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Several porcupine taxa are reported from the middle Miocene to the early Holocene in the Old World. Among these, five species of the subfamily Hystricinae occurred in Africa approximately in the last 6 Ma: the extinct Hystrix makapanensis, Hystrix leakeyi, and Xenohystrix crassidens and the still living Hystrix africaeaustralis and Hystrix cristata. The large-sized H. makapanensis is reported from numerous sites in East and South Africa between the early Pliocene and Early Pleistocene. In this paper, we describe a new mandible of H. makapanensis from the world-renowned Tanzanian paleontological and archeological site of Olduvai Gorge (HWK West; lowermost Bed II; ca. 1.8–1.7 Ma). The discovery of the new mandible triggered a comprehensive review of the entire African record of H. makapanensis. In particular, we describe or re-analyze the samples from South Africa (Makapansgat Limeworks, Gondolin, Kromdraai, Swartkrans, and Sterkfontein), Tanzania (Olduvai and Laetoli), Ethiopia (Omo Shungura and Hadar), and Kenya (Chemeron), enriching the quantity of specimens confidently referable to this species and above all improving the information on its craniodental anatomy. On this basis, we: (1) propose an emended diagnosis of H. makapanensis; (2) point out the morphological and biometric differences between H. makapanensis and other African Hystricinae (also in terms of body mass); and (3) broaden the knowledge on the geographical and chronological distribution of this extinct species. Supplementary Information The online version contains supplementary material available at 10.1007/s10914-021-09588-z.

International audience; The expedition commanded by the Frenchman Nicolas Thomas Baudin aboard the ships Le Géographe and Le Naturaliste (and Le Casuarina for the return journey) to the southern hemisphere between 1800-1804 collected specimens from numerous locations including the Canary Islands (Tenerife), Île de France (Mauritius), Cape Town (South Africa), Australia and Timor. Additionally, specimens were donated or purchased from locations not visited including the Comoros, Madagascar and Sumatra. Unfortunately, Baudin died at Île de France on the return trip so the responsibility of the account of the voyage was given to other members of the expedition. Responsibility for writing up the primary account of the voyage was granted to François Péron, who published the first volume of the narrative of the Voyage de Découvertes aux Terres Australes in 1807. Following his death in 1810, the second volume of the narrative was completed by Louis de Freycinet and published in 1816. The other four volumes of the Voyage included three atlases (the first by expedition artists Charles-Alexandre Lesueur and Nicolas-Martin Petit in 1807 and the second and third by Freycinet in 1811 and 1812), and a volume entitled Navigation et Géographie by Freycinet in 1815. Based on recent and on-going research, a review of many of the original documents is presented here, revealing hitherto unpublished details about who collected and donated mammals to the expedition. Research was conducted mainly in the collections of the Muséum national d'Histoire naturelle (MNHN) in Paris and their associated acquisition books. The Baudin expedition was responsible for an unprecedented collection of over 100 000 specimens of natural history, which remains the single largest collection of natural history specimens from Australia. A total of 101 mammal taxa relating to the Baudin expedition were identified during this study, which included 51 species described as a result of the expedition and 50 species that were described either before or subsequently, but not associated with the expedition. Of the taxa described, 20 species and three subspecies are currently recognised valid. During this study 43 museum specimens that were referable to 29 taxa were identified and at least five specimens seem to have been misplaced based on the information available. These specimens were derived from 24 holotypes, four paratypes, three syntypes, five lectotypes, and six paralectotypes and one topotype that are currently held at the MNHN. Some of these specimens are part of the estimated 51 mammals that were brought back alive to France on the boats. Charles-Alexandre Lesueur's illustrations complemented this study; of the 177 that are held at the Muséum d'Histoire naturelle du Havre, 149 could be attributed to one or more species and 28 were of unidentified species.

Hystrix capensis Grill, 1858 Hystrix capensis Grill, 1858: 19. COMMON NAME. — Cape Porcupine. CURRENT NAME. — Hystrix africaeaustralis Peters, 1852. See Woods & Kilpatrick (2005: 1543) and Barthelmess (2016: 327). COLLECTOR/S. — Two from the Cape collected alive recorded by Pierre Bernard Milius (Péron 1804a). See also Milius (1987: 57) and Jangoux et al. (2010: 269, 276). COLLECTION LOCALITY. — Cape Town, South Africa. COLLECTION DATE. — Before 24 Jan 1804. SPECIMEN NUMBER/S. — No specimen identified in MNHN collection. COMMENTS. — Two specimens, male and female, were collected alive and taken back to France on board Le Géographe under the name of “porcs-epics vivants” (É. Geoffroy Saint-Hilaire 1804c: 172; Girard 1856: 105). Upon arrival in France the animals were kept in the menagerie of the Muséum and fed bread and rice (Jangoux et al. 2010: 276). ILLUSTRATIONS. — No data., Published as part of Jackson, Stephen M., Jansen, Justin J. F. J., Baglione, Gabrielle & Callou, Cécile, 2021, Mammals collected and illustrated by the Baudin Expedition to Australia and Timor (1800 - 1804): A review of the current taxonomy of specimens in the Muséum national d'Histoire naturelle de Paris and the illustrations in the Muséum d'Histoire naturelle du Havre, pp. 387-548 in Zoosystema 43 (21) on page 425, DOI: 10.5252/zoosystema2021v43a21, http://zenodo.org/record/5142972, {"references":["GRILL J. W. 1858. - Zoologiska Anteckingar under en resa I sodra delarne af Caplandete aren 1853 - 1855. Af J. F. Victorin. Ur den a flidnes papper samlade och ordnade at J. W. Grill. Kong. Svenska Vetenska. Kongliga Svenska Vetenskaps-Akademiens Handlingar, Stockholm 2 (2): 1 - 62. https: // www. biodiversitylibrary. org / page / 34317295","PETERS W. 1852. - Naturwissenschaftliche Reise nach Mossambique. Zoologie. I. Saugethiere. Druck und Verlag von Georg Reimer, Berlin, 202 p.","WOODS C. A. & KILPATRICK C. W. 2005. - Infraorder Hysticognathi, in WILSON D. A. & REEDER D. M. (eds) Mammal Species of the World. A Taxonomic and Geographic Reference. Johns Hopkins University Press, Baltimore. Third Edition: 1538 - 1600.","BARTHELMESS E. L. 2016. - Family Hystricidae, in WILSON D. E., LACHER T. E., MITTERMEIER R. A. (eds) Handbook of the Mammals of the World. Volume 6. Part 1. Lagomorphs and Rodents. Lynx Edicions, Barcelona: 304 - 329.","PERON F. 1804 a. - Diarium Zoographicum. No. XV. Ans XI et XII. Observationes generales de Collectionibus factis in Zoologia ex profectura nostra de Portu Jackson (27 Brumaire an XI) adusque Promontorium Bonae Spei inclusive (30 Pluviose an XII). Museum d'Histoire naturelle du Havre, no. 21001.","MILIUS P. B. 1987. - Recit du voyage aux terres australes / par Pierre- Bernard Milius, second sur le Naturaliste dans l'expedition Baudin (1800 - 1804); transcription du texte original par Jacqueline Bonnemains, Pascale Hauguel, avec l'autorisation de Monsieur Shand Kydd. Societe Havraise d'Etudes diverses Museum d'Histoire naturelle du Havre, 82 p.","JANGOUX M., JOUANIN C. & METIVIER B. 2010. - Les animaux embarques vivants sur les vaisseaux du voyage de decouvertes aux Terres australes. Actes du colloque Portes par l'air du temps: les voyages du capitaine Baudin, Bruxelles, septembre 2009. Etudes sur le XVIIIe Siecle 38: 265 - 282.","GIRARD M. 1856. - F. Peron, naturaliste, voyageur aux Terres Australes; sa vie, appreciation de ses travaux, analyse raisonnee de ses recherches sur les animaux vertebres et invertebres d'apres ses collections depos e es au Museum d'Histoire naturelle. J. - B. Bailliere, Paris. https: // www. biodiversitylibrary. org / page / 46101762"]}

