Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) vol. 126(1): 157-187. March 2020 FOSSIL LAGOMORPHA (MAMMALIA) OF ITALY: SYSTEMATICS AND BIOCHRONOLOGY CHIARA ANGELONE1,2,3 *, BLANCA MONCUNILL-SOLÉ1,4 & TASSOS KOTSAKIS1 1* Corresponding author. Dipartimento di Scienze, Università Roma Tre, Largo San Leonardo Murialdo 1, 00146 Roma (Italy). E-mail: angelone@uniroma3.it 2 Institute of Vertebrate Palaeontology and Palaeoanthropology, Chinese Academy of Sciences, 142 Xi Zhi Men Wai Da Jie, 100044 Beijing (China). 3 Institut Català de Paleontologia Miquel Crusafont, Edifici Z ICTA−ICP, Carrer de les Columnes s/n, Campus de la Universitat Autònoma de Barcelona, 08193 Cerdanyola del Vallès, Barcelona (Spain). 4 Centro de Investigacións Científicas Avanzadas (CICA), Facultade de Ciencias, Universidade da Coruña, As Carballeiras s/n, Campus de El- viña, 15071 A Coruña (Spain). To cite this article: Angelone C., Moncunill-Solé B. & Kotsakis T. (2020) - Fossil Lagomorpha (Mammalia) of Italy: systematics and biochrono- logy. Riv. It. Paleontol. Strat., 126(1): 157-187. Keywords: Stem lagomorphs; Ochotonidae; Leporidae; Neogene; Quaternary; endemism. Abstract. Revisions performed in the last 15 years added remarkable novelties to the taxonomy and biochronology of Italian fossil lagomorphs. Several new taxa have been erected basing on new materials and on the revision of old materials. This paper aims to illustrate the state of the art of such researches. The lagomorph diversity in the Italian Neogene and Pleistocene is quite high, accounting 9 ochotonids, 14 leporids, and 3 stem lagomorphs. Among the lagomorph taxa recorded in Italy, quite a high number are insular or continental endemics. The oldest Italian lagomorphs are the insular endemic Paludotona aff. minor, P. etruria and P. minor from the early-middle Turolian of the Tusco-Sardinia palaeobioprovince, and Prolagus apricenicus and P. imperialis from the late Turolian of the Abruzzi-Apulia palaeobioprovince. In the Italian peninsula, lagomorphs are known since the late Turolian (early Messinian). Prolagus appears in pre-evaporitic Messinian (Prolagus sorbinii) and gives rise to species endemic to the Italian peninsula (Prolagus italicus, late Villányian, central Italy; P. savagei/P. aff. sorbinii, early Villányian-early Biharian, NW and central Italy). The presence of Prolagus was probably constant until the early Biharian, age of its latest report in the Italian peninsula. The gap in the Pliocene fossil record is apparent and due to the extreme scarcity of fossil localities bearing continental vertebrates in the Italian Pliocene. The Asian genus Ochotona appeared at least twice in the Middle-Late Pleistocene of NE Italy, in correspondence of cold climatic phases. Prolagus of the latest Miocene-Early Pliocene of Italy represents the stock which originated the Sar- dinian lineage of Prolagus. The Sardinian species of Prolagus form an endemic insular anagenetic lineage whose oldest element is the earliest Late Pliocene P. aff. figaro, and which persists till historical times. The main turnover in the lineage, i.e. the transition P. figaro-P. sardus, is an event possibly related to the Mid-Pleistocene Transition. Minor morphological and dimensional changes can be detected among the population of both P. figaro and P. sardus, allowing a relative chronological arrangement of the Prolagus-bearing fossil sites. The first Italian leporids appear in the late Turolian. The continental endemic Alilepus meini is limited to the pre-evaporitic Messinian of Tuscany, and Trischizolagus sp., a taxon that still lacks a detailed systematic revision, is known from a couple of evaporitic and post-evaporitic Messinian fossil localities of northern-central Italy. There are no leporids in the Italian fossil record between the earliest Pliocene and the Late Pliocene (early Villányian), i.e. before the oldest Italian record of Hypolagus. This genus is represented by scanty remains pertaining to different species, and is distributed all along the peninsula (H. petenyii, early Villányian, NW Italy; H. brachygnathus, earliest Biharian, NE Italy; the continental endemic H. aff. brachygnathus, early Biharian, SE Italy). In Sicily, Hypolagus is present with an endemic insular species (H. peregrinus, ?late early Biharian). Also Sardinia records an endemic insular leporid, Sardolagus obscurus (Early Pleistocene). Its morphological features underwent significant modifications due to the permanence in an insular environment, preventing for the moment reliable phylogenetic inferences aimed to link it to known European continental genera. Extant lagomorph genera arrive in Italy with some delay with respect to the rest of Europe. In Europe, Oryctolagus is known since the Late Pliocene, whereas the oldest Italian species, O. valdarnensis, is known from the Early Pleistocene (late Villányian-early Biharian) of central and probably SE Italy. After a gap in the late Biharian, Oryctolagus is recorded in the earliest Toringian of NW and western-central Italy with O. burgi. The extant species O. cuniculus appears in the Italian peninsula in the late Middle Pleistocene. Its presence is attested until MIS 3 in several Italian fossil sites, and in the MIS 2 and early Holocene in southernmost Italy, Sicily, and southern Italian minor islands. It can not be excluded that those relict populations interacted with the rabbits introduced by humans during the Holocene. As for Lepus, the oldest reliable Italian report dates back to the late early Biharian of central Italy (originally classified as L. terraerubrae). Extant species of Lepus appear in Italy since the early Toringian. Among them, Lepus corsicanus has the oldest record; this species expands its geographical distribution to Sicily in the latest Pleistocene. Follows L. europaeus, whose oldest reliable Italian record is in the earliest late Toringian; this species expanded in the Italian peninsula at the expenses of L. corsicanus. During the Last Glacial Maximum, L. timidus makes sporadic appearance(s) in Italy, even in southern regions. Though showing endemic characters, the Sardinian hare L. mediterraneus is the result of human introduction in historical times. Received: May 10, 2019; accepted: October 31, 2019 158 Angelone C., Moncunill-Solé B. & Kotsakis T. Introduction In the text, we used the following abbreviations: BV0, BV1, BV2, BV3: Baccinello local biochron V0, V1, V2, V3 FC, FsC: Faunal Complex, Faunal sub-Complex Lagomorphs represent a very important, di- FU: Faunal Unit verse, and abundant component of European small IsIPU: Istituto Italiano di Paleontologia Umana mammal assemblages since Miocene on. In the last NHMB: Naturhistorisches Museum Basel PB: palaeobioprovince two decades, a renewed interest in European fos- sil lagomorphs led to the revision of several Italian lagomorph taxa (Angelone 2007, 2008a, 2013; An- Taxonomic overview of fossil gelone & Rook 2011, 2012; Angelone et al. 2008, lagomorphs of Italian mainland 2015, 2017, 2018 among others). We provide here: (1) an update of the systematics of Italian lago- Early-middle Turolian (MN11-12, Late morphs since their first appearance in the Italian Miocene) territory (Late Miocene) to the Holocene; (2) their Paludotona Dawson, 1959 phylogenetic affinities, when possible; (3) a synthe- The endemic insular genus Paludotona is the sis of the biochronological history of Italian lag- oldest known Italian lagomorph. Paludotona was omorphs of each different bioprovinces, starting considered an ochotonid in older literature, but re- from the early-middle Turolian; and finally (4) we cent data suggest instead its relationship with stem pinpoint the research perspectives and the open lagomorphs, the basal group which includes primi- questions emerged after this review. tive lagomorphs (Angelone et al. 2017). Paludotona etruria Dawson, 1959 (Fig. 1.a) was Material and methods reported from Tuscany (at that time part of the in- sular domain known as the Tusco-Sardinia PB), in The sources of the data here illustrated are published refer- horizons referable to BV1 (MN11). Another spe- ences or unpublished material currently under study. An iconograph- cies of the genus, Paludotona minor Angelone, Čer- ic overview of the Italian lagomorph taxa is given in Figs. 1-2. mák & Rook, 2017 (Fig. 1.b) was recognized in BV2 A list of late Turolian-Biharian fossil lagomorphs of Italian mainland, supplied with taxonomic details, geographical distribution, (MN12, and possibly earliest MN13, magnetostrati- age, and references is given in Tab. 1. For Toringian, we selected graphically correlated to ~7.1-6.7 Ma; Rook et al. among the numerous reports available in literature, those publica- 2011; Benvenuti et al. 2015). The scanty remains tions in which there is at least a description/discussion of the lago- morph taxa in the text. As for insular assemblages, the list of fossil of Paludotona recently recovered from BV0 (MN11, lagomorphs retrieved in insular endemic domains which are today magnetostratigraphically correlated to ~8.3-8.1 Ma; part of mainland Italy (the Tusco-Sardinia and the Abruzzi-Apulia Rook et al. 2011; Benvenuti et al. 2015) were clas- palaeobioprovinces; Kotsakis et al. 1997; Rook et al. 2006) is report- sified as Paludotona sp. by Cirilli et al. (2016). Ange- ed in Tab. 1, whereas fossil lagomorphs of Italian islands are listed in Tab. 2. lone et al. (2017) noticed that Paludotona from BV0 The taxonomy of lagomorph remains recorded in insular appeared morphologically similar to P. minor from palaeobioprovinces is illustrated together with the fossil lagomorphs BV2, rather than to P. etruria from BV1 which is of Italian mainland. The lagomorphs from Italian islands are dis- chronologically closer, and decided to ascribe those cussed in separate sections. The chronological distribution of Italian lagomorph taxa is remains to P. aff. minor. Paludotona is absent from illustrated in Fig. 3 (mainland Italy), and in Fig. 4 (Sicily and Sardin- BV3 continental assemblages, deposited after the ia). The partition in Mammal Ages and Faunal Units of the Italian re-establishment of the connection with mainland, Plio-Pleistocene follows Gliozzi et al. (1997), integrated with Petro- nio and Sardella (1999). Latest criticisms and proposals of modifi- which persists up to date. cation/integration (e.g. Raia et al. 2005, 2006; Palombo & Sardella Though abundant in Tuscany, Paludotona is 2007; Petronio et al. 2007, 2019; Sardella & Palombo 2007; Palombo not known from Fiume Santo, the sole fossiliferous 2014, 2018a; Bellucci et al. 2015) are to be accepted only after a revi- locality of the Tusco-Sardinia PB located in Sardin- sion of the huge amount of new data collected in the XXI century, combined with the revision of older data. ia, possibly coeval to V2 horizon of the Baccinel- The age of the fossiliferous sites of peninsular Italy follows lo-Cinigiano Basin (Abbazzi et al. 2008a; Casano- Kotsakis et al. (2003), Masini and Sala (2007, 2011), Sala and Masini vas-Vilar et al. 2011). (2007), or specific publications cited in the text. For Sardinia, we fol- low Palombo (2006, 2009, 2018b), and Palombo and Rozzi (2014). For Sicily, we follow the scheme originally proposed by Kotsakis Late Turolian (MN13, latest Miocene) (1979), eventually modified by Bonfiglio and Kotsakis (1987) and Prolagus Pomel, 1853 Bonfiglio et al. (2002), in its latest version proposed by Marra (2013). Remains of the ochotonid Prolagus sorbinii Fossil Lagomorpha (Mammalia) of Italy 159 Fig. 1 - Fossil stem lagomorphs and ochotonids of Italy. a) Paludotona etruria, p3 dx (Baccinello V1; fig. 2C, p. 461 in Angelone et al. 2017); b) Paludotona minor, p3 dx (Baccinello V2; fig. 2I, p. 461 in Angelone et al. 2017); c) Prolagus sorbinii, p3 sin (Monte Castellaro; fig. 3c, p. 410 in Angelone 2007); d) Prolagus apricenicus, p3 sin (Gargano fissure infilling F8; pl. 1.26, p. 235 in Mazza 1987, reversed); e) Prolagus imperialis, p3 sin (Gargano fissure infilling F32; pl. 3.4, p. 239 in Mazza 1987); f) Prolagus cf. sorbinii, p3 dx (Borro Strolla; fig. 3e, p. 132 in Angelone & Rook 2012); g) Prolagus aff. sorbinii, p3 dx (Arcille; fig. 3o, p. 132 in Angelone & Rook 2012); h) Prolagus savagei, p3 dx (Arondelli; CA, unpublished data); i) Prolagus italicus, p3 sin (Montagnola Senese; fig. 2a, p. 448 in Angelone 2008a); j) Prolagus aff. figaro, p3 sin (Capo Mannu D1; fig. 2A, p. 290 in Angelone et al. 2015); k) Prolagus aff. figaro, p3 dx (Capo Mannu D1; fig. 2B, p. 290 in Angelone et al. 2015); l) Prolagus figaro, p3 dx (Monte Tuttavista X4; CA, unpublished data); m) Prolagus sardus, p3 sin (Monte Tuttavista VI6; fig. 3e, p. 113 in Angelone et al. 2008). Masini, 1989 (Fig. 1.c), or of closely