavtseva et al. (2014) analyzed the Cytochrome b gene of pikas from south of Russian Far East (genetic lineage F in our study) and Hokkaido Island and concluded that pikas from Hokkaido, Sakhalin, and Sikhote-Alin represent the same phylogenetic clade. From the position of bioacoustics, it is known that pikas from Hokkaido belong to the eastern acoustic race (Kawamichi 1981; Lissovsky 2005; this study). Thus, there is no information about any taxonomic peculiarities with respect to the Japanese pika, and we consider it to belong to the same subspecies as pikas from mainland Sikhote-Alin Range and Sakhalin Island. The name Ochotona yesoensis Kishida, 1930 is the senior synonym for the pikas from the distribution range of this subspecies., Published as part of Lissovsky, Andrey A., Obolenskaya, Ekaterina V., Dokuchaev, Nikolai E. & Okhlopkov, Innokentiy M., 2021, Intraspecific variation and taxonomy of northern pika Ochotona hyperborea (Mammalia, Lagomorpha), pp. 28-53 in Journal of Mammalogy 102 (1) on page 40, DOI: 10.1093/jmammal/gyaa150, http://zenodo.org/record/7850862, {"references":["KISHIDA, K. 1930. Diagnosis of a new piping hare from Yeso. Lansania 2: 45 - 47.","KISHIDA, K. 1932. A new species of Piping Hares from Saghalien. Lansania 4: 149 - 152.","KARTAVTSEVA, I. V., I. N. SHEREMETYEVA, V. YU. GUS'KOV, A. A. VAKURIN, E. V. KUMAKSHEVA, AND L. V. FRISMAN. 2014. Taxonomic status of northern pika (Ochotona hyperborea) from the Sikhote-Alin. Vestnik of the Far East Branch of the Russian Academy of Sciences Issue 2: 79 - 85.","KAWAMICHI, T. 1981. Vocalizations of Ochotona as a taxonomic character. Pp. 324 - 339 in Proceedings of the World Lagomorph Conference (K. Myers and C. D. MacInnes, eds.). International Union for the Conservation of Nature and Natural Resources. Gland, Switzerland.","LISSOVSKY, A. A. 2005. Comparative analyses of the vocalization of pikas (Ochotona, Mammalia) from alpina-hyperborea group. Byulleten' Moskovskogo Obshchestva Ispytateley Prirody. Otdel Biologicheskii 110: 12 - 26."]}

Ochotona hyperborea hyperborea (Pallas, 1811) Lagomys hyperboreus var. ferruginea Schrenk, 1858 Lagomys hyperboreus var. normalis Schrenk, 1858 Lagomys litoralis Peters, 1882 Ochotona kolymensis Allen, 1903 Ochotona hyperborea minima Sokolov et al., 1994 Ochotona hyperborea shamani Sokolov et al., 1994 Type.— Specimen unknown; the nominal taxon Lepus hyperboreus was described by P. S. Pallas on the basis of specimens collected by Carl Heinrich Merck (Pallas 1811:152). Type locality.— Chukchi Peninsula “terris Tschuktschicis” (Pallas 1811:152). This toponym can describe large territory; however, C. Merck only visited the northern part of Chukchi Peninsula along the trek from St. Laurent Bay–Mechigmen Bay (13–26 of September 1791)–Kolyutchin Bay (14 October)– Amguema River (18 November)–Chaun River (19 January 1792)–Anyuyskiy Ostrog at the estuary of Angarka River (14 February; 66.85°N, 164.25°E; Merck 1782–1792 [1980]). The detailed map of the route was published by Sarychev (1811). Merck mentioned Anyuyskiy Ostrog as the final point of expedition, hence probably did not collect during the journey to Yakutsk through Nizhnekolymskiy Ostrog. Thus, the route from St. Laurent Bay to Anyuyskiy Ostrog should be considered as the type locality of O. h. hyperborea. Description.— Pikas of the A genetic lineage; northern acoustic race. Besides genetics, this subspecies can be distinguished from neighboring O. h. uralensis by the shape of parietal suture of the skull (Fig. 5). There are differences in the shape of the frequency modulation curve of alarm call with another spatial neighbor— O. h. cinereoflava (Fig. 6). Distribution.— The most northeastern part of the distribution range of the species: Chukchi and Kamchatka Peninsulas, Kolyma and Koryak Uplands, and Indigirka River basin. Provisionally, the distributional border with O. h. uralensis corresponds roughly to the watershed between the Yana and Indigirka River basins (at least its eastern part). Junction with distribution of O. h. cinereoflava is possible to the north of the Dzhugdzhur Range. Nomenclatural notes.