Hystrix cristata Linnaeus, 1758: 56 Rey (1693: 206) Hystrix cristata europaea Kerr, 1792: 213 Daubenton (1764: 412 Buffon (1764: 407) Daubenton 1764: 412 H. cristata Miller (1912: 543) Hystrix senegalica Cuvier, 1823: 430 H. cristata Misonne (1974: 8) Acanthion Cuvier, 1823 Maguire (1976: 47) H. cristata Acanthion Daubentonii Cuvier, 1823: 431 Daubenton 1764: 435–436 Ellerman (1940: 219) Acanthion Cuvieri Gray, 1847: 102 Sclater (1865: 356) Hystrix galeata Thomas, 1893: 230 Corbet & Jones (1965: 295) Hystrix galeata ambigua Lönnberg, 1908: 29 Misonne (1974: 8) Hystrix galeata conradsi F. Müller, 1910b: 314 Moreau et al. (1946: 430) Allen & Loveridge (1933: 26) Misonne (1974: 8) Hystrix galeata lademanni F. Müller, 1910b: 314 H. galeata ludemanni H. galeata lademanni Müller, 1910b: 314 Moreau et al. (1946: 430) Misonne (1974: 8 Hystrix galeata lönnbergi F. Müller, 1910b: 315 Misonne (1974: 8) Hystrix galeata somalensis Lönnberg, 1912: 109 Moreau et al. (1946: 430) Misonne (1974: 8) Hystrix occidanea Cabrera, 1924: 220 Ellerman (1940: 219) Ellerman et al. (1954: 520) H. cristata cristata Linnaeus, 1758 H. cristata Misonne (1974: 8) Hystrix aerula Thomas, 1925: 196 Misonne (1974: 8) Hystrix cristata Linnaeus, 1758 H. cristata Hystrix senegalica Cuvier, 1823 H. cristata Acanthion Cuvier, 1823 H. cristata H. cristata H. cristata Acanthion daubentonii Cuvier, 1823 H. cristata europaea Kerr, 1792 Acanthion cuvieri Gray, 1847 Hystrix galeata Thomas, 1893 H. galeata H. cristata H. africaeaustralis Hystrix galeata ambigua Lönnberg, 1908 H. galeata Thomas, 1893 H. galeata H. africaeaustralis H. galeata H. galeata H. cristata Hystrix galeata somalensis Lönnberg, 1912 H. galeata Thomas, 1893 Hystrix occidanea Cabrera, 1924 H. cristata H. senegalica H. cristata H. cristata H. cristata cristata Linnaeus, 1758 Hystrix aerula Thomas, 1925 H. senegalica Cuvier Subspecific taxonomy of African porcupines Hystrix spp.: is there anything beyond the species level? Mori, Emiliano Sogliani, Davide Senini, Caterina Laurenzi, Alessandro Viano, Andrea Vi- Cianferoni, Fabio Zootaxa 2021 2021-10-04 5047 5 501 519   Hystrix cristata  Linnaeus, 1758: 56   7VHZZ Linnaeus, 1758 Linnaeus 1758 [151,535,459,485] Mammalia Hystricidae Hystrix Animalia Rodentia 4 505 Chordata species cristata      Hystrix cristata Linnaeus, 1758: 56. Type/s: HT: specimen mentioned by  Rey (1693: 206); Typelocality: “Asia”; then restricted by Thomas (1911)to Italy, mountains near Rome – “Italis […] Romae […] in montibus vicinis”. Repository: unknown (not known if preserved).     Hystrix cristata europaea Kerr, 1792: 213. Type/s: HT: specimen figured by Smellie (1785: pl. CCV): “iconotype”; based on the figure by L.  Daubenton (1764: 412, pl. LI), mentioned by  Buffon (1764: 407). Typelocality: Italy, Rome – “Rome” (L.  Daubenton 1764: 412). Repository: unknown (not known if preserved). Denominated “variety” by Kerr (1792). Unnecessary renaming of  H. cristata. Syn.  Miller (1912: 543).     Hystrix senegalica Cuvier, 1823: 430. Type/s: HT: young specimen (molar germs attached to portions of head). Typelocality: Senegal– “ Sénégal”. Repository: unknown. Downgraded by Ellerman (1940)to subspecies of  H. cristata. Syn.  Misonne (1974: 8). Re-upgraded and placed into the genus  Acanthion Cuvier, 1823by  Maguire (1976: 47), although it is an unofficial nomenclatural act since included in a thesis dissertation. Revalidated as subspecies of  H. cristataby Angelici et al.(2020). Here again synonymized (see below). Syn. rev.     Acanthion Daubentonii Cuvier, 1823: 431. Nomen dubium. Type/s: HT: probably young specimen (skeleton with lacking some teeth; see L.  Daubenton 1764: 435–436). Typelocality: unknown. Repository: unknown. Syn.  Ellerman (1940: 219).     Acanthion Cuvieri Gray, 1847: 102. Type/s: HT: adult specimen (skull). Typelocality: unknown. Repository: NHML. Syn. P.  Sclater (1865: 356).     Hystrix galeata Thomas, 1893: 230. Type/s: HT: subadult specimen (skull). Typelocality: Kenya, LamuIsland, Lamu– “ Lamu, [British] East Africa”. Repository: unknown. Syn.  Corbet & Jones (1965: 295).     