ON SUBSPECIFIC TAXONOMY OF MICROTUS SAVII (RODENTIA, ARVICOLIDAE)
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1 Hystrix It. J. Mamm. (n.s.) 14 (1-2) (2003): ON SUBSPECIFIC TAXONOMY OF MICROTUS SAVII (RODENTIA, ARVICOLIDAE) LONGINO CONTOLI Via Arno, 38, Roma, Italy. RIASSUNTO - Sulla tassonomia sottospecifica di Microtus savii (Rodentia, Arvicolidae). Viene riveduta e riassunta la situazione tassonomica sottospecifica di Microtus (Terricola) savii, anche tramite la descrizione di due nuovi taxa: Microtus (Terricola) savii tolfetanus, dei Monti della Tolfa e Microtus (Terricola) savii niethammericus, del Gargano. Parole chiave: Tassonomia, sottospecie, Microtus (Terricola) savii, Italia. Microtus (Terricola) savii (de Sélys Longchamps, 1838) is probably the most widespread species of its subgenus in Italy; moreover, it is near the sole known Terricola species occurring in the middle southern range of the Italian peninsula, with the exception of M. (T.) brachycercus in Calabria (Galleni, 1995; Galleni et al., 1992, 1994 and 1998) and a problematic population of M. (T.) multiplex in the Abruzzi highlands (Amori et al., 1986); note that the living populations of Salento (Puglia) show some character morphae (e. g., in the M 1 fore part) quite similar to those of the fossil taxon M. (T.) tarentinus Brunet Lecomte, 1988 from Villa Castelli (Taranto, Puglia). The differences among local populations of M. (T.) savii, even if quite important among biotopes at the boundaries of the species range, usually appeared to be continuous and gradual (Contoli, 1980; Krapp, 1982). In fact, the frequency of the persimplex morpha of M 3 increases towards the southernmost boundaries of the geographic range (e. g. Contoli, 1980 and Battisti, 1990); even the proportions in the fore complex of M 1 (e.g. Brunet Lecomte and Chaline, 1990) seem to change progressively, at least along the Adriatic coast, showing an increase of some mediterranean morphae (e. g., Pasa, 1951 and 1959). We still lack an exhaustive caryologic and genetic knowledge in this field. Nevertheless, there are some local differencies that seem to be relatively more sharp and discrete, with a possible systematic meaning at the subspecific rank. It is well known that the importance of subspecific taxonomy is still questionable; nevertheless, in the specific context of M. (T.) savii some forms were signalled under various taxonomic meanings. Thereafter, one of them was considered at species level, when others were not adequately characterized from a taxonomic perspective.trying to summarize and reviewing this situation, this work has a nearly exclusively heuristic pur- 107
2 Contoli pose, in order to stimulate new researches in this field. THE SUBSPECIES of Microtus (Terricola) savii (de Sélys Longchamps, 1838). Microtus (Terricola) savii savii (de Sélys Longchamps, 1838). SYNONIMY: Pitymys savii savii, Ellerman and Morrison-Scott, Checklist of Palaearctic and Indian Mammals 1758 to 1946, Trustees of Brit. Mus.: 688. Microtus (Pitymys) savii savii, Krapp, in Niethammer and Krapp, Handbuch der Säugetiere Europas, 2/1: 432. Microtus savii savii, Galleni, Stanyon, Tellini, Giordano and Santini, Cytogenetics and Cell Genetics, 59: LOCUS TYPICUS: surroundings of Pisa (Italy). DISTRIBUTION: Italian peninsula, perhaps with exception of the Calabria and Puglia subpeninsulae. Microtus (Terricola) savii nebrodensis (Minà Palumbo, 1868). SYNONIMY: Arvicola nebrodensis, Minà- Palumbo Ann. Agric. Sc. Sicil. (sez. 2): 61. Pitymys nebrodensis, Miller, Catalogue of the Mammals of Western Europe, Trustees of Brit. Mus.: 770. Pitymys savii nebrodensis, Miller, Proc. Soc. Biol. Washington, 26: 81. Pitymys savii nebrodensis, Ellerman and Morrison-Scott, Checklist of Palaearctic and Indian Mammals 1758 to 1946, Trustees of Brit. Mus.: Microtus (Pitymys) savii nebrodensis, Krapp, in Niethammer and Krapp, Handbuch der Säugetiere Europas, 2/1: 432. LOCUS TYPICUS: Madonie mountains, Sicily. DISTRIBUTION: Sicily. Microtus (Terricola) savii