— The nominal taxon Ochotona hyperborea shamani provisionally is considered herein a junior synonym of O. h. hyperborea. Ochotona h. shamani was described on the basis of a sample of pikas from the Indigirka River basin (near Shamanovo: 69.95°N, 147.57°E); we had no genetic data from this location. Our distributional analysis confirmed the proximity of the Indigirka River basin to the distribution O. h. hyperborea, although this factor alone cannot be used as a strong argument to select between O. h. hyperborea and O. h. uralensis. The prevailing shape of the parietal suture in the studied sample also suggests this population belongs to O. h. hyperborea., Published as part of Lissovsky, Andrey A., Obolenskaya, Ekaterina V., Dokuchaev, Nikolai E. & Okhlopkov, Innokentiy M., 2021, Intraspecific variation and taxonomy of northern pika Ochotona hyperborea (Mammalia, Lagomorpha), pp. 28-53 in Journal of Mammalogy 102 (1) on pages 36-37, DOI: 10.1093/jmammal/gyaa150, http://zenodo.org/record/7850862, {"references":["PALLAS, P. S. 1811. Zoographia Rosso-Asiatica, sistens omnium animalium in extenso Imperio Rossico et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum. Officina Caes. Acadamiae Scientiarum Impress, Petropoli (St. Petersburg) 1: 1 - 572.","SCHRENK, L. 1858. Reisen und Forschungen im Amur-Lande in den Jahren 1854 - 1856. Erste lieferung. Einleitung. Saugethiere des Amur-Landes. Kaiserlichen Akademie der Wissenschaften. St. Petersburg, Russia.","SOKOLOV, V. E., E. YU. IVANITSKAYA, V. V. GRUZDEV, AND V. G. HEPTNER. 1994. Млikopitaющii Рoссii i сopрidiльныk рigioнov. Zaйtiobрazныi [Mammals of Russia and adjacent regions: lagomorphs]. Nauka. Moscow, Russia.","SARYCHEV, G. 1811. Putisiсtvii kapitaнa Biллiнgсa cрiz Cukotсkuю ziмлю ot Biрiнgova pрoлiva do Нizнikoлымсkogo oсtрoga i pлavaнii kapitaнa Gaллa нa сudнi Ciрнoм Oрлi po Сiviрovoсtocнoмu okiaнu v 1791 godu; с pрiлoziнiiм слovaрi 12 - ti нaрiciй dikik нaрodov, нabлюdiнii нad сtuziю v Viрkнikoлымсkoм oсtрogi, i нaсtavлiнii daннogo kapitaнu Biллiнgсu iz goсudaрсtviннoй Adмiрaлtiйсtv- Koллigii [Travels of Captain Billings across the Chukchi Land from the Bering Strait to Nizhenakolymsk Fort, and voyage of Captain Gal on the ship Black Eagle across the Northeastern Sea in 1791; with the addition of a dictionary of 12 indigenous languages, observations on cold temperatures at the Verkhnekolymsk Fort, and the instructions given to the Captain Billings by the State Naval Department]. Morskaya Tipografiya. St. Petersburg, Russia.","FORMOZOV, N. A. 1991. Acoustic signalling of pikas (Ochotona, Lagomorpha): geographical, taxonomical and ecological aspects. Ph. D. dissertation, Biological Faculty, Moscow State University. Moscow, Russia [in Russian]."]}

Ochotona hyperborea uralensis Flerov, 1927 Ochotona hyperborea naumovi Formozov et Yakhontov, 2003 Lectotype (designated by).— Ognev 1940 ZIN 16107, adult female, skull and skin. Collected by K. K. Flerov 10.8.1926. Paralectotypes (Baranova et al. 1981): ZIN 16106, 16110, 16111. Type locality (according to museum label).— Eastern slope of Bolshoy Ural, Kharuta River, Synya River tributary (65.79°N, 62.78°E). Description.— Pikas of the B genetic lineage; northern acoustic race. The majority of the representatives of this subspecies (except for pikas from the Ural Mountains) have a clear V-shaped parietal suture (Fig. 5B). Accordingly, besides genetic identification, this subspecies can be distinguished from neighboring O. h. hyperborea by the shape of the parietal suture of the skull (Fig. 5). There are differences in the shape of the frequency modulation curve of alarm call with another spatial neighbor— O. h. cinereoflava (Fig. 6). Distribution.— The subspecies has a notably disjunct distribution—the northern Ural Mountains, Putorana Plateau and adjacent uplands, as well as Yakutia between the middle and lower Aldan River basin (to the north of Aldan Plateau) and Lena River basin (right bank, to the north of the Buotama River), including mountains of the Verkhoyansk region. The provisional distribution border with O. h. hyperborea corresponds roughly to the course of the Indigirka River. The case of mitochondrial introgression from the latter subspecies was found in the Adycha River basin. This subspecies should contact O. h. cinereoflava along the foothills of the Aldan Plateau and Dzhugdzhur Range, but an exact contact zone remains unknown. Nomenclatural notes.