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Quaternary Hystrix (Rodentia, Mammalia) from North China: Taxonomy, stratigraphy and zoogeography, with discussions on the distribution of Hystrix in Palearctic Eurasia

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Abstract

Today, Hystrix is a genus that only occurs in warm areas. However, during the Pleistocene and even the early Holocene, this species frequently appeared in the northern part of China, which belongs to the Palearctic Region. A total of 22 fossil Hystrix-bearing localities have been reported in North China, the time span of which covers almost all the stages of Quaternary Period. Among those localities, Tianyuan Cave, a recently discovered locality near Zhoukoudian, contains abundant Hystrix materials. This is the latest Hystrix record in North China. Currently, two Hystrix species have been recognized in North China, Hystrix subcristata and Hystrix lagrelii. The small porcupine from the Zhoukoudian area should be referred to H. lagrelii instead of Atherurus as previously proposed by other authors, and therefore the occurrence of Atherurus in the Palearctic Region is denied. Based on both fossil and extant records, it seems that Hystrix can be generally used as indicator of warm climate.

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... These are comparable molars, but the lower-crowned V8833 is longer (rectangular), reflecting its earlier stage of wear refossa. Tong (2008) also noted two porcupines in Pleistocene sites of China, but came to the conclusion that appropriate names are H. lagrelli and H. subcristata. The discrepancy in the identification of the larger porcupine may be explained by different comparative material. ...
... The discrepancy in the identification of the larger porcupine may be explained by different comparative material. Except for a Nihewan premolar, all of Tong's (2008) comparative material postdates considerably the Haiyan porcupine. For example, V8834 is over 1 m.y. ...
... For example, V8834 is over 1 m.y. older than and larger than the Gongwangling porcupine (see Hu and Qi 1978) that Tong (2008) discussed. Hystrix subcristata may be the appropriate name for large late Pleistocene porcupines in North China. ...
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Surface finds of small mammal fossils tend to be large body-size species; small species are retrieved more readily by applying special collecting techniques. The bias against small fossil recovery from Yushe Basin was greatly diminished by screen washing, beginning in 1987. Prior to that time, the historical monographic study by Teilhard de Chardin on the rodents of Yushe Basin focused on relatively large body-size species, especially beavers and zokors . Two other large kinds of rodents, bamboo rats and porcupines, were known to Teilhard and are reviewed here. Of these, the bamboo rats include Brachyrhizomys shansius Teilhard de Chardin, 1942 named for fossils from the Pliocene of Yushe, and a smaller late Miocene bamboo rat recovered by the Sino-American collaborative field team in 1991. This species is the oldest rhizomyine in North China, and the oldest species assignable to the extant Rhizomys group. Specimens representing the Old World porcupine Hystrix have stratigraphic importance, with a higher crowned species replacing Hystrix gansuensis by the early Pleistocene.
... В среднемпозднем плейстоцене в области распространения фауны Stegodon-Ailuropoda отмечены H. b. punungensis Van Weers, 2003 (вымерший подвид) на Яве [2], H. b. subcristata в Китае (как H. subcristata, H. cf. subcristata) [3,4] и Таиланде (как H. hodgsoni subcristata) [5], H. brachyura ssp. во Вьетнаме и на Суматре [6-8; Van Weers, 1985, Ява) и H. lagrelli Lönnberg, 1924 (Китай и Ява) [2]. ...
... С этой точкой зрения не согласен Х. Тун [4], отстаивающий принадлежность обсуждаемых ископаемых форм из Китая к рецентному H. subcristata как отдельному виду, который по строению черепа сближается им с подродом Hystrix, а не Acanthion, и характеризуется в том числе видовыми признаками H. kiangsenensis по Веерсу, включая длинные носовые кости и высокий череп. Кроме того, Тун указывает в числе диагностических признаков этой формы крупный удлиненный верхний премоляр [4], а по приведенным им морфометрическим данным [3: табл. ...
