taxonID	type	description	language	source
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	John Rey described “ Hystrix, The Porcupine ” (Rey 1693: 206) on the basis of a specimen captured from mountains near Rome (Italy), whilst Charles Perrault and Denis Dodart in their description of the “ Porc-Epics ” and “ Herissons ” (Perrault & Dodart 1731: 233) did not define the origin of the animals examined. Linnaeus (1758) reported “ Habitat in Asia ” since he included in the references after H. cristata also “ Hystrix orientalis cristata ” by Seba (1734: 79). The latter described “ Hystrix, Orientalis, cristata ” from Sumatra and Java (Seba 1734), today belonging to other species (see Barthelmess 2020). However, the first quotation of Linneaus (1756: 9) is to Ray (1693), therefore Thomas (1911) restricted to Italy (mountains near Rome) the “ locus typicus ” of H. cristata (see also Miller 1912). Curiously, the species was described on the basis of a specimen from Italy, where the species most likely has been introduced in historical times (see Trucchi & Sbordoni 2009; Masseti et al. 2010; Mori et al. 2013, 2014; Trucchi et al. 2016; see also the discussion below on published and available nominate “ trinomen ” H. cristata cristata Linnaeus, 1758, as opposed to the other variety he named H. cristata indica (currently a valid species: H. indica Kerr, 1792). According to the Code (ICZN 1999, Art. 45.6), this trinomen must be treated as subspecies. Miller (1912: 543) correctly synonymized it with H. cristata Linnaeus, 1758. The name coined by Robert Kerr refers to the specimen figured by Smellie (1785: pl. CCV) in its English translation of the “ Histoire Naturelle ” by Georges-Louis Leclerc, Comte de Buffon. Although William Smellie redrawed the figure (cf. Smellie 1785: pl. CCV), he based them on the original plate by Louis-Jean-Marie Daubenton (cf. L. Daubenton 1764: 412, pl. LI) and mentioned by Buffon (1764: 407). Thus, the specimen portrayed is the same one mentioned by Buffon (1764: 407) and L. Daubenton (1764: 412), which was sent to him from Rome (Italy) and then the type locality is not unknown as asserted by Allen (1939). Therefore, also the type locality of this taxon coincides with that of H. cristata Linnaeus, 1758 (see above).	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	Although no material from Senegal is available also for us, this country falls within the range of H. cristata and, according to nuclear sequences, western Africa material is in the same clade with part of the specimens from north Africa (cf. Trucchi & Sbordoni 2009, Trucchi et al. 2016). Therefore, no reason exists, at the present state of knowledge, for considering the populations south of the Sahara deserving to be classified in a separate taxon. Accordingly, Trucchi & Sbordoni (2009) showed that samples of H. cristata from Eastern Africa, Western Africa, Libya and Tunisia belong to the same molecular clade (see Fig. 3); RAD sequencing (Trucchi et al. 2016) also seems to confirm this pattern. According to the low recorded genetic distance (global native population F ST value = 0.66; matrices available in Trucchi 2008) and to the potential lack of reproductive isolation, molecular differences between the Mediterranean and the Sub-Saharan population of H. cristata have to be considered as elements of intraspecific variation, with no clear subspecific separation (Trucchi 2008). Furthermore, net genetic distances based on the GTR + Γ + I model of sequence evolution (Tavarè 1986) were calculated on the total mtDNA fragments of both H. cristata and H. africaeaustralis (Trucchi & Sbordoni 2009) and were all lower than 0.008 amongst different populations of H. cristata (net genetic distance H. cristata - H. africaeaustralis = 0.057 ± 0.002), therefore showing only intraspecific variability, without confirming the existence of any subspecies (see Trucchi 2008). Albeit, as also suggested by Angelici et al. (2021), further research is needed especially on sub-Saharan populations of H. cristata. Then, cautiously and pending further research, we revalidate here the synonymy Hystrix senegalica Cuvier, 1823 = Hystrix cristata Linnaeus, 1758. Syn. rev.	