— Only two nominal taxa were described from the distribution range of this subspecies. The senior synonym appeared to be Ochotona hyperborea uralensis Flerov, 1927, which described a specimen from the very remote isolate in Ural Mountains., Published as part of Lissovsky, Andrey A., Obolenskaya, Ekaterina V., Dokuchaev, Nikolai E. & Okhlopkov, Innokentiy M., 2021, Intraspecific variation and taxonomy of northern pika Ochotona hyperborea (Mammalia, Lagomorpha), pp. 28-53 in Journal of Mammalogy 102 (1) on pages 37-38, DOI: 10.1093/jmammal/gyaa150, http://zenodo.org/record/7850862, {"references":["OGNEV, S. I. 1940. Zviрi СССР i pрiлizaщik сtрaн [Mammals of the USSR and adjacent countries]. Izdatel'stvo AN SSSR, Moscow - Leningrad 4: 1 - 616. (Translated as: Ognev, S. I. 1966. Mammals of the USSR and adjacent countries: rodents. Israel Program for Scientific Translations, Jerusalem 4: 1 - 429.)","BARANOVA, G. I., A. A. GUREEV, AND P. P. STRELKOV. 1981. Kataлog tipovыk ekziмpлiрov koллiktii Zooлogiciсkogo iнсtituta AН СССР. Млikopitaющii (Mammalia) Vыp. 1. Insectivora, Chiroptera, i Lagomorpha [Catalog of the type specimens from the collection of the Zoological Institute of the USSR Academy of Sciences. Mammals (Mammalia) Vol. 1. Insectivora, Chiroptera, and Lagomorpha]. Nauka. Leningrad, Russia."]}

Ochotona hyperborea cinereofusca (Schrenk, 1858) Ochotona svatoshi Turov, 1924 Ochotona hyperborea davanica Sokolov et al. 1994 Ochotona hyperborea stenorostrae Sokolov et al. 1994) Neotype (designated herein, see below [Nomenclatural notes] for the rationale for designation of a neotype, as per Art. 75 of the ICZN [1999]).— ZMMU S-175370, adult female, skull (Supplementary Data SD9). Collected by A. A. Lissovsky and S. V. Kruskop 8 April 2003. Right lower jaw is broken. The specimen identification (O. hyperborea versus O. mantchurica) was carried out on the basis of morphology and bioacoustics (Lissovsky 2005; Lissovsky et al. 2008). Type locality.— Russia, Zabaykalskiy Territory, left bank of Shilka River, 18 km below the mouth of the Dzheltuga River, 158 km above the mouth of the Shilka River; 53.421°N, 119.871°E. Description.— Pikas of the C genetic lineage; southern acoustic race. Specimens of O. h. cinereofusca can be distinguished from O. h. cinereoflava on the basis of genetic traits only. Differences with another spatial neighbor O. h. fedoseevi include the shape of the frequency modulation curve of alarm call (Fig. 6). Distribution.— Wide range from the Sayan and Khangai Mountains, Tannu-Ola Range in Tuva to the North Baikal Plateau in the north, and Khentiyn Nuruu, as well as mountains along the left bank of Shilka River in the east. Although O. h. cinereofusca seems to be separated from other subspecies by unsuitable habitats (Supplementary Data SD6 and SD7), there are two zones of introgression of O. h. cinereofusca genes into other subspecies. One of them, in Tukuringra Range, involves O. h. fedoseevi ssp. n. The second, in Aldan Plateau, involves O. h. cinereoflava. Nomenclatural notes.— The name cinereofusca was not used stably during the 20th century. Different authors have applied this name to different pikas of the Manchurian region. Leaving aside the problem of incorrect taxonomic identification of pikas from this region (Lissovsky et al. 2008; Formozov and Baklushinskaya 2011), the majority of authors have applied the name cinereofusca to O. mantchurica, mainly as “ O. alpina cinereofusca ” (Kuznetsov 1929; Ognev 1940; Feng and Zheng 1985; Smith et al. 1990; Hoffmann and Smith 2005). Yakhontov and Formozov (1992) proposed using this name for O. hyperborea. Sokolov et al. (1994) did not use this name at all, although they placed it in the list of synonyms of O. alpina (considering O. mantchurica) and stated that the name belongs to O. hyperborea. The nominal taxon Lagomys hyperboreus var. cinereofusca was described by L. Schrenk on the basis of two specimens collected by R. Maak in the “ Amur River.” The specimens were lost