... С этой точкой зрения не согласен Х. Тун [4], отстаивающий принадлежность обсуждаемых ископаемых форм из Китая к рецентному H. subcristata как отдельному виду, который по строению черепа сближается им с подродом Hystrix, а не Acanthion, и характеризуется в том числе видовыми признаками H. kiangsenensis по Веерсу, включая длинные носовые кости и высокий череп. Кроме того, Тун указывает в числе диагностических признаков этой формы крупный удлиненный верхний премоляр [4], а по приведенным им морфометрическим данным [3: табл. 3] видно, что относительно крупным и удлиненным является и нижний премоляр (μOL P 4 /M 1-2 = 1.176, ...
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The dental remains of the porcupine from the Pleistocene deposits of the Lang Trang cave in northern Vietnam are described. Previously, this porcupine was identified as Hystrix brachyura Linnaeus, 1758 or H. indica Kerr, 1792. Based on the analysis of morphometric data, the systematic position of the form from Lang Trang and other large Pleistocene porcupines of Vietnam is determined as Hystrix kiangsenensis Wang, 1931.
... Middle to Late Pleistocene forms that occurred within the range of the Stegodon-Ailuropoda fauna include H. b. punungensis Van Weers, 2003 (fossil subspecies) in Java [2], H. b. subcristata in China (as H. subcristata, H. cf. subcristata) [3,4] and Thailand (as H. hodgsoni subcristata) [5], H. brachyura ssp. in Vietnam and Sumatra [e.g., [6][7][8] [2]. Systematic identification of isolated teeth of fossil porcupines is to a large extent based on their morphometric characteristics. ...
... Noting the relatively large size, relatively high skull and long nasals, van Weers and Zheng [11] assigned Middle Pleistocene porcupines from China to the species H. kiangsenensis within the subgenus Hystrix, Fig. 1 This view is rejected by Tong [4], who argues that these fossil Chinese forms belong to the recent H. sub-cristata as a separate species for which this author suggests an affinity with Hystrix, rather than Acanthion, and is characterized, among other things, by species characters of H. kiangsenensis according to van Weers, including long nasals and a high skull. In addition, Tong indicates the large elongated upper premolar as one of the diagnostic characters for this form [4], while his morphometric data [3, Table 3] show that the lower premolar is also relatively large and elongated (μOL P 4 /M 1-2 = 1.176, ...
... Noting the relatively large size, relatively high skull and long nasals, van Weers and Zheng [11] assigned Middle Pleistocene porcupines from China to the species H. kiangsenensis within the subgenus Hystrix, Fig. 1 This view is rejected by Tong [4], who argues that these fossil Chinese forms belong to the recent H. sub-cristata as a separate species for which this author suggests an affinity with Hystrix, rather than Acanthion, and is characterized, among other things, by species characters of H. kiangsenensis according to van Weers, including long nasals and a high skull. In addition, Tong indicates the large elongated upper premolar as one of the diagnostic characters for this form [4], while his morphometric data [3, Table 3] show that the lower premolar is also relatively large and elongated (μOL P 4 /M 1-2 = 1.176, P 4 /M 1 = 1.329, ...
Article
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The dental remains of the porcupine from the Pleistocene deposits of the Lang Trang cave in northern Vietnam are described. Previously, this porcupine was identified as Hystrix brachyura Linnaeus, 1758 or H. indica Kerr, 1792. Based on the analysis of morphometric data, the systematic position of the form from Lang Trang and other large Pleistocene porcupines of Vietnam is determined as Hystrix kiangsenensis Wang, 1931.
... Dong and Li (2008) assigned the larger cervid remains to Megaloceros ordosianus. van der Made and Tong (2008), however, prefer the assignment to the genus Sinomegaceros, an extinct genus of deer known from the Early to Late Pleistocene of East Asia. It is considered to be part of the group of "giant deer" taxa (often referred to collectively as members of the tribe Megacerini), with a probable close relationship to Megaloceros. ...
... Equus) to migrate southward into SE Asia during glacial periods and warm-humid adapted taxa (e.g. Ailupropoda, Macaca, Bubalus) to expand their range to areas north of the Qinling Mountain Range (Tong, 2007(Tong, , 2008Norton et al., 2010). ...
