Taxonomic Revision of the Cretan Fauna of the Genus Temnothorax Mayr, 1861 (Hymenoptera: Formicidae), with Notes on the Endemism of Ant Fauna of Crete
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ANNALES ZOOLOGICI (Warszawa), 2018, 68(4): 769-808 TAXONOMIC REVISION OF THE CRETAN FAUNA OF THE GENUS TEMNOTHORAX MAYR, 1861 (HYMENOPTERA: FORMICIDAE), WITH NOTES ON THE ENDEMISM OF ANT FAUNA OF CRETE SEBASTIAN SALATA1*, LECH BOROWIEC2, APOSTOLOS TRICHAS3 1Institute for Agricultural and Forest Environment, Polish Academy of Sciences, Bukowska 19, 60-809 Poznań, Poland; e-mail: [email protected] 2Department of Biodiversity and Evolutionary Taxonomy, University of Wrocław, Przybyszewskiego 65, 51-148 Wrocław, Poland; e-mail: [email protected] 3Natural History Museum of Crete, University of Crete, Greece; e-mail: [email protected] *Corresponding author Abstract.— We revise the Cretan species of the ant genus Temnothorax Mayr, 1861. Sixteen species are recognized, including seven new species which are possiblyendemic to Crete: T. crassistriatus sp. nov., T. daidalosi sp. nov., T. ikarosi sp. nov., T. incompletus sp. nov., T. minotaurosi sp. nov., T. proteii sp. nov., and T. variabilis sp. nov. A new synonymy is proposed, Temnothorax exilis (Emery, 1869) =Temnothorax specularis (Emery, 1916) syn. nov. An identification key to Cretan Temnothorax, based on worker caste is given. We provide a checklist of ant species described from Crete and discuss their status, distribution and endemism. Ë Key words.— Key, checklist, Myrmicinae, new species, Mediterranean Subregion, new synonymy INTRODUCTION 2000 mm in the high White Mountains range (Lefka Ori) (Grove et al. 1993). Temperature on mountains Crete is the fifth largest island in the Mediterranean seems to fall at a rate of about 6°C per 1000 m (Rack- ham & Moody 1996). Above 1600 m most of the precipi- Sea and the biggest island of Greece. It’s situated on tation falls as snow that covers the ground from late the southernmost part of the Aegean Sea, with an area October until May (or locally even July). The high of 8400 km2 (Vogiatzakis et al. 2008). The landscape is mountains are limestone. Crete shows significant tem- mountainous, defined by 15 high mountain ranges perature gradients in west-east and north-south direc- crossing from west to east, three of them exceeding tions (Vogiatzakis et al. 2008). 2000 m of altitude. The main island is also surrounded Crete, next to Cyprus, is one of two large Mediter- by about 36 larger or smaller offshore islets. The moun- ranean islands that has not been subject of intense tains create huge regional variations in Crete’s myrmecological studies. Almost all knowledge of Mediterranean climate, with rain-excesses and rain myrmecofauna of this island comes from papers pub- shadows. In general, aridity increases from west to lished before World War II, including faunistic notes east and from north to south. Annual precipitation (Lucas 1854, Forel 1886, 1889, 1910, Emery 1894, 1921, ranges from about 240 mm in the south-east to at least 1925, Müller 1923, Santschi 1927a, Stitz 1928) and PL ISSN 0003-4541 © Museum and Institute of Zoology PAS doi: 10.3161/00034541ANZ2018.68.4.004 770 S. SALATA, L. BOROWIEC and A. TRICHAS a few descriptions of new, mostly endemic, taxa (Forel method was a direct sampling (hand collecting). Indi- 1886, 1889, 1910, Emery 1894, 1906, 1908a, 1908b, Ka- vidual specimens and nests were collected on the ravaiev 1912, 1927, Santschi 1929). Consequently, they ground, in leaf litter and rock rubble, under stones and do not take into account many taxa that have been tree trunks. This method was occasionally supplement- divided into groups of species as a result of modern ed by litter sifting and collecting material with an ento- revisions. In recent years we have observed an mological umbrella. All specimens were preserved in increase in myrmecological research worldwide, 75% EtOH. Study was supported with material deposit- including Crete as a study target, and they are mainly ed in the Natural History Museum of Crete (University regional checklists (Legakis 2011, Borowiec & Salata of Crete, Heraklion) and the collection of G. Bračko 2012, 2013), and modern revisions (Radchenko 1997, (Ljubljana, Slovenia). All studied type specimens of Boer 2013, Borowiec & Salata 2014a, Csősz et al. 2015, taxa mentioned in differential diagnoses or character- Salata & Borowiec 2015a, 2017). istics are listed in the Appendix 2. Examined speci- Based on literature, 106 ant species were recorded mens are housed in the following collections: from Crete (Legakis 2011, Borowiec & Salata 2012, BMNH – Natural History Museum, London; 2013, Csősz et al. 2015, Salata & Borowiec 2015a, 2017, CASC – California Academy of Sciences, San Francis- 2018). There are 20 taxa described from Crete, among co, California, USA; which 7 have an