The present study is based on the revision of key diagnostic features of the problematic species “Mustela” ardea Gervais, 1848–1852 in comparison to other mustelids of the subfamily Ictonychinae of Eurasia, particularly of those of the tribe Galictini (sensu Baskin, 1998). The examined and described fossils are housed in the Museum of Natural History, Geological and Palaeontological section, the University of Florence (IGF; see abbreviations below) and in the collections of the “Musée des Confluences”, in the collection of the “Université Claude-Bernard” (Lyon, France). As a comparative fossil sample, the collections of the IGF, “Museo Archeologico Nazionale”, Nuoro (Italy), “Museo di Geologia e Paleontologia” of the “Università degli Studi di Torino”, American Museum of Natural History (New York, United Stated of America) and Hungary Museum of Natural History (Budapest Hungary) were studied, and all the relevant literature on the Plio-Pleistocene Ictonychinae was inspected (Colombero et al., 2012, Fejfar et al., 2012, Ficcarelli and Torre, 1967, García and Howell, 2008, García et al., 2008, Geraads, 2016, Jin and Liu, 2009, Morlo and Kundrat, 2001, Ogino and Otsuka, 2008, Peters and de Vos, 2012, Qiu et al., 2004, Rook, 1995 and Viret, 1954). Extant specimens from the “La Specola” Zoology section, Museum of Natural History, University of Florence (Italy) were also used for morphological and morphometric comparisons. Craniodental measurements were taken to the nearest 0.1 mm with a digital caliper, following Driesch von den (1976) also for anatomical nomenclature.

The fossil comparative sample includes specimens of Pannonictis pliocaenica Kormos, 1931 from Villany-Kalkberg; Pannonictis pilgrimi Kormos, 1933 from Villany and Beremend; Pannonictis nestii (Martelli, 1906) from Upper Valdarno and Pirro Nord; Enhydrictis galictoides Major, 1901 from Monte S. Giovanni; and Pannonictis sp. and Enhydrictis sp. from Monte Tuttavista. The extant comparative sample includes specimens of Galictis cuja, G. vittata, Lutra lutra (Linnaeus, 1758), Lontra longicaudis (von Olfers, 1818), Lontra provocax (Thomas, 1908), Mustela putorius Linnaeus, 1758, Mustela nivalis Linnaeus, 1758, and Eira barbara housed in the MZUF.

A cladistic analysis was carried out based on fossil and extant genera of the Ictonychinae from Eurasia and the American continent. For this purpose, thirty cranial and dentognatic characters for nine genera were selected. The freeware software Mesquite version 3.31 (build 765) (Maddison and Maddison, 2017) was used to build the data matrix of these unpolarised, non-additive characters. The character descriptions and the resulting data matrix are provided in Appendix A and B of the Supplementary Material. The analysis used TNT version 1.5 (Golobov and Catalano, 2016) to provide a 50% major cutoff tree using the traditional search setting (2000 random addition sequences and the TBR algorithm). Subsequently, Mesquite 3.31 (built 765) was used to manage the tree resulting from the analysis and to export it in a graphic format.

AMNH, American Museum of Natural History, New York, United States of America; HMHN, Hungarian Museum of Natural History, Budapest, Hungary; IGF, Museum of Natural History, Geological and Palaeontological section, the University of Florence (Italy); MAN, Museo Archeologico Nazionale, Nuoro (Italy); MNHL, Musée des Confluences, Lyon (France); MZUF, Museum of Natural History, “La Specola” Zoology section, University of Florence (Italy).