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The Chinese open-air site Xuchang-Lingjing (Henan) is located near the border between the Palearctic and the Oriental biozone in a lowland depression at the southern edge of the North China Plain. The site yielded a Middle Palaeolithic assemblage that includes the two fragmented, incomplete human (possibly Denisovan) skulls (Xuchang 1 and Xuchang 2), more than 15,000 artefacts and more than 40,000 mammalian remains representing 20 taxa. The composition of the faunal assemblage is biased by hominin hunting activities; it is, however, diverse and “natural” enough to be used as a proxy to reconstruct the paleoenvironmental condition during the human occupation of the site. The Palearctic faunal assemblage indicates a grassland-dominated palaeoecological environment, with a mosaic of scattered forest and mixed forest vegetation as well as along rivers and/or lakes the occurrence of swampy areas and with bushes. The OSL-dates and the assumed correlation with the last interglacial (MIS 5) palaeosol strongly suggest that we are dealing with an interglacial fauna. However, the fact that the Lingjing assemblage represents the southernmost Palearctic fauna questions the assumption that the Lingjing Middle Palaeolithic finds date from an interglacial period. A late Middle Pleistocene (MIS 6) or Late Pleistocene glacial or stadial phase (MIS 4) seems more likely.
... Bone remains of H. brachyura vinogradovi were also found in the MIS3 deposits of the Altai caves (Strashnaya, Razboynichiya and Ust-Kanskaya) (Kuzmin et al., 2017). In the Palaearctic the majority of the fossil porcupine sites are located outside its modern range (Tong, 2008;Kuzmin et al., 2017). Currently, the Makhnevskaya Ledyanaya Cave is the most northern site where the porcupine remains have been discovered. ...
... Cohabiting Malayan and Indian porcupines (North-East of India) prefer humid tropical broad-leaved and evergreen forests, and have a wide temperature tolerance range(from 0°to 38°С) (Talukdar et al., 2019). Based on the modern geographic distribution of representatives of the family Hystricidae, fossil remains of the Pleistocene porcupines can be used as indicators of a warm climate (Tong, 2008). ...
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Pollen and fossil mammals of the terrigenous deposits in the Makhnevskaya Ledyanaya Cave (59°26′ N 57°41’ E), located in the northern part of the Middle Urals and on the border with the Northern Urals, were studied. The spore-and-pollen spectra are divided into four pollen zones. In all the spectra, pollen concentrations of tree species amount to 50–70%. The presence of pollen in thermophilic (Quercus robur-type, Tilia cordata-type, Ulmus sp., Carpinus betulus, Corylus avellana-type) and moderate boreal (Abies, Picea, Pinus, Betula, Alnus) species and a lack of any tundra flora elements indicate their interglacial nature. The following have been identified as small mammal fauna: 8 taxa of insectivorous mammals, 4 taxa of bats, 2 taxa of lagomorphs, 4 species of carnivores, 16 species of rodents, 1 species of odd-toed ungulates and 4 species of even-toed ungulates. The remains of cryoxerophilous (Dicrostonyx sp., Lasiopodomys gregalis) and steppe (Ochotona sp.) species are scarce in the studied fauna. Species typical of forests (genus Sorex, genus Craseomys, genus Myodes, Myopus schisticolor, Microtus agrestis) are dominant. Remains of Erinaceus sp., Talpa sp., Crocidura leucodon, Sciurus vulgaris, Hystrix brachyura, Dryomys nitedula, Apodemus flavicollis, Stephanorhinus kirchbergensis have been found. The morphology and dimensions of the teeth of Panthera spelaea fossilis, Hystrix brachyura, Dryomys nitedula, Arvicola amphibius, mice of genus Apodemus are described. The compositions of the palynocomplexes and mammal fauna as well as the SDQ values for m1 of Arvicola amphibius allow us to date the deposits from the Makhnevskaya Ledyanaya Cave to the Eemian interglacial period (MIS 5e).
... Angelici et al. (2003) found some similarities between Italian and African porcupines. Status, taxonomy, distribution and specific characters of the crested porcupine were discussed in many ways (Setzer, 1957b;Corbet and Jones, 1965;Angelici, 1992, 1999;Pigozzi, 1992;Olson and Lewis, 1999;Tong, 2008). Five skulls (three males and two females) of this species were recently obtained in this work from an animal dealer in Misurata city in the northwestern Libya. ...