tolfetanus, subspecies nova. DERIVATIO NOMINIS: from the popular name of people inhabiting the locus typicus of this subspecies. SYNTYPI: 25 skulls preserved in microtheriological collection of Longino Contoli), presently at the Zoological Museum, Department of Animal and Human Biology, University of Rome La Sapienza : La Farnesiana locality, included Allumiere station (Contoli legit), Bufalareccia and Saracinesca (Aloise legit) sites; ; specimens nr.: 0a 0f, 378, 379, 425, 427, 675, 731, 732, 761, 766, 767, 879, 885, 1211, 1124, 2861, 2864, 2866, 2868, Figurated in Contoli, LOCUS TYPICUS: La Farnesiana, Tolfa hills, Allumiere, Roma Province. DISTRIBUTION: Monti della Tolfa (Tolfa hills) and their surroundings, a volcanic district NW of Rome, in the antiapenninic and coastal range of NE Latium. DESCRIPTION AND DIAGNOSTIC CHARAC- TERS: a relatively big Savi s pine vole (22,9 < CB < 26,8 mm (Tab. 1; Contoli, 1980), with some uncommon morphological features. In fact, three M 3 cha- 108
3 On subspecific taxonomy of Microtus savii racters mainly show morphae that are usually uncommon in the nominal subspecies: Typica, joined 2 th fields and M 3 > M 2 in length ; while morphological variation in the species is customarily quite gradual, a marked step of size decreasing is observed at the borders of the subspecies range. Nevertheless, in the above area the characteristic M. (T.) savii caryotype was recorded (Contoli et al., 1978). Microtus (Terricola) savii niethammericus, subspecies nova. DERIVATIO NOMINIS: from J. Niethammer, which first recorded some outstanding features of demes from the Gargano. TAXONOMIC NOTE: due to an unfortunate misprint, the taxon was firstly indicated as Microtus (Terricola) savii niethammeri (Contoli, 2000); note that the subspecific name niethammeri was already utilized for a subspecies of Microtus multiplex [Microtus (Terricola) multiplex niethammeri Brunet - Lecomte and Volobouev, 1994]; moreover, according to ICZN (2000, art. 16.4), the name Microtus (Terricola) savii niethammeri is not available because not accompanied by the fixation of name - bearing types (the paper was submitted and accepted before 2000). For this reason, in the present paper I proposed to name this subspecies as Microtus (Terricola) savii niethammericus ssp. nova. SYNTYPI: series of 21 specimens; J. Niethammer legit, 1980 (1981. Säugetierk. Mitt., 29: 45-48; Museum A. Koenig, Bonn: ). LOCUS TYPICUS: Foresta Umbra woodland and its surroundings (m.t Sant Angelo etc.). DISTRIBUTION: Foresta Umbra, on Gargano highlands, Apulia, Italy. According Galleni et al. (1998) the range of this form could extend southwards, at least as far as Metaponto (Potenza). DESCRIPTION AND DIAGNOSTIC CHARAC- TERS: according to Niethammer (1981), a middle sized Savi s pine vole; main differences with respect to M. s. savii are in relation to the breast (2 fore and 1 hind pairs), the M 3 often showing a Table 1 - Skull measurements (0.1 mm) and character s prevailing forms (in %) of M. (T.) savii tolfetanus (see Contoli, 1980) at the locus typicus. N: number of specimens; min: minimum value; x: average; max: maximum value; SD: standard deviation. 109
4 Contoli trend towards the Typica morpha, highly variable M 1, acrocentric "X" chromosome (2N = 54; n.f. = 60; elsewhere, this caryotype was recorded for this species in a specimen from the surroundings of Metaponto, not so far from the Gargano); it is noticeable that, in the neighbouring districts of Taranto and Foggia, M 3 "simplex" or, even, with a good frequence, "persimplex" morphae were found (Battisti, 1990), while M 1 seems more constantly of a "mediterranean" type (cfr. Brunet- Lecomte and Chaline, 1990); moreover, while the morphology of the fore complex of M 1 seems to vary continuously from the north to the south, (Contoli, unpub. data) according to a nearly hysometric trend, the Gargano demes seems to be less aligned to the general pattern. Unfortunately, there are no hybridological nor genetical data on this form. So, it is