related forms an, according to Abbazzi et al. 2008b), was less for- (P. cf. sorbinii), are present in Velona (pre-evaporit- tunate. The presence of Lagomys G. Cuvier, 1800 ic Messinian, Tuscany), Monte Castellaro (evapo- at Casino was first reported by Major (1875, part ritic Messinian, Marches, type locality of the spe- I: 39). In part II of the same paper (Major 1875, cies), and in the post-evaporitic Messinian localities part II: 238), the author introduced Myolagus elsanus Brisighella (Romagna), Ciabòt Cagna (Piedmont), (=Prolagus elsanus). Those remains were not figured Moncucco (Piedmont), Verduno (Piedmont) (see nor described in the above mentioned paper. A Masini 1989; Ghetti et al. 2002; Angelone 2007; An- short morphological description of P. elsanus was gelone & Cavallo 2010; Angelone et al. 2011; Ange- given in Major (1899). However, such description lone & Rook 2012; Colombero et al. 2014, 2017). mentions features which have been lost after the Prolagus remains from the post-evaporitic Messinian earliest Miocene in the lineage leading to the genus locality Cava Stingeti (Molise; Cosentino et al. 2018) Prolagus (Angelone & Rook 2012), thus deepening do not have a specific attribution yet. the mystery around the lagomorph from Casino. At Recent surveys in the NHMB (CA, October present, the remains of P. elsanus seem to be lost. 2015), led to the re-discovery of an upper first inci- In the late Turolian, two endemic insular spe- sor collected in levels referable to BV3 (pre-evapo- cies of Prolagus were present in the Abruzzi-Apulia ritic Messinian), whose size and overall appearance PB (now part of the Italian peninsula): Prolagus apri- are compatible with Prolagus. Such specimen was cenicus Mazza, 1987 (Fig. 1.d), and Prolagus imperia- first mentioned in Rook and Masini (1990), quot- lis Mazza, 1987 (Fig. 1.e). Remains of both species ing a personal communication of B. Engesser. The are yielded in several karst fissure infillings located specimen was not found in a previous survey of in the Gargano peninsula (Apulia). However, they NHMB material (see Angelone & Rook 2012: 140), have been recovered together only in the most re- thus, its existence could not be officially re-con- cent among the Gargano infillings accumulated firmed until now. during the Neogene (Masini et al. 2010). The dental The search of the remains attributed to Pro- morphology and size of P. imperialis are highly mod- lagus elsanus Major, 1899, recovered in the second ified following a specific pattern recognized in in- sedimentary cycle of Casino (Tuscany, late Messini- sular endemic congeneric species (Angelone 2005). 160 Angelone C., Moncunill-Solé B. & Kotsakis T. The available populations of P. apricenicus, on the sinian assemblages corresponding to BV3 (mag- other hand, show quite variable dental morpho-di- netostratigraphically dated at ~6.7-6.4 Ma; Rook mensional traits, ranging from smaller forms whose et al. 2011). In spite of belonging to a balanced, morphology is almost non-modified with respect continental faunal assemblage, A. meini, as well to continental ones, to larger specimens in which as other elements of BV3, is a taxon endemic to marked insular endemic morphological modifica- southern Tuscany not inherited from the endem- tions can be observed (Mazza 1986a, 1986b, 1987; ic Tusco-Sardinia PB. Alilepus meini is likely to be Mazza & Zafonte 1987; Angelone 2005). There is related to Alilepus turolensis López Martínez, 1977, no final accordance about the phylogenetic relation- a species reported from NE Spain and Greece in ships between the two species: they may be either MN12-14 (Angelone & Rook 2011 and references related by an ancestor-descendant relationship, or therein). The erection and description of A. meini they may be two sister species developed from a allowed to definitively discard the presence of Hy- common ancestor in different islands of the Gar- polagus Dice, 1917 in BV3, hypothesized in litera- gano palaeoarchipelago. Indeed, the “sudden” ap- ture older than Angelone and Rook (2012: 135, see pearance of P. imperialis and the lack of a transition- references in synonymic list). No other report of al form between the two species, are elements in Alilepus is known in the Italian peninsula after the favour of the second hypothesis. Cladistic analyses early Messinian. confirm the hypothesis raised by Mazza (1987), and supported by Angelone and Čermák (2015), that the Trischizolagus Radulesco & Samson, 1967 continental ancestor of P. apricenicus and P. imperialis A leporid reported from Brisighella was orig- may be a still not described species close to Prola- inally classified as Trischizolagus cf. maritsae Bruijn, gus oeningensis (König, 1825) pertaining to an eastern Dawson & Mein, 1970 (De Giuli 1989; Rook et European clade of Prolagus (Angelone et al. 2015). al. 2015 and references therein). No description It is worth to remark that P. apricenicus has nor illustration of the material is available at pres- been the starting point of a palaeobiological ap- ent. Pending a systematic revision, we address such proach to the study of European lagomorphs. The materials as Trischizolagus sp. basing on preliminary results illustrated in Moncunill-Solé et al. (2015, observations by CA (2019). Remains ascribed to 2016a) predict a mass of 280 g for P. apricenicus Trischizolagus sp. (Fig. 2.b) recovered in the low- from Cava Dell’Erba (F1 site), and of 600 g for the er sub-synthem of Borro Strolla (Tuscany, latest population of the same species from Cava Fina (F8 Messinian), consist of unworn, juvenile teeth, and site). The skeletochronology of the former popu- were not determinable at the specific level due to the lation (F1) indicates a longevity of at least 7 years, scantiness and bad preservation of the specimens suggesting for P. apricenicus a slower life history than (Angelone & Rook 2012). In the Italian peninsula, expected from body size, when compared with ex- Trischizolagus seems not to survive the Mio-Pliocene tant ochotonids. boundary. An articulate skeleton of Prolagus was recov- ered in marine sediments in Capo di Fiume (Abruz- Leporidae indet. zo), an area that in the Late Miocene pertained to Undetermined leporids have been reported the Abruzzi-Apulia PB (Mazza et al. 1995). A lower from the late Messinian fossiliferous sites of Mon- age constraint to the lower Messinian is provided by cucco and Cava Stingeti (Angelone et al. 2011; Co- forams and molluscs. The remains have been tenta- lombero et al. 2017; TK, pers. data). tively attributed to P. cf. apricenicus on the basis of palaeogeographical evidences (Mazza et al. 1995). Ruscinian-early Villányian (MN14-16, In fact, observations of the teeth occlusal surface Pliocene) are not possible at present. Prolagus Prolagus cf. sorbinii (Fig. 1.f) is present in the Alilepus Dice, 1931 upper sub-synthem of Borro Strolla, dated to the Alilepus is the oldest leporid recorded in the Messinian/Zanclean transition (earliest MN14; Italian peninsula. Alilepus meini Angelone & Rook, Abbazzi et al. 2008b; Angelone & Rook 2012). The 2011 (Fig. 2.a) has been reported from early Mes- following reports of Prolagus in the Italian Penin- Fossil Lagomorpha (Mammalia) of Italy 161 Fig. 2 - Fossil leporids of Italy. a) Alilepus meini, p3 dx (Baccinello V3 localities; fig. 2a, p. 153 in Angelone & Rook 2011); b) Trischizolagus sp., p3 dx (Borro Strolla, lower sub-synthem; fig. 5c, p. 136 in Angelone & Rook 2012); c) Hypolagus petenyii, P2 dx (Arondelli; CA, unpublished data); d) Oryctolagus valdarnensis, p3 sin (Upper Valdarno; fig. 3h, p. 132 in Angelone & Rook 2012); e) Hypolagus brachygnathus, p3 sin (Monte La Mesa, fig. 3.8, p. 92 in Marchetti & Sala 2002); f) Hypolagus aff. brachygnathus, p3 dx (Pirro Nord, unknown fissure; fig. 2e, p. 97 in Angelone 2013); g) Oryctolagus sp., p3 sin (Monte Peglia, upper levels, fig. 9a, p. 27 in Van der Meulen 1973); h) Lepus terraerubrae, p3 dx (Monte Peglia, lower levels, fig. 8 dx, p. 27 in Van der Meulen 1973); i) Oryctolagus burgi, p3 sin (Borgio Verezzi, fig. 2-Val 90 27a/3b, p. 21 in Nocchi & Sala 1997a); j) Oryctolagus cuniculus, p3 sin (San Sidero 3; fig. 2-MP232-sx, p. 173 in Nocchi & Sala 1997b); k) Lepus corsicanus, p3 dx (La Grave; fig. 5A, Kotsakis et al. in press); l) Leporidae indet., right upper molariform (Capo Mannu D1; fig. 2V, p. 290 in Angelone et al. 2015); m) Sardolagus obscurus, p3 sin (Monte Tuttavista IVm; fig. 3.1, p. 510 in Angelone et al. 2018); n) Hypolagus peregrinus, p3 dx (Monte Pellegrino; fig. 2d and 2f, p. 42 in Fladerer & Fiore 2003). sula are dated to MN16a. They are P. aff. sorbinii put it in synonymy with any of the species reputed from Arcille (Tuscany) (Fig. 1.g) and Prolagus savagei as valid. This latter point was questioned by Azzar- Berzi, 1967 from Arondelli (Piedmont) (Fig. 1.h). oli (1977), who instead supported the validity of P. Prolagus aff. sorbinii, though showing mor- savagei. Prolagus savagei was eventually synonymyz- phological similarities to P. sorbinii, is an independ- ed with Prolagus calpensis Major, 1905 (Alberdi et ent species whose formalization depends on the al. 1997). Angelone and Rook (2012), however, re- reassessment of P. savagei. Actually, P. aff. sorbinii jected the synonymy with P. calpensis, listing several and P. savagei are likely to pertain to the same spe- characters to differentiate P. savagei + P. aff. sorbinii cies (Angelone & Rook 2012), however, P. savagei, from P. calpensis (ibid: 135). which should have nomenclatorial priority, does not have a clearly stated diagnosis and its supposed Hypolagus Dice, 1917 distinctive characteristics are confusing and contra- A tooth referable to the genus Hypolagus dictory (see Berzi 1967). Its validity was questioned found in Arondelli was at first classified as Hypola- by López Martínez and Thaler (1975), who listed gus sp. (Berzi 1967), and eventually ascribed to Hy- the species of Prolagus known at the time (ibid.: 851- polagus petenyii Čermák & Fladerer in Čermák, 2009 852), commenting also on the nomenclatorial/tax- (Fig. 2.c). The specimen from Arondelli represents onomic problems related to some of them. In the the sole report of the genus in the Italian Peninsula case of P. savagei, they argued that the species was before Pleistocene times. based on a fossilization anomaly. Indeed, they did not list it in the species of Prolagus they considered Late Villányian (MN17, Early Pleistocene) as valid (ibid.: 852). Nevertheless, they did not lit- Prolagus erally declare P. savagei as invalid, and they did not The continental endemic species Prolagus 162 Angelone C., Moncunill-Solé B. & Kotsakis T. Fig. 3 - Chronological distribution of lagomorph taxa in mainland Italy. Integrated chronology (Geochronology/Magnetostratigraphy, Large Mammal Ages, Small Mammal Ages, MN Zones, Italian FU, and selected fossil localities) modified from Mein (1975), Sala and Masini (2007), Masini and Sala (2011), Hilgen et al. (2012), Colombero et al. (2014). Fossil Lagomorpha (Mammalia) of Italy 163 Fig. 4 - Chronological distribution of lagomorph taxa recorded in Sicily and Sardinia. A relative chronological arrangement of some infillings of Monte Tuttavista has been proposed on the basis of the lagomorph content (Angelone et al. 2008, 2009). However, such arrange- ment takes into account only a part of the fissures containing Prolagus. Thus, we decided to group the selected localities in clusters without an internal chronological arrangement. The clusters are arranged in relative chronological order and, when possible, next to the correspondent FsC following Palombo (2009). The relative chronological position of most Capo Figari infillings is doubtful due to taphonomic mixing/sampling bias, thus we do not report them here. Integrated chronology (Geochronology/Magnetostratigra- phy, Large Mammal Ages, Small Mammal Ages, MN Zones, Sardinian and Sicilian FC, and selected fossil localities) modified from Mein (1975), Bonfiglio et al. (2002), Masini et al. (2008), Palombo (2009), Hilgen et al. (2012), Marra (2013). GST-Pianetti: Grotta San Teodoro-Pianetti. FC Monte Pell.: Monte Pellegrino FC. italicus Angelone, 2008 (Fig. 1.i) has been erected ivola FU; Rook & Masini 1990), were tentatively basing on fossil material from Montagnola Senese referred to P. aff. sorbinii (Angelone & Rook 2012). (Tuscany, Coste S. Giacomo FU; Fondi 1970). In This specific attribution was mainly based on some the same paper, were attributed to P. italicus also the mandibular features shared by P. aff. sorbinii and Prolagus remains from Torre di Picchio (Umbria, the Solava specimen, which lacks species-diagnos- Olivola FU; Girotti et al. 2003). Recent researches tic teeth. highlight the presence of