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The crested porcupine Hystrix cristata is the biggest rodent in Libya. It has a high value from the faunistic point. It is known in Libya from only one specimen killed in a suburb of Tripoli in 1962. With aids of an animal dealer in Misurata, this work will provide information on the morphology,feeding habits,habitat, cranial and dental characters of this animal.
... The discovery of rodent fossils in the Upper Pliocene Irrawaddy sediments fills a chronological and geographical gap in the Pliocene Figure 13. Distribution of four genera of rodents in the Indochinese Division with their fossil records from the Late Pliocene/Early Pleistocene localities (IUCN Red List data; Chaimanee 1998; Tong 2008; Jin et al. 2009). A, Hapalomys; B, Hystrix; C, Maxomys; D, Cannomys. ...
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The Upper Pliocene Irrawaddy sediments in the Gwebin area of central Myanmar recently yielded a rodent assemblage that contains nine species belonging to four families: four species of Muridae, three of Hystricidae, one of Spalacidae, and one of Sciuridae. The murids consist of Hapalomys cf. longicaudatus, Maxomys pliosurifer sp. nov., Rattus jaegeri and cf. Rattus sp. indet., which include both extinct and extant forms. Maxomys pliosurifer is relatively similar to Maxomys surifer that lives in South-East Asia in terms of tooth morphology but retains plesiomorphic features shared with the ancestral rat, Karnimata, and possible sister genera of Maxomys, such as Ratchaburimys and Millardia. The three hystricids belong to the genus Hystrix and consist of two extinct brachydont species (Hystrix paukensis and Hystrix sp. indet.) and one hypsodont species similar to living form (Hystrix cf. brachyura). This finding indicates that primitive brachydont species and derived hypsodont species of Hystrix had likely coexisted in the locality, but the brachydont species are significantly more common amongst specimens collected from the Gwebin area. The spalacid species is Cannomys cf. badius and the sciurid species is Menetes sp. indet. These two rodents are similar to living species in continental South-East Asia although they show minor differences in tooth characteristics compared to the living forms. Some species and genera of the fossil rodent assemblage from the Gwebin area also occur in Upper Pliocene localities of Thailand, suggesting chronological correlation between these two faunas. Moreover, these fossil rodent assemblages are composed primarily of the species distributed endemically in continental South-East Asia. Late Pliocene rodents of continental South-East Asia were affected by river barriers that formed during the Mio-Pliocene, and they were probably not able to disperse from South-East Asia into South and East Asia.
... Porcupine (genus Hystrix) remains have been found at more than 100 Pleistocene fossil localities in Europe and Asia (Baryshnikov, 2003;Tong, 2008); however, they are very rare in Siberia and the Urals (Fig. 1). Therefore, 14 C dates on these specimens are extremely important for palaeoecological reconstructions. ...
... Hystrix (Hystrix) depereti is considered an eurytopic species, based on the present occurrence of Hystrix in many different habitats (Santini, 1980). Subgenus Hystrix is currently spread in areas with an average annual temperature above 10° C (Tong, 2008). ...
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The systematic analysis of more than 20,000 fossils (Vertebrata and Mollusca), recovered from the post-evaporitic Messinian (5.41-5.33 Ma) succession of Moncucco Torinese (NW Italy), resulted in the identification of 90 vertebrate and 65 mollusk taxa that provide additional information about the paleoecological context and the paleoenvironmental settings of NW Italy slightly before the Mio-Pliocene boundary. Our analyses indicate a landscape dominated by open woodlands within a mosaic environment also including closed canopy forests, grasslands, rocky outcrops and limited water edges. The wide spectrum of habitats may have had a prominent role in determining the high paleobiodiversity observed in the paleocommunity of Moncucco Torinese. Slight variations in the abundances of the most common rodent species over the investigated succession are probably related to local changes in the paleolandscape. From a paleoclimatic point of view, the overall information provided by the fauna indicates mesic conditions in a subtropical climate, which is also consistent with the interpretation derived from paleobotanical and sedimentological analyses for the latest Messinian of Northern Italy.