convenient to follow Niethammer (1981) and Krapp (1982) and rely the above Gargano demes to M. (T.) savii. KEY OF THE SUBSPECIES 1 - Acrocentric X chromosome (NF = 60); normally, 6 breasts; morphae of M 3 "simplex" to "typica": niethammericus. - Metacentric "X" chromosome (NF = 62); normally, 4 breasts: M 3 mainly in the "persimplex" morpha; braincase quite high and rounded; skull quite short, expecially in its post rostral portion: nebrodensis. - M 3 mainly not in the "persimplex" morpha; skull not as above: M 3 "simplex" to "typica", up to complex s. l.; skull quite large (normally: CB > 24 mm, DS > 7 mm, Md > 15 mm): tolfetanus. - M 3 normally "simplex"; skull smaller (normally: CB < 24 mm, DS < 7 mm, Md < 15 mm): savii. ACKNOWLEDGEMENTS This work could not be accomplished without the kind contribution of many friends, among which P. Agnelli, G. Aloise, G. Amori, C. Battisti, P. Brunet Lecomte, G. Carpaneto, M. Cristaldi, R. Hutterer, T. Kotsakis and its coworkers, F. Krapp, A. Nappi, J. Pignatti, C. Prigioni, L. Santini, the staff of the Museum A. Koenig (Bonn) and some contributors to the microtheriological collection of Longino Contoli. REFERENCES Amori G., Cristaldi M. and Contoli L Sui Roditori (Gliridae, Arvicolidae, Muridae) dell' Italia peninsulare ed insulare in rapporto all' ambiente bioclimatico mediterraneo. Animalia, 11(1-3): (1984). Battisti C Sistemi trofici "Mammiferi terragnoli - Tyto alba (Scopoli, 1769) nel Salento: aspetti faunistico - biogeografici ed ecologici. Tesi di laurea, Fac. Sc. M.F.N., Un. "La Sapienza", Roma, 90 pp. Brunet Lecomte P Les campagnols souterrains (Terricola, Arvicolidae, Rodentia ) actuels et fossiles d Europe occidentale. Thèse de Doctorat, Univ. De Bourgogne, 146 pp Brunet Lecomte P Description de trois nouvelles espèces fossiles de campagnols souterrains du Quaternaire d Europe occidentale (Rodentia, Arvicolidae). C. R. Acad. Sci. Paris, 307 (s. II):
5 On subspecific taxonomy of Microtus savii Brunet-Lecomte P. and Chaline J Rélations phylogénétiques et évolution des campagnols souterrains d'europe (Terricola, Arvicolidae, Rodentia). C. R. Acad. Sci. Paris, 311 (2): Brunet Lecomte P. and Volobouev V Comparative morphometry and cytogenetic of Microtus (Terricola) multiplex (Arvicolidae, Rodentia) of the western French Alps. Zeitschr. Säugetierkunde, Hamburg und Berlin, 59: Contoli L., Les Pitymys de l'italie centrale-occidentale (Rodentia, Arvicolidae). Données craniometriques et dentaires. Mammalia, 44(3): Contoli L Rodents of Italy: species richness maps and Forma Italiae. Hystrix, It. J. Mamm. (n.s.), 11 (2): Contoli L., Amori G. and Cristaldi M Geographic variability of Pitymys in central Italy. II Congr. Theriol. Internaz., Brno; R. Orbtel, C. Folk, J. Pellantova eds., Brno: 100. Galleni L Speciation in the Savi pine vole, Microtus savii (De Sel. L.) (Rodentia, Arvicolidae): a theoretical biology approach. Boll. Zool., 62: Galleni L., Stanyon R., Contadini L. and Tellini A Biogeographical and karyological data of the Microtus savii group (Rodentia, Arvicolidae) in Italy. Bonn. Zool. Beitr., 47 (3 4): Galleni L., Stanyon R., Tellini A., Giordano G. and Santini L Karyology of the Savi pine vole: Microtus savii (De Sélys Longchamps, 1838) (Rodentia, Arvicolidae): C. G. DA/DAPI and Alu - 1 bands. Cytogenetics and Cell Genetics, 59: Galleni L., Tellini A., Stanyon R., Cicalò A. and Santini L Taxonomy of Microtus savii (Rodentia, Arvicolidae) in Italy: cytogenetic and hybridization data. J. Mammalogy, 75(4): Krapp F., Microtus savii (de Sélys - Longchamps, 1838) - Italienische Kleinwuehlmaus. In: J. Niethammer and F. Krapp (eds): Handbuch der Säugetiere Europas, 2/1 (Nagetiere): Niethammer J., Über Microtus (Pitymys) savii (De Sèlys Longchamps, 1838) vom Monte Gargano, Italien. Saugetierk. Mitt., 29: Pasa A Alcuni caratteri della Mammalofauna pugliese. Mem. Biog. Adriat., 2: Pasa A Alcuni caratteri delle Mammalofaune sicule. Mem. Mus. Civ. St. Nat. Verona, 7:
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