P. italicus in the Coste S. The genus Lagomys was mentioned by Falcon- Giacomo fossil locality (Latium, Coste S. Giaco- er (1868) among the vertebrates of Upper Valdarno mo FU; Bona et al. 2015). This discovery slightly (Tuscany). Its presence was eventually confirmed pre-dates the appearance of P. italicus with respect by Major in Stoppani (1873). However, Lagomys was to older literature. Cladistic analyses suggest that not reported in subsequent faunal lists (Major 1875, P. italicus developed from P. sorbinii “immediately” 1876, 1884). Due to the lack of descriptions, fig- after its arrival in the Italian peninsula in the ear- ures, and references to repositories and to inventory ly Messinian (Angelone et al. 2015). Being derived numbers, it is not possible to know if such speci- from P. sorbinii, P. italicus is the sister species of P. mens were lost, or if their taxonomic assignment savagei/P. aff. sorbinii. was later reconsidered. Indeed, in the lists of ver- Other remains of Prolagus of the late Villány- tebrates of the Upper Valdarno reported by Major ian of the Italian peninsula (Solava; Tuscany, Ol- (1875, 1876, 1884), Lagomys was substituted with 164 Angelone C., Moncunill-Solé B. & Kotsakis T. Lepus Linnaeus, 1758 (see also next paragraph). Two namely those from Torre di Picchio, Casa Sgherri more specimens of Lagomys from Upper Valdarno (Tuscany; Marcolini et al. 2000), and Upper Val- were described by Bosco (1899a, 1899b). Tobien darno localities of Olivola FU (Angelone & Rook (1935) argued that those specimens had to be as- 2012 and references therein). The morphological cribed to the genus Prolagus. Those specimens were differences noticed among the populations of O. not figured in any of the cited papers, and they valdarnensis recognized in the Italian peninsula are are probably lost (Angelone & Rook 2012). It is due to different factors: some changes may be re- impossible to guess neither their biochronological lated to temporal trends, others to geographical position (i.e. whether they can be assigned to the trends probably linked to ecological factors (Ange- late Villányian Olivola FU, or to the early Biharian lone & Rook 2012: 142). Tasso FU), or their specific (or even generic) attri- Oryctolagus appears in Italy much later than bution. the European first appearance of the genus (~3.5 Ma) and slightly after the first appearance of O. la- Oryctolagus Lilljeborg, 1873 costi, a species with western-central European dis- The presence of a leporid in the Upper Val- tribution, which is dated at ~2.5 Ma (López Martí- darno was known since the late XIX century. In nez 2008; Angelone & Rook 2012). We exclude the the oldest publications, faunal lists reported the presence of O. lacosti in the Italian peninsula during presence of Lepus sp. (Major 1875, 1876, 1884). It the late Villányian (contra Pelletier et al. 2015: fig. must be remarked, though, that in previous publi- 1.E). In fact, in our opinion, all the late Villányian cations the faunal lists of Upper Valdarno reported Oryctolagus remains up to now collected in Italy be- Lagomys, not Lepus (see previous paragraph). Even- long to O. valdarnensis. tually, Weithofer (1889) erected the species Lepus valdarnensis Weithofer, 1889 to include such mate- Leporidae indet. rials. Bosco (1899a, 1899b) split the leporid from Faunal lists in Biddittu et al. (1979) and Cas- Upper Valdarno in two taxa (L. valdarnensis and L. soli and Segre Naldini (1984) report the presence of etruscus Bosco, 1899), leaving undetermined post- Lepus sp. in the Fontana Acetosa fossiliferous local- cranial remains in open nomenclature (Lepus sp.). ity (Anagni, Latium) dated to the latest Villányian On the other hand, Major (1899) reputed that the or the to earliest Biharian basing on a controversial leporid material from Upper Valdarno pertained report of Hippopotamus (see Bellucci et al. 2014). in its entirety to the Asian genus Caprolagus Blyth, Segre Naldini et al. (2009), probably basing on the 1845. Decades later, Viret (1954) assigned the same same material taken into account by Biddittu et al. material to Oryctolagus lacosti (Pomel, 1853). An un- (1979), mention instead the presence of “Oryctola- successful effort of revalidation of L. valdarnensis gus cf. O. etruscus”. Such material was not described was attempted by Forteleoni (1971). Angelone and nor figured. Rook (2012), agreeing with Viret (1954), consid- As at the present state of the art all the lep- ered the Upper Valdarno leporid remains as per- orids of the Villányian of central Italy have been taining to the genus Oryctolagus. They validated and ascribed to O. valdarnensis, it is quite likely that the redescribed the species O. valdarnensis (Weithofer, leporid from Acqua Acetosa may be referred to 1889) (Fig. 2.d), including as a synonym L. etruscus this species. (Angelone & Rook 2012). Indeed, the very large teeth and the presence of several “leporine” char- Biharian (Early Pleistocene-early Middle acters in the maxilla of O. valdarnensis (see Ange- Pleistocene) lone & Rook 2012: 138) may be misleading, and Prolagus explain its eventful taxonomic history. A time span of ~1.4 Ma separate the early Montagnola Senese records the oldest Italian Villányian P. aff. sorbinii from Arcille and the con- Oryctolagus. The remains from Montagnola Senese specific remains found at Case Inferno (Tuscany, were originally assigned to Oryctolagus cuniculus (Lin- Tasso FU; Rook & Masini 1990; Angelone & Rook naeus, 1758) (Fondi 1972). Eventually, they were 2012). Such discontinuity does not represent the ascribed to O. valdarnensis, as well as the other late actual absence of Prolagus in central Italy, but is due Villányian remains of Oryctolagus of central Italy, to the poor Italian Pliocene record of fossil locali- Fossil Lagomorpha (Mammalia) of Italy 165 ties bearing continental vertebrates. A fragment of Oryctolagus maxilla of Prolagus, discovered in a fissure infilling During the Biharian, O. valdarnensis is still of uncertain age in Colleferro (Latium), shows mor- present in Tuscany, Umbria, and Apulia (Casa Frata phological characters of P4-M2 compatible with P. and other Upper Valdarno localities of Tasso FU; aff. sorbinii, but a size comparable to P. italicus (CA Pietrafitta and Monte Argentario, Farneta FU; and and TK, unpublished data). Pirro Nord, Pirro FU) (Borselli et al. 1980; Argenti Remains of Prolagus have been reported also & Kotsakis 2009; Angelone & Rook 2012 and ref- from Monte Riccio (Latium, Tasso FU; Mazzini erences therein; Rook & Angelone 2013; Siori et al. et al. 2000), but they consist in fragmentary speci- 2014). It is worth to mention that in a younger pa- mens, non-diagnostic at the species level. The Tasso per, Oryctolagus from Pirro Nord is still reported as FU records the latest reports of the genus Prolagus O. aff. lacosti (Angelone 2013). This is not due to a in the Italian peninsula. subsequent reconsideration of the taxonomic attri- bution of the taxon, but to simply to a long delay Hypolagus (2008-2013) between the manuscript approval and In the earliest Biharian, Hypolagus appears the publication of the paper and the impossibility to again in northern Italy, this time in the eastern Al- modify the text according to the new highlights oc- pine region (Monte La Mesa, Veneto, Tasso FU; curred in between (see foreword in Angelone 2013: Marchetti et al. 2000) (Fig. 2.e). The dental morpho- 96). dynamic indexes and the comparison with other Eu- Oryctolagus sp. from the upper levels of Monte ropean populations of Hypolagus, indicate that the Peglia (latest early Biharian, Colle Curti FU) (Fig. sample from Monte La Mesa, formerly attributed 2.g) was originally classified as Lepus sp. (Van der by Marchetti et al. (2000) to Hypolagus beremendensis Meulen 1973). Angelone and Rook (2012) and (Petényi, 1864), fits in the lower range of Europe- Angelone (2013), basing on a lower third premo- an populations of Hypolagus brachygnathus (Kormos, lar figured in Van der Meulen (1973: fig. 9), agree 1930) (Angelone 2013: fig. 3; see Čermák 2009 for with López Martínez (1989), who reconsidered the nomenclatorial details about the genus Hypolagus). taxonomical position of the specimen ascribing it Indeed, the temporal and geographical distribution to the genus Oryctolagus. Indeed, the measurements of H. brachygnathus (see Čermák 2009) support the and the morphological features observable in Van taxonomic assignment of the remains from Monte der Meulen (1973), fit with their attribution to O. La Mesa based on the tooth morphology. valdarnensis. However, the current unavailability Another early Biharian report of Hypolagus, of the specimens curated at the IsIPU, prevents slightly younger than the one from Monte La Mesa, well-founded taxonomic statements. is H. aff. brachygnathus from Pirro Nord (Apulia, Oryctolagus sp. from Monte Peglia represents Pirro FU; Angelone 2013) (Fig. 2.f). Unfortunate- the last report of the older stock of Italian pop- ly, the most recent revision of Hypolagus from Pir- ulations of Oryctolagus defined as Older Oryctolagus ro Nord revealed that a large part of the material Populations in Angelone (2013). During the late Bi- collected in the oldest surveys may be lost (Ang- harian there is no report of Oryctolagus in the Italian elone 2013). Nevertheless, the remains available Peninsula. up to date, coupled with older literature data, were In the first alluvial unit of Castagnone (Pied- sufficient to make some interesting considerations. mont, Colle Curti FU), it was reported a leporid Hypolagus from Pirro Nord differs from the coeval classified as Oryctolagus cf. lacosti (Giraudi et al. 2003). central European H. brachygnathus by having high- The same remains have been classified as Leporidae er dental morphodynamic indexes, coupled with a indet. by Siori and Sala (2007). smaller dental size and a poor cursorial structure of Caloi and Palombo (1994) reported fragmen- the limbs (Fladerer & Fiore 2003; Angelone 2013). tary cranial remains of a leporid from the lacustrine Such mix of advanced and primitive characters ob- sediments of Capena (Latium), and assigned them served in H. aff. brachygnathus was explained hypoth- to Lepus etruscus (synonym of Oryctolagus valdarnensis esizing the existence of an endemic, conservative according to Angelone & Rook 2012). The fossil- Italian lineage of Hypolagus derived from a basal iferous site of Capena was previously assigned to stock of H. brachygnathus of central Europe. Pirro FU (Gliozzi et al. 1997), and eventually to an 166 Angelone C., Moncunill-Solé B. & Kotsakis T. age range between Pirro and Slivia FUs (Marra et al. to; Bartolomei et al. 1982; Sala 1990; Bon et al. 1991). 2018). The specimens were not figured, and were eventually lost. Oryctolagus The genus Oryctolagus is absent in the Italian Lepus Linnaeus, 1758 fossil record during the late Biharian (Slivia FU and The oldest reliable record of the genus Lepus Ponte Galeria FU). Oryctolagus reappears in the ear- in the Italian peninsula occurs in the lower levels liest Toringian (Borgio Verezzi, Liguria, Isernia FU; of Monte Peglia (late early Biharian, Colle Curti Nocchi & Sala 1997a, 1997b) with a new species, FU). The remains were ascribed to Lepus terraeru- O. burgi Nocchi & Sala, 1997 (Fig. 2.i). Oryctolagus brae Kretzoi, 1956 (now nomen dubium, pers. comm. cf. burgi may be present also at Ponte Galeria 3 S. Čermák 2019) in the first and only paper dealing (Latium, Isernia FU; Kotsakis & Barisone 2008, and with the lagomorphs from Monte Peglia (Van der currently under revision by Sardella and colleagues). Meulen 1973) (Fig. 2.h). As well as Oryctolagus sp. This species is unknown at present in the rest of from the upper levels of the same locality, this fossil Europe, and in Italy it is the oldest of the Modern hare is currently unavailable for updated compari- Oryctolagus Populations, a stock of Italian Oryctolagus sons. characterized by morphological features similar to Italian reports of Lepus older than the above extant species (Angelone 2013). At the present state mentioned one are probably the result of an error of the art, O. burgi seems to be exclusive of the Iser- in the generic attribution. This may be the case, for nia FU. The phylogenetic relationships of O. burgi example, of Lepus sp. from Soave Cava Sud (Vene- with congeneric species is still a matter of debate to, late early Biharian, Pirro FU) mentioned in a no- (Nocchi & Sala 1997a, 1997b). menclatorial update (not a taxonomic revision) of A large leporid found at Isernia (Molise, ear- old faunal lists of Plio-Pleistocene assemblages of liest Toringian, Isernia FU) was formerly reported NE Italy (Bon et al. 1991 and references therein). as Lepus sp. (Sala 1983). Eventually, its taxonomic position was reconsidered as cf. Oryctolagus sp. (Sala Leporidae indet. 