... Similarly, 21% of the bones fragments (Figure 4 a) were gnawed by large rodents, with the majority of remains reduced to cubic or "diamond tip" remains (Figure 4 b and d). This is comparable to the marks described by several authors (Pei 1938;Rabinovich and Horwitz 1994;Tong 2008;Tong et al. 2008;Filoux et al. 2015) or to the 16% of gnawed teeth in the (Figure 4 i, j, k). This is followed by further gnawing of these dental elements by Hystricidae, which then attack the dentin (Figure 4 l, m, n). ...
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Since the initial description of the complex Ailuropoda-Stegodon as a faunal association with biochronological significance for the Southeast Asian area, few sites have provided paleontological data allowing for an extensive documentation of past fauna. Biodiversity and paleo-environmental reconstructions of Pleistocene fauna are still generally based on bone assemblages whose taphonomy and dating are not or badly documented. However, in order to be useful in a paleo-eclogical perspective, the dating of collected assemblages should be associated with periods of times corresponding to climatic episodes. In this study, we provide a detailed stratigraphic, taphonomical, paleontological and ESR dating studies concerning the site of Ban Fa Suai II discovered near the cave of the Monk in northern Thailand. Our results demonstrate the changes in the taxonomic composition of the fauna over time and argue against the use of the currently available regional dataset for paleo-ecological reonstructions.
... 的动物群名单 [3,10] ,大额牛的研究成果也已发表 [11] ,但对整个动物群的系统古生物学研究 尚未开展,因此而影响到了对动物群性质及时代的准确判断和深入对比,造成现今对白龙 洞遗址时代认识的不一致 , 如有学者认为动物群时代为中更新世时期 , 大约与北京猿人时 代相当 [1,12] ;也有学者指出 , 它属于早更新世晚期阶段 [13] 。 由于发掘次数多且间隔时间长,动物化石保管分散,这是造成动物化石一直未得到系 统研究的客观原因。就目前所知,白龙洞化石主要保存于郧西县文管所和中科院古脊椎动 物与古人类研究所 ( 主要是 1977 和 1982 年发掘标本 ) [25] 。牙齿均为高齿冠,但上颊齿更高冠些;在上、下 臼齿中,齿冠前侧面均高出后侧面;上臼齿牙体中段凸向前舌侧,且自上而下逐渐变粗; 由此引发的问题是齿长的测量数据会因不同作者而有差异 [24,26] 。白龙洞的雄性颊齿,测量 数据均大于早更新世的繁昌竹鼠 [26] ;而与盐井沟的咬洞竹鼠相似,也应当归入中华竹鼠。 比较与讨论:南方第四纪的竹鼠演化阶段分明,由早到晚,牙齿大小及冠高都在逐渐 增大;繁昌竹鼠 (R. fanchangensis Wei et al., 2004) 和拟低冠竹鼠 (R. brachyrhizomyoides Zheng, 1993) 代表我国第四纪最早的竹鼠,后来便是早中更新世常见的咬洞竹鼠 (Rhizomys troglodytes Matthew and Granger, 1923),到了距今约 50 万年,才出现现生种中华竹鼠 [24] 描述:牙齿大小和形状与我国现生种无颈鬃豪猪一致 [27] 。 比较与讨论:豪猪是我国南方第四纪动物群中最常见动物之一,甚至在华北平原北端 的周口店地区也很常见 [28] ,但白龙洞的豪猪较少,其他骨骼上也少见豪猪的咬痕,这与其 所处地理位置很不一致, 这其中一定另有原因;不过, 在盐井沟动物群中, 豪猪化石也很少 [29] 。 食肉目 Carnivora Bowdich, 1821 犬科 Canidae Fischer de Waldheim, 1817 变异狼 Canis mosbachensis variabilis (Pei, 1934) 1934 Canis lupus var. variabilis-Pei, p.13~18, text fig. 1, Pl. I, 2~4, PL II, 1, 3~4 [31] 1941 Canis variabilis-Teilhard de Chardin, Pei, p.8~11, text figs. ...
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... Tong (2007) notes that Hystrix, Macaca, Palaeoloxodon, Dicerorhinus and Bubalus are the most frequently recorded warm-adapted elements that appeared in north China. Because Hystrix has been found at 22 localities in north China, it is probably one of the best faunal indicators of a warm climate in this region (Tong, 2008). ...