A camera trap survey was conducted in the recently protected Cleopatra’s Needle Critical Habitat (CNCH) in Puerto Princesa City, Palawan, Philippines from February to May 2015 at 39 camera trap sites. A bait of common pig’s blood was used at 36 sites, while the three remaining sites were surveyed without a bait and monitored a stream with a latrine site or mud bath with tracks. Seven native species were detected and three of these species were endemic to the island province. Species included: Common Palm Civet Paradoxurus philippinensis, Palawan Porcupine Hystrix pumila, Collared Mongoose Urva semitorquata, Palawan Stink Badger Mydaus marchei, Palawan Leopard Cat Prionailurus bengalensis heaneyi, Asian Small-clawed Otter Aonyx cinereus, and Malay Civet Viverra tangalunga. Analysis of the activity patterns of the three most commonly captured species revealed predominantly nocturnal activity for the Common Palm Civet, Palawan Porcupine, and Palawan Stink Badger. The Philippine Palm Civet showed occasional diurnal activity. The seven photo-captured species appeared most common, or were at the least recorded, below 750m. Five species (the Philippine Palm Civet, Palawan Porcupine, Collared Mongoose, Palawan Stink Badger, and Palawan Leopard Cat) were also recorded above 1000m. The CNCH supports two threatened species, the Palawan Porcupine and the Asian Small-clawed Otter, which are listed as Vulnerable by the IUCN, and the Collared Mongoose is listed as Near Threatened. The Palawan Leopard Cat is considered Vulnerable within the Philippines, although it has yet to be assessed by the IUCN. This documentation highlights the biodiversity significance within the newly protected critical habitat and the need to support ongoing conservation efforts within the critical habitat.

Taxonomy is a mistreated matter, but its role in ecology, behaviour and conservation studies is pivotal. Disentangling amongst different subspecies is challenging given the high arbitrariness level in determining thresholds of genetic and morphological distances. Splitting frenzy trends have increased the number of animal taxa and for most of them a critical redefinition is required. In this work, we reviewed knowledge and validity of subspecific taxa identified for African crested porcupines Hystrix cristata and Hystrix africaeaustralis. In the past, several subspecies were recognized for both the species, but successive works suggested H. cristata and H. africaeaustralis as monotypic species with no clear explanation. Recently, the validity of the taxon H. cristata senegalica has been claimed again. We analysed all available data and discussed all the subspecific taxa in light of both genetic and morphological data. We revalidated here the synonymy Hystrix senegalica Cuvier, 1823 = Hystrix cristata Linnaeus, 1758. Syn. rev. Two names are treated as nomina dubia: Acanthion daubentonii Cuvier, 1823 (formalization) and “Hystrix capensis Gr.”. Hystrix cristata var. alba de Sélys-Longchamps, 1839 has been deleted from the synonymic list of H. cristata. Neither mitochondrial nor nuclear DNA data militate for the existence of any subspecific taxon, although further data are required for H. cristata from East Africa (e.g., Kenya and Tanzania). Similarly, morphology seems to play for a clinal variation in both species. For available data, we thus strongly recommend to keep both H. cristata and H. africaeaustralis as monotypic species.

Family Hystricidae This family includes the Old-World porcupines. The head and neck are covered with a crest of long bristles. The dorsal side is covered with spines of various sizes. Porcupines are nocturnal animals that feed entirely on roots, bulbs and other cultivated crops (Wilson et al., 2016)., Published as part of Amr, Zuhair S., Abu, Mohammad A., Qumsiyeh, Mazin & Eid, Ehab, 2018, Systematics, distribution and ecological analysis of rodents in Jordan, pp. 1-94 in Zootaxa 4397 (1) on page 71, DOI: 10.11646/zootaxa.4397.1.1, http://zenodo.org/record/1202783

Distributional and ecological data were given to all rodents of Jordan. The rodent fauna of Jordan consists of 28 species with 20 genera in eight families (Cricetidae, Dipodidae, Gliridae, Hystricidae, Muridae, Myocastoridae, Sciuridae,and Spalacidae), including four introduced species.Keys for families and species were provided, along with diagnosis for each species and cranial illustrations for most species. Habitat preference and zoogeographic affinities of rodents in Jordan wereanalyzed, as well as their status and conservation.Threat categories and causes of threats on the rodents of Jordan were also analyzed. The distribution of rodents in Jordan represents a reflection of their global distribution ranges and habitat preferences. Species associated with the temperate forest of northern Jordan includes Sciurus anomalus and two wood mice, Apodemus mystacinus and A. flavicollis, while non-forested areas are represented by Nannospalax ehrenbergi and Microtus guentheri. Strict sand dwellers include Gerbillus cheesmani and G. gerbillus. Petrophiles associated with sandstone or black lava deserts are exemplified by Acomys russatus, A. r. lewsi, H. indica and S. calurus. Others including: Jaculus jaculus, G. nanus, G. henleyi, Meriones crassus, and M. libycus are all desert-adapted species with wider ranges of distribution where scarce vegetation, wadibeds, and marabs with clay, loess, or gravel surfaces provide foraging grounds and shelter. A single species, Gerbillus dasyurus, exhibits a wide range of distribution over diverse habitat types. The rodent fauna of Jordan consists of assemblages of different zoogeographical affinities. Nine, three, and seven were restricted or had most of its range within the Mediterranean, Irano-Turanian, and Saharo Arabian, respectively. Sciurus anomalus, Apodemus sp., Nannospalax ehrenbergi, and Microtus guentheri reached their most southern range of distribution in the Mediterranean regions of Jordan. The distribution of Gerbillus cheesmani extends from Asian deserts in India westwards into the Arabian Peninsula crossing Jordan as its most western range of distribution. Typical rodents of Saharo-Arabian affinities are represented by desert jerboas, gerbils, and jirds. North African species such as G. andersoni, G. gerbillus reached their most eastern distribution in southern Jordan. Both G. henleyi and G. nanus are widely-distributed species across North Africa reaching as far as India to the east, representing most northern outpost for these two species. Sekeetamys calurus is a nearly endemic to the Eastern Mediterranean region within southern Jordan and Sinai. Relicts are represented by Eliomys melanurus and Acomys russatus lewisi. Several threats affecting the rodent biodiversity in Jordan were identified including habitat loss and degradation, human disturbance and related activity, legislative and public awareness. The global conservation status of the rodents of Jordan according to the IUCN Red List include 22 species as least concern, one as near threatened (Allactaga euphratica), and one as data deficient (Nannospalax ehrenbergi). According to the regional assessment, one species is critically endangered, three species are considered endangered, one vulnerable.

Mörs, Thomas, Hugueney, Marguerite (2017): A Hystrix Linnaeus, 1758 incisor (Mammalia, Rodentia) from the Early Pleistocene of Senèze, France. Geodiversitas 39 (4): 797-802, DOI: 10.5252/g2017n4a8, URL: http://dx.doi.org/10.5252/g2017n4a8