Hystrix galeata ambigua Lönnberg, 1908: 29. Type/s: HT: adult old specimen (description based on skull only). Typelocality: Tanzania, Kilimanjaro, Kibong’oto – “[ Deutsch-Ostafrika, Kilimandjaro:] Kibonoto”. Repository: unknown. Syn.  Misonne (1974: 8).     Hystrix galeata conradsiF. Müller, 1910b: 314. Type/s: STS: probably two specimensbut not specified (skulls). Typelocality: Tanzania, Mwanzaand “Neuwied” [catholic mission station] on Ukerewe Island on Lake Victoria – “[ Deutsch-Ostafrika,] Muansa und […] Neuwied auf Ukerewe”. Restricted to Ukerewe Island by  Moreau et al.(1946: 430)since  Allen & Loveridge (1933: 26)found topotypic material of this taxon on indication by the collector A. Conrads. Repository: ZMB. Syn.  Misonne (1974: 8).     Hystrix galeata lademanniF. Müller, 1910b: 314. First spelled “ H. galeata ludemanni”, then twice spelled “ H. galeata lademanni”; since dedicated to Lademann (  Müller, 1910b: 314), first spelling can be considered an inadvertent error and must be corrected (ICZN 1999, Art. 32.3). Type/s: STS: probably two specimensbut not specified (skulls). Typelocality: Tanzania, Kondoa Irangi and lower Bubu – “[ Deutsch-Ostafrika,] Kondoa Irangi und […] unteren Bubu”. Restricted to “Kondoa lrangi” by  Moreau et al.(1946: 430). Repository: ZMB. Syn.  Misonne (1974: 8, misspelled “ ladermaniMüller, 1910”).     Hystrix galeata lönnbergiF. Müller, 1910b: 315. Type/s: HT: skull given to the SBM by von der Marwitz. Typelocality: Tanzania, Kilimanjaro– “[ Deutsch-Ostafrika,] Kilimandjaro”. Repository: ZMB. Syn.  Misonne (1974: 8).     Hystrix galeata somalensis Lönnberg, 1912: 109. Type/s: STS: 1 maleand 1 female, old specimens (skulls from Burao [ Somalia]; NHML), 1 adultfemale specimen (Nyoro [ Kenya]; not specified if preserved; possible repository unknown). Typelocality: Somalia, Burco [= Burao] – “[ British East Africa,] Burao [and] Nyoro”; restricted to “Burao, British Somaliland” by  Moreau et al.(1946: 430). Repository: NHML (in part). Syn.  Misonne (1974: 8).     Hystrix occidanea Cabrera, 1924: 220. Type/s: HT: adult (old) male. Typelocality: Morocco, Essaouira– “[ Marruecos,] Mogador”. Repository: MNCN. Downgraded to subspecies by  Ellerman (1940: 219). Syn. (with doubt)  Ellerman et al.(1954: 520)with  H. cristata cristata Linnaeus, 1758; included under  H. cristataby  Misonne (1974: 8).     Hystrix aerula Thomas, 1925: 196. Type/s: HT: young specimen (skull only). Typelocality: Niger, Aouderas (Asben [Aïr]) – “Aouderas, Asben”. Repository: NHML. Syn.  Misonne (1974: 8).   Hystrix cristata Linnaeus, 1758 Carl Linnaeus in the 10 thedition of the Systema Naturae ( Linnaeus 1758), after the description of  H. cristata, refers to the 9 thedition of his work ( Linnaeus 1756). Although, the latter was published before the starting point of zoological nomenclature ( 1 stJanuary 1758) fixed by the International Code of Zoological Nomenclature (ICZN 1999), it is in this work ( Linnaeus 1756: 9) that he clearly refers to two pre-Linnean works: Rey (1693)and Perrault & Dodart (1731).  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. cristataalso “  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 typelocality is not unknown as asserted by Allen (1939). Therefore, also the typelocality of this taxon coincides with that of  H. cristata Linnaeus, 1758(see above).    Hystrix senegalica Cuvier, 1823  Cuvier 1823(430) described as a different species the first molar germs attached to portions of the head of specimens from Senegal. Maintained as a separated species by Allen (1939)and downgraded to subspecies by Ellerman (1940), finally synonymized with  H. cristataby Misonne (1974). Maguire (1976: 47)re-upgraded the name at species level and placed it into the genus  Acanthion Cuvier, 1823, but this is an unofficial nomenclatural act since included in a thesis dissertation. Recently, Angelici et al.(2021)revalidated this taxon as subspecies of  H. cristata, claiming skull morphological differences between North African and sub-Saharan populations of  H. cristata, “which seem re-conductible mostly to size difference”. Angelici et al.