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Suppl. Al 52 2005 5-135 Finito Di Stampare ISSN: 0365-1576 Dicembre 2006

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Suppl. Al 52 2005 5-135 Finito Di Stampare ISSN: 0365-1576 Dicembre 2006 ISSN: 0365-1576 ATTI DEL MUSEO CIVICO DI STORIA NATURALE DI TRIESTE SUPPLEMENTO AL VOL. 52 - 2005 MUSEO CIVICO DI STORIA NATURALE - TRIESTE In memory of Augusto Cesare Ambrosi (Casola in Lunigiana, 30.V.1919 – Firenze, 29.III.2003) eminent Tuscan historian and archaeologist, discove- rer of the stelae-statues of Lunigiana and of Speleomantes ambrosii. In memoria di Augusto Cesare Ambrosi (Casola in Lunigiana, 30.V.1919 – Firenze, 29.III.2003) eminente storico e archeologo toscano, scopritore delle statue-stele lunigianesi e dello Speleomantes ambrosii. DIRETTORE RESPONSABILE: Sergio Dolce Autorizzazione del Tribunale di Trieste N. 491 Reg. Periodici del 16.02.1976 Finito di stampare Stampa Tipo/Lito Astra S.r.l. Atti Mus. Civ. Stor. Nat. Trieste Suppl. al 52 2005 5-135 Finito di stampare ISSN: 0365-1576 dicembre 2006 A REVIEW OF SYSTEMATICS, TAXONOMY, GENETICS, BIOGEOGRAPHY AND NATURAL HISTORY OF THE GENUS SPELEOMANTES DUBOIS, 1984 (AMPHIBIA CAUDATA PLETHODONTIDAE) BENEDETTO LANZA (1), CHRISTIAN PASTORELLI (2),PAOLO LAGHI (3),ROBERTA CIMMARUTA (4) (1) Dipartimento di Biologia Animale e Genetica & Museo di Storia Naturale (sezione zoologica "La Specola") – Università di Firenze – Via Romana, 17 – 50125 Firenze – Italia <[email protected]> (2) Via Cerchia di Sant’Egidio, 2205 – 47023 Cesena (FC) – Italia <[email protected]> (3) Via Bruno Costante Garibaldi, 22 – 47100 Forlì – Italia <[email protected]> (4) Dipartimento di Ecologia per lo Sviluppo Economico Sostenibile – Università della Tuscia – Largo dell’Università – 01100 Viterbo – Italia <[email protected]> Abstract - A review of systematics, taxonomy, genetics, biogeography and natural history of the genus Speleomantes Dubois, 1984 (Amphibia Caudata Plethodontidae). Outside the American continent the family Plethodontidae includes only the south Korean Karsenia koreana Min, Yang, Bonett, Vieltes, Brandon et Wake, 2005, and the European genus Speleomantes Dubois, 1984, whose systemat- ics and taxonomy have rather recently been object of a morphologic and genetic review, which allowed to recognize 3 continental species [S. strinatii (Aellen, 1958); S. ambrosii ambrosii (Lanza, 1955) and S. ambrosii bianchii Lanza, Cimmaruta, Forti, Bullini et Nascetti, 2005; S. italicus (Dunn, 1923)] and 4 Sardinian ones [S. flavus (Stefani, 1969); S. supramontis (Lanza, Nascetti et Bullini, 1986); S. imperialis imperialis (Stefani, 1969) and S. imperialis sarrabusensis Lanza, Leo, Forti, Cimmaruta, Caputo et Nascetti, 2001); S. genei (Temminck et Schlegel, 1838) with subsp A and subsp. B]. Until recently the knowledge of Speleomantes’ eco-ethology has been widely lacunose and specific research on this topic started only in the nineties. The present paper summarizes the current knowledge on the European plethodontid salamanders, genus Speleomantes. The authors deal with systematics, taxonomy, geo- graphical and altitudinal distribution, genetics (two keys to species and subspecies are given: one based on morpholo- gy and geographical distribution, the other based on genetic characters), biogeography, ecology (habitats, herpeto- coenosis, predators, parasites, food), ethology (feeding behaviour, activity, habitat use, displacement, antipredatory adaptations, communication), reproduction (sexual dimorphism, gametogenesis, mating behaviour, oviparity, ovovi- viparity, parental cares), development, population dynamics, conservation (abundance, threats, conservation guide- lines), as well as with research prospects. The quoted bibliography includes 564 titles. Key words: Amphibia, Plethodontidae, Speleomantes, systematics, taxonomy, keys, genetics, biogeography, biology, conservation, France, Italy. Riassunto: Revisione della sistematica, tassonomia, genetica, biogeografia e storia naturale del genere Speleomantes Dubois, 1984 (Amphibia Caudata Plethodontidae). Al di fuori del continente americano la famiglia Plethodontidae conta solo la specie sud coreana Karsenia koreana Min, Yang, Bonett, Vieltes, Brandon et Wake, 2005, e il genere europeo Speleomantes Dubois, 1984, la cui sistemati- ca e tassonomia, oggetto di una relativamente recente revisione su basi genetiche e morfologiche, ha portato a riconoscere 3 specie continentali [S. strinatii (Aellen, 1958); S. ambrosii ambrosii (Lanza, 1955) e S. ambrosii bianchii Lanza, Cimmaruta, Forti, Bullini et Nascetti, 2005; S. italicus (Dunn, 1923)] e 4 sarde [S. flavus (Stefani, 1969); S. supramontis (Lanza, Nascetti et Bullini, 1986); S. imperialis imperialis (Stefani, 1969) e S. imperialis sarrabusensis Lanza, Leo, Forti, Cimmaruta, Caputo et Nascetti, 2001); S. genei (Temminck et Schlegel, 1838) con le sottospecie A e B]. Sino a