2. African Brush-tailed Porcupine Atherurus africanus French: Athérure africain / German: Afrikanischer Quastenstachler / Spanish: Puercoespin africano de cola de cepillo Taxonomy. Atherurus africanus Gray, 1842, “West Africa, Sierra Leone.” Distribution of A. africanus largely overlaps the West and Central African distribution of Hystrix cristata, and it might overlap somewhat with the distribution of H. africaeaustralis in Kenya and Uganda. Monotypic. Distribution. W & C Africa, in the Rainforest Biotic Zone and rainforest-savanna mosaic from Guinea to E & S DR Congo and Rwanda and S to the Republic of the Congo and NW & N Angola (Cabinda and Carumbo Lagoon in Lunda Norte Province), also on Bioko I (Equatorial Guinea), and small populations in W Gambia, S South Sudan, Uganda, and E Kenya; also recorded in E Tanzania (Lake Tanganyika). Descriptive notes. Head-body 365-600 mm, tail 100-260 mm, ear 38-39 mm, hindfoot 71-73 mm; weight 1-5—4-3 kg. The African Brush-tailed Porcupine is large and dark, with relatively long body and short, stout limbs; it appears almost entirely spiny. Dorsal pelage is dark brown and thickly spined. Hairs are modified into coarse, thick spines that are colored off-white at bases and gradually darken along shafts toward middles, appearing blackish brown at sharply pointed tips; fine hairs grow between spines. Softer spines occur on underside,legs, and head. Lengths of spines are variable and depend on body region; they are shortest (c.20 mm) on neck, longer (25-45 mm) on flanks and mid-dorsal region, and up to 90 mm on rump. Head of the African Brush-tailed Porcupine is sparsely covered with short, dark, coarse hairs, and ears are darkly pigmented and mostly naked. Mystacial vibrissae are long and black. Relative to species of Hystrix, skull of the African Brush-tailed Porcupine is elongated, with noninflated nasal bones that end anteriorly to zygomatic arch. Postorbital processes are either lacking or very weak. Incisors are smooth and without grooves on outer surfaces. Ventral pelage is off-white to pale brown. Ventral hairs are softer and less spiny than dorsal hairs and are ¢.10-15 mm. Forefeet and hindfeet are covered with coarse dark brown hairs. Each foot has five digits, and feet are partially webbed and armed with blunt claws, except on first digit of forefoot. The African Brush-tailed Porcupine has a short tail; ratios of tail to head-body length are ¢.25-50%. Tail is thick, swollen at base, tapers toward tip, and covered with short, black spines. Tail ends with a brush-like cluster of off-white or yellow, “platelet-bristles.” Each is a hollow, flattened bristle with 4-5 bead-like “platelets” or shafts that swell like rice grains at regular intervals. When shaken, they produce a rattling sound. Tail breaks easily and is often lost. Female African Brush-tailed Porcupines have two pair of lateral, thoracic mammae. Habitat. Evergreen and gallery forests with hollow trees, buttress roots, and soft soil, especially close to streams in small valleys from sea level to elevations of ¢.3000 m. The African Brush-tailed Porcupine also occurs on farmlands adjacentto forest. Food and Feeding. The African Brush-tailed Porcupine is primarily frugivorous and eats fruits from forest trees, most of which have fallen to the ground from the tree canopy. It also eats flowers, leaves, and some roots. When adjacent to farms,it will eat maize, manioc (cassava), bananas, and palm nuts. Occasionally, it eats carrion and earthworms. Breeding. Young African Brush-tailed Porcupines are found during most months of the year in the DR Congo, and this pattern likely occurs throughout the Rainforest Biotic Zone. In the wild, females have 2-3 litters/year. Gestation lasts 100-120 days, followed by birth oflitters of 1-2 young (occasionally up to four). Female African Brushtailed Porcupines are likely polyovular, so the common litter size of one observed in captivity may be an artifact of holding conditions. Females have a bicornuate uterus, with two separated uterine horns, a uterine body, and a cervix. Females in captivity show spontaneous estrous cycles in a polyestrous pattern, with an average estrous cycle of 27-1 days. Young are born weighing ¢.100-175 g and are precocious; their eyes are open, and they have soft spine-like hairs on their back and flanks and can walk on all four legs within a few hours of birth. Young nurse for about two months although they begin to eat solid food at 2-3 weeks of age. The African Brush-tailed Porcupine grows slowly; adult size is reached after ¢.300 days, and sexual maturity at about two years of age or older. Activity patterns. The African Brush-tailed Porcupine is a strictly nocturnal and primarily terrestrial. It hides during the day in dens located in hollow trees, hollow logs, caves, or underbrush piles or roots of large trees (it does not dig its own burrow). It defecates regularly at the same place, either under a rock or in a den. It walks and trots along the forest floor, swims readily, and can climb trees to a limited extent to escape pursuit threat. While moving, it carriesits tail held slightly upward and back from its body; sometimes rustling of rattle-quills can be heard even when it cannot be seen. African Brush-tailed Porcupines in the wild in Gabon typically left their hiding places at ¢.17:00 h, returning to their dens at c¢.05:30 h. Nightly activity, particularly among females, usually followed a trimodal pattern with two rest periods during the night, although the pattern shifted to bimodal on moonlit nights. Female rest periods clustered around 20:00-21:00 h and again at 02:00-04:00 h. Males spent more time immobile and had more irregular patterns, resting in the middle of the night. Individuals also took rest periods of about ten minutes every hour; these rests tended to be longer in the middle of the night than at the start or end, and longer in the presence of bright moonlight. On dark nights, females did not normally take a rest in the middle of night, although males rested some of the time. Average inactive time on dark nights was 1-9 hours. On nights close to full moonlight, period of inactivity increased to an average of 3-4 hours. When alarmed or threatened, an African Brush-tailed Porcupine will raise spines on its back and shakeits tail, rustling the bristles. If, after this warning display,it remains threatened, it will move quickly sideways and backward with its sharp, pointed quills facing forward toward the opponent. Movements, Home range and Social organization. While foraging at night, African Brush-tailed Porcupines move rapidly across the landscape, traveling ¢.100 m every four minutes. They follow well-established trail networks as they move through undergrowth and generally travel alone. Average nightly distances moved varied with sex and moon phase. Males averaged 1697 m/night with moonlight and 2333 m on dark nights, and females averaged 1646 m/night with moonlight and 1953 m on dark nights. Home ranges of males in Gabon averaged slightly larger than those of females, with means of 15-6 ha and 10-2 ha, respectively. Male and female home ranges overlap. Although total area used and distance traveled did not differ significantly between males and females, pattern of space use did. Male African Brush-tailed Porcupines made single, complete circuits of their home ranges and typically rested in the middle of the night at the end of the home range opposite the sleeping den and did not display territorial behavior. Females followed no regularitinerary while active, wandering back and forth unpredictably. Although African Brush-tailed Porcupines travel alone at night in search of food, they are thought to be gregarious. They occasionally meet at feeding places and have a tendency to share dens during the day. Of 22 dens investigated in Gabon, twelve contained one individual, seven each held two individuals, one held three individuals, and one held four individuals. Associations in the dens were all male, all female, or males and females. During the period of the study, males used an average of eight dens, and females used six dens. Some dens were used repeatedly by the same individual, but others were used only once. In captivity, African Brush-tailed Porcupines exhibit mutual grooming along with auditory displays of dominance and submission. They may form family groups of one male, one female, and offspring, although social organization appears fluid, without formation of monogamous pairs or cohesive groups. Breeding system is currently unknown. Populations of African Brush-tailed Porcupines can be locally abundant, with densities of 2:4-13-2 ind/km? in the Central African Republic, 55 ind/km? in Equatorial Guinea, and 58 ind/km? in Gabon. In Gabon, sex ratio was male biased at 2-4:1 for adults and 2-1:1 for subadults, although this bias may have been an artifact of differences in capture probabilities between sexes. Status and Conservation. Classified as Least Concern on The IUCN Red List. Main predators of the African Brush-tailed Porcupine are humans and Leopards (Panthera pardus). It is vigorously hunted by humansfor its succulent flesh and is one of the most common bushmeat species in markets across West and Central Africa. It does not appear to be overhunted, although its relatively low recruitment rate, due to fairly long generation times, seems to contradict that conclusion. Loss of forest habitat is a cause for concern in parts ofits distribution. It is also killed as a crop pest. Bibliography. Albrechtsen et al. (2005), Anadu et al. (1988), Carr et al. (2013), Colell et al. (1994), Cordeiro (1998), Emmons (1983), Fa et al. (2002), Fischer et al. (2002), Foley et al. (2014), Happold (2013a), Hoffmann & Cox (2008), Huntley & Francisco (2015), Mayor et al. (2003), Nowak (1999a), Storch (1990)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 306-307, DOI: 10.5281/zenodo.6612213