The species Martellictis ardea has been extensively described by other authors (among others, Schaub, 1949 and Viret, 1954). Here, we only provide a revised punctual comparison with other Plio-Pleistocene Eurasian species of the tribe Galictini. In size, Martellictis ardea is similar to Pannonictis nestii from Pietrafitta, P. pilgrimi from Villany and Beremend, E. galictoides from Monte San Giovanni and Enhydrictis sp. of Monte Tuttavista (Abbazzi et al., 2004, Ficcarelli and Torre, 1967 and García and Howell, 2008). Compared to these latter species, the postorbital constriction of M. ardea is less marked, although more prominent than in P. pliocaenica from Villany-Kalkberg and P. cf. nestii from Atapuerca TE (García and Howell, 2008) or Eirictis (Qiu et al., 2004). The morphology of the tympanic bullae differs from the medial inflation of P. pliocaenica and P. pilgrimi from Hungary and especially from their sharp and marked notch on the mesial side the bullae of this species. It possesses a button-like P1 placed close to the C, as in Eirictis robusta Qiu et al., 2004 from Longdan and in some specimens of P. pilgrimi and P. nestii (Rook, 1995), whereas it is lacking in Enhydrictis spp. (Ficcarelli and Torre, 1967) and only rarely present in P. pliocaenica from Villany-Kalkberg. In contrast to E. galictoides, the mesial margin of P4 is lobed, as in some species of Pannonictis (e.g., P. nestii, P. pilgrimi or P. pliocaenica). The P4 protocone and the lingual cingulum that departs from it are reduced in comparison to those seen in P. nestii of Pirro Nord. Moreover, the protocone area is cup-shaped as that of Enhydrictis, Oriensictis and Pannonictis, unlike that of Eirictis, which possesses a prominent conical cusp. The P4 hypocone, although present, is not as developed as it is in Oriensictis nipponica Ogino and Otsuka, 2008 from Matsugae cave (Ogino and Otsuka, 2008) or P. pliocaenica and P. pilgrimi from Hungary. It resembles more closely the condition visible in E. galictoides from Sardinia (Ficcarelli and Torre, 1967). The M1 shows an enlarged lobed talon, distally directed, unlike P. pliocaenica and similar to P. nestii, P. pilgrimi and E. galictoides to some extent, although less large than in the latter species. Unlike in E. galictoides, a narrowing occurs between the trigon cusps and the talon of M1, as can be found in Pannonictis species. Furthermore, the M1 in Martellictis ardea shows the buccal side that is smaller than the lingual one, in a condition similar to P. nestii from Pietrafitta and unlike Enhydrictis spp. or P. pliocaenica. The mandible corpus of Martellictis ardea from Olivola and St. Vallier is shallow, slender and it is rather shortened rostrocaudally if compared to other Galictini, e.g., most of all Eirictis spp. from China but also P. pliocaenica Villany-Kalkberg, P. pilgrimi from Beremend, P. nestii Upper Valdarno and Enhydrictis spp. from Sardinia. The lower carnassial of Martellictis ardea possesses a short and enlarged paraconid and a large metaconid that ends posteriorly compared to the distal wall of the protoconid. The talonid has a large hypoconid, larger than that of O. nipponica from Matsugae cave, E. robusta from Longdan and Eirictis variabilis Jin and Liu, 2009 from Renzidong and E. galictoides from Monte San Giovanni. It is individualised by a low furrow from a distal accessory cuspulid unlike Eirictis, Enhydrictis or Oriensicitis. This distal accessory cuspulid is followed lingually by a high distal cristid, which borders the distal and distolingual portion of the talonid. The distal cristid is not so elevated in P. pliocaenica from Villany-Kalkberg, but unworn specimens of P. nestii from Pirro Nord and Upper Valdarno and of P. pilgrimi from Beremend also show an accessory cuspulid distal to the hypoconid.

The analysis of the anatomical features of the highly disputed species “Mustela” ardea Gervais, 1848–1852 has revealed numerous cranial and dentognatic characteristics that, altogether, do not fit with the diagnosis of the Galictini genera (sensu Baskin, 2011) described in literature, namely in comparison to Enhydrictis Forsyth Major, 1901, Eirictis Qiu et al., 2004, Oriensictis Ogino and Otsuka, 2008, Pannonictis Kormos, 1931. Among these different morphologies there is the degree of embayment of the mesial side and the medial inflation of the tympanic bullae, the prominence of the postorbital constriction, morphology of the braincase, the cup-like P4 protocone, the P4 hypoconid; the development and morphology of the cuspids on M1 and the outline and cuspulids of m1 in occlusal view. From the morphological analysis, it is clear that some traits of M. ardea recall those of Pannonictis spp. but other those of Enhydrictis spp.: this is the reason why many scholars assigned the taxon to the former genus (Fejfar et al., 2012, Rabeder, 1976 and Schaub, 1949) whereas others to the latter one (Ficcarelli and Torre, 1967, Rook, 1995 and Viret, 1954). Nevertheless, neither attribution accords with the set of features possessed by Martellictis ardea. This is particularly true for the cranial ones, e.g., the morphology of the tympanic bullae, which in Carnivora is among the most systematically relevant characteristics (as noted by Fejfar et al., 2012).