endemic status (Appendix 1). Except DBET – Department of Biodiversity and Evolutionary one Lasius species, all valid endemic Cretan species Taxonomy, University of Wrocław, Poland; are members of the subfamily Myrmicinae. The high- DSAB – Dipartimnto di Scienze Agrarie, Universita di est number of endemics belongs to Aphaenogaster Bologna, Bologna, Italy; (3 species). Other endemic species are members of GBC – G. Bračko collection, Ljubljana, Slovenia; Tem nothorax (1), Monomorium (1) and Oxyopo - HNHM – Hungarian National History Museum, Buda - myrmex (1) respectively. Nevertheless, our study pest, Hungary; shows that there is at least one more endemic species MHNG – Museumd’Histoire Naturelle,Geneva, Switzer- of Formicinae subfamily as well: Cataglyphis cretica land; (Forel, 1910), which is currently considered as a junior MNHN – Muséum National d’Histoire Naturelle, Paris, synonym of C. aenescens (Borowiec & Salata 2013). France; The degree of endemism of Cretan myrmecofauna was MNHW –Museum of Natural History, University of estimated by Legakis (1983) at 16%. Nevertheless, in Wro cław, Wrocław, Poland; the same study, only 45 ant species were considered as MSNG – Museo Civico di Storia Naturale, Genova, present on Crete. Legakis & Kypriotakis (1994) pub- Italy; lished a review of endemic Cretan flora and fauna MZLS – Museum of Zoology, Lausanne, Switzerland; showing that 56% of all endemic animals were found in NHMB – Naturhistorisches Museum Basel, Switzer- mountainous areas, a fact mainly attributed to the land; complex geological history of the island. A more recent NHMC – Natural History Museum of Crete, University study (Triantis & Mylonas 2009) indicated that land of Crete, Heraklion; snails endemism can reach up to 50%. These results UASK – Institute of Zoology, National Academy of Sci- have shown that myrmecofauna of Crete is poorly ences of Ukraine, Kiev; investigated and needs more detailed study. ZMHB – Museum für Naturkunde, Zentralin-stitut der Material and data collected between 2007 and 2014, Humboldt-Universität, Berlin, Germany. contributed to confirm that the species diversity of Cre- Moreover, to determine a distribution range and tan ants ant fauna is rich and comprises several unde- a morphological variability of all Cretan populations scribed, and probably endemic species. Below we pres- we compared them with material collected from other ent a review of Cretan species of the genus Temnotho- Greek regions. Data concerning samples used in the rax. Distribution and biology of nine known species are comparison is provided in series of regional checklists discussed and seven species are described as new to (Borowiec & Salata 2012, 2013, 2014b, 2017a, b, 2018a, science. As Cretan populations of a few wide distrib- b, Bračko et al. 2016). Therefore, we see no reason to uted species differ from these known from other repeat this information. Mediterranean regions, we decided to redescribe also Specimens were compared using standard methods T. exilis, T. recedens and T. semiruber. of comparative morphology. Photos were taken using a Nikon SMZ 1500 stereomicroscope, Nikon D5200 photo camera and Helicon Focus software. All given MATERIAL AND METHODS label data are in original spelling, presented in square brackets; a vertical bar (|) separates data on different Most of the material was sampled between 2007 and rows and double vertical bars (||) separate labels. 2014 from sites in different parts of Crete. The main Images of type specimens are available online on A REVISION OF CRETAN TEMNOTHORAX (HYMENOPTERA: FORMICIDAE) 771 AntWeb (www.AntWeb.org) and are accessible using SI2 – SL/HW * 100; the unique specimen identifier, CASENT or FOCOL. MI – HTL/ML * 100; Pilosity inclination degree applies to this used in EI1 – EW/EL * 100; Hölldobler & Wilson (1990). The adpressed (0–5°) hairs EI2 – EW/HL * 100; run parallel, or nearly parallel to the body surface. TI – HW/HTL * 100; Decumbent hairs stand 10–15°, subdecumbent hair PI – PL/PH * 100; stands 30°, suberect hairs stand 35–45°, and the erect PPI – PPL/PPH * 100. hairs stand more than 45° from the body surface. Abbreviations Measurements q. – gyne; EL – eye length; measured along the maximum verti- w. – worker; cal diameter of eye; m. – male. EW – eye width; measured along the maximum hori- HC – hand collecting; zontal diameter of eye; EU – entomological umbrella; HL – head length; measured in straight line from LS – litter sifting. mid-point of anterior clypeal margin to mid- point of posterior margin in full-face view; We list all other ant species collected from the same HTL – hind tibia length; maximum length of hind tibia; localities as species new to science. In our opinion it HW – head width; measured in full-face view directly provides valuable information about ecosystem struc- above the eyes; ture and species diversity characteristic