7. Malayan Porcupine Hystrix brachyura French: Porc-épic de Malaisie / German: Malaiisches Stachelschwein / Spanish: Puercoespin de Malasia Other common names: Himalayan Crestless Porcupine, Hodgson's Porcupine Taxonomy. Hystrix brachyura Linnaeus, 1758, “Habitat in Asia.” Identified by A. Grote in J. E. Gray in 1866 as “Malacca,” Malaya. Hystrix brachyura is in subgenus Acanthion. Many different subspecies have been described (some based on single specimens) including four recent subspecies in China, butD. J. van Weers in 1979 concluded after careful measurements of many specimens that although some clearly different populations can be recognized,full subspecific designation was unwarranted due to a high degree of intergradation. Further examination of a larger number of specimens, especially a larger number from China, will help resolve subspecific taxonomy. Northern limit of H. brachyura in China is in southern Gansu and southern Shaanxi from where Pleistocene fossils have also been recorded as H. subcristata. Three subspecies recognized. Subspecies and Distribution. H.b.brachyuraLinnaeus,1758 — MalayPeninsula,PenangI(offWcoastofPeninsularMalaysia),Sumatra,andBorneo. H.b.hodgsoniGray,1847 — SofHimalayasinENepal,NIndia(Sikkim). H. b. subcristata Swinhoe, 1870 — S Bhutan, NE India, C & S China (including Hainan I), Bangladesh, Burma (= Myanmar), and mainland South-east Asia. Descriptive notes. Head—body 455-930 mm,tail 60-170 mm, ear 25-38 mm, hindfoot 75-95 mm; weight 8-27 kg. The Malayan Porcupine is large, stocky, and blackish brown, with short tail. In some individuals, spines along back of neck form rudimentary dorsal crest. Quills of back, shoulders, and sides are brown without white tips. Dorsal quills are square anteriorly and become round posteriorly. Long defensive quills on lower back are white, with single brownish black band in middle of each quill and white tips ¢.200 mm in length. Diameters of thickest quills on back are 5-7 mm. Ears and eyes are relatively small. Nasals are long (41-60% of occipito-nasal length) and broad (37-62% of zygomatic breadth). Skull has inflated pneumatic cavities. Quills on hindquarters are white, with fairly narrow sub-terminal black bands, and quills under tail are entirely white. Length of short tail is less than 20% of head-body length. Tail carries hollow, goblet-shaped rattlequills, extending 200-300 mm in length that rattle when shaken. Lengths of hollow parts of rattle-quills are 15-34 mm. Channels in network of medulla in quills might serve to transport pheromones. There are three pairs of lateral mammae. Diploid number is 2n = 60. The Malayan Porcupine can be differentiated from the Indian Crested Porcupine (H. indica), with which it overlaps in India, by having rudimentary crest, if any, and having quills with only single brown bands. The Asiatic Brush-tailed Porcupine (Atherurus macrourus) is smaller than the Malayan Porcupine, with shorter brown quills, a longer scaly tail with a brush on the tip, and beaded tail quills. Habitat. Forests and open fields where large burrows are dug into banks and under rocks. Malayan Porcupines are often seen crossing open areas at night. In India, they prefer forests and grasslands of eastern Himalaya, especially near cultivated land. Food and Feeding. The Malayan Porcupine is a generalist herbivore. It eats roots, tubers, bark, grains, vegetation, and fallen fruit including African oil palm (Elaeis guineensis, Arecaceae). It craves calcium and seeks out bones and antlers to gnaw. Breeding. Gestation of the Malayan Porcupine is c.110 days. Two (sometimes three) precocial young are born, and two litters can be produced per year. Activity patterns. The Malayan Porcupine is nocturnal and primarily terrestrial. In a camera trap survey of Indonesian rainforest mammals, 100% of photographs of Malayan Porcupines were at night. It digs extensive burrows beneath the forest floor. Movements, Home range and Social organization. The Malayan Porcupine lives alone or in groups of 2—4 individuals. Family groups can occupied burrows together and emerge at night to forage together along well-defined runways. Malayan Porcupines can reach very high densities, resulting in problems with vegetable farms and tree plantation when it feeds in agricultural areas. Frequent visual sightings and detections in camera-trap surveys suggest an increase in numbers around Hong Kong in the last 20 years, although the Malayan Porcupine was described as rare in that area in pre-1990 publications. When alarmed, an individual rattlesits tail quills, snorts, and stomps its feet. If the threat does not subside, it might attack by running backward, quills erect, into the enemy. Status and Conservation. Classified as Least Concern on The IUCN Red List. Overhunting is the greatest threat to the Malayan Porcupine. It is hunted for meat and also farmed in Vietnam where populations have declined 20% since the 1990s. The Malayan Porcupine is one of three species of native rodents that remains in Hong Kong. Its presence and increase there is somewhat surprising given that, until recently, it was persecuted as an agricultural pest and hunted for food. Habitat loss and poaching are the main threats to populations of Malayan Porcupines in India. Bibliography. Agrawal (2000), Brooks et al. (2010), Chung & Corlett (2006), Corbet & Hill (1992), Francis (2008), Gray (1866), Lunde, Aplin & Molur (2008), Medway & Harrisson (1963), Menon (2009), Nowak (1999a), Raha et al. (2015), van Schaik & Griffiths (1996), Smith & Yan Xie (2008), Srinivasulu & Srinivasulu (2012), Storch (1990), Tempa et al. (2013), van Weers (1979)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 308, DOI: 10.5281/zenodo.6612213

5. Thick-spined Porcupine Hystrix crassispinis French: Porc-épic de Bornéo / German: Borneo Stachelschwein / Spanish: Puercoespin de espinas gruesas Other common names: Borneo Short-tailed Porcupine Taxonomy. Hystrix crassispinis Gunther, 1877, “Borneo.” A. Gunther in 1877 cross referenced an earlier paper of his in the same journal in which the type locality was given as “Borneo, opposite of Labuau.” H. crassispinis is in subgenus Thecurus. Monotypic. Distribution. Borneo. Descriptive notes. Head-body 450-665 mm, tail 65-190 mm, ear 35-42 mm, hindfoot 80-90 mm; weight 3.8-5.4 kg. The Thick-spined Porcupine is medium-sized and brown, with extremely thick quills. It is the largest species in the subgenus Thecurus, with the largest quills, tactile bristles, and rattle-quills. It appears distinctly speckled by white tips of spines on cheeks, neck, shoulders, flanks, and sometimes anterior of back. Some large defensive quills on posterior of back have white tips ¢.25-65 mm long, while others are blackish up to their tips. Number of white-tipped quillsis larger than number of black quills; number of tactile bristles is small. As with other species in the subgenus Thecurus, there is no crest on head. Thick quills reach up to 8:3 mm in diameter, making them thicker than those in the largest specimens of subgenus Acanthion and as thick as those in subgenus Hystrix. Tail is short, with ratio of tail-to-head-body length less than 20%, and it has rattle-quills. Lengths of hollow capsule-like parts of rattle-quills are 12-16 mm. Long quills of hindquarters are dark brown, with fairly narrow white tips and bases, and quills beneath tail are brown and white. The Thick-spined Porcupine can be differentiated from the Sumatran Porcupine (H. sumatrae) byits larger skull with slightly longer nasals, longer rattle-quills, thicker quills, and largertactile bristles. Habitat. Primary lowland rainforest, old and young secondary forest, and open areas up to elevations of 1200 m in East Kalimantan, Indonesia. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Least Concern on The IUCN Red List. Endemic island rodents such as the Thick-spined Porcupine may be particularly at risk of extinction and require further study. Bibliography. Amori, Gippoliti & Helgen (2008), Corbet & Hill (1992), Glinther (1877b), Helgen & Lunde (2008), Medway & Harrisson (1963), Nowak (1999a), Rustam et al. (2011), Storch (1990), van Weers (1978)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 308, DOI: 10.5281/zenodo.6612213

6. Philippine Porcupine Hystrix pumila French: Porc-épic des Philippines / German: Palawan-Stachelschwein / Spanish: Puercoespin de Filipinas Other common names: Indonesian Porcupine, Palawan Porcupine, Philippine Short-tailed Porcupine Taxonomy. Hystrix pumila Gunther, 1879, “Puerto Princesa, in the island of Paragua [= Palawan],” Philippines. Placed in subgenus Thecurus. Monotypic. Distribution. SW Philippines, endemic on Palawan and adjacent Busuanga and Balabac Is. Descriptive notes. Head-body 450-665 mm, tail 64-190 mm, ear 31 mm (one individual), hindfoot 69 mm (one individual); weight 3.8-5.4 kg. The Philippine Porcupineis the smallest species of Hystrix. Spiny covering of bodyis poorly developed, with very small quills,tactile bristles, and rattle-quills and only a small number of quills on back. Like other species of Hystrix, tail has rattle-quills. Lengths of hollow capsulelike parts of rattle-quills are 10-11 mm. There is no crest on head. Maximum diameters of thickest defensive quills on back are 4-5 mm, and they are colored as single dark bands with no white tips. Absence of white tips on long quills of back differentiates Philippine Porcupines from Sumatran Porcupines and Thick-spined Porcupines. Habitat. There is no information available for this species. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Vulnerable on The IUCN Red List. Population of the Philippine Porcupine is expected to decline by more than 30% in the next three generations, based on rate of forest loss and numbers of individuals taken for bushmeat and the pet trade. The Philippine Porcupine is the most important game species of the Tagbanwa people and is often dug out of subterranean dens. Endemic island rodents such as the Philippine Porcupine may be particularly at risk of extinction and require further study. Bibliography. Amori, Gippoliti & Helgen (2008), Corbet & Hill (1992), Esselstyn et al. (2004), Heaney et al. (2008), Nowak (1999a), Storch (1990), van Weers (1978), Woods & Kilpatrick (2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 308, DOI: 10.5281/zenodo.6612213