1996). In correspondence of the Fontana Ranuccio Ambrosetti et al. (1979) reported Lepus euro- FU, Oryctolagus disappears again in the Italian fossil paeus Pallas, 1778 from Slivia (Friuli-Venezia-Giulia, record. late Biharian, Slivia FU), but such attribution was In the Italian peninsula, the oldest report of made only on a dimensional basis. Bon et al. (1992) Oryctolagus cuniculus (Fig. 2.j) occur in correspond- revised the faunal assemblage from Slivia, but did ence of the lower levels of Torre in Pietra (Latium, not include the leporid. Thus, for the moment, it is late early Toringian, Torre in Pietra FU; Caloi & preferable to leave such remains in open nomencla- Palombo 1978). Since late Toringian on, O. cunicu- ture as Leporidae indet. lus becomes quite common in the Italian peninsula. A few remains of an undetermined leporid Some early Toringian findings do not have a specific have been reported from San Lorenzo (Basilica- attribution and are reported in literature as Oryctola- ta, latest Biharian, Ponte Galeria FU; Masini et al. gus sp. (Tab. 1). 2005). Callou (2003) argues that Oryctolagus may be absent from the Late Pleistocene and Early Hol- Toringian (late Middle Pleistocene-Late ocene fossiliferous sites of southern France. We Pleistocene) checked this possibility in the Italian fossil record: Ochotona Link, 1795 Oryctolagus is present in several localities of south- Fossil lagomorphs collected in the middle/up- ern Italy assigned to MIS 5 and MIS 3 (see Tab .1, per levels of Visogliano A (Veneto; early Toringian, and López Martínez 2008: fig. 10), and in the MIS 4 Isernia FU/Fontana Ranuccio FU) were at first at- of Arma delle Manie (Liguria; El Guennouni 2001). tributed to Ochotona sp. (Bartolomei et al. 1977; Bar- Thus, there is evidence of the survival of Oryctolagus tolomei & Tozzi 1978), and eventually classified as in peninsular Italy in the Late Pleistocene at least Ochotona pusilla (Pallas, 1769) by Abbazzi et al. (2000). until MIS 3. Anyway, the presence of Oryctolagus in In the late Toringian, Ochotona cf. pusilla is re- southernmost Italy during the Last Glacial Maxi- ported in the lower levels of Tagliente Shelter (Vene- mum and the early Holocene can be inferred also Fossil Lagomorpha (Mammalia) of Italy 167 by indirect evidences, and a recent report suggests to the mountain hare Lepus timidus (or, in older lit- the survival of the wild rabbit in peninsular Italy erature, to its synonym Lepus variabilis Pallas, 1778). also in the early Holocene (see below). In some cases, L. timidus is reported together with L. europaeus. Lepus timidus is a species associated Lepus to colder climates, so it is noteworthy its report Toringian Italian fossil remains of Lepus at Grotte delle Striare (in the largest cave, lev. h, need a complete revision. The specifically deter- Apulia; Di Stefano et al. 1992), which represents mined remains are attributed to four extant spe- the southernmost record in the Italian peninsula. cies: Lepus europaeus, Lepus corsicanus De Winton, Additional reports of L. timidus are listed in Ca- 1898 (Fig. 2.k), Lepus timidus Linnaeus, 1758, Lep- passo Barbato et al. (1991). Minieri et al. (1995 us capensis Linnaeus, 1758. However, such specific with references) lists 11 Toringian sites with L. determinations are often based on non-diagnostic timidus, 41 reports of L. europaeus (including L. cor- remains, on dimensional features only, or rely on sicanus), and 19 fossiliferous sites with O. cuniculus the known (or better said: supposed) biochrono- (19). Most of the reports in Capasso Barbato et logical and geographical Toringian distribution of al. (1991) and Minieri et al. (1995) need a reassess- the above mentioned extant species, which in its ment and are not listed in Tab. 1. turn needs to be thoroughly revised, generating Several remains of leporids attributed by a dangerous circular reasoning. For this reason, Pasa (1947) and Bartolomei (1964) to Lepus mediter- pending a taxonomic revision of Italian fossil Lep- raneus Wagner, 1841 and L. europaeus were collected us, we prefer to list in open nomenclature (Lepus from a few localities of NE Italy (Cengelle 1, Cas- sp.) several Toringian reports previously classified tello, Viatelle, Spessa II; late Biharian or Toringi- to the species level (see Tab. 1). They go to enlarge an after Bon et al. 1991 with references). Also for the list of Italian Toringian reports of Lepus which these remains is absolutely necessary a systematic currently lack a determination at the species level revision. (see Tab. 1, and Kotsakis et al. 2003 and references therein). Holocene At present, Lepus corsicanus from the lower Oryctolagus levels of Visogliano A (late Galerian, Isernia FU/ The presence of the genus Oryctolagus in the Fontana Ranuccio FU; Abbazzi et al. 2000; Visma- early Holocene was reported in old faunal lists, but ra 2012) represent the oldest reliable record of the such material was not figured and needs a restate- species, followed by L. cf. corsicanus from Cretone ment. Oryctolagus was introduced in Italy by humans (Latium, Fontana Ranuccio FU; Di Canzio et al. for hunting and as livestock in the late Holocene 2003). (Angelici et al. 2008; Masseti 2008). After Rustioni The oldest reliable Italian reports of L. euro- et al. (2007), the rabbit is present in the late Ne- paeus seems to date back to the late Toringian. Nev- olithic (VI millennium BP) of Offida (Marches). ertheless, the remains of L. europaeus from central However, it is quite likely that the rabbit found in and southern Italy may pertain to L. corsicanus (cfr. southernmost Italy at Grotta di Latronico 3 (Basil- Kotsakis 2008). Remains from tardiglacial assem- icata, ~8000-7400 ky; Dini et al. 2008) may be not blages of Calabria, classified as L. europaeus, have a human introduction, but a relict population. been eventually attributed to Lepus sp. (Mangano 2007; Marra 2009 both with references). Lepus The endemic Italian hare, L. corsicanus, con- At the beginning of the Holocene, L. eu- sidered as a subspecies of L. europaeus by Miller ropaeus can be considered the predominant spe- (1912), was eventually revalidated on the basis of cies of the genus in northern and central Italy, dental and cranial characters by Palacios (1996). whereas L. corsicanus was probably distributed in Several authors refer to L. corsicanus the remains southern-central and southern Italy (Angelici et from Late Pleistocene fossiliferous sites of south- al. 2008). It is worth to mention that, although its ern-central and southern Italy (see Tab. 1). name may suggest an origin from Corsica, the spe- In northern and northern-central Italy, late cies L. corsicanus is not autochthonous of Corsica. Toringian remains of Lepus are frequently referred In fact, it has been introduced there by humans 168 Angelone C., Moncunill-Solé B. & Kotsakis T. Age Locality Region Species References MN11 Baccinello V0 Tuscany Paludotona aff. minor Cirilli et al. 2016; Angelone et al. 2017 Turolian middle early- ?latest MN11-early MN12 Baccinello V1 Tuscany Paludotona etruria Dawson 1959; Hürzeler & Engesser 1976; Angelone et al. 2017 late MN12-early MN13 Baccinello V2 Tuscany Paludotona minor Angelone et al. 2017 pre-evaporitic Messinian, Capo di Fiume Abruzzi Prolagus cf. apricenicus Mazza et al. 1995; Angelone 2007 MN13 Gargano (several fissure MN13 Apulia Prolagus apricenicus Masini et al. 2013; Moncunill-Solé et al. 2016a infillings Gargano (several fissure Prolagus apricenicus, Prolagus Mazza 1986a, 1986b, 1987; Mazza & Zafonte 1987; Angelone 2007; MN13/?MN14 Apulia infillings) imperialis Masini et al. 2013; Moncunill-Solé et al. 2016a pre-evaporitic Messinian, Rook & Ghetti 1997; Ghetti et al. 2002; Angelone 2007; Angelone & Velona Tuscany Prolagus cf. sorbinii MN13 Rook 2012 pre-evaporitic Messinian, Baccinello V3 Tuscany ?Prolagus sp., Alilepus meini Rook & Masini 1990; Angelone & Rook 2011, 2012 MN13 Emilia- evaporitic Messinian, MN13 Monte Castellaro Prolagus sorbinii Masini 1989 late Turolian Romagna post-evaporitic Messinian, Ciabòt Cagna Piedmont Prolagus sorbinii Cavallo et al. 1993 as Prolagus michauxi; Angelone & Cavallo 2010 MN13 post-evaporitic Messinian, Moncucco Piedmont Prolagus sorbinii, Leporidae indet. Angelone et al. 2011; Colombero et al. 2017 MN13 post-evaporitic Messinian, Verduno Piedmont Prolagus sorbinii Colombero et al. 2014 MN13 post-evaporitic Messinian, De Giuli 1989 as Prolagus cf. sorbinii and Trischizolagus cf. maritsae; Brisighella Romagna Prolagus sorbinii, Trischizolagus sp. MN13 Masini 1989; Angelone 2007; Rook et al. 2015 with references post-evaporitic Messinian, Major 1875, 1899, 1905; Rütimeyer 1876; López Martínez & Thaler [Casino] Tuscany [Prolagus elsanus] MN13 1975; Azzaroli et al. 1986; Angelone & Rook 2012 with references post-evaporitic Messinian, Cava Stingeti Molise Prolagus sp., Leporidae indet. Cosentino et al. 2018 MN13 Borro Strolla lower synthem latest Messinian Tuscany Trischizolagus sp. Abbazzi et al. 2008b; Angelone & Rook 2012 (=Cava Sefi) Borro Strolla upper synthem earliest MN14 Tuscany Prolagus cf. sorbinii Abbazzi et al. 2008b; Angelone & Rook 2012 Ruscinian- Villányian (=Cava Sefi) early MN16a, Triversa FU Arcille Tuscany Prolagus aff. sorbinii Hürzeler & Engesser 1976; Angelone & Rook 2012 Prolagus savagei, Hypolagus Berzi 1967; Berzi et al. 1970; Azzaroli et al. 1986; Čermák 2009; MN16a, Triversa FU Arondelli Piedmont petenyii Angelone & Rook 2012 with references Prolagus italicus, Oryctolagus Fondi 1970, 1972; Azzaroli et al. 1986; Angelone 2008a, 2013; MN17, Coste S. Giacomo FU Montagnola Senese (=Rosia) Tuscany valdarnensis Angelone & Rook 2012 MN17, Coste S. Giacomo FU Coste S. Giacomo Latium Prolagus italicus Bona et al. 2015 Prolagus italicus, Oryctolagus MN17, Olivola FU Torre di Picchio Umbria Girotti et al. 2003; Angelone 2008a, 2013; Angelone & Rook 2012 late Villányian valdarnensis MN17, Olivola FU Casa Sgherri Tuscany Oryctolagus valdarnensis Marcolini et al. 2000; Angelone & Rook 2012; Angelone 2013 MN17, Olivola FU Solava Tuscany Prolagus aff. sorbinii Angelone & Rook 2012 with references; Rook & Angelone 2013 Upper Valdarno (several MN17, Olivola FU Tuscany Oryctolagus valdarnensis Angelone & Rook 2012 with references; Rook & Angelone 2013 localities) Biddittu et al. 1979, Cassoli & Segre Naldini 1984; Segre Naldini et al. latest Villànyian/earliest Fontana Acetosa Latium ?Oryctolagus valdarnensis 2009; alternatively reported as Oryctolagus cf. O. etruscus or Lepus Biharian sp. [Upper Valdarno (several Tasso FU/Olivola FU Tuscany [?Prolagus] Angelone & Rook 2012 with references localities)] Tasso FU Case Inferno Tuscany Prolagus aff. sorbinii Rook & Masini 1990; Angelone & Rook 2012 Tasso FU Casa Frata Tuscany Oryctolagus valdarnensis Borselli et al. 1980; De Giuli & Masini 1986 Upper Valdarno (several Tasso FU Tuscany Oryctolagus valdarnensis Angelone & Rook 2012 localities) Tasso FU Monte Riccio Latium Prolagus sp. Mazzini et al. 2000 Tasso FU Monte La Mesa Veneto Hypolagus brachygnathus Marchetti et al. 2000; Angelone 2013 ?Early Pleistocene Colleferro Latium Prolagus sp. CA and TK unpublished data Biharian Farneta FU Pietrafitta Umbria Oryctolagus valdarnensis Argenti & Kotsakis 2009; Angelone & Rook 2012; Angelone 2013 Farneta FU Monte Argentario Tuscany Oryctolagus valdarnensis Angelone & Rook 2012; Siori et al. 2014 Hypolagus aff. brachygnathus, Pirro FU Pirro Nord Apulia Fladerer & Fiore 2003; Angelone & Rook 2012; Angelone 2013 Oryctolagus valdarnensis Pirro FU Soave Cava Sud Veneto ?Lepus sp. Bon et al. 1991 Colle Curti FU Castagnone Piedmont Leporidae indet. Giraudi et al. 2003 as Oryctolagus cf. lacosti; Siori & Sala 2007 Colle Curti FU Monte Peglia lower levels Umbria Lepus terraerubrae Van der Meulen 1973; López Martínez 1989 Colle Curti FU Monte Peglia upper levels Umbria Oryctolagus sp. Van der Meulen 1973; López Martínez 1989; Angelone 2013 Colle Curti FU [Capena] Latium ?Oryctolagus valdarnensis Caloi & Palombo 1994 as [Lepus etruscus] Slivia FU Slivia Venezia Giulia Leporidae indet. Ambrosetti et al. 1979 as Lepus europaeus Ponte Galeria FU San Lorenzo Basilicata Leporidae indet. Masini et al. 2005 Isernia FU Isernia Molise cf. Oryctolagus sp. Sala 1983 as Lepus sp.; Sala 1996 Borgio Verezzi lower level Isernia FU Liguria Oryctolagus burgi Nocchi & Sala 1997a, b (=Valdemino) Isernia FU Ponte Galeria 3 (=Casal Selce) Latium Oryctolagus cf. burgi, Lepus sp. Kotsakis & Barisone 2008; Sardella and coll. under study Isernia FU Spessa II Veneto Lepus sp. Bon et al. 1991 Isernia FU/Fontana Ranuccio Visogliano A lower levels Veneto Lepus corsicanus Abbazzi