before S. Ognev (1940) wrote his revision. Published dates of the collected specimens allowed several authors (Yakhontov and Formozov 1992; Lissovsky et al. 2008; Formozov and Baklushinskaya 2011) to find exact localities where the specimens were collected. On the first day, 18 May, Maak collected a specimen in a site near the mouth of Tontokoy Spring (53.194°N, 119.419°E); the fact of collecting a pika is mentioned in the report (Maak 1859). The second day, 20 May, Maak landed at a meadow near the lower Anikino River (53.444°N, 120.287°E) and near the mouth of the Grishkina River (53.479°N, 120.760°E); no information about pikas was recorded for this day. All three sites are located on the left bank of the Shilka River. Lissovsky explored pika settlements along the entire length of Shilka River and found that the river separates O. hyperborea and O. mantchurica (Lissovsky et al. 2008). Northern pikas sparsely inhabit only the left bank, while the Manchurian pikas live on the right bank only. Lissovsky explored all three places that comprise the terra typica of O. h. cinereofusca (Tontokoy, Anikino, and Grishkina) and did not find pikas in any area. The meadow near the lower Anikino River has good taluses suitable for pikas, while stony habitats near the Tontokoy Spring and near mouth of the Grishkina River were found unsuitable for pikas at the moment of observation. It is highly likely that the stones were covered by vegetation during the last 150 years and abandoned by pikas. During the expeditions along the Shilka, one specimen (ZMMU S-175370) was collected between Tontokoy and Anikino. Several years later, Formozov and Baklushinskaya (2011) designated the lost specimen collected by R. Maak near the mouth of Tontokoy Spring, as a lectotype, acting under the Art. 74.4 (ICZN 1999). The conclusion of this publication includes the statement that allocation of the name O. h. cinereofusca to O. hyperborea or O. mantchurica cannot be considered as definitively solved because pikas from the terra typica (Tontokoy Spring at the moment) had not yet been studied. Such a detailed explanation is necessary because the taxonomic allocation of the nominal taxon Lagomys hyperboreus var. cinereofusca is crucial for the arrangement of pika names under discussion. The name cinereofusca was described long before the name mantchurica. Thus, if the name belongs to O. mantchurica, the latter species and the subspecies of O. hyperborea under discussion will be renamed. Until there is some probability of allocating cinereofusca to O. mantchurica, the nomenclature of the two species remains unstable. The nomenclatural action of Formozov and Baklushinskaya destabilize the situation because it deprives us of a chance to analyze material from the terra typica (because there is no more available habitat for pikas in this location). The lectotype (and paralectotype) of the nominal taxon Lagomys hyperboreus var. cinereofusca was collected on the border of the distribution of two pika species; this lectotype (and the paralectotype) was lost many years ago, while the type specimen is urgently needed to identify taxonomic allocation of the name-bearer; hence, we designate a neotype here. The neotype is selected among the specimens collected as close as possible to the localities collected by Maak. This action should resolve the use of unstable names of two species of pikas from Amur region., Published as part of Lissovsky, Andrey A., Obolenskaya, Ekaterina V., Dokuchaev, Nikolai E. & Okhlopkov, Innokentiy M., 2021, Intraspecific variation and taxonomy of northern pika Ochotona hyperborea (Mammalia, Lagomorpha), pp. 28-53 in Journal of Mammalogy 102 (1) on page 38, DOI: 10.1093/jmammal/gyaa150, http://zenodo.org/record/7850862, {"references":["SCHRENK, L. 1858. Reisen und Forschungen im Amur-Lande in den Jahren 1854 - 1856. Erste lieferung. Einleitung. Saugethiere des Amur-Landes. Kaiserlichen Akademie der Wissenschaften. St. Petersburg, Russia.","SOKOLOV, V. E., E. YU. IVANITSKAYA, V. V. GRUZDEV, AND V. G. HEPTNER. 