The genus Pannonictis is considerably widespread across Eurasia (Fig. 2), with a broad geographic range spanning from Spain (García and Howell, 2008 and Madurell-Malapeira et al., 2014) to Great Britain (Stuart, 1982), southeastern Russia and Moldova (Sotnikova et al., 2002), China and Mongolia (Sotnikova, 1980). Fig. 2 resumes the most important and verified localities that record Pannonictis (as well as of the other palearctic genera Eirictis, Enhydrictis, Oriensictis). Its earliest record is most probably the one reported by Zdansky (1927) as Mustelidae gen. indet. sp. indet. from the late Miocene Hipparion beds from China (as pointed out by numerous authors, e.g., García and Howell, 2008, Kormos, 1931, Pilgrim, 1932, Rook, 1995 and Schreuder, 1935). Therefore, it could be possibly referred to as Pannonictis sp. indet. Zdansky (1927) (n.b. not as P. pachygnatha as García and Howell, 2008 suggested).

One of the most extensive records of Galictini of the Asian Pliocene is that the Yushe Basin (Teilhard De Chardin and Leroy, 1945) in China, that of Shamar in Mongolia (Sotnikova, 1980) and that of the early Pleistocene Nihewan Basin (Teilhard De Chardin and Piveteau, 1930). These fossils were ascribed to the large-sized species Pannonictis pachygnatha. Qiu et al. (2004) described the material from the early Pleistocene of Longdan (Dongxiang, Gansu, China) and erected a different genus, Eirictis Qiu et al., 2004, that included the “P.” pachygnatha of Nihewan and a new species from Longdan, E. robusta. García and Howell (2008) recognised the validity of the diagnostic features used by the authors to discriminate this new genus from Pannonictis (especially the absence of a hypocone on P4). More recently, Jin and Liu (2009) included the taxon from the early Pleistocene site Renzidong Cave (Eastern China) in this genus, ascribing the sample to a new species, E. variabilis. Another Asian taxon of Galictini comes from the middle Pleistocene Matsugae cave deposits (northern Kyushu) of West Japan, the genus Oriensictis with the species O. nipponica (Naora, 1968). These authors propose the inclusion of the species Lutra melina Pei, 1934 from Zhoukoudian 1 (middle Pleistocene, China) in the genus Oriensictis.

Of the numerous genera of Ictonychinae of North America, the one most probably related to Pannonictis is the early Pleistocene Trigonictis (Rook, 1995). Numerous authors have proposed synonymising these two genera (e.g., Kurtén and Anderson, 1980 and Repenning, 1967), but the presence of primitive features and some characters (e.g., the morphology of the M1 and the development of its cusps), which cannot be found in either Pannonictis or in Enhydrictis, would suggest maintaining Trigonictis as a separate genus.