et al. 2000; Vismara 2012 FU Isernia FU/Fontana Ranuccio Bartolomei et al. 1977; Bartolomei & Tozzi 1978 as Ochotona sp.; early Toringian Visogliano A middle/upper levels Veneto Ochotona pusilla FU Abbazzi et al. 2000 Fontana Ranuccio FU Fontana Ranuccio Latium Lepus sp. Biddittu et al. 1979 as Lepus cf. capensis Fontana Ranuccio FU Bristie I Venezia Giulia Lepus sp. Lugli & Sala 2000 as Lepus europaeus Torre in Pietra FU Castel di Guido Latium Lepus sp. Sala & Barbi 1996 as Lepus cf. europaeus Torre in Pietra FU Torre in Pietra, lower levels Latium Oryctolagus cuniculus, Lepus sp. Caloi & Palombo, 1978; Caloi et al. 1998 Torre in Pietra FU Cretone Latium Lepus cf. corsicanus Di Canzio et al. 2003 Tozzi 1969 as Lepus europaeus and reported as found in lower levels; ?Torre in Pietra FU Borgio Verezzi upper level Liguria Lepus sp. Nocchi & Sala 1997a "Vitinia FU" Torre in Pietra, upper levels Latium Oryctolagus cuniculus Caloi & Palombo 1978; Caloi et al. 1998 Grotta Maggiore di San ?early or late Toringian Veneto Lepus sp. Bartolomei 1960; for the age: López-García et al. 2017 Bernardino (level unknown) Caverna degli Orsi Venezia Giulia Lepus europaeus, Lepus timidus Bon et al. 1991 with references; Berto & Rubinato 2013 Caverna Pocala Venezia Giulia Lepus europaeus, Lepus cf. timidus Fabiani 1919, Bon et al. 1991 with references late Toringian ?late Toringian Montorio Veneto Lepus europaeus Pasa 1952; Bon et al. 1991 Tagliente Shelter lower levels Veneto Ochotona cf. pusilla Bartolomei et al. 1982; Sala 1990; Bon et al. 1991 Tagliente Shelter upper levels Veneto Lepus europaeus, Lepus cf. timidus Capuzzi & Sala 1980; Bon et al. 1991 Grotta del Broion Veneto Lepus europaeus Sala 1980; Bon et al. 1991 Mezzena Shelter Veneto Lepus europaeus Bartolomei et al. 1980; Bon et al. 1991 Tab. 1 - List of lagomorph-bearing localities of continental Italy detailed with geographical location. See next page for caption. Fossil Lagomorpha (Mammalia) of Italy 169 Age Locality Region Species References MN11 Baccinello V0 dei Covoli di Grotta inferiore Tuscany Paludotona aff. minor Cirilli et al. 2016; Angelone et al. 2017 Turolian middle Veneto Lepus europaeus Bona et al. 2007 early- ?latest MN11-early MN12 Velo Baccinello V1 Tuscany Paludotona etruria Dawson 1959; Hürzeler & Engesser 1976; Angelone et al. 2017 late MN12-early MN13 Grotta della Baccinello V2Volpe Veneto Tuscany Lepus timidus Paludotona minor Pasa 1950a; Angelone et Bon et al. 1991 al. 2017 Grotta A del Ponte di Veja Veneto Lepus cf. europaeus, Lepus sp. Pasa 1950b; Bon et al. 1991 pre-evaporitic Messinian, Capo di Fiume Zandobbio Abruzzi Lombardy Prolagus cf. apricenicus Lepus cf. europaeus Mazza1957 Vialli et al. 1995; Angelone 2007 MN13 Caverna Generosa Lombardy Lepus sp. Bona et al. 2009 Gargano Arma delle (several Maniefissure Liguria Oryctolagus cuniculus El Guennouni 2001Moncunill-Solé et al. 2016a MN13 Apulia Prolagus apricenicus Masini et al. 2013; infillings Grotta del Principe lower levels Liguria Oryctolagus cuniculus Barral & Simone 1971 Gargano (several fissure Arene Candide Liguria Prolagus apricenicus, Lepus europaeus, Prolagus Lepus timidus Mazza 1986a, Cassoli 1986b, 1994 & Tagliacozzo 1987; Mazza & Zafonte 1987; Angelone 2007; MN13/?MN14 Apulia infillings) Monte Croara Emilia imperialis Lepus cf. timidus Masini1969 Pasini et al. 2013; Moncunill-Solé et al. 2016a pre-evaporitic Messinian, Grotta Parignana lev. B Tuscany Lepus timidus Rook & Ghetti Farina 2012, 2014 1997; Ghetti et al. 2002; Angelone 2007; Angelone & Velona Tuscany Prolagus cf. sorbinii MN13 Manciano Tuscany Lepus cf. europaeus Rook 2012 Petronio et al. 2014 pre-evaporitic Messinian, Oryctolagus cuniculus, ?Lepus Baccinello V3 Grotta di Sant’Agostino Tuscany Latium ?Prolagus sp., Alilepus meini Rook 1970 Tozzi & Masini 1990; Angelone & Rook 2011, 2012 MN13 europaeus Grotta Barbara Emilia- Latium Lepus europaeus Caloi evaporitic Messinian, MN13 Monte Castellaro Prolagus sorbinii Masini&1989 Palombo 1989 late Turolian Cava Muraci Romagna Latium Lepus sp. Gatta et al. 2019 post-evaporitic tardiglacial Messinian, Grotta Cagna Mora Cavorso (Jenne) Latium Lepus corsicanus Salari Ciabòt Piedmont Prolagus sorbinii Cavalloetetal.al.2011 1993 as Prolagus michauxi; Angelone & Cavallo 2010 MN13 tardiglacial Riparo Salvini Latium Lepus europaeus Tang & Kotsakis 2008 post-evaporitic Messinian, Grotta Cola Abruzzi Lepus europaeus Di Canzio & Moncucco Piedmont Prolagus sorbinii, Leporidae indet. Angelone etPetronio al. 2011;2001 Colombero et al. 2017 MN13 Grotta Rutina Abruzzi Lepus sp. Saccà 2016 post-evaporitic Messinian, Grotta di Scario level A Verduno Campania Piedmont Lepus europaeus Prolagus sorbinii Ronchitelli Colomberoet etal.al.1998 2014 MN13 Grotta di Castelcivita Campania Lepus europaeus Cioni et al.1979; Masini & Abbazzi 1997 post-evaporitic Messinian, De Giuli 1989 as Prolagus cf. sorbinii and Trischizolagus cf. maritsae; San Sidero 3 Brisighella Apulia Romagna Oryctolagus Prolagus cuniculus sorbinii, Trischizolagus sp. De Giuli 1983 MN13 Masini 1989; Angelone 2007; Rook et al. 2015 with references Oryctolagus cuniculus, Lepus post-evaporitic Messinian, Melpignano Apulia Bologna et al.1899, Major 1875, 19941905; Rütimeyer 1876; López Martínez & Thaler [Casino] Tuscany europaeuselsanus] [Prolagus MN13 1975; Azzaroli et al. 1986; Angelone & Rook 2012 with references Oryctolagus cuniculus, Lepus post-evaporitic Messinian, Cava Donno Apulia Pandolfi et al. 2017 Cava Stingeti Molise corsicanussp., Leporidae indet. Prolagus Cosentino et al. 2018 MN13 Oryctolagus cuniculus, Lepus Avetrana Borro Strolla lower synthem Apulia Pandolfi et al. 2013; Kotsakis et al. in press latest Messinian Tuscany corsicanus Trischizolagus sp. Abbazzi et al. 2008b; Angelone & Rook 2012 (=Cava Sefi) Tana delle Iene Apulia Lepus sp. Conti et al. 2012 Borro Strolla upper synthem earliest MN14 Cava Spagnuolo Tuscany Apulia Prolagus cf. sorbinii Oryctolagus cuniculus Abbazzi et Mecozzi et al. al. 2008b; 2018 Angelone & Rook 2012 Ruscinian- Villányian (=Cava Sefi) Capasso Barbato et al. 1992; Petronio & Sardella 1998 as L. early MN16a, Triversa FU Arcille Ingarano Tuscany Apulia Prolagus aff.cuniculus, Oryctolagus sorbinii Lepus sp. Hürzeler & Engesser 1976; Angelone & Rook 2012 europaeus; for the age: Bedetti & Pavia 2007 Prolagus savagei, Hypolagus Berzi 1967; Berzi et al. 1970; Azzaroli et al. 1986; Čermák 2009; MN16a, Triversa FU Arondelli Sternatia Piedmont Apulia Oryctolagus cuniculus Rustioni et al. 1994 petenyii Angelone & Rook 2012 with references Oryctolagus cuniculus, Lepus Cardamone Apulia Prolagus italicus, Oryctolagus Fondi 1970, Rustioni 1972; et al. 2003Azzaroli et al. 1986; Angelone 2008a, 2013; MN17, Coste S. Giacomo FU Montagnola Senese (=Rosia) Tuscany europaeus valdarnensis Angelone & Rook 2012 Grotte delle Striare Apulia Lepus timidus Di Stefano et al. 1992 MN17, Coste S. Giacomo FU Coste S. Giacomo Latium Prolagus italicus Bona et al. 2015 Oryctolagus cuniculus, Lepus Grotta Romanelli Apulia Prolagus italicus, Oryctolagus Blanc 1920; Fiore 2003; Tagliacozzo 2003; Riga et al. 2003 MN17, Olivola FU Torre di Picchio Umbria corsicanus Girotti et al. 2003; Angelone 2008a, 2013; Angelone & Rook 2012 late Villányian valdarnensis tardiglacial Grotta Paglicci Apulia Lepus corsicanus Vismara 2012 MN17, Olivola FU Casa Sgherri Tuscany Oryctolagus valdarnensis Marcolini et al. 2000; Angelone & Rook 2012; Angelone 2013 transition late MN17, Olivola FU Solava Grotta delle Mura Tuscany Apulia Prolagus aff. sorbinii Lepus corsicanus Angelone Bon & Rook & Boscato 2012 1993; with references; Vismara 2012 Rook & Angelone 2013 Toringian/Holocene Upper Valdarno (several MN17, Olivola FU Grotta di Torre Nave Tuscany Calabria Oryctolagus Lepus sp. valdarnensis Angelone 1972; Bulgarelli & Rook 2012 with Mangano references; 2007; Rook & Angelone 2013 Marra 2009 localities) tardiglacial Grotta di Romito Calabria Lepus sp. Boscato et al. 1998; Mangano 2007; Marra 2009 Biddittu et al. 1979, Cassoli & Segre Naldini 1984; Segre Naldini et al. Holocene latest Villànyian/earliest ~8000-7400 ky Grotta di Latronico 3 Basilicata Oryctolagus cuniculus Dini et al. 2008 Fontana Acetosa Latium ?Oryctolagus valdarnensis 2009; alternatively reported as Oryctolagus cf. O. etruscus or Lepus Biharian sp. Tab. 1 - List of lagomorph-bearing localities of continental Italy detailed with geographical location, age, lagomorph content (=the latest taxonomic attribution of the considered taxa in case of discrepancies), and main related bibliographical references. For Late Pleisto- cene reports of Lepus and Oryctolagus are reported only some selected findings. The fossiliferous sites older than the late Toringian are ordered, as far as possible, in a relative chronological order. The sites of late Toringian age (=Late Pleistocene) are instead grouped according to a geographical criterion (Italian regions). Material that has been lost, unavailable for study, doubtful reports, and obsolete namings are highlighted between square brackets […]. In shaded lgrey are highlighted fossil endemic insular domains now located in continental areas: Tusco-Sardinia PB in light grey; Abruzzi-Apulia PB in darker grey. Faunal Units after Gliozzi et al. (1997) and Pe- tronio and Sardella (1999). Age of fossiliferous sites after Kotsakis et al. (2003), Masini and Sala (2007, 2011), Sala and Masini (2007). shortly before the XVI century (Vigne 1992). Taxonomic overview of fossil As for L. timidus, there are not sufficient data lagomorphs of Italian islands: Sardinia about its Italian distribution during the Holocene. Prolagus Sylvilagus Gray, 1867 The oldest report of the genus Prolagus in The American (Palaearctic and Nearctic) Sardinia dates back to the earliest part of the Late species Sylvilagus floridanus (J.A. Allen, 1890) has Pliocene (P. aff. figaro, Capo Mannu D1; Angelone been introduced in Piedmont (NW Italy) in 1966 et al. 2015) (Fig. 1.j, k). Such remains are likely to for hunting purposes. In Italy, contrarily to other be the “missing link” between the continental spe- western and central-western European countries cies P. sorbinii and Prolagus figaro López Martínez in in which it was also introduced, it became natural- López Martínez & Thaler, 1975, the oldest of the ized, and eventually expanded in both territory and two Sardinian insular endemic species. Actually, P. number in the north-west of the Italian peninsula aff. figaro shows the basic morphology of P. sorbinii, (Gallo et al. 2005 and references therein; Angelici but also the incipient modifications typical of insu- & Spagnesi 2008). At present, Sylvilagus represents lar endemic Prolagus (see Angelone 2005), later fully a pest for agriculture and is a biological competitor developed in P. figaro. No other reports of Prolagus of Lepus europaeus (Gallo et al. 2005). are known in Sardinia before the latest Pliocene/ 170 Angelone C., Moncunill-Solé B. & Kotsakis T. earliest Pleistocene, age of the oldest reports of P. The species turnover at the onset of the figaro in the Monte Tuttavista fissure infillings (An- Dragonara FsC (~0.8/0.7 Ma; inferred from Pa- gelone et al. 2015). Prolagus figaro (Fig. 1.l) character- lombo 2009: fig 2) is the most remarkable among izes the Capo Figari/Orosei 1 subcomplex of the those recorded in the Quaternary of Sardinia (Ab- Nesogoral FC and the Orosei 2 subcomplex of the bazzi et al. 2004; Palombo 2009), and is probably Microtus (Tyrrhenicola) FC (latest Pliocene/earliest related to the climatic changes occurred during the Pleistocene to late Early Pleistocene). Mid-Pleistocene transition (Moncunill-Solé et al. Prolagus sardus (Wagner, 1829) (Fig. 1.m) 2016b). It is very interesting to notice that Prolagus, characterizes instead the Sardinian Middle Pleisto- in countertrend to other Sardinian small mammal cene-Holocene (Angelone et al. 2008, 2015 and ref- lineages, does not experience an abrupt dental size erences therein). This species appears in the Drag- increase in correspondence of the turnover. In fact, onara FsC of the Microtus (Tyrrhenicola) FC, and is in the oldest population of P. sardus can be observed reported in almost all Pleistocene and Holocene fos- a drastic reduction of the dental size (about 30%) siliferous sites of Sardinia. The Monte Tuttavista fis- relative to its ancestor P. figaro (Moncunill-Solé et al. sure infillings yield an impressive record of P. sardus 2016b). Such decrease is not coupled with a body of different ages. Among the best known Sardinian mass decrease: conversely, the body mass increases fossiliferous sites bearing P. sardus, we mention Mon- throughout the lineage P. figaro-P. sardus. te San Giovanni, Grotta dei Fiori, Is Oreris, Bonaria, Prolagus