4. Sumatran Porcupine Hystrix sumatrae French: Porc-épic de Sumatra / German: Sumatra-Stachelschwein / Spanish: Puercoespin de Sumatra Other common names: Sumatran Short-tailed Porcupine Taxonomy. Thecurus sumatrae Lyon, 1907, “Aru Bay, east coast of Sumatra.” H. sumatraeis in subgenus Thecurus. Monotypic. Distribution. Sumatra. Descriptive notes. Head-body 455— 560 mm, tail 90-110 mm, hindfoot 68-75 mm; weight 3.8-5.4 kg. The Sumatran Porcupine is intermediate in size, and its quill development is between the Philippine Porcupine (H. pumila) (smaller) and the Thick-spined Porcupine (H. crassispinis) (larger). The Sumatran Porcupine is distinctly speckled from white tips of spines on cheeks, neck, shoulders, flanks, and anterior part of back. Posterior of back has many defensive quills (more than the Philippine Porcupine and fewer than the Thick-spined Porcupine); number of quills with white tips is either equal to or less than number of quills with black tips. On quills with white tips, length of white is 10-30 mm and is almost always shorter than on the Thickspined Porcupine. On tail of the Sumatran Porcupine, rattle-quills are very small, no larger than on the Philippine Porcupine and always shorter than on the Thick-spined Porcupine. Lengths of hollow capsule-like parts of rattle-quills are 12-16 mm. There is no crest on head. Maximum diameters of thickest defensive quills on back are 5—7 mm and colored as a single dark band with a white tip ¢.70 mm in length. Skull of the Sumatran Porcupine is relatively broader and nasals shorter than those of the Thickspined Porcupine. ~ Habitat. Variety of primary and secondary habitats from sea level to elevations of at least 300 m. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. The Sumatran Porcupine is common and can occur at higher densities in secondary or disturbed habitats than in primary habitats. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Sumatran Porcupine is hunted for food, but hunting it not a serious conservation threat. Endemic island rodents such as the Sumatran Porcupine may be particularly at risk of extinction and require further study. Bibliography. Amori, Gippoliti & Helgen (2008), Aplin, Frost et al. (2008), Corbet & Hill (1992), Nowak (1999a), Storch (1990), van Weers (1978), Woods & Kilpatrick (2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 307, DOI: 10.5281/zenodo.6612213

10. Cape Porcupine Hystrix africaeaustralis French: Porc-épic du Cap / German: Stidafrikanisches Stachelschwein / Spanish: Puercoespin de El Cabo Other common names: CapeCrested Porcupine, South African Porcupine, Southern African Porcupine Taxonomy. Hystrix africaeaustralis Peters, 1852, “Querimba” and “Tette.” Restricted by R. E. Moreau and colleagues in 1946 to Querimba coast (ca. 10° 30’ to 12° 00’S, 40° 30’E, sea level) Mozambique. Hystrix africaeaustralis is in the subgenus Hystrix. It is sympatric with the similar H. cristata, in parts of southern Uganda, Kenya, and Tanzania. Up to three subspecies have been identified, but none was recognized by J. A. J. Meester in 1986 or D. C. D. Happold in 2013. Monotypic. Distribution. S one-half of Africa, from S of the Republic of the Congo and S DR Congo E to S Uganda, SW Kenya, and Tanzania (not in Zanzibar I), and S to South Africa, excluding Namibian coast. Descriptive notes. Head-body 630-805 mm, tail 105-130 mm, ear 39-48 mm, hindfoot 89-114 mm; weight 10-24.1 kg. The Cape Porcupine and the similarly sized Crested Porcupine (H. cristata) are the largest rodents in Africa. The Cape Porcupine is stout, heavily built, and well-bristled, with relatively short legs and short, well-hidden tail. There is no sexual dimorphism. Quills of several types on the Cape Porcupine are most notable. True quills are defensive quills; they are the thickest of modified hairs, circular in cross section, long (up to 300 mm) with very sharp points, and relatively inflexible. Their diameters are greatest at mid-lengths of quills and taper toward both ends. True quills carry microstructural “feathering” along tips, analogous to barbed quills seen in the North American Porcupine (Erethizon dorsatum), which may aid in reducing force required to allow muscle penetration by quills. Tactile bristles, also circular in cross section, are very flexible. Their diameters are greatest near bases and smaller than diameters of defensive quills. Transitional quills are intermediate between true quills and tactile bristles in length, diameter, and flexibility. Spines are long (up to 500 mm), sharp,stiff, flattened, and grooved bristly hairs that cover most of the body. Rattle-quills, located only on tail, are circular in cross section, hollow, capsule shaped, and open at ends. They attach to tail by stalks at bases of quills. Spines and quills are distributed across posterior two-thirds of dorsum and flanks but not on ventrum. Quills along dorsum are banded black and white; black bands are broader with white tips. Longer quills have multiple repeating bands of black and white, with up to six white sections per quill. There is an erectile crest of coarse hairs, up to 500 mm in length, occurring posteriorly from top of head to top of shoulders. Hairs at front of crest are black at bases with white tips. Some Cape Porcupines have triangular patches of white bristles (flattened hair) on either side of neck. Head is rounded, with grayish brown, bristled face and stout, movable vibrissae. Small eyes are situated far back on head, and ears are inconspicuous and humanlike in shape. Nasal bones are 51-58% of occipito-nasal length, wide, and posteriorly extended almost to anterior ends of orbits. Frontal-to-nasal bone ratio is 49-68%. Upper incisors are smooth, and folds of enamel and dentine characterize cheekteeth. Rump is covered with short, flat, white bristles on dorsal and ventral sides. Shorttail is covered with short quills, mostly invisible beneath defensive quills on lower back and has rattle-quills that rattle when tail is shaken. There are 4-6 mammae. In the field, it is difficult to distinguish Cape Porcupines from Crested Porcupines; short quills on lower rump of the Cape Porcupine are white, but they are black on the Crested Porcupine. Skull differences are clear and well described by van Weers. Habitat. [.owland and montane tropical forests, savanna grassland, brushland, Acacia and Brachystegia (miombo) woodlands, and semiarid deserts, with broad daily and seasonal temperature changes, from sea level to elevations of ¢.2000 m. In South Africa, Cape Porcupines prefer Burkea—Acacia savanna, probably due to concentration of preferred foods. They are not found in swampy areas. Food and Feeding. The Cape Porcupine is omnivorous and eats subterranean tubers, bulbs, and roots; shoots; fruits; and stems. Osteophagy is common, particularly in phosphorous-limited areas. It might eat carrion. It is considered a pest by farmers; it will raid melon, pumpkin, and maize crops; uproot young trees; and damage young plantations by debarking and feeding on innerliving tissue of trees from ground level to c.0-5 m up the trunk. Cape Porcupines are ecosystem engineers because their foraging has landscape-level effects through food choice and physical effects of digging. Selective bark feeding in some natural savanna ecosystems may influence patterns of succession. Cape Porcupines dig circular, conical holes in the soil surface in search of tubers, insect larvae, and seeds, which creates sites for seedlings and allows water to infiltrate the soil crust. Digging by Cape Porcupines annually disturbs 0-34% ofthe soil surface and displace 1-6 m®/ha/year ofsoil acrossits distribution. Breeding. The Cape Porcupine,like the Crested Porcupine and the Indian Crested Porcupine (H. indica), is monogamous. Sexual maturity is reached at 8-18 months of age for males and in the second year oflife for females. Males and females are reproductive throughout the year. In both captive and wild individuals in South Africa, most litters are born in August-March, corresponding to periods of summer rain; this timing probably maximizes neonatal survival, especially in arid regions that experience cold, dry winters. Females are polyestrous, with cycle lengths of 17-42 days. Sexual activity (lordosis, mounting, and ejaculation) occurs throughout estrus, pregnancy, and lactation, but actual copulation outside of estrus is hindered by presence of a vaginal closure membrane. This physical contact between sexes maintains cyclic ovarian activity and may strengthen pair bonds. Estrus begins when vaginal membrane opens, coinciding with a surge in 17B-estradiol. Copulation is typically preceded by the female approaching the male and occurs ¢.2-8 days after onset of estrus. The female adopts a receptive posture, with tail and rump raised and quills pointed away from the male. He mounts the female from behind, with his forepaws resting on her back. Intromission only occurs when the female is in estrus. After mating, a copulatory plug is formed. Gestation lasts 93-105 days, and females average onelitter per year, with 1-3 precocial young; 59% oflitters produce singletons and 30% twins. Newborns weigh 300-400 g and have soft defensive quills and closed, pointed rattle-quills at birth. Young nibble on solid foods by 9-14 days of age and begin to feed alone at 4-6 weeks of age, although they continue to nurse for 13-19 weeks when they weigh 2.5-4.7 kg. Lactation averages 101 days and includes c.110 days of anestrus during lactation. The father provides paternal care by accompanying young on foraging trips, grooming and huddling with them, and protecting them from threats. Although Cape Porcupines live in family groups, only the adult pair reproduces. Females are almost continuously exposed to sexually mature males. Subordinate females experience cyclic ovarian activity, but only the dominant female conceives. Mechanism of reproductive suppression of subordinate females is unknown, but progesterone concentrations, despite cyclic changes, remain insufficient for implantation. Social factors may impact ovarian function. Females separated from males in captivity experience impairment of ovarian function. Physical contact with males may be required to sustain regular follicular and luteal activities of females. Densities might influence whether or not females reproduce during their first year. Conception in sexually mature females is suppressed until after dispersal from her natal group. Cape Porcupineslive for about ten years in the wild and up to 20 years in captivity. Activity patterns. Cape Porcupines are nocturnal. They spend their days at rest in shelters in rock crevices, caves, or abandoned burrows of Aardvarks (Orycteropus afer), which they modify to suit their needs with their front claws. They may excavate their own holes or use termite mounds as dens. Cape Porcupines often bring bones into their shelters. They forage alone at night, although pairs forage near one another, spending less than 25% of their active periods more than 400 m apart and spending more than 50% of active time within 200 m of one another. A pair of Cape Porcupines foraging in proximity of one other could be mate guarding because it is not apparent that they cooperate in foraging or anti-predator defense. When alarmed, defensive action is to raise quills and shake the rattle-quills. If a threat does not subside, a Cape Porcupine might attack by backing rapidly toward the threat, stabbing it with long defensive quills that detach easily from skin. Movements, Home range and Social organization. Cape Porcupines are monogamous and live in family groups typically consisting of an adult male-female pair and subadult and juvenile offspring. Mature offspring may remain in their natal groups when dispersal opportunities are limited; groups of up to 14 individuals have been observed sharing a single burrow. One family unit will irregularly use up to six different burrows. Den entrances can be 10 m apart, and main tunnels can be 20 m long. A living chamber is found c.2 m below ground. Home range is a large, non-exclusive area surrounding a smaller, exclusive core area (territory) that is defended by an adult pair. Males scent-mark more frequently than females and more often in feeding than non-feeding areas, and scent marking may thus play a larger role in territory defense. Territorial boundaries are neither permanent nor rigidly defined. Space use changes with season, but home ranges of adult pairs tend to include the same general area throughout the year, and most activity occurs in a fairly small proportion of the home range. Home ranges of adjacent groups overlap little; neighbors rarely encounter one another in areas of overlap. Home range size is smaller for individuals that forage on natural food vs. individuals that raid crops. Cape Porcupines that feed in natural areas also have larger home ranges in winter (c.116 ha) than summer (c.67 ha). On average, home ranges of males and females in mated pairs overlap ¢.75%. Cape Porcupines exhibit bi-parental care of offspring. Adult males accompany young on foraging excursions and defend against territory intruders. Group huddling among parents and offspring while sleeping in the burrow also provides thermoregulatory benefits, particularly for newborns that do not leave burrows for about nine weeks. Assistance with thermoregulation allows newborns more energy to allocate toward growth. The monogamous pair bond, unusual in mammals, may allow for territory maintenance. There is immediate burrow take-over by a new pair upon the death of one member of a pair. Group living and a monogamous mating system appear to increase territory holding and juvenile success. In absence of natural predation, population age structure is variable, changing between seasons and years. Population regulation is densitydependent. High-density populations of 25 ind/km? have been observed on riverine plains in the South African Karoo. Eight Cape Porcupines per square kilometer is a high density for semiarid regions. Population size appears to be limited by available den sites and territory size. Populations can also be limited by extrinsic factors such as predation; Leopards (Panthera pardus) and Lions (Panthera leo) are primary predators of Cape Porcupines. Status and Conservation. Classified as Least Concern on The IUCN Red List. Cape Porcupines damage crops and are a source of accessible meat protein to indigenous people, leaving them vulnerable to hunting, trapping, and snaring. Their role as an ecosystem engineer indicates that their protection should be included in conservation plans. Although the Cape Porcupine can hybridize with the Crested Porcupine in captivity, hybridization does not occur in their natural zone of overlap in East Africa. Bibliography. van Aarde (1985, 1987, 1998), van Aarde & Skinner (1986), van Aarde & van Wyk (1991), Alexander (1956), Barthelmess (2006), Bragg et al. (2005), Corbet & van Aarde (1996), Dean & Milton (1991), De Graaff (1981), Duthie & Skinner (1986), Ferguson et al. (2010), Findlay (1977), Foley et al. (2014), Gooden & Augee (2015), Greaves & Khan (1978), Grubb (2008b), Haim et al. (1992), Happold (2013a), Lawton & Jones (1995), Macdonald (2009), Meester et al. (1986), Mohr (1964), Moreau et al. (1946), Morris & van Aarde (1985), Nowak (1999a), Roberts (1951), Skinner & Chimimba (2005), Skinner et al. (1984), Storch (1990), Thomson (1974), de Villiers & van Aarde (1994), van Weers (1979), Yeaton (1988)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 310-311, DOI: 10.5281/zenodo.6612213