Among the earliest European representatives of Pannonictis is the record from the early Pliocene (MN 15) of Wölfersheim (Germany) (Fig. 2). From this site, Morlo and Kundrat (2001) described two dental specimens of Pannonictis sp. and few others of E. ardea. As the authors pointed out, the material is considerably old for the Eurasian Galictini record, probably the earliest record of Pannonictis-like mustelids in Europe. The partial right P4 (SMF 2000/212) and the right m2 (SMF 2000/213) seems reasonably ascribed to the genus Pannonictis, but the material of Enhydrictis possesses several features, which significantly contrast with the type specimen of the genus. The assumption of a primitive state of the taxon for its old age cannot compensate the “lack of some typical features” (Morlo and Kundrat, 2001: 170) or the prominence of other characters. This eventually casts doubts on the generic and specific attribution of the material, even if the specimens are close in size to M. ardea from St. Vallier. Among the most evident discrepancies, there is the high conical cuspid on the P4 protocone area, a feature absent in the majority of the Galictini of Eurasia, as they generally possess a cup-like concavity bounded by a swelling cingulum. Nevertheless, a similar condition is found in Eirictis, which, according to Qiu et al. (2004), possesses a conical cusp in the protocone area. Furthermore, the lack of a hypocone on the lingual side of the P4 as in the Wölfersheim specimen is another diagnostic feature of Eirictis (García and Howell, 2008 and Qiu et al., 2004). The morphology of the right M1 SFM2000/217 from Wölfersheim differs from that of other Galictini taxa, as it is proportionately shortened buccolingually, with an enlarged buccal cingulum, expanded buccally and undivided between the paracone and the metacone. In contrast, the species of Pannonictis, Enhydrictis, Martellictis, Eirictis and Oriensictis have the cingulum divided into two lobes, a larger one for the paracone and the smaller for the metacone (Abbazzi et al., 2004, Ficcarelli and Torre, 1967, García and Howell, 2008, Ogino and Otsuka, 2008 and Qiu et al., 2004). As in Enhydrictis from Monte San Giovanni and Monte Tuttavista, the M1 is not so curved distally, in occlusal view. Nevertheless, when compared closely to the M1 of Enhydrictis spp., the M1 metacone of SMF 2000/217 is more developed than that in the Sardinian taxa. The development of the cusp is more similar to that of some Pannonictis species (e.g., P. pilgrimi and P. pliocaenica from Hungarian localities) or to M. ardea from St. Vallier. Furthermore, the hypocone is missing or greatly reduced in the German specimen, whereas it is generally enlarged in Pannonictis and Eirictis. The area of the hypocone lobe differs greatly even from M. ardea from St. Vallier as the latter shows a prominent distal extension of the M1, giving the tooth a peculiar morphology in occlusal view. The upper and lower canines (respectively, SMF 2000/214 and SMF 2000/215) are short mesiodistally. The left m1 SMF 2000/218, and particularly its paraconid, is shorter mesiodistally compared to that of Enhydrictis from Monte San Giovanni and Monte Tuttavista (Ficcarelli and Torre, 1967) and Eirictis robusta from Longdan (Qiu et al., 2004), but similar to the m1 of the holotype of M. ardea from Côte de Ardé. In occlusal view, the metaconid is more developed and stouter than in Enhydrictis and Eirictis spp. but not as individualized as in Pannonictis and Martellictis. Proportionately, the metaconid is more developed than in Eirictis from Longdan (Qiu et al., 2004). Among the Galictini of Eurasia, Oriensictis has the largest and the most individualised m1 metaconid (Ogino and Otsuka, 2008). In occlusal view, the m1 talonid of SMF 2000/218 shows a round outline of the cristid starting from the hypoconid and reaching the correspondent point on the lingual side, similar to some species of Pannonictis (e.g., P. nestii from the Upper Valdarno and Pirro Nord) or Oriensictis from Matsugae Cave (Ogino and Otsuka, 2008). Enhydrictis, Eirictis and Martellictis have more elongated m1 talonids. Considering all these features, the attribution to the genus Enhydrictis or the species M. ardea are rather unlikely. Some features are suggestive of an affinity with the genus Eirictis (e.g., the P4 protocone, the absence of a P4 hypocone) even if the specimens are considerably smaller and thinner than the average Eirictis individuals are. Morlo and Kundrat (2001) also point out some similarities with the taxon Martes wenzensis Stach, 1959 (e.g., in the “crenulation” of the enamel of the hypocone lobe of the m1). In conclusion, the record from Wölfersheim reports possibly the earliest European specimens of a member of the genus Pannonictis and a few other specimens of an undetermined taxon not related to the genus Enhydrictis nor with the species M. ardea. Future revision and additional cranial and dentognatic material could help clear out these doubts on Eirictis. sp. indet.