corsicanus Giebel, 1847, reported in Cava Sant’Elia, Su Concali, Bolotana, Tavolara (To- Sardinia by Vaufrey (1929), and in the neighbouring bien 1935; Comaschi Caria 1959, 1968; Gliozzi et al. island of Tavolara by Piveteau (1961), is a synonym 1984; Anfossi et al. 1996; Melis et al. 2013; Zoboli of P. sardus (see Major 1899). & Pillola 2016a, 2016b, 2017; Zoboli 2017). A spe- cial mention is deserved by Grotta del Campanac- Leporidae indet. cio, a cave in which two articulated skeletons of P. Fossil leporids were unknown in Sardinia be- sardus have been recovered (Zoboli & Caddeo 2016). fore the discovery of the Monte Tuttavista karst Prolagus sardus has been reported also from several complex (Rook et al. 2003). The oldest Sardinian Sardinian archaeological sites (e.g. Grotta Corbeddu; leporid record (earliest Late Pliocene, ~3.6 Ma) Dawson 1969; Sondaar et al. 1984), and was the sub- consists of a sole, ill-preserved, non-diagnostic ject of a palaeopathological study of the postcranial tooth from Capo Mannu D1 (Angelone et al. 2015), bones (Zoboli et al. 2018). left in open nomenclature as Leporidae indet. (Fig. The turnover P. figaro/P. sardus is not due to 2.l). Hypotheses about the origin of Leporidae in- the dispersal of a new immigrant in the insular do- det. from Capo Mannu D1 and its relationships main, but to the anagenetic replacement of the first with other Sardinian leporids are illustrated below. species by the latter. The striking morphological and dimensional similarities of the juvenile specimens of Sardolagus Angelone, Čermák, Moncuni- the younger popolations of P. figaro and the adult ll-Solé, Quintana, Tuveri, Arca, & Kotsakis, 2018 specimens of the older samples of P. sardus led An- Sardolagus obscurus Angelone, Čermák, Mon- gelone et al. (2015) to hypothesize that P. sardus may cunill-Solé, Quintana, Tuveri, Arca, & Kotsakis, have been the direct descendant of P. figaro through 2018 (Fig. 2.m) is an endemic insular leporid report- a neoteny phenomenon. It is worth to mention that ed from several fissures infillings of the Monte Tut- López Martínez and Thaler (1975) report P. figaro to- tavista karst complex referable to the Capo Figari/ gether with P. sardus in the fissures of Capo Figari. Orosei 1 and Orosei 2 faunal sub-complexes (Early However, the presence of both species in the old Pleistocene, ~2.1/1.9-1.1 Ma) (Abbazzi et al. 2004; collections of Capo Figari is likely to be due to sam- Angelone et al. 2018). It is quite likely that the Early pling bias or taphonomical mixing. The presence of Pleistocene species S. obscurus may be phylogenet- both P. figaro and P. sardus in the same sample has ically linked to Leporidae indet. from the earliest been reported also from Monte Tuttavista infilling Late Pliocene of Capo Mannu D1. However, such XIp, but it is explained by Angelone et al. (2008) a statement can not be fully supported at present arguing that the infilling of XIp lasted longer than due to the extreme scantiness and bad preservation other studied samples of Monte Tuttavista. state of the remains from Capo Mannu D1. The Fossil Lagomorpha (Mammalia) of Italy 171 mosaic of primitive and advanced morphological lecular studies, even considering the Sardinian hare characters of S. obscurus (see differential diagnosis as an independent species, refer to it as Lepus sp. in Angelone et al. 2018: 510), and the consequent (Suchentrunk et al. 2006; Scandura et al. 2007). difficulty to unravel its phylogenetical affinities, On a morphological basis, Palacios (1998) were responsible for the choice of its specific name individuate the source of Sardinian hares in pop- (Angelone et al. 2018). Its morphological similarity ulations of the coastal region of northern Africa. with Oryctolagus misleaded in the preliminary analy- Molecular data analyses (Scandura et al. 2007) sug- ses Angelone in Rook et al. (2003) and in Abbazzi gest Tunisia as a potential source area for the intro- et al. (2004), who classified the Sardinian leporid as duction. Oryctolagus aff. lacosti. Erecting the new genus Sardo- The earliest Sardinian reports of the hare oc- lagus to include the leporid materials from Monte cur during the Bronze Age (Nuraghe Santu Antine, Tuttavista, Angelone et al. (2018) proposed two al- ~4000-3000 years BP; Masala 2015), and during the ternative hypotheses to explain the mix of archaic Bronze/Iron Age (Santa Niedda, ~3000 years BP; and modern morphological features resulting in an Wilkens 2003, see also Wilkens & Delussu 2003). Oryctolagus-like appearance: (1) either a convergent evolution from an archaeolagine/leporine ancestor, or (2) a selective reversal morphocline from an Oryc- Taxonomic overview of fossil tolagus-like leporine. If S. obscurus is linked to Lepor- lagomorphs of Italian islands: Sicily idae indet. from Capo Mannu D1, the first hypothe- sis is the one which better fits the fossil record. Prolagus De Gregorio (1925) mentions the presence Oryctolagus of Myolagus Hensel, 1856 (=Prolagus) at Grotta del- Oryctolagus cuniculus was introduced by humans la Castellana, a fossil assemblage attributed to the in Sardinia, perhaps in Roman times (Vigne 1992: Spinagallo FC (=“Elephas falconeri” FC in Bonfiglio fig. 2), but its presence is attested with certainty et al. 2002), or to the Maccagnone FC (=“Elephas since the Middle Ages (Delussu 1996; Vigne 1999). mnaidriensis” FC in Bonfiglio et al. 2002). Indeed, the specimen figured in De Gregorio (1925: pl. 3, n. Lepus 33-34) has a size and an overall morphology com- The extant Sardinian hare was at first quot- parable to Prolagus. However, the specimen is lost ed by Rüster (1835, fide Wagner 1841: 439) as L. and the figures are not detailed enough to allow a timidus?. Eventually, Wagner (1841) considered it as confident generic attribution. a separate species with the name of Lepus mediter- raneus. Miller (1912) and Palacios (1998) regarded Hypolagus the Sardinian hare as an independent, endemic spe- A couple of upper jaws and a few teeth of cies. Other authors (see Ellerman & Morrison-Scott a leporid were reported from Monte Pellegrino 1951; Mitchell-Jones et al. 1999) considered the (Monte Pellegrino FC), and classified as Lepus sp. Sardinian leporid as a subspecies of L. capensis, i.e. by De Gregorio (1886). Some decades later, the re- the taxon that, according to Petter (1959, 1961), in- mains were assigned to Lepus (Oryctolagus) (De Gre- cludes almost all Old World hares. It is worth to gorio 1925). Eventually, Thaler (1972) attributed the mention a provisional assignment of the Sardinian remains to the genus Hypolagus. Until 2002, subse- hare as a subspecies of Lepus granatensis Rosenhauer, quent publications reprised this taxonomic assign- 1856 (Hoffmann 1993; Hoffmann & Smith 2005). ment (Bonfiglio et al. 2002 and references therein). In the latest revision of Italian extant mammals, A specific attribution was made by Fladerer and Angelici and Spagnesi (2008 and references there- Fiore (2003), who erected the insular endemic taxon in) considered the Sardinian hare as a subspecies of Hypolagus peregrinus Fladerer & Fiore 2003 (Fig. 2.n). L. capensis. Genetic analyses seem to point out that Basing on the structure of the limbs, Fladerer and the Sardinian hare constitutes a separate species Fiore (2003) argue that H. peregrinus stemmed out (Pierpaoli et al. 1999; Alves et al. 2003). However, from populations of the Early Pleistocene conti- in both cited papers the Sardinian hare continues to nental European species H. brachygnathus older than be addressed as L. capensis mediterraneus. Other mo- 1.2 Ma. Local conditions (insularity and a rocky, 172 Angelone C., Moncunill-Solé B. & Kotsakis T. steep environment) led to forelimb modifications 1987; Bonfiglio et al. 2002). Those remains were resulting in the ambulatory locomotion that char- originally attributed to L. europaeus (ibid.). All Late acterized H. peregrinus (Fladerer & Fiore 2003). As Pleistocene and Holocene leporids of Sicily require for the teeth structure, it is interesting to compare a systematic review, however, in our opinion it is H. peregrinus to other Italian species. Whereas the extremely probable that the materials from Grotta dental morphodynamic indexes of H. petenyii from del Castello may pertain to L. corsicanus, the species Arondelli and H. brachygnathus from Monte La Mesa currently living throughout the island (Angelici & fit into the cloud of the corresponding European Spagnesi 2008). populations, those of H. aff. brachygnathus from Pir- Lepus has also been reported from the II Ne- ro Nord and H. peregrinus clearly stand out, appar- olithic phase of Grotta dell’Uzzo (Holocene; Ta- ently forming a side branch (Angelone 2013: fig. 3). gliacozzo 1993). These observations suggest a close relationship be- tween the peninsular endemic H. aff. brachygnathus from the late early Biharian of south-eastern Italy, Taxonomic overview of fossil but deserve further investigations. lagomorphs of Italian islands: minor islands Oryctolagus The hypothesis commonly accepted by zool- Fossil remains of lagomorphs have been ogists is that the introduction of rabbit in Sicily by discovered in some minor Italian islands. Most of humans probably occurred in Roman times (Flux the remains listed below date back to the first half 1994; Masseti & De Marinis 2008), but according of the XX century and need a taxonomic revision. to Lo Valvo et al. (2014) the oldest remains of Oryc- All the faunal assemblages are balanced (mainland tolagus in the island date back to the Middle Ages. type sensu Dermitzakis & Sondaar 1978), and can However, there are some reports of the presence be referred to the Toringian (see Tab. 2 for details). of O. cuniculus in Sicily in more ancient times. Lag- They reached the islands taking advantage of dif- omorph remains recovered in the upper layer of ferent low stands of the sea level occurred in cor- the Grotta di San Teodoro were classified as Lepus respondence of cold climatic phases. cuniculus by Anca (1867). According to Mangano In the Grotta dei Colombi, on the island of and Bonfiglio (1998), Grotta San Teodoro 2 is a Palmaria (Liguria), Regàlia (1893, 1900) reported level of tardiglacial age, but it is not to be excluded three species of leporids, namely Lepus cuniculus, L. that part of the fossils collected in this layer may be timidus, and L. variabilis in a Late Pleistocene faunal reworked. Masini et al. (2008: fig. 4) report the ap- assemblage. Cigna et al. (1967) assign to L. europae- pearance of O. cuniculus in Sicily during the transi- us the fossils classified as L. timidus (or L. variabilis) tion from the Castello FC to the Holocene faunas. by Regàlia (1893). Villari (1986) reports remains of wild rabbits from Del Campana (1910) reports from Grotta di Grotticella di Porto Palo and from Grotta Corrug- Reale, in the Elba Island (Tuscany), two species of gi, both dated to the early post-glacial, and from Lepus: L. timidus and Lepus meridionalis Géné in Ger- the Basile Caves in strata of the early or middle vais, 1859. The latter species is at present reputed Bronze Age. These evidences suggest that Sicily, as nomen nudum and put in synonymy with L. europaeus. well as southernmost Italy and some Italian minor Mochi (1911) assigns all the remains of leporids islands, represented a refugium for O. cuniculus during from Grotta di Reale to L. europaeus. the Last Glacial Maximum and the earliest Holo- On the island of Pianosa (Tuscan archipela- cene. go), De Stefano (1913) reports the presence of L. timidus together with the rodent Marmota. In more Lepus recent papers, the presence of Marmota has not The genus Lepus is reported in Sicily since been confirmed. Azzaroli (1978 with references) the Late Pleistocene of Grotta del Castello (Regàlia and Azzaroli et al. (1990) argue that the Pianosa 1907a), the type locality of the Castello FC (the fossils belong to two faunal assemblages of differ- most recent Sicilian FC), characterized by the ab- ent age, both assigned to the Late Pleistocene. The sence of endemic elements (Bonfiglio & Kotsakis one containing Lepus is the youngest. Fossil Lagomorpha (Mammalia) of Italy 173 Island Age Faunal complex Locality Species References earliest Late Pliocene Capo Mannu D1 FsC Capo Mannu D1 (=Mandriola) Prolagus aff. figaro, Leporidae indet. López Martínez & Thaler 1975; Angelone et al. 2015 with references (?latest Pliocene-) Early Monte Tuttavista (several karst Rook et al. 2003; Abbazzi et al. 2004; Angelone et al. 2008; Moncunill- Capo Figari/Orosei 1 FsC Prolagus figaro, Sardolagus obscurus Pleistocene infillings) Solé et al 2016b; Angelone et al. 2018 Monte Tuttavista (several karst Rook et al. 2003; Abbazzi et al. 2004; Moncunill-Solé et al. 2016b; late Early Pleistocene Orosei 2 FsC Prolagus figaro, Sardolagus obscurus infillings) Angelone et al. 2018 Monte Tuttavista (several karst Rook et al. 2003; Abbazzi et al. 2004; Angelone et al. 2008; Moncunill- Dragonara FsC Prolagus sardus infillings) Solé et al. 2016b Dragonara FsC Grotta dei Fiori Prolagus sardus Melis et al. 2013 Dragonara FsC Is Oreris Prolagus sardus Gliozzi et al. 1984 Sardinia Middle Pleistocene- Dragonara FsC Monte San Giovanni Prolagus sardus Tobien 1935; Anfossi et al. 1996 ?Holocene Dragonara FsC Dragonara Prolagus sardus Tobien 1935 Dragonara FsC Su Concali Prolagus sardus Zoboli & Pillola 2017 Dragonara FsC Bolotana Prolagus sardus Zoboli & Pillola 2016a Dragonara FsC Grotta del Campanaccio Prolagus sardus Zoboli & Caddeo 2016 Dragonara FsC Cava S. Elia Prolagus sardus Comaschi Caria 1959 Dragonara FsC Bonaria Prolagus sardus Zoboli & Pillola 2016b Dragonara FsC Grotta Corbeddu Prolagus sardus Dawson 1969; Sondaar et al. 1984 latest Pleistocene Dragonara FsC Tavolara Prolagus sardus Tobien 1935; Piveteau 1961; Anfossi et al. 1996 Holocene Bronze Age Nuraghe Santu Antine Prolagus sardus, Lepus mediterraneus Masala 2015 Early Pleistocene Monte Pellegrino FC Monte Pellegrino Hypolagus peregrinus De Gregorio 1886 as Lepus sp.; Thaler 1972; Fladerer & Fiore 2003 Early/Middle Spinagallo FC or Grotta della Castellana ?