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	Cuvier (1823) based his description on that skeleton figured by L. Daubenton (1764: 413, pl. LIII) and hypothesized that could be one of the porcupines dissected by Perrault (1676: 113) and that its origin could be Africa (Réaumur 1729). There is actually no evidence in confirmation of the hypotheses formulated by Cuvier (1823). Moreover, Georges-Louis Leclerc, Comte de Buffon did not mention the skeleton in his part to “ Le Porc-épic ” in the “ Histoire naturelle ” (Buffon 1764), but only the two specimens from Italy and Indies. Thus, the skeleton may have been extracted from one of these as well. This taxon was set as synonym, with doubt, of H. cristata cristata Linnaeus, 1758 by Ellerman (1940: 219). He wrote that, since the locality of origin is unknown, perhaps it would be “ best regarded as unidentifiable ”. Therefore, although it is not possible to know to which of the currently available species of Hystrix to refer this taxon, it is quite safe that this is a synonym and we leave it as a synonym of H. cristata Linnaeus, 1758, since it is the most likely scenario. A petition to the ICZN Commission to set aside this name under its plenary power (see ICZN 1999, Arts. 75.5, 81) seems not necessary in this case. Moreover, the skeleton described and figured by L. Daubenton (1764) lacks three teeth: in particular, the first molars on each side of the upper jaw and the third molar on the left side of the same jaw, and it could be a young specimen with some milk teeth fallen out (see L. Daubenton 1764: 435 – 436). Accordingly, in future the provenance of this specimen might also be reconstructed and its origin identified.	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	However, the provenance of the skull examined by John Edward Gray is unknown (Gray 1847: 102). Then, P. Sclater (1865: 356) synonymized it under H. cristata Linneaus, 1758 and Gray (1866) left it in the group of Italian and African porcupines (see also Cabrera 1924). Therefore, even if the provenance of the skull studied by Gray (1847) remains unknown, its synonymization with H. cristata remains the most plausible one.	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	However, Corbet & Jones (1965) compared more data and found “ mean differences in all these characters between the East and North African groups [of Hystrix spp.] ” and “ in every case ” they found “ wide overlap ” and “ no reason to consider ” H. galeata specifically distinct from H. cristata, “ nor a justification for the recognition of a subspecific difference ”. Thus, they synonymized this taxon with H. cristata (Corbet & Jones 1965: 295). Both mtDNA and nuclear sequences available from Tanzania (southernmost population, compared to Kenya) fall in the clade of H. cristata, with a quite high genetic flux with other African populations (F ST value = 0.34 - 0.46: Trucchi 2008). Trucchi & Sbordoni (2009) showed that samples of H. cristata from Eastern and Western Sub-Saharan Africa, as well some samples from Libya and Tunisia, belong to the same molecular clade, as also supported by the genetic structure obtained with RAD sequencing (Trucchi et al., 2016). Therefore, we are able to confirm the view of Corbet & Jones (1965).	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	It is difficult to pronounce in this case, since it could be assigned as a synonym of H. cristata or H. africaeaustralis, both occurring in Tanzania (Barthelmess 2020). However, since the descriptor found strong affinities with H. galeata (= H. cristata), we follow the view of Misonne (1974). However, it has been suggested that the taxon H. galeata ambigua might be related to potential hybrid specimens between H. cristata and H. africaeustralis (Trucchi & Sbordoni 2009; Mori et al. 2013; Trucchi et al. 2016). Hystrix galeata conradsi F. Müller, 1910 Ferdinand Müller described this taxon as new subspecies of H. galeata Thomas, 1893 on the basis of differences on skull, i. e., by the size of the premaxillary nasal process and the thickness of the maxillary zygomatic arch (Müller 1910 b: 314). This taxon was included under H. cristata by Misonne (1974: 8), hence informally synonymizing it, but without further explanation. According to nuclear sequences (see Trucchi et al. 2016) specimens from Tanzania fall in the clade of H. cristata. Therefore, we agree with Misonne (1974). Hystrix galeata lademanni F. Müller, 1910 Müller (1910 b: 314) described a further taxon as a new subspecies of H. galeata Thomas, 1893 on the basis of the same features examined for H. galeata conradsi F. Müller, 1910. The name was first spelled “ H. galeata ludemanni ”, then twice “ H. galeata lademanni ” (Müller, 1910 b: 