Next to the record from Wölfersheim, the early Pleistocene species P. nestii, from the Upper Valdarno in Tuscany (Martelli, 1906) and the younger site of Pietrafitta in Umbria (Rook, 1995) is among the first Pannonictis species to be described. Several specimens from Sima del Elefante (Sierra de Atapuerca) have been referred to Pannonictis cf. nestii by García and Howell (2008). García et al. (2008) grouped P. nestii with “E.” ardea, with no discussion of the morphological grounds for this placement. On this matter, other scholars disagree (see, among others, Ficcarelli and Torre, 1967, Rook, 1995, Sotnikova et al., 2002 and Viret, 1954). Another record is that of P. pliocaenica from the complex of cavities of Villany-Kalkberg, Hungary. In particular, two of the richest localities for Galictini remains are Villany 3 and 5, which span between 2 and 1.5 Ma (Jánossy, 1986 and Spassov, 2000). From these localities, Kormos (1933) re-described some small specimens as P. pilgrimi. In the scientific literature, various synonyms have been provided for this taxon: in addition to those reported above regarding the disputed synonymy to “E.” ardea (Viret, 1954), Ficcarelli and Torre (1967) suggested uniting this species with P. nestii rather than with “E.” ardea. By contrast, García and Howell (2008) proposed the inclusion of P. pilgrimi within P. pliocaenica, addressing the issue of the large size difference as a strong sexual dimorphism (where P. pliocaenica would represent the larger male individuals, whereas P. pilgrimi would represent the smaller females). The genus Pannonictis has also been recovered from Mediterranean islands: Pannonictis arzilla (De Gregorio, 1886) from Sicily, in the endemic fauna of the Monte Pellegrino, a taxon probably close to P. nestii (Burgio and Fiore, 1997 and Rook, 1995); and Pannonictis sp. from the early-middle Pleistocene levels of Monte Tuttavista (Nuoro, Sardinia; Abbazzi et al., 2004). Recently, Peters and de Vos (2012) described some dentognatic remains from the Dutch locality of Langenboom (Noord-Brabant) attributing them to “Pannonictis” ardea. The specimens show several peculiarities worth noting, as described by Peters and de Vos (2012). For instance in the upper teeth, the presence of a cusp surrounded by a marked cingulum in the protocone area on the P4, the P4 hypocone absent or greatly reduced, the buccolingually large and slightly distally curved M1 in the talon area, the expanded and continuous buccal cingulum on the M1 paracone and metacone, the large M1 protocone and protoconule but no metaconuleand absence of the M1 hypocone. On the lower teeth, the p3 and p4 are strongly elongated mesiodistally in occlusal view, the p4 possesses a prominent distal accessory cuspulid, visible in buccal, lingual, and occlusal views. Furthermore, the m1 has a mesiodistally elongated paraconid, it possesses an enlarged and strongly individualized metaconid and a round talonid. Many of these features are similar to those of the fossils from Wölfersheim (Morlo and Kundrat, 2001; discussed above), although in size the specimens from Langenboom are larger compared to those from the German sample. As for the German sample, the attribution to the genus Pannonictis and to the species M. ardea seems unlikely. As the Langenboom fauna resembles that of St. Vallier and Tegelen (Peters and de Vos, 2012), the affinity between the German and the Dutch finds would suggest the occurrence of a previously unknown form of mustelid, possibly related to Eirictis, in the Pliocene-earliest Pleistocene of central-western Europe,.

The most peculiar taxon within the European record of Galictini is Enhydrictis galictoides, described by Forsyth Major (1901) from the late Pleistocene Sardinian locality of Monte San Giovanni (Ficcarelli and Torre, 1967) (Fig. 2). In the fossil record, the genus Enhydrictis is more elusive than Pannonictis and, apart from E. galictoides, is only known from few other Sardinian and Corsican localities (e.g., the older Enhydrictis sp., from the fissure filling of Monte Tuttavista, Abbazzi et al., 2004). The peculiar features that typify this genus (e.g., strong postorbital constriction, shorter muzzle, etc.) have been regarded by many authors as adaptations to an aquatic lifestyle (Ficcarelli and Torre, 1967, Forsyth Major, 1901 and Pilgrim, 1932).

The first fossils of a taxon of the tribe Galictini from northern Africa were recently reported by Geraads (2016). The author described the sample from Tighennif (or Ternifine, Algeria; Fig. 2) and ascribed the scarce sample to the new species Enhydrictis hoffstetteri Geraads, 2016. This discovery greatly expands our knowledge and the areal of Plio-Pleistocene Galictini, up to then unknown in the African continent. Unfortunately, no complete cranial or mandibular specimens have been found other than these three, so it is difficult to precisely ascribe the material to a species because the major diagnostic characters are missing (e.g., the postorbital constriction, the height of the cranium, the shape of the braincase, the presence and/or the development of hypocone on the P4 and features of the m1 talonid; Table 1). Nevertheless, there are several features displayed by the holotype and the referred specimens (Fig. 1 in Geraads, 2016: page 447) that cast doubts on the attribution of the Algerian material to the genus Enhydrictis. Among these, we can briefly discuss here the morphology of M1 that tends to be distally curved and stretched and possesses a thick talon, enlarged compared to the trigon cusps. In Enhydrictis, the M1 is not curved distally in occlusal view. Moreover, the M1 metacone of E. hoffstetteri is smaller than the paracone but still prominent and it is evidently bounded by a large and round cingulum especially on the distal side, unlike the specimens of Enhydrictis from Monte S. Giovanni or Monte Tuttavista in which the metacone is greatly reduced and does not possess a prominent cingulum around it (Abbazzi et al., 2004 and Ficcarelli and Torre, 1967). The positions of the enlarged M1 protocone and protoconule differ from the condition visible in Enhydrictis spp. because in the specimens from Tighennif they lie in the centre of the tooth, whereas in latter taxa they are displaced towards the lingual side of the M1, very close to the hypocone. The talon tends to narrow towards the hypocone area, unlike the condition seen in M. ardea. The features of the M1 of Tighennif taxon are similar to those retained by some species of Pannonictis (e.g., P. pliocaenica from Villany). The considerable height and breadth of the mandible corpus of TER-2008 contrast with the slender and shallow mandibles of M. ardea and resemble those of robust forms like Pannonictis pliocaenica or even Eirictis, even though it is hard to say as the author did not include a table of measurements of the specimen with mandibular measures. In the lower teeth, the prominent and high wrinkled pattern of the basal cingulid of the canine is similar to that shown by some species of Pannonictis (e.g., P. pliocaenica from Villany or P. cf. nestii from Atapuerca, García and Howell, 2008). Furthermore, the stout and shortened m1, especially considering the talonid, and the enlarged metaconid extended lingually differ from the principal features evident in specimens of Enhydrictis spp. (e.g., large metaconid but close to the protoconid). By contrast, M. ardea possesses a buccolingually wider m1, with a shorted and stouter paraconid, compared to that of TER-2008. Altogether, these morphologies are suggestive of a closer affinity between the taxon from Tighennif (Ternifine) and the genus Pannonictis rather than to Enhydrictis. Probably the species “E.” hoffstetteri should be referred to as Pannonictis hoffstetteri. The misinterpretation might have arisen as a consequence of the underestimation of the diversity of Plio-Pleistocene Galictini of Eurasia as the Geraads (2016) suggests to include the cluster of all the genera in Enhydrictis sensu lato, which seems cumbersome considering the numerous differences between these taxa (for further discussion on the variability of Galictini see also Colombero et al., 2012 and García and Howell, 2008).