1. Long-tailed Porcupine Trichys fasciculata French: Porc-épic a longue queue / German: Pinselstachler / Spanish: Puercoespin de cola larga Other common names: Asian Long-tailed Porcupine Taxonomy. Hystrix fasciculata Shaw, 1801, “Malacca.” This species is monotypic. Distribution. W Peninsular Malaysia, Sumatra, and Borneo. Descriptive notes. Head-body 350-480 mm, tail 175-235 mm, ear 28-32 mm, hindfoot 61-67 mm; weight 1.5-2.2 kg. The Long-tailed Porcupine is the least specialized of the Old World porcupines. Tail is long, brown, and scaly with soft spines, giving it a superficial resemblance to members of the genus Rattus (Muridae). Dorsum varies from blackish to light brown. Head, neck, and shoulders are somewhat lighter in color than posterior region of back. Ventrum is whitish or sometimes very light brown. Body is covered with short stiletto-like spikes and individual bristle-like spines in between. Underfur is woolly. Dorsal spines of Long-tailed Porcupines are short and flattened, whitish at bases and dark brown toward tips. Longest spines on adults are 33-46 mm; longesttactile bristles are 60-80 mm. Tactile bristles have same dark color as spines, with white bases 10-30 mm in length. Soft, flexible bristles on head have essentially same form as larger spines on back. Skull has well-marked postorbital processes. Broad forefeet have four well-developed digits, each with thick claw. Long tail (more than 35% of head-body length including brush, ¢.50% without brush) ends with brush of light colored, flattened bristles that do not bristle or rattle when shaken. Length oftail bristles depends greatly on wear; on study skins, largest tail bristles are 55-220 mm. Bristles are parallel-sided, 1-1 mm broad and 0-3 mm thick. Part or all of tail is often missing from specimens, suggesting that it breaks easily from body. One suggestion is that light-colored brush tail distracts predators from the head; when a Long-tailed Porcupine is seized,tail easily breaks off, allowing escape. Diploid number is 2n = 46. Habitat. [Lowland forest and secondary and degraded habitats and cultivated areas. Long-tailed Porcupines occur in lowland and submontane localities throughout Borneo, reaching elevations of 1220 m on Kinabalu and in the Kelabit uplands. It is semicommensal with humans. In a survey of primary lowland rainforest, old and young secondary forest, and open areas in East Kalimantan, Indonesia, the Long-tailed Porcupine was one of only four species detected only in primary forest. Food and Feeding. The Long-tailed Porcupine is often seen feeding on the ground where it consumesfallen tree seeds and bamboo shoots. It also climbs well and forages from tops oftrees and shrubs. It is reported to destroy pineapples in some areas. Breeding. There is no information available for this species. Activity patterns. The Long-tailed Porcupine is active mainly at night. It sleeps during the day in an underground burrow. It is primarily terrestrial but is an agile climber. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Longtailed Porcupine has been extinct from Bukit Soeharto Grand Forest Park in Borneo (East Kalimantan) since 1992. Bibliography. Aplin & Lunde (2008b), Corbet & Hill (1992), Francis (2008), Medway & Harrisson (1963), Nowak (1999a), Rustam et al. (2011), Storch (1990), van Weers (1976, 1993), Woods (1984), Woods & Kilpatrick (2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 306, DOI: 10.5281/zenodo.6612213