1994. Млikopitaющii Рoссii i сopрidiльныk рigioнov. Zaйtiobрazныi [Mammals of Russia and adjacent regions: lagomorphs]. Nauka. Moscow, Russia.","LISSOVSKY, A. A. 2005. Comparative analyses of the vocalization of pikas (Ochotona, Mammalia) from alpina-hyperborea group. Byulleten' Moskovskogo Obshchestva Ispytateley Prirody. Otdel Biologicheskii 110: 12 - 26.","LISSOVSKY, A. A., Q. YANG, AND A. E. PILNIKOV. 2008. Taxonomy and distribution of the pikas (Ochotona, Lagomorpha) of alpina-hyperborea group in North-East China and adjacent territories. Russian Journal of Theriology 7: 5 - 16.","KUZNETSOV, B. A. 1929. Gрыzuны Voсtocнogo Zabaйkaльi [Rodents of East Transbaikalia]. Proceedings of the Association of Scientific-Research Institute at the Faculty of Mathematics and Physics I MSU 2: 59 - 106.","OGNEV, S. I. 1940. Zviрi СССР i pрiлizaщik сtрaн [Mammals of the USSR and adjacent countries]. Izdatel'stvo AN SSSR, Moscow - Leningrad 4: 1 - 616. (Translated as: Ognev, S. I. 1966. Mammals of the USSR and adjacent countries: rodents. Israel Program for Scientific Translations, Jerusalem 4: 1 - 429.)","SMITH, A. T., N. FORMOZOV, R. HOFFMANN, C. ZHENG, AND M. ERBAJEVA. 1990. The pikas. Pp. 14 - 60 in Rabbits, hares and pikas: status survey and conservation action plan (J. A. Chapman and J. E. C. Flux, eds.). IUCN. Gland, Switzerland.","MAAK, R. 1859. Putisiсtvii нa Aмuр, сoviрsiннoi po рaсpoрiziнiю Сibiрсkago Otdiлa Iмpiрatoрсkago Рuссkago Giogрaфiciсkago Obщiсtva, v 1855 g [A travel to Amur in 1855, committed by order of the Siberian Division of the Russian Geographical Society]. Izdaniye S. F. Solovyeva. St. Petersburg, Russia."]}

Ochotona hyperborea fedoseevi, ssp. nov. Lissovsky Holotype.— Zoological Museum of Moscow State University S-194595, subadult male, skin and skull (Figs. 8 and 9). Collected by A. A. Lissovsky and E. V. Obolenskaya 12 September 2014 at Russia: Khabarovsk Territory; Bureinskiy Range, Bolshoy Suluk Lake; 51.301 N, 134.333 E. Etymology.— The name is given in honor of famous land surveyor G. A. Fedoseev, who made a great contribution to the description of the Russian Far East; his popular books gave impulse to several generations of investigators to explore this territory. Diagnosis.— The new subspecies can be distinguished from the rest O. hyperborea (except for Ochotona hyperborea yesoensis) by the specific shape of the frequency modulation curve of alarm call: the frequency grows first, then decreases (Fig. 6); ascending branch of the curve prevails over descending. Maximum of basal frequency is above 5500 Hz. Representatives of the new subspecies form separate clade E in the molecular phylogenetic analysis (Figs. 1 and 3). Description.— The new subspecies cannot be distinguished from remaining O. hyperborea by morphological traits; the identification should be carried out on the basis of bioacoustic and genetic features described in the diagnosis. The dorsal parts of pikas in summer fur are reddish brown, ventral parts are ochraceous. The dorsal parts of winter specimens are gray brown with darker median stripe, sides are sandy, ventral parts are gray or sandy. Fur near eyes is gray. Condylobasal length 37.3 mm (35.4–39); zygomatic breadth 20.0 mm (18.9–20.7). Distribution.— Inhabits a number of mountain ranges in the Russian Far East—from the left bank of the lower Amur River to Tukuringra Range in the west: Djaki-Unakhta-Yanbiyana, Bureinskiy, Turana, Ezop, Tukuringra–Dzhagdy, and closely situated mountain ranges and uplands. The northern limit of the distribution goes along the Uda River Valley and Verkhnezeyskaya Plain. The Tukuringra Range population bears mitotypes of O. h. cinereofusca. The contact zone with O. h. cinereoflava has not yet been found, our model suggests this zone should be situated in the area of the upper Aldan River. Nomenclatural notes.— No available names have been previously described from the distribution range of this subspecies. We therefore describe a new subspecies herein., Published as part of Lissovsky, Andrey A., Obolenskaya, Ekaterina V., Dokuchaev, Nikolai E. & Okhlopkov, Innokentiy M., 2021, Intraspecific variation and taxonomy of northern pika Ochotona hyperborea (Mammalia, Lagomorpha), pp. 28-53 in Journal of Mammalogy 102 (1) on page 40, DOI: 10.1093/jmammal/gyaa150, http://zenodo.org/record/7850862