As shown in Table 1, the peculiar morphological pattern of the cranial and dentognatic features of M. ardea differ quite prominently from those of other Eurasian taxa of Galictini, thereby supporting the recognition of a new genus. Furthermore, this has important implications in the complex taxonomic dispute as it forces a reconsideration of the Plio-Pleistocene palaeodiversity of Galictini and their morphological affinities and relationships.

The cladistic analysis was carried out based on the character matrix for fossil and extant genera of Eurasian and American Galictini, as described in the Appendix B of the Supplementary material. The genera analysed were selected among the holoarctic fossil members of the tribe Galictini based on the completeness of their fossil record and the number of characters effectively recognizable in the type specimens of these taxa. According to recent molecular phylogenies (e.g., Sato, 2016 and Sato et al., 2012) the two closest subfamilies to Ictonychinae are Lutrinae and Mustelinae. As the members of the former subfamily possess several marked adaptations to an aquatic lifestyle, the genus chosen as outgroup in the performed cladistic analysis is Mustela Linnaeus, 1758.

The analysis used TNT ver. 1.5 to provide a 50% major cutoff tree (Fig. 3) based on the six shortest trees (Appendix C of the Supplementary material). This first attempt to investigate the phylogeny of fossil Eurasian Galictini at a generic level, produced relatively well resolved tree, with a consistency index of 0.645 and a retention one of 0.352. As is visible in Fig. 3, there are different degrees of possible affinities previously underestimated in scientific literature. Lyncodon and Oriensictis equally differ from the two clustered groups, i.e. Pannonictis-Eirictis and Enhydrictis-Trigonictis. The displayed arrangement – with the marked distinction of the Eurasian “pannonictine” group (i.e. Pannonictis-Eirictis) from Martellictis, on the one side, and from Enhydrictis and the American Trigonictis, on the other – indeed testifies to previously unnoticed relationships in the tribe Galictini. The resulting tree emphasizes the peculiarly of the set of morphological features of Martellictis for the position of this genus in the phylogenetic tree (Fig. 3): together with the South American Galictis, it is the sister taxon of all the other Eurasian genera of Galictini and the American Lyncodon and Trigonictis. In this configuration, the affinity of Martellictis to Pannonictis or to Enhydrictis is not as close as expected from literature (sections 1.1 The intricate “Mustela” ardea issue in scientific literature and 4.2 A review of the diversity of the Plio-Pleistocene Galictini) and Martellictis possesses a set of features suggestive of a probably more primitive state among Galictini. This supports the distinction of these taxa into different genera. Fig. 3 also points out the difference between the endemic genus Enhydrictis and the European taxa of Galictini, suggesting that its evolution was limited to the insular context of Sardinia and Corsica (Fig. 2) from mainland ancestors like Martellictis or Pannonictis (the latter is notably present in the record of Sardinia, i.e. Monte Tuttavista see Abbazzi et al., 2004). The schematic representation in Fig. 4, relates the phylogenetic relationship between the Galictini genera considered in the cladistics analysis here performed and the fossil and extant distribution of these taxa.