[Prolagus] De Gregorio 1925 as Myolagus Pleistocene Maccagnone FC Late Pleistocene Castello FC Grotta del Castello Lepus corsicanus Regàlia 1907a; Masini et al. 2008 with references as L. europaeus latest Pleistocene tardiglacial Grotta San Teodoro 2 Oryctolagus cuniculus Anca 1867 as Lepus cuniculus; Mangano & Bonfiglio 1998 Sicily latest Castello FC/Holocene - Oryctolagus cuniculus Masini et al. 2008 with references Pleistocene/Holocene Grotticella di Porto Palo and Holocene early post-glacial Oryctolagus cuniculus Villari 1986 Grotta Corruggi Holocene II Neolithic phase Grotta dell’Uzzo Lepus sp. Tagliacozzo 1993 Oryctolagus cuniculus, Lepus europaeus, Palmaria latest Pleistocene Grotta dei Colombi Regàlia1893, 1900; Cigna et al. 1967 Lepus timidus Elba latest Pleistocene Grotta di Reale Lepus europaeus Del Campana 1910; Mochi 1911 Pianosa latest Pleistocene Lepus timidus De Stefano 1913; Azzaroli 1978 with references; Azzaroli et al. 1990 late Middle Pleistocene Quisisana-Certosa Oryctolagus cuniculus Capasso Barbato & Gliozzi, 1995 Capri Holocene (2ky BP)* Marina Grande Oryctolagus cuniculus Albarella,1992; Masseti & De Marinis 2008 Nisida Holocene (2ky BP)* Oryctolagus cuniculus Barrett-Hamilton 1912 San Domino Cala degli Inglesi and Grotta del (Tremiti "post-thyrrenian" Oryctolagus cuniculus Pasa 1953 Sale Islands) Levanzo latest Pleistocene (Aegadian probably Castello FC Grotta di Cala Genovesi Oryctolagus cuniculus Cassoli & Tagliacozzo 1982 (Mesolithic) Islands) Trocchi & Riga 1995; Masseti & De Marinis 2008 both citing Wilkens 1987 Pantelleria Bronze Age* ?[Oryctolagus cuniculus] who, however, does not report the presence of this species in the text Tab. 2 - List of lagomorph-bearing localities of Italian islands detailed with age, lagomorph content (=the latest taxonomic attribution of the considered taxa in case of discrepancies), and main related bibliographical references. For Late Pleistocene Sardinian reports of Prolagus we list only some selected localities. Material that has been lost, unavailable for study, and doubtful reports/obsolete namings are highlighted between square brackets […]. For the biochronology of continental Plio-Pleistocene fossiliferous sites of Sardinia, we follow the scheme proposed by Palombo (2006, 2009, 2018b) and Palombo and Rozzi (2014). For Sicily, we follow the scheme proposed by Kotsakis (1979), modified by Bonfiglio and Kotsakis (1987) and Bonfiglio et al. (2002). For minor islands we apply the biochronological schemes followed for mainland Italy (see Material and Methods and Tab. 1), and the listed reports are arranged in latitudinal order. *: introduced by humans. In the Quisisana-Certosa fossil site, located in ports of wild rabbits (=not introduced by humans) the island of Capri (Campania), and biochronolog- in some fossiliferous sites of Italian minor islands: ically dated to ~250-150 ky, Capasso Barbato and - Pasa (1953) reports the presence of O. cu- Gliozzi (1995) report the presence of the rabbit, O. niculus in the island of San Domino (Cala degli In- cuniculus. glesi and Grotta del Sale, Tremiti Islands, Apulia), The rabbit is present in Capri also in archae- in coastal deposits of “post-tyrrhenian” age. If we ological deposits, introduced by humans in Ro- consider the Tyrrhenian as a chronostratigraphic man age (~2 ky BP) (Albarella 1992; Masseti & De unit for the Late Pleistocene whose boundaries are Marinis 2008 and references therein). In the neigh- at 130 ky and 11.5 ky (Cita et al. 2005), the Oryc- bouring island of Nisida (Campania), Barrett-Ham- tolagus-bearing fossil sites of the Tremiti Islands can ilton (1912) reports the introduction of O. cuniculus be assigned to the Holocene. This attests to their during the Roman Imperial Age. The presence of arrival into the archipelago during the Last Glacial O. cuniculus, introduced by humans, in an archaeo- Maximum, an indirect evidence of their persistence logical layer of Bronze Age in the Pantelleria Island in the Italian peninsula after MIS 3. (Sicily), was mentioned by Trocchi and Riga (2005) - Cassoli and Tagliacozzo (1982) mention O. and Masseti and De Marinis (2008) purportedly cuniculus in the Grotta di Cala Genovesi, on the island basing on a former report (Wilkens 1987). How- of Levanzo (Aegadian Islands, Sicily). The fossil as- ever, Wilkens (1987) does not mention the rabbit semblage includes Equus hydruntinus Regàlia, 1907, a among the studied remains. taxon characteristic of the Castello FC (Marra 2013 Of particular interest are the following re- and references therein). 174 Angelone C., Moncunill-Solé B. & Kotsakis T. Discussion and conclusions and this means that the lineage originated from the Italian peninsula, and not from SW Europe as pre- Taxonomic and phylogenetic novelties viously thought (López Martínez & Thaler 1975). The principal novelties relative to Italian lag- Moreover, the fact that P. sardus has been recognized omorph taxonomy arisen in the last 15 years are the as a descendant of P. figaro, and not of P. calpensis following: (see López Martínez & Thaler 1975), eliminates the - after several decades, Paludotona, the oldest long-debated enigma relative to the age and mo- known Italian lagomorph genus, known from the dalities of its dispersal to Sardinia (Angelone et al. insular Turolian domain of the Tusco-Sardinia PB, 2015). ceased to be monospecific, due to the description - The discovery of leporids in the Late Plio- of P. minor and P. aff. minor (Angelone et al. 2017), cene and Early Pleistocene of Sardinia (Sardolagus which join Paludotona etruria erected in the 50ties of obscurus and Leporidae indet.) is a quite important the last century (Dawson 1959). novelty, even if at present their phylogenetic affini- - For the first time Italy records stem lago- ties are unknown. morphs. In fact, the features that led some authors - The 8 new lagomorph species described, re- to classify Paludotona as an ochotonid are instead the validated, or individuated without formalization in result of a convergent evolution (Angelone et al. the last 15 years in the Italian territory are endemic. 2017). Several of them are endemic to the Italian peninsula - The teeth morphology of Paludotona indi- (continental endemisms), whereas others are insu- cates that its continental ancestor should be sought lar endemics. At present, Italy records 3 endemic among pre-MN2 European lagomorphs (Angelone species of stem lagomorphs (Paludotona aff. minor, P. et al. 2017). This phylogenetic datum provides a etruria, and P. minor, all of them insular endemics); minimum age constraint to the onset of the Tus- 9 species of ochotonids, of which 5 insular endem- co-Sardinia PB to the earliest Miocene. ics (Prolagus apricenicus, P. imperialis, P. aff. figaro, P. - Recent revisions confirmed the presence figaro, P. sardus) and 2 continental endemics (Prolagus of Prolagus in BV3; the sole specimen from BV3, italicus, P. savagei/P. aff. sorbinii); and 14 species of unfortunately not diagnostic at the species level, is leporids, of which 5 continental endemics (Alilepus among the oldest Prolagus-bearing localities of pen- meini, Hypolagus aff. brachygnathus, Oryctolagus valdar- insular Italy. nensis, O. burgi, Lepus corsicanus) and 2 insular endem- - The presence of Hypolagus in BV3 was in- ics (Hypolagus peregrinus, Sardolagus obscurus/Lepori- stead definitively discarded, as those remains were dae indet.); the status of Trischizolagus sp. must be ascribed to the genus Alilepus (Angelone & Rook verified. 2011). - In older literature, Plio-Pleistocene Prolagus Biochronology findings of the Italian peninsula did not have a clear The oldest lagomorphs recorded in the Ital- specific attribution (as an example, see synonymic ian territory are stem lagomorphs (Paludotona spp.), list of P. aff. sorbinii in Angelone & Rook 2012: 133- exclusive of the early-middle Turolian of the Tus- 134). At the present state of the art, Plio-Pleisto- co-Sardinia PB. Every species of Paludotona char- cene Prolagus of peninsular Italy are assigned to the acterizes a precise time interval, corresponding to species P. savagei/P. aff. sorbinii and to P. italicus. a local biochron (BV0, BV1, BV2). Nevertheless, - The taxonomic mayhem relative to the Val- the biochronological distribution of Paludotona is darno leporid (Oryctolagus valdarnensis), known in not limited to BV0-BV2 (~8.3-6.7 Ma). Its conti- literature since the XIX century, has been ended nental ancestor has to be sought among European revalidating the species, and excluding any affinity species with hourglass-shaped lower third premolar, with the genus Lepus (Angelone & Rook 2012 and a morphology that disappeared in the fossil record references therein). after MN1-2 (Angelone et al. 2017). Thus, Paludoto- - Prolagus aff. figaro, P. figaro, and P. sardus have na possibly existed for ~12.6 Ma. Paludotona, which been recognized as forming an anagenetic lineage became extinct in the earliest Messinian, is the last (Angelone et al. 2015). Prolagus aff. figaro is now of stem lagomorphs to survive. In fact, the last con- reputed to be phylogenetically close to P. sorbinii, tinental stem lagomorphs did not survive after the Fossil Lagomorpha (Mammalia) of Italy 175 early Late Miocene (~9 Ma; López Martínez 2008: polagus disappears from the Italian fossil record for figs 3, 5, 6). An endemic insular stem lagomorph, ~1.4 Ma. When it is recorded again, the new Italian Gymnesycolagus gelaberti Mein & Adrover, 1982, per- species of Hypolagus are phylogenetically related to sisted after the early Late Miocene in Western Me- H. brachygnathus, but as in the case of H. petenyii, there diterranean insular environments (Balearic Islands; is one sole report for every taxon: Hypolagus brachy- Quintana & Agustí 2007 and references therein). gnathus at ~1.8 Ma (earliest Biharian, Tasso FU), H. However, it became extinct in the early Tortonian, aff. brachygnathus at ~1.3-1.2 Ma (late early Bihari- earlier than Paludotona. an, Pirro FU), and H. peregrinus in correspondence Ochotonids appear quite late in the Italian ter- of the Monte Pellegrino FC in Sicily. The termi- ritory with respect to European mainland. In fact, nal part of the Monte Pellegrino FC approximately Prolagus, abundant and widespread in Europe since corresponds to the late early Biharian (see Masini et the Early Miocene, reaches the Abruzzi-Apulia PB al. 2008: fig. 5), with an upper limit at ~0.9 Ma or and the newly formed Italian peninsula only in the ~1 Ma (Masini et al. 2008; Marra 2013). Estimating early Messinian. Even if morphological and palae- the persistence of H. peregrinus in Sicily is more obiogeographical evidences point out that it must complicated. Masini et al. (2008: fig. 6) estimate it in have arrived in the insular domain already in the ear- ~0.7 Ma, which means a dispersal at ~1.8-1.7 Ma. liest Messinian (Angelone 2007), the endemic insu- According to Fladerer and Fiore (2003), the disper- lar Prolagus apricenicus is reported in the fossil record sal of the forerunner of H. peregriinto the island only in the latest Miocene-?earliest Pliocene infill- occurred at ~1.3-1.2 Ma. We know that the earliest ings of Gargano (Abruzzi-Apulia PB). The submer- expansion of the distribution area of H. brachygna- sion of the Gargano Archipelago at the Mio/Plio- thus occur since the latest Villányian (see Čermák cene boundary caused