314). However, also since it was clearly dedicated to Lademann (Müller, 1910 b: 314), first spelling can be considered an inadvertent error and must be corrected (ICZN 1999, Art. 32.3). This taxon, misspelled as “ ladermani Müller, 1910 ”, was included under H. cristata by Misonne (1974: 8), hence informally synonymized, but without further explanation. According to mtDNA and nuclear sequences (Fig. 3), specimens from Tanzania fall in the clade of H. cristata. Also in this case, the differences highlighted (see Müller 1910 b) seem to fall within population variability. Therefore, we agree with Misonne (1974). Hystrix galeata loennbergi F. Müller, 1910 Müller (1910 b: 315) named a new “ rasse [= race] ” from Mount Kilimanjaro (locality not specified) on the basis of a skull given to the Zoologische Museum of Berlin [now Natural History Museum, Berlin]). He stated that the ratio of anterior to posterior width of the nose is completely different from that of H. galeata ambigua Lönnberg, 1908 but he leaves open the possibility that it coincides with this taxon or with H. galeata Thomas, 1893. As the previous taxa, this was included too under H. cristata by Misonne (1974: 8), hence informally synonymized, but without further explanation. Then, this taxon was indicated as synonym by subsequent author (e. g., Woods & Kilpatrick 2005 as “ lonnebergi [sic!] Müller, 1910 ”). However, according to the Code (ICZN 1999, Art. 32.5.2.1) this must be corrected transcribing the vowel with umlaut “ ö ” as “ oe ”. According to mtDNA and nuclear sequences (Fig. 3) specimens from Tanzania fall in the clade of H. cristata. Moreover, morphological differences between individuals of Tanzania and those from other African countries seem to fall within the intraspecific variability (Müller 1910 b). Therefore, we agree with Misonne (1974).	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	The author highlighted some minor differences in the skull. Misonne (1974: 8) listed this taxon under H. cristata, hence informally synonymizing it, but without further explanation. According to molecular sequences (Fig. 3) specimens from East Africa (e. g., Tanzania and Ethiopia) fall in the clade of H. cristata. Also in this case, the differences highlighted (see Lönnberg 1912) seem to fall within population variability. Therefore, we agree with Misonne (1974).	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	Molecular analyses conducted both on mtDNA and nuclear genes (Fig. 3), confirmed that populations from northern and western Morocco are genetically close to the central and southern Italian ones (type locality of H. cristata is Rome). The populations from Italy were very likely introduced just from Maghreb (see e. g., Trucchi & Sbordoni 2009; Masseti et al. 2010; Mori et al., 2015, 2016; Trucchi et al., 2016; Viviano et al. 2020). Therefore, no doubts on the correctness of this synonymy seem to exist and the differences observed by Cabrera (1924) are very likely due to the variability of populations at the extremes of the range of H. cristata.	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF95FFA2FF31B5ADF3E2F9BC.taxon	description	Misonne (1974: 8) listed this taxon under H. cristata, hence informally synonymizing it, but without further explanation. Thomas (1925) considered this “ a diminished Aïr form of the ordinary W. - African porcupine ”; however, there are no reasons to think to a good taxon, considering also it was a young specimen difficult to reliably compare, thus we agree to Misonne (1974).	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF98FFA0FF31B2C5F3A8FB22.taxon	description	“ Hystrix capensis Gr. ” Grill (1860: 9, 19) listed this name “ Hystrix capensis Gr. ” without any description associated. He merely wrote that the Swedish zoologist and explorer Johan Fredrik Victorin does not seem to have seen this animal alive, but of a dead specimen, which he found at Saltriver near Knysna (October 23 [1954]), he has brought home the skull ” [translated from Swedish]. In a following contribution, Grill (1863) reported the notes by J. F. Victorin who wrote that at Saltriver (the same locality of the first mentioned record) he “ found a dead porcupine (Hystrix capensis), and, though the animal smelled disgustingly, however, [he] took off the skull and cleaned that for [his] collection ” [translated from Swedish]. This name has been reported by some subsequent authors without authorship (e. g., Leche 1883) or