(2021)do not specify the exact origin of the African specimens analysed, however the populations north and south of the Sahara resulted not separable morphologically on the basis of their results. Moreover, data seem partial and not providing a complete overall vision of the range of  H. cristata, especially in the border areas between areas north and south of Sahara (e.g., western north Africa), which are data deficient and may have only recently been separated (see discussion below).  Although no material from Senegalis available also for us, this country falls within the range of  H. cristataand, 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. cristatafrom Eastern Africa, Western Africa, Libyaand Tunisiabelong 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 STvalue = 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. cristatahave 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. cristataand  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.    Acanthion daubentonii Cuvier, 1823  Cuvier (1823: 431)added a new species, providing a brief description on the basis of a skeleton previously described and figured by Louis-Jean-Marie Daubenton (L. Daubenton 1764: 413, pl. LIII), to whom he dedicated it. Daubenton referred to a specimen which was sent to him from Rome ( Italy) (L. Daubenton 1764: pl. LI; Buffon 1764: 407; the same specimen on which  H. cristata europaea Kerr, 1792was based: see discussion above), another from “Indes” [= Indies] (L. Daubenton 1764: pl. LII; Buffon 1764: 407), and to the aforementioned skeleton without giving additional information about its origin (L. Daubenton 1764: 413, pl. LIII).   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 specimensfrom Italyand 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, 1758by 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  Hystrixto 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.    Acanthion cuvieri Gray, 1847  Gray (1847: 102)described a new species based on the observation of a skull of an adult specimen preserved in “Museum of the Zoological Society” of London (today NHML), stating that it agrees with that figured by Brandt (1835, pl. VIII, Figs. 1, 2) and by Cuvier (1823, pl. I, fig. 1) on a specimen from Italy.  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. cristataLinneaus, 1758and 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. cristataremains the most plausible one.    Hystrix galeata Thomas, 1893  Thomas (1893: 230)described a new species from Kenya( Lamu) on the basis of a skull of a subadult specimen. He noticed “conspicuous differences between  H. galeataand its allies [i.e.,  H. cristataand  H. africaeaustralis] in the relative proportions of the two interorbital breadths, anterior and posterior, in the shape of the nasals, and in the height of the skull.  However, Corbet & Jones (1965)compared more data and found “mean differences in all these characters between the East and North African groups [of  Hystrixspp.]” and “in every case” they found “wide overlap” and “no reason to consider”  H. galeataspecifically 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 STvalue = 0.34-0.46: Trucchi 2008). Trucchi & Sbordoni (2009)showed that samples of  H. cristatafrom Eastern and Western Sub-Saharan Africa, as well some samples from Libyaand 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).    Hystrix galeata ambigua Lönnberg, 1908  Lönnberg (1908: 29)described a subspecies of  H. galeata Thomas, 1893from an adult old specimen (only the skull was presented and described) collected in Tanzania, on the slopes of Kilimanjaro(“Kibonoto” = Kibong’oto). He found features which agree with  H. galeata, others more with  H. africaeaustralisand “in still others was intermediate or differs from both”. However, he likely does not consider that the specimen in question was a hybrid, but he believed that it could be “a representative of an intermediate geographic race” and he assigned it to a new subspecies of  H. galeata. It is curious that from the same locality he listed other two specimenshe assigned to the “typical”  H. galeata. Subsequently, this taxon was synonymized with  H. cristataby Misonne (1974: 8), who included it under the Linnaeus’ species, but without further explanation.  