Hystrix cristata Linnaeus, 1758

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), pp. 501-519 : 505-510

Hystrix cristata Linnaeus, 1758

Hystrix cristata Linnaeus, 1758: 56 . Type /s: HT: specimen mentioned by Rey (1693: 206); Type locality: “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). Type locality: 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). Type locality: 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, 1823 by Maguire (1976: 47), although it is an unofficial nomenclatural act since included in a thesis dissertation. Revalidated as subspecies of H. cristata by 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). Type locality: unknown. Repository: unknown. Syn. Ellerman (1940: 219).

Acanthion Cuvieri Gray, 1847: 102 . Type /s: HT: adult specimen (skull). Type locality: unknown. Repository: NHML. Syn. P. Sclater (1865: 356).

Hystrix galeata Thomas, 1893: 230 . Type /s: HT: subadult specimen (skull). Type locality: Kenya, Lamu Island, 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). Type locality: Tanzania, Kilimanjaro, Kibong’oto – “[ Deutsch-Ostafrika, Kilimandjaro:] Kibonoto”. Repository: unknown. Syn. Misonne (1974: 8).

Hystrix galeata conradsi F. Müller, 1910b: 314 . Type /s: STS: probably two specimens but not specified (skulls). Type locality: Tanzania, Mwanza and “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 lademanni F. 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 specimens but not specified (skulls). Type locality: 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 “ ladermani Müller, 1910”).

Hystrix galeata lönnbergi F. Müller, 1910b: 315 . Type /s: HT: skull given to the SBM by von der Marwitz. Type locality: Tanzania, Kilimanjaro – “[ Deutsch-Ostafrika,] Kilimandjaro ”. Repository: ZMB. Syn. Misonne (1974: 8).

Hystrix galeata somalensis Lönnberg, 1912: 109 . Type /s: STS: 1 male and 1 female, old specimens (skulls from Burao [ Somalia]; NHML), 1 adult female specimen (Nyoro [ Kenya]; not specified if preserved; possible repository unknown). Type locality: 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. Type locality: 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. cristata by Misonne (1974: 8).

Hystrix aerula Thomas, 1925: 196 . Type /s: HT: young specimen (skull only). Type locality: Niger, Aouderas (Asben [Aïr]) – “Aouderas, Asben”. Repository: NHML. Syn. Misonne (1974: 8).

Hystrix cristata Linnaeus, 1758

Carl Linnaeus in the 10 th edition of the Systema Naturae ( Linnaeus 1758), after the description of H. cristata , refers to the 9 th edition of his work ( Linnaeus 1756). Although, the latter was published before the starting point of zoological nomenclature (1 st January 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. 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).

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. cristata by 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 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.

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, 1792 was 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 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.

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. 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.

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. galeata and its allies [i.e., H. cristata and 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 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).

Hystrix galeata ambigua Lönnberg, 1908

Lönnberg (1908: 29) described a subspecies of H. galeata Thomas, 1893 from 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. africaeaustralis and “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 specimens he assigned to the “typical” H. galeata . Subsequently, this taxon was synonymized with H. cristata by 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. 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 1910b: 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 (1910b: 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, 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 “ 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 1910b) seem to fall within population variability. Therefore, we agree with Misonne (1974).

Hystrix galeata loennbergi F. 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, 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 1910b). Therefore, we agree with Misonne (1974).

Hystrix galeata somalensis Lönnberg, 1912

Lönnberg (1912: 109) described a new subspecies of H. galeata Thomas, 1893 from Somalia and 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., 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).

Hystrix occidanea Cabrera, 1924

Cabrera (1924: 220) described a new species from Morocco ( Essaouira). He differentiated that specimen from H. cristata and H. senegalica (= H. cristata ) mainly on the basis of skull shape. The species was downgraded to subspecies of H. cristata by 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 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 .

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. senegalica Cuvier ), finding differences in size (smaller than Senegal specimens), 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).

Figures

FIGURE 1. Distribution of the subgenus Hystrix: 1, Hystrix cristata; 2, Hystrix indica; 3, Hystrix africaeaustralis (map based on Mori et al. 2019 and IUCN data, updated with further publications: Altorabi 2018; Vale et al. 2016; Bauer et al. 2018). Published distribution data were also integrated with occurrences uploaded on online platforms, i.e., iNaturalist (data checked on 08.03.2021). Dotted areas indicate areas where no record of H. cristata after 2000 is available, question marks indicate nonsampled areas after 2000.

FIGURE 2. Presumed distribution of the subspecies of African Hystrix porcupines according to Mohr (1965).

FIGURE 3. Maximum-likelihood trees on mtDNA sequences of (a) cytochrome-b and (b) D-loop of Hystrix cristata and Hystrix africaeaustralis (from Trucchi & Sbordoni 2009). Numbers at main nodes indicate bootstrap values. (c) Maximumlikelihood tree of genome-wide RAD nuclear markers (taken from Trucchi et al. 2016, readapted). Available bootstrap values are shown. Red colour refers to H. cristata, grey to H. africaeaustralis; a sequence of H. indica was included as outgroup for mtDNA trees.