attributed to “ Grill, 1958 ” (correct year would be however 1960) as synonym (without further explanation) of H. africaeaustralis Peters, 1852 (e. g., Sclater 1901; Allen 1939; Ellerman 1940; Ellerman et al. 1954; Maguire 1976; Woods & Kilpatrick 2005; Barthelmess 2006). However, Grill (1860) clearly indicated and described the new species in the work and specified them also in the introduction (i. e., Rhinolophus auritus Vict. [= Victorin, 1860] and Delphinus Victorini nobis [D. victorini Grill, 1860]). Therefore, he possibly described this taxon in a previous work (before October 23, 1954 when J. F. Victorin wrote his zoological notes during his journey in the Cape region) or it has been described by another author abbreviated as “ Gr. ”. Despite the attempts, however, we have not been able to find it and thus we listed this name as nomen dubium. If instead it will be proved that this name is attributable to Grill (1860) who did not provide a description, it will have to be considered as nomen nudum. It is interesting to note that Cuvier (1823) mentioned “ le porc-épic du Cap de Bonne-Espérance ” that he states is different from H. cristata Linneaus; however, without any description or latin name. In any case, no reason exist to consider this taxon as distinct from H. africaeaustralis since the specimens from southern Africa in the phylogenetic tree are clustered together (Fig. 3). Hystrix stegmanni F. Müller, 1910 In the description of this new taxon, Müller (1910: 186) states that it is most similar to H. galeata ambigua Lönnberg, 1908 and H. africaeaustralis Peters, 1852 but “ shows essentially different peculiarities of both ” (i. e., measurements and details of the skull). Corbet & Jones (1965) studying topotypical specimens, found that these are intermediate between H. cristata Linnaeus, 1758 and H. africaeaustralis for some features, but for several others they match well with H. africaeaustralis. Therefore, they synonymized H. stegmanni with H. africaeaustralis, but they added also they did not express opinion as a possible validity of this taxon at subspecific level (Corbet & Jones 1965: 295). The type locality, in the Rwandan side of Lake Kivu, is where ranges of H. cristata and H. africaeaustralis seem to meet (see Mohr 1965; Trucchi & Sbordoni 2009) and further research should be useful to deepen if hybrid specimens exist (see Mohr 1965). However, we believe that no reasons exist to consider this taxon as distinct from the two currently valid species. Hystrix africaeaustralis prittwitzi F. Müller, 1910 Soon after, Müller found some minor differences in the skull of a specimen from Tabora, in present day Tanzania, and described a new taxon close to H. africaeaustralis Peters, 1852, placing it as a subspecies. In particular, he observed a different development of some skull bones: i. e., nasal process of premaxilla andzygomatic (see Müller 1910 b: 313 for details). Corbet & Jones (1965) showed variability in these features among different populations, but they did not deal with subspecies. Misonne (1974: 8) included this taxon under H. africaeaustralis, but without further explanation, and this line has been followed until today considering it a synonym (e. g., Meester et al. 1986, Woods & Kilpatrick 2005, Happold 2013). We agree with the view since the variability evidenced by Corbet & Jones (1965). This population is probably only an extreme one in the range of the species and it is quite obvious that differences can emerge from comparison with more southern populations.	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
039F87D5FF98FFA0FF31B2C5F3A8FB22.taxon	description	We agree with Ellerman et al. (1954: 237) who synonymized this taxon under H. africaeaustralis Peters, 1852 (without further explanation), since it appears very likely these features fall in the variability of the species (populations at extreme of range) and because both mtDNA and nuclear DNA analyses showed all the specimens from Southern Africa clustered together (Trucchi & Sbordoni 2009; Trucchi et al. 2016). For completeness, Woods & Kilpatrick (2005) recognized this subspecies, but soon after Barthelmess (2006) returned to consider it as synonym of H. africaeaustralis.	en	Mori, Emiliano, Sogliani, Davide, Senini, Caterina, Laurenzi, Alessandro, Viano, Andrea Vi-, Cianferoni, Fabio (2021): Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Zootaxa 5047 (5): 501-519, DOI: 10.11646/zootaxa.5047.5.1