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The goal of this paper is to describe a single upper molar and a fragmented radius of Old World porcupines recently discovered in the latest Messinian localities of Moncucco Torinese and Verduno in the Tertiary Piedmont Basin, NW Italy. The available material can be assigned to the large-sized species Hystrix (Hystrix) depereti, rarely found in the late Turolian and early Ruscinian of Europe. A combined comparative and morphofunctional analysis of the fragmented radius suggest that Hystrix (Hystrix) depereti was characterized by a generalized terrestrial locomotory behaviour thus being very similar to extant Old World porcupines. Paleobiogeography and paleoecological consequences are also discussed.
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This paper describes the first fossil porcupine remains from Iran. Four upper cheek teeth and two fragmentary lower incisors present sufficient characters for identification as Hystrix aryanensis, a species previously known from the late Miocene locality of Molayan (Afghanistan) estimated at ca. 7–8Ma. The dental features of porcupines are discussed to show their systematic value and highlight evolutionary trends in late Miocene and Pliocene porcupines. This study also discusses the dispersal history of fossil porcupines in relation to paleobiogeographic provinces and environmental changes during late Miocene to late Pliocene time. Diese Arbeit beschreibt die ersten Stachelschweinfossilfunde aus dem Iran. Vier Oberkieferzähne und zwei fragmentäre untere Frontzähne zeigen Merkmale, die zur Bestimmung von Hystrix aryanensis ausreichen, eine Art, die bisher aus dem jüngeren Miozän aus Molayan (Afghanistan) bekannt ist und auf ca. 7 bis 8Ma geschätzt wird. Die Zahnmerkmale der Stachelschweine werden diskutiert, um deren systematische Aussagekraft hervorzuheben, und um die Evolutionstrends der Stachelschweine im jüngeren Miozän und Pliozän zu beleuchten. Diese Arbeit diskutiert auch die Entwicklung der Verteilung von fossilen Stachelschweinen in Verbindung mit den paläobiogeographischen Provinzen und Umweltveränderungen während des jüngeren Miozäns und Pliozäns. KeywordsHystricidae-Rodentia-Late Miocene-Iran-Biogeography SchlüsselwörterHystricidae-Rodentia-Jüngeres Miozän-Iran-Biogeographie
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Molars of 35 skulls of Hystrix brachyura brachyura Linnaeus, 1758 have been studied. Problems with the currently used dental nomenclature are discussed. A positional, numerical system for the indication of occlusal elements, and a system of wear classes for all the different occlusal patterns are presented. Dimensions of the teeth in relation to wear are given, the variation in the occlusal patterns is described and their changes with progressive wear are discussed.
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From nocturnal activity durations of free-ranging animals and food intake rates of captive subjects, we predict that Indian crested porcupines (Hystrix indica Kerr) will not occur at latitudes where minimal night duration is less than 7 h. Available range records support that hypothesis. They indicate that the most northern populations of the species occur in the Aralo-Caspian region of the U.S.S.R. at about 44⚬ N, where night duration during the summer solstice is 7.3 h. To exist north of this postulated 'fitness boundary', crested porcupines would be forced to forage during daylight hours or reduce food intake. Presumably the costs associated with either of these alternatives would not promote fitness. The northern species range limits do not coincide with broad bioclimatic or vegetation boundaries.
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Fossil Hystrix material from a large number of localities is biometrically analysed and compared with the extant species. H. major, H. etrusca, H. makapensis, and H. angressi are shown to be junior synonyms of H. refossa. H. bessarabica and H. trofimovi are shown to be junior synonyms of H. primigenia, and H. schaubi and H. cristata minor of H. vinogradovi. The dental characteristics of H. refossa, H. primigenia and H. vinogradovi are discussed.
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The characters used for specific diagnosis in the subgenus Hystrix have been studied to determine their constancy as specific characters and their geographical variation within the species. As a result three species are recognized, namely H. indica, H. cristata and H. africaeaustralis. H. galeata is considered a synonym of H. cristata, and H. stegmanni a synonym of H. africaeaustralis. The morphological differences between cristata and africaeaustralis in East Africa are not continued clinally within these species. Between cristata and indica some of the diagnostic characters vary clinally, others change abruptly.
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