It is difficult to pronounce in this case, since it could be assigned as a synonym of  H. cristataor  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 ambiguamight be related to potential hybrid specimens between  H. cristataand  H. africaeustralis( Trucchi & Sbordoni 2009; Mori et al. 2013; Trucchi et al. 2016).   Hystrix galeata conradsiF. Müller, 1910 Ferdinand Müller described this taxon as new subspecies of  H. galeata Thomas, 1893on 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 1910b: 314). This taxon was included under  H. cristataby Misonne (1974: 8), hence informally synonymizing it, but without further explanation. According to nuclear sequences (see Trucchi et al.2016) specimens from Tanzaniafall in the clade of  H. cristata. Therefore, we agree with Misonne (1974).   Hystrix galeata lademanniF. Müller, 1910  Müller (1910b: 314)described a further taxon as a new subspecies of  H. galeata Thomas, 1893on the basis of the same features examined for  H. galeata conradsiF. Müller, 1910. The name was first spelled “ H. galeata ludemanni”, then twice “  H. galeata lademanni”( Müller, 1910b: 314). However, also since it was clearly dedicated to Lademann ( Müller, 1910b: 314), first spelling can be considered an inadvertent error and must be corrected (ICZN 1999, Art. 32.3). This taxon, misspelled as “ ladermaniMüller, 1910”, was included under  H. cristataby Misonne (1974: 8), hence informally synonymized, but without further explanation. According to mtDNA and nuclear sequences ( Fig. 3), specimens from Tanzaniafall in the clade of  H. cristata. Also in this case, the differences highlighted (see Müller 1910b) seem to fall within population variability. Therefore, we agree with Misonne (1974).   Hystrix galeata loennbergiF. Müller, 1910  Müller (1910b: 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, 1908but 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. cristataby Misonne (1974: 8), hence informally synonymized, but without further explanation. Then, this taxon was indicated as synonym by subsequent author (e.g., Woods & Kilpatrick 2005as “ 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 Tanzaniafall in the clade of  H. cristata. Moreover, morphological differences between individuals of Tanzaniaand those from other African countries seem to fall within the intraspecific variability ( Müller 1910b). Therefore, we agree with Misonne (1974).    Hystrix galeata somalensis Lönnberg, 1912  Lönnberg (1912: 109)described a new subspecies of  H. galeata Thomas, 1893from Somaliaand Kenya(see list above for more information).  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., Tanzaniaand 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).    Hystrix occidanea Cabrera, 1924  Cabrera (1924: 220)described a new species from Morocco( Essaouira). He differentiated that specimen from  H. cristataand  H. senegalica(=  H. cristata) mainly on the basis of skull shape. The species was downgraded to subspecies of  H. cristataby Ellerman (1940: 219)and then synonymized (with doubt and without further explanations) by Ellerman et al.(1954: 520)with  H. cristata cristata Linnaeus, 1758.  Molecular analyses conducted both on mtDNA and nuclear genes ( Fig. 3), confirmed that populations from northern and western Moroccoare genetically close to the central and southern Italian ones ( typelocality of  H. cristatais Rome). The populations from Italywere 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.    Hystrix aerula Thomas, 1925  Thomas (1925: 196)described a new species on the basis of a skull of a young specimen from Nigerian Sahara. He compared this specimen with porcupines from Senegal(he called  H. senegalicaCuvier), finding differences in size (smaller than Senegalspecimens), while nasal and frontal bones show about the same proportions.   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).