poco tempo fa le conoscenze sull’eco-etologia dello Speleomantes sono rimaste ampia- mente lacunose e ricerche approfondite in proposito sono state avviate solo a partire dagli anni ‘90. Il presente con- tributo riassume le attuali conoscenze sui Pletodontidi europei del genere Speleomantes. Gli autori trattano sistemati- ca, tassonomia, distribuzione geografica ed altitudinale, genetica (vengono fornite due chiavi per l’identificazione delle specie e sottospecie: una basata sulla morfologia e la distribuzione geografica, l’altra basata sulle caratteristiche genetiche), biogeografia, ecologia (habitat, erpetocenosi, predatóri, parassiti, dieta), etologia (comportamento pre- datòrio, attività, uso dell’habitat, spostamenti, adattamenti antipredatòrî, comunicazione), riproduzione (dimorfismo sessuale, gametogenesi, corteggiamento, oviparità, ovoviviparità e cure parentali), sviluppo, dinamica di popolazione, conservazione (abbondanza, minacce, linee-guida per la conservazione) e prospettive di ricerca. La bibliografia citata comprende 564 titoli. Parole chiave: Amphibia, Plethodontidae, Speleomantes, sistematica, tassonomia, chiavi, genetica, biogeografia, biologia, conservazione, Francia, Italia. 6 BENEDETTO LANZA, CHRISTIAN PASTORELLI , PAOLO LAGHI , ROBERTA CIMMARUTA CONTENTS 1. INTRODUCTION ....................................................................... p. 7 2. MATERIALS AND METHODS ................................................ p. 7 3. SYSTEMATICS, TAXONOMY AND GENETICS .................. p. 8 3.1 Systematics, taxonomy and geographical distribution..…….... p. 8 3.2 Altitudinal distribution......................................................……. p. 12 3.3 Morphological-geographical key to species and subspecies... p. 13 3.4 Genetics..............................................................................…... p. 19 3.4.1 Diagnostic loci and biochemical key.............................…... p. 19 3.4.2 Genetic differentiation (allozymes and mitochondrial DNA) p. 24 3.4.3 Karyology and repetitive DNA.....................................….... p. 26 4. BIOGEOGRAPHY...................................................................... p. 27 5. ECOLOGY................................................................................... p. 37 5.1 Habitats .................................................................................. p. 37 5.2 Herpetocoenosis ...................................................................... p. 41 5.3 Predators ................................................................................. p. 42 5.4 Parasites .................................................................................. p. 42 5.5 Food ......................................................................................... p. 43 6. ETHOLOGY................................................................................ p. 45 6.1 Feeding behaviour ................................................................... p. 45 6.2 Activity, habitat use, and displacement ................................... p. 47 6.3 Antipredatory adaptations ....................................................... p. 63 6.4 Communication........................................................................ p. 65 7. REPRODUCTION AND DEVELOPMENT............................. p. 67 7.1 Sexual dimorphism................................................................... p. 67 7.2 Gametogenesis......................................................................... p. 69 7.3 Mating behaviour .................................................................... p. 71 7.4 Oviparity, ovoviviparity and parental cares ............................ p. 73 7.5 Development ............................................................................ p. 78 8. POPULATION DYNAMICS ...................................................... p. 81 9. CONSERVATION........................................................................ p. 83 9.1 Abundance ............................................................................... p. 83 9.2 Threats and conservation guidelines ....................................... p. 86 10. RESEARCH PROSPECTS....................................................... p. 90 11. ADDENDUM.............................................................................. p. 91 ACKNOWLEDGMENTS ............................................................... p. 91 DISTRIBUTIONAL MAPS AND SYNONYMS........................... p. 93 COLOUR PLATES (Figs. 8-23) ..................................................... p. 107 BIBLIOGRAPHY............................................................................ p. 115 A REVIEW OF SYSTEMATICS, TAXONOMY, GENETICS, ecc. 7 1. INTRODUCTION Plethodontidae is the largest family of the order Urodela, including about 362 species in 28 genera, corresponding to nearly 70% of the 535
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