the extinction of the whole 2009), and that H. peregrinus and H. aff. brachygna- insular endemic assemblage of which P. apricenicus thus from Pirro Nord are phylogenetically close and was part, including its possible sister species P. impe- originated from a basal form of H. brachygnathus rialis. Thus, it is possible to estimate that the species (Fladerer & Fiore 2003). Thus, the first hypothesis P. apricenicus lasted ~1.7-1.8 Ma, whereas we do not implies that one of the oldest dispersal waves of H. have enough information to estimate the time span brachygnathus reached southern Italy and Sicily. The lived by P. imperialis. In the territories which formed latter hypothesis implies instead a two-step model: the Italian peninsula after its connection to Euro- H. brachygnathus dispersed to southern Italy first, and pean mainland, the oldest available fossil record of the dispersal to Sicily was delayed at ~1.3-1.2 Ma. A Prolagus (early Messinian) is roughly coincident with dispersal from Europe to southern Italy and Sicily the time of its dispersal. Prolagus thrives in peninsu- at ~1.3-1.2 Ma fits too tight with the proposed phy- lar Italy for ~5Ma, until the early Biharian, and its logenetic setting which excludes from the ascend- absence in the Ruscinian fossil record of mainland ence of H. peregrinus and H. aff. brachygnathus from Italy is probably due to the lack of data relative to Pirro Nord the advanced forms of European H. continental Early Pliocene sediments in Italy. brachygnathus, which appear at ~1.2 Ma. Also leporids disperse to Italy during the As well as Hypolagus, also Oryctolagus dispersed Messinian. The first occurrence of leporid genera several times into the Italian peninsula. The Italian in Italy is always delayed with respect to continental endemic species O. valdarnensis appeared at ~2.1 Europe. The first genus to reach the Italian penin- Ma (MN17, late Villányian, Coste S. Giacomo FU), sula is Alilepus, followed by Trischizolagus. They do much later than the first appearance of the genus not last long: Alilepus meini is exclusive of the early in Europe, and persisted until ~1.2 Ma (late early Messinian of BV3 (~0.3 Ma), and Trischizolagus sp. Biharian, Pirro FU). It is worth to notice that, in probably lasts even less, as it is known only in the Pirro Nord, O. valdarnensis is reported in faunal lists post-evaporitic Messinian of northern-central Italy. together with Hypolagus aff. brachygnathus. However, Leporids re-appear in the Italian peninsula at the most recent review of the lagomorphs of Pir- ~3.2 Ma (early Villányian, Triversa FU; see Masini ro Nord evidences that the two genera were never & Sala 2007) with Hypolagus petenyii. With this sole present in the same fissure (Angelone 2013). This datum, it is impossible to make any inference about datum may be of biochronological importance, but the temporal distribution of H. petenyii in Italy. Hy- it has to be further investigated. Oryctolagus fossil 176 Angelone C., Moncunill-Solé B. & Kotsakis T. remains dated at ~1 Ma (latest early Biharian, Colle is more palaeoecological/palaeoclimatical than bio- Curti FU) must be revised. At ~0.6 Ma, Oryctolagus chronological. re-appears in the Italian fossil record in the earliest Prolagus flourishes in Sardinia for ~3.6 Ma, i.e. phases of the early Toringian (Isernia FU) with O. since the earliest late Pliocene to historical epoch, burgi, another Italian endemic species not directly re- but its record is not continuous. In fact, the Sardin- lated to O. valdarnensis. The extant rabbit, O. cuniculus, ian lagomorph record has a gap of at least ~1.5 Ma appears in the Italian peninsula at ~0.3 Ma (late early between Capo Mannu D1 and the older remains of Toringian, Torre in Pietra FU), delayed with respect Prolagus from Monte Tuttavista, dated at ~2.1/1.9 to the first appearance of O. cuniculus in the Iberian Ma. The minimum time span for the persistence of peninsula (~0.6 Ma; López Martínez 2008). Oryctola- P. figaro is ~1.4 Ma, whereas P. sardus lasts ~0.8 Ma. gus cuniculus persists as a wild species in southernmost The remains of P. figaro and P. sardus are so extremely sectors of Italian mainland during the Last Glacial abundant that it was possible to perform morphodi- Maximum and during the Holocene (at least until mensional analyses of the populations that resulted in ~8-7.4 ky). Sicily and Southern Italian islands (Trem- a preliminary relative chronological arrangement of iti Islands, Sicily, and the Aegadian Islands) host wild the Prolagus-bearing fissure infillings of Monte Tut- rabbits during the latest Pleistocene/early Holocene. tavista (Angelone et al. 2008, 2009). The proposed This represents a very important biochronological order of the karst fissures containing P. figaro is, from and palaeobiogeographical datum, as the rabbit was older to younger: X3-IVm-X4-X5. For the infillings thought to become extinct during the Last Glacial containing P. sardus, instead, the proposed order is, Maximum anywhere except for the Iberian peninsula. from older to younger: IV5-IV20-XIr-VIb6. These Lepus appears much later with respect to Oryc- arrangements are in slight discordance with the rel- tolagus in the Italian peninsula (~1 Ma vs. ~2.1 Ma ative chronological order of the infillings based on respectively), and with a noticeable delay with respect several other mammalian taxa of Monte Tuttavista to continental Europe, where the oldest dispersals of proposed by Palombo (2009: fig. 2). Such results are Lepus occurs at ~2 Ma. The so-called Lepus terraeru- quite encouraging because they demonstrate that lag- brae from the lower levels of Monte Peglia (late early omorphs can be used for biochonological purposes Biharian, Colle Curti FU) is the oldest reliable report with a very high resolution (Angelone 2008b; Ange- of the genus Lepus in Italy (~1.0 Ma) or slightly earli- lone et al. 2008, 2009). The biochronological history er. The report of Lepus in the Slivia FU is not certain, of leporids in Sardinia follow the same pattern of in the Ponte Galeria FU there is only a report of an ochotonids, except that it is shorter. In fact, after undetermined leporid, and in the Isernia FU there is the first report at ~3.6 Ma (Leporidae indet.), lep- no Lepus report. The extant species of Lepus appear orids “appear” again in the fossil record at ~2.1/1.9 in Italy between Isernia FU and Fontana Ranuccio Ma (Sardolagus obscurus). Studied remains of Sardola- FU. This means a discontinuity of ~0.6 Ma in the gus cover ~1 Ma. However, it is quite likely that: (1) Italian record of Lepus. According to the available slightly younger leporid remains recovered in the fossil record (which is pending a massive systemat- Monte Tuttavista infillings pertain too to Sardolagus, ic revision, though), the oldest reliable report of the and that (2) Leporidae indet. from Capo Mannu D1 extant species of Lepus pertains to L. corsicanus, fol- is related to Sardolagus. In this case, the persistence of lowed in the earliest late Toringian by L. europaeus, this genus would be noticeably longer (~3.6-0.8 Ma). and by L. timidus. This latter species appears in Italy in correspondence of the MIS 2 (24-11.6 ky). In the Open questions and further investigations Castello FC (late MIS 3/early MIS 2, ~25 ky), the As usual, new data solve some questions but genus Lepus appears for the first time in Sicily. open new problems or new perspectives of study. We The Asian genus Ochotona, the sole extant survi- envisage a focus of the future studies about Italian vor among the once flourishing family Ochotonidae, fossil lagomorphs in the following topics: is reported twice in NE Italy. Those appearances are - the lower Messinian Prolagus cf. apricenicus related to the expansion of the distribution area of from Capo di Fiume (Abruzzi-Apulia PB) is one of Ochotona occurred in two different cold phases: MIS the very few articulated, almost complete skeletons 12/MIS 10 (474-427 ky or 364-334 ky), and MIS4 of Prolagus available at present. A morpho-function- (74-59 ky). The importance of this datum, though, al study would clarify several questions about the Fossil Lagomorpha (Mammalia) of Italy 177 life-style of Prolagus. The analysis of its teeth, now Trieste, Italy). Journal of Archaeological Sciences, 27: 1173- hidden, will provide insights about the taxonomic 1186. position of this specimen, inferred on a palaeogeo- Abbazzi L., Angelone C., Arca M., Barisone G., Bedetti C., Delfino M., Kotsakis T., Marcolini F., Palombo M.R., graphical basis (Mazza et al. 1995). Pavia M., Piras P., Rook L., Torre D., Tuveri C., Valli A. - Trischizolagus sp. from the evaporitic Messin- & Wilkens B. (2004) - Plio-Pleistocene fossil vertebrates ian of Brisighella needs to be studied in order to of vertebrates of Monte Tuttavista (Orosei, Eastern understand its phylogenetic affinity with other Eu- Sardinia, Italy), an overview. Rivista Italiana di Paleontologia ropean species, and to reconstruct the dispersal path e Stratigrafia, 110: 681-706. followed by the genus to achieve its pan-European Abbazzi L., Delfino M., Gallai G., Trebini L. & Rook L. (2008a) - New data on the vertebrate assemblage of Fiu- distribution. me Santo (north-west Sardinia, Italy), and overview on - Apart from the taxonomic revision, neces- the Late Miocene Tusco-Sardinian palaeobioprovince. sary to confirm that Prolagus aff. sorbinii and P. savagei Palaeontology, 51: 425-451. pertain to the same species, it will be necessary to Abbazzi L., Benvenuti M., Ceci M.E., Esu D., Faranda C., clarify their relationship with P. depereti López Martín- Rook L. & Tangocci F. (2008b) - The end of the La- ez in López Martínez & Thaler 1975, distributed in go-Mare time in the SE Valdelsa basin (Central Italy): in- terference between local tectonism and regional sea-lev- the MN15 of southern France (See Angelone & el rise. Geodiversitas, 30: 611-63 Rook 2012: 140, 141). The result will have noticeable Albarella U. (1992) - La fauna. In: Arthur P. (Eds) - L’isola e il palaeobiogeographical repercussions (see Angelone santo. La Chiesa di San Costanzo alla Marina Grande di et al. 2019). Capri, scavi 1990: 53-58. Editoriale Scientifica, Napoli. - The reports of remains attributed to the ge- Alberdi M.T., Cerdeño E., López Martínez N., Morales J. nus Lepus in Italy need a complete revision, necessary & Soria M.D. (1997) - La fauna villafranquiense de El Rincón-1 (Albacete, Castilla-La Mancha). Estudios Geo- to understand the distribution of the genus in the lógicos, 53: 69-93. Italian Pleistocene. Allen J.A. (1890) - Descriptions of a new species and a new These tasks are important not only for the ac- subspecies of the genus Lepus. Bulletin of the American counting of palaeobiodiversity and the updating of Museum of Natural History, 3: 159-160. biochronological data, but also to better understand Alves P.C., Ferrand N., Suchentrunk F. & Harris D.J. (2003) the dispersal events and the palaeobiogeographical - Ancient introgression of Lepus timidus mtDNA into L. granatensis and L. europaeus in the Iberian Peninsula. framework of western central Europe during the late Molecular Phylogenetics and Evolution, 27: 70-80. Neogene-Pleistocene. Ambrosetti P., Bartolomei G., De Giuli C., Ficcarelli G. & Torre D. (1979) - La breccia ossifera di Slivia (Aurisina Acknowledgements: We are thankful to the reviewers S. Čermák - Sistiana) nel Carso di Trieste. Bollettino della Società Pale- and A. Savorelli for their careful revision of the manuscript and for ontologica Italiana, 18: 207-220. the helpful insights and suggestions, and to the Editors C. Lombardo Anca F. (1867) - Paleoetnologia sicula. Giovanni Lorsnaider, and L. Rook for the quick and clear handling of the manuscript. S. Čermák in particular is acknowledged also for sharing helpful personal Palermo, 16 pp. knowledge and data. Anfossi G., Santi G. & Ticozzi M.G. (1996) - I resti di lago- CA has been supported by a Visiting Professor grant of the morfi, insettivori e roditori quaternari della Sardegna President’s International Fellowship Initiative of the Chinese Acad- e della Corsica del Museo di Scienze Naturali di Pavia emy of Science; the Spanish Agencia Estatal de Investigación and (Lombardia-Italia): studio osteologico. Atti Ticinensi di the European Regional Development Fund of the European Union Scienze della Terra, 38: 149-169. (CGL2016-76431-P); the CERCA Program, Generalitat de Catalunya; Angelici F.M. & Spagnesi M. (2008) - Ordine Lagomorpha a Grant to Department of Science, Roma Tre University (MIUR-Italy Brandt, 1855. In: Amori G., Contoli L. & Nappi A. Dipartimenti di Eccellenza, ART. 1, C. 314-337 L. 232/2016). (Eds) - Fauna d’Italia, Mammalia II. 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