Family HYSTRICIDAE (OLD WORLD PORCUPINES) • Large nocturnal, terrestrial, herbivorous rodents, with bodies covered in long, sharp spines for defense. • • 50-110 cm. • Afrotropical, Palearctic, and Indo-Malayan Regions. • Mediterranean coastal shrubland, warm and cold savanna grasslands, dry semi-deserts, lowland and montane tropical forest, forested hills and steppes, acacia and miombo woodland, agricultural, and urbanized habitats. • 3 genera, 11 species, 13 taxa. • 1 species Vulnerable; none Extinct since 1600., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 304, DOI: 10.5281/zenodo.6612213

3. Asiatic Brush-tailed Porcupine Atherurus macrourus French: Athérure asiatique / German: Asiatischer Quastenstachler / Spanish: Puercoespin asiatico de cola de cepillo Other common names: Asian Brush-tailed Porcupine Taxonomy. Hystrix macroura Linnaeus, 1758, “Habitat in Asia.” Restricted by M. W. Lyon, Jr. in 1907 to “Malacca,” Malay Peninsula. Some authors recognize two subspecies in China, hainanus on Hainan Island and the nominate macrourus of Sichuan, Yunnan, Guizhou, Hubei, Hunan, and Guangxi, while others have argued that A. macrourus is monotypic. In a 1977 review of the genus, D. J. van Weers found only minor differences among purported subspecies insufficient to warrant subspecific designation. [ts distribution overlaps substantially with Hystrix brachyura. Monotypic. Distribution. SE Asia, from NE India (Assam, Meghalaya, and Mizoram) E to C & S China (S Sichuan, Guizhou, E Hubei, N Hunan, Yunnan, Guangxi, and including Hainan I) and S through Burma (= Myanmar) and mainland South-east Asia to Peninsular Malaysia, also on Tarutao I (SW coast of Thailand), and on Aur, Pemanggil, and Tioman Is (E coast of Peninsular Malaysia); possibly present in Bangladesh. Recorded from Sumatra but needsverification. Descriptive notes. Head—body 380-570 mm, tail 160-195 mm, ear 33-42 mm, hindfoot 68-80 mm; weight 1-5—4-3 kg. The Asiatic Brush-tailed Porcupine is small to medium-sized, with long,scaly tail that ends in brush of white bristles more than 200 mm in length. Body is dark brown above and covered with short, rigid, flattened, whitishtipped spines, not unlike those on a hedgehog. Neck, shoulders, and undersurface of body are covered with soft flattened spines. Spines of lumbar region are longer than others and mixed with a few long white bristles. Spines are grooved. Largest spines in a study skin are 1-2-2-2 mm thick and 1-8-3 mm wide. There is subtle to more distinct pattern of light and dark longitudinal stripes along back and sides of adults. Dorsal color varies due to different frequencies of visible white parts of spines and tactile bristles on body. Chin, throat, and chest between forelegs are whitish. Ventral color varies among individuals; some are mostly white on belly, and others range toward light brown and carry spines with pale brown midsections and whitish tips. The Asiatic Brush-tailed Porcupine does not have a crest on its head. Nasal bones are narrow and short, less than 30% of greatest skull length, and preorbital fossa is smaller than temporal fossa. Molars are completely rooted, and upperincisors are orange and nongrooved. Onfeet, pollux is vestigial, and hallux is clawed. There are 2-3 pairs oflaterally placed mammae. Diploid number is 2n = 54. Tail of the Asiatic Brush-tailed Porcupine is perhaps the most notable aspect ofits appearance. It is brown and scaly and spineless for the first one-third ofits length, and it ends in brush of quills shaped like strings of beads that rattle when shaken; sometimestail is broken off. Tails are 25-50% of head-body length and covered with scales, interspersed with spiny bristles. It ends in a cluster ofalternately expanded and contracted papery bristle hairs, termed “platelet bristles,” that is 80-100 mm in length. Each has 2-6 expansions or “beads” that are 1-3 mm wide and 10-15 mm long. Lengths oflargest tail bristles are 50-130 mm. The Asiatic Brush-tailed Porcupine differs from the African Brush-tailed Porcupine (A. africanus) by having a relatively flatter skull and lacking true quills on its body. It differs from the Long-tailed Porcupine (Trichysfasciculata) in the number of vertebrae: 14 dorsal vertebrae and 19-23 caudal vertebrae in Atherurusvs. 16 and 22-25 in Trichys. Habitat. In general, dense forests and plantations, preferring rocky areas. The Asiatic Brush-tailed Porcupine is rarely seen in the open. Food and Feeding. The Asiatic Brush-tailed Porcupine is primarily herbivorous and eats roots, tubers,fruits, tree bark, and some cultivated crops and grains. It will also eat insects and carrion and is known to chew on bones. Breeding. Female Asiatic Brush-tailed Porcupines typically give birth to one (sometimes two) precocial young after 100-110 days of gestation. Two litters may be produced per year. Young leave the nest after about one week—the time it takes for quills to harden after birth. Activity patterns. The Asiatic Brush-tailed Porcupine is nocturnal and generally terrestrial, but it is known to climb trees. During the day, it takes shelter in burrows among rocks in hilly areas. It constructs its own burrows that might connect and hold several individuals at a time. Movements, Home range and Social organization. The Asiatic Brush-tailed Porcupine 1s gregarious and lives in small groups of 6-8 individuals, using the same burrow system. Collection of six individuals within two nights at one location indicated thatit might not be as rare as previously thought. It is confirmed prey of the Indochinese Clouded Leopard (Neofelis nebulosa). Status and Conservation. Classified as Least Concern on The [UCN Red List. The Asiatic Brush-tailed Porcupine is considered near threatened in India, primarily due to habitat loss and harvest for food by humans. Bibliography. Agrawal (2000), Corbet & Hill (1992), Francis (2008), Grassman et al. (2005), Lunde & Molur (2008a), Lyon (1907), Menon (2009), Molur et al. (2005), Nowak (1999a), Smith & Yan Xie (2008), Srinivasulu & Srinivasulu (2012), Storch (1990), van Weers (1977), Woods & Kilpatrick (2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 307, DOI: 10.5281/zenodo.6612213

9. Crested Porcupine Hystrix cristata French: Porc-épic a créte / German: Gewohnliches Stachelschwein / Spanish: Puercoespin crestado Other common names: North African Crested Porcupine Taxonomy. Hystrix cristata Linnaeus, 1758, “Habitat in Asia.” Restricted by O. Thomas in 1911 to “near Rome,Italy.” Hystrix cristata is in subgenus Hystrix. Two species, H. cristata and H. galeata, had been described from Africa north of the Equator, but wide overlap in all characteristics led G. B. Corbet and L. A. Jones in 1965 to regard them as a single species. Several subspecies had been described in Italy and North Africa, although D. C. D. Happold recognized none in 2013. Monotypic. Distribution. S Europe (mainland Italy and Sicily Is), N Africa (along the coast and in parts of the Atlas Mts in Morocco, Algeria, Tunisia, parts of Libyan and Egyptian coasts, and isolated populations in the semiarid parts of Mauritania, Mali, and Niger), E Africa (S Sudan, South Sudan, Eritrea, S Djibouti, Ethiopia, and Somalia), and across a narrow band extending from Senegal to Tanzania (and including Zanzibar I). Introduced in the 19" century into Elba I (but now possibly extinct there) and from 2005 into Sardinia from mainland Italy. Descriptive notes. Head-body 455-930 mm, tail 60-170 mm, ear 39 mm (one individual), hindfoot 86 mm (one individual); weight 8-27 kg. The Crested Porcupine is the second largest rodent in Europe (second to the Eurasian Beaver, Castorfiber) and, along with the similarly sized Cape Porcupine (H. africaeaustralis), is the largest rodent in Africa. It is thickset, with coarse blackish brown fur and well-bristled body, relatively short legs, and short, well-hidden tail. Male-biased and reverse sexualsize dimorphism has been reported, but two recent studies found no difference in body size between adult males and females. Quill type and distribution are similar to the Cape Porcupine. Head, neck, feet, and belly of the Crested Porcupine are covered with short, sturdy, flattened spines and bristles (20-45 mm long, 2 mm wide). Back and sides are covered with 300-400 mm long,stiff true quills, banded in black and white. Well-developed integumentary smooth muscles and subcutaneous muscles reflect quill arrangement on back; these muscles contract to raise quills. Microscopic spiny bristles are present on quill surfaces and oriented in opposite direction from quills, resulting in a “harpoonlike” effect. Legs and ventrum are covered with short, coarse black hairs. Short legs are stout, and feet have large pads and well-developed claws. Erectile, well-developed crest of coarse brownish white hair up to 500 mm in length spreads from top of head to shoulders and characterizes morphology of the subgenus Hystrix. White chin patch is often present. Yellow-gray head of the Crested Porcupine is large and blunt, with small dark eyes and darkly pigmented ears, and has long, dark mystacial vibrissae. Long, wide nasal bones are more than 57% of occipito-nasal length and extend posteriorly almost to anterior end of orbits. Frontal:nasal ratios are 23-38%. Upper incisors are smooth, and folds of enamel and dentine characterize cheekteeth. Rump is covered with short, flat, dark bristles on dorsal and ventral sides. Short tail is covered with short quills, mostly invisible beneath defensive quills on lower back, and has rattle-quills that rattle when tail is shaken. There are 4-6 mammae. Perianal glands are well developed. In the field, it is difficult to distinguish the Crested Porcupine from the Cape Porcupine; short quills on lower rump of the Crested Porcupine are black, whereas they are white on the Cape Porcupine. Skull differences are clear and were well described by van Weers. Compared with the Indian Crested Porcupine (H. indica), short spines on rump of the Crested Porcupine form conspicuous white patches when raised, and crest is predominantly white. Premaxillae of medium width, and frontal-to-nasal ratio 23-38%. Habitat. Woodlands, warm and cold savanna grasslands, dry semi-deserts, and warm Mediterranean coastal shrublands. In Algeria and Morocco, the Crested Porcupine lives in forested hills and steppes. In the Atlas Mountains and on Kilimanjaro,it lives at elevations of up to 3500 m. It prefers rocky areas. Food and Feeding. The Crested Porcupine is a generalist herbivore. It eats subterranean bulbs, roots,fruits, and bark. It can be a pest to farmers, damaging cassava, sweet potato, and groundnut crops in savanna farmlands. In Italy, the Crested Porcupine changesits diet seasonally, consuming roots as the predominant food year-round, supplemented by herbs in winter and spring, grass inflorescences and fruits in summer, and fruits in autumn. Breeding. The Crested Porcupine, like the Cape Porcupine and the Indian Crested Porcupine, is monogamous. Breeding takes place throughout the year, with a female giving birth to 1-2 precocial offspring once or twice per year. In captivity, there is a nightly rhythm of mounting. Multiple mountings per night are typical, but copulation takes place only during estrus (a difference from the Indian Crested Porcupine and the Cape Porcupine in which mounting and copulation take place independent of stage of estrus). The female Crested Porcupine presses her quills against her back to allow mating. Gestation lasts 7-8 weeks,after which litters of 1-4 young are born in a grass-lined burrow. Longevity is up to 20 years. Activity patterns. The Crested Porcupine is nocturnal. Individuals spend their days at rest in burrows. In Italy, a Crested Porcupine might share its burrow with a European Badger (Meles meles). The Crested Porcupine is active across a wide range of temperatures, with mean duration of nocturnal and crepuscular activity in central Italy lasting on average 9-2 hours/night, varying little throughout the year. In cold months when nights were longer, individuals left burrows after sunset and returned some hours before sunrise. In the warm season, they left at sunset or shortly before and returned at or shortly after sunrise. Moonlight avoidance is slight. Diurnal activity is scarce. When alarmed, a Crested Porcupine first raises its quills and erects its crest; second,it rattles its tail; and third, it stomps its hindfeet and growls. If these displays fail to deter a threat, it may attack by moving rapidly backward and sideways toward it, stabbing it with its long defensive quills that detach easily from the skin. Movements, Home range and Social organization. Crested Porcupines live in family groups. In captivity, males mark feeding sites with secretions from perianal glands. Chemical profiles from secretions of perianal glands may aid in individual recognition. Published studies of space use, home range, and social behavior report only on Italian populations. Home range size does not differ between males and females, although in one study in Tuscany, upper-limit of home range size was significantly higher for females than males. Home range size varies with habitat and season. Crested Porcupines in agricultural areas have larger home ranges than those in natural maquis shrubland (densely growing evergreen shrubs), and home range size decreases with increasing habitat richness. Home ranges were significantly larger in the warm season than the cold season in both habitats. Average minimum convex polygon home range size in agricultural areas was 151 ha in summer and 128 ha in winter vs. 50 ha in summer and 34 ha in winter in natural areas. For each male-female pair, median home range overlap is ¢.75%, with no change between seasons. Pair members show a strong and consistent spatial overlap throughout the year. Crested Porcupines prefer covered habitats by day. At night, those foraging in agricultural habitat spent 42-5% of time in ecotone areas, 28-5% in fields or fallows, and 28% in woods; those foraging in natural habitats stayed in cover 69% ofthe time, using the ecotone only 19% and open habitat 12% of the time. In terms of distance moved per night, Crested Porcupines moved greater distances in pine woods habitat (460-746 m) than grazed areas (210-225 m). Those living in pinewoods did not move to agricultural fields to forage, but those living in grazed areas did. Crested Porcupines will visit feeding areas as far as 10-12 km from their dens. Very little is known about densities and population dynamics of Crested Porcupines. In Italy, the Crested Porcupine has been found in feces of Red Foxes (Vulpes vulpes) and, rarely, Gray Wolves (Canis lupus). Status and Conservation. Classified as Least Concern on The IUCN Red List. The Crested Porcupine is a nationally protected species in Italy, although its eradication from Sardinia, where it has been recently introduced, has been recommended. Some recommend listing the Crested Porcupine as an endangered species in Europe, due to destruction of maquis,its optimal habitat, and because of poaching (especially in recently colonized areas). Payments of up to €19,500/year are claimed for crop damage by farmers in southern Tuscany. Distributional expansion in Italy would suggest that the Crested Porcupine is in fact doing well. Lack of data on abundance and densities regionally and nationally in Italy makes management a challenge. In Africa, the Crested Porcupine is one of the most important species for subsistence and income in the Albertine Rift where it is used for human food and wearing apparel. Almost nothing is written about ecology or behavior of the Crested Porcupine from anywhere in Africa. Although there remains some debate about how the Crested Porcupine came to be in Italy, its presence there is likely the result of an introduction from North Africa. Although there are fossil remains from the genus Hystrix in Europe beyond Italy, recent molecular analysis of three genes from Italian and African specimens shows a pattern of diversity congruent with an introduction from northern Africa during the Roman Age; one of three clades revealed in the analysis grouped Italian specimens together with specimens from Tunisia. A large number of DNA haplotypes present in Italian specimens may indicate several introductions over historical time. Since the 1970s, distribution of the Crested Porcupine in Italy has been expanding northward and eastward, crossing the Apennines from the Tyrrhenian coast to Marche. Historical and social factors, including progressive urbanization and consequent abandonment of traditional land use in mountainous areas, probably have contributed to distributional expansion. Despite suggestions otherwise, the Crested Porcupine has never occurred on the Ionian and Aegean islands, the Balkans, or the Iberian Peninsula in the Holocene, even in the very recent past. Bibliography. Amori & Angelici (1992), Angelici, Cabras & Trucchi (2009), Angelici, Capizzi et al. (2003), Bruno & Riccardi (1995), Carr et al. (2013), Corbet (1978), Corbet & Jones (1965), Corsini et al. (1995), Felicioli et al. (1997), Fischer et al. (2002), Foley et al. (2014), Giotto (2011), Grubb et al. (2008), Happold (2013a), Hoath (2009), Lovari et al. (2013), Masseti (2012), Massolo et al. (2009), Mohr (1965), Mori & Lovari (2014), Mori et al. (2013), Nowak (1999a), Pigozzi (1987, 1992), Pigozzi & Patterson (1990), Sonnino (1998), Storch (1990), Thomas (1911c¢), Trucchi & Sbordoni (2009), van Weers (1979)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Hystricidae, pp. 304-312 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 309-310, DOI: 10.5281/zenodo.6612213