Calosoma maderae (Fabricius, 1775)

(Figs 2B, 3, 9)

Dispersal power: Macropterous, flight active (Lindroth 1945).

Habitat: Epigeic species of open habitats (arable fields, steppes, batha, wadis in the Mediterranean, dunes) (Lindroth 1985; Bruschi 2013, pers. obs. in Europe and in the Levant). In Israel, from the north (Mt Hermon, Golan Heights, Upper Galilee) to the steppe zone in northern Negev (e.g. Nature Reserve Pura, Ma'agar Yeroham), although the majority of the collected specimens is from the Mediterranean part of the country (pers. obs.; SMNHTAU). A similar climate niche is reported from Morocco, where the species occurs in the Mediterranean climate zone to the semi-arid steppe zone close to Marrakesh (Gourves 1997). In both Morocco and the southern Levant, C. maderae co-occurs with C. olivieri.

Phenology: Spring breeder with summer larvae. Adults hibernate and can live up to three years (Larsson 1939; Lindroth 1945, 1985).

Material examined (all SMNHTAU, if not stated otherwise): Israel: 1♀ ‘ Israel’; Har Hermon: 1♂ Har Hermon, 9.vi.1992, Shney-Dor?; 1♂ Har Hermon, 2200 m, 2.vi.1993, V. Chikatunov; 1♀ Har Hermon, 2000 m, 8.vi.1993, V. Chikatunov; 1♀ Har Hermon, Nahal Guveta, 1.5 km W Majdal Shams, 1250 m, 28.iv.1995, E. Orbach; Hula Valley: 1♀ Hula Valley, 17.iii.1969, M.P. Pener, Y. Ayal; 1♂ Dan, 26.iv.1974, E. Horovitz; 1♀ Dan, 30.iv.1974, E. Horovitz; 1♂ Dan, 16.vi.1974, E. Horovitz; 1♂ Dan, cotton field, vi.1979, E. Tzhori; 1♂ Hula Nature Reserve, 9.iv.1987, G. Muller; 1♀ Sede Nehemya [Huliyyot], 28.iii.1957; 1♀ Sede Nehemya, 16.iv.1960, Z. Shoham; 1♂ Sede Nehemya, 23.iv.1960, Z. Shoham; 2♀ Sede Nehemya, 2.ii.1962, 3.vii.1962; 1♀ Sede Nehemya, 1.vi.1968, H. Bytinski-Salz, light; 2♂ Sede Nehemya, 20.vi.1968, H. Bytinski-Salz, light; 1♂ Sede Nehemya, 20.iv.1967, Z. Shoham; 2♂ 1♀ Sede Nehemya, 15.v.1969, Z. Shoham; 1♂ 1♀ Sede Nehemya, Hawat Ramatayyim, 4.vi.1969, Z. Shoham, light trap; 3♂ Sede Nehemya, 8.vi.1972, Z. Shoham; Upper Galilee: 1♀ Ziv'on, meadow, 23.iv.2011, Th. Assmann (CAB); Northern Coastal Plain: 1♂ Nahsholim, sea coast, 19.v.2012, A. Orlov; 1♂ Kefar Glikson, 29.vii.1959; 1♂ Binyamina, 1.v.1965, M. Kaufshtein; Central Coastal Plain: 1♂ Kefar Vitkin, 18.iii.1940, A. Shulov; 1♀ Shefayyim, 4.iii.1990, E. Orbach; 1♂ Hod haSharon, 14.v.1993, A. Traub; 1♂ Hod haSharon, 1996, A. Traub; 1♀ Hod haSharon, 21.iii.1998, A. Traub; 1♂ Tel Aviv, Xii.1960, Z. Ilan; Southern Coastal Plain: 1♀ Giv'at Brenner, 19.vii.1970, D. Gerling, light trap; 1♂ Neta'im, 8.vii.1972, Ch. Levinson; Judean Foothills: 1♂ 'Adullam, 3.iv.2003, U. Columbus, T. Levanony; Judean Hills: 1♂ Yerushalayim [Jerusalem], ii.1956; 1♀ Zur Hadassa, 31.iii.2001, Y. Mandelik; Northern Negev: 1♀ 1♂ Pura Nature Reserve, 210 m, 31.472°N 34.774°E, 1–18.iv.2010, C. Drees, pitfall (1♂ in CAB); 1♀ Gevulot, vii.2018, M. Zaytzov-Raz; 1♀ Hazerim, 16.v.1992, E. Orbach; 1♂ Be'er Sheva', 7.iv.2015, A. Orlov; Central Negev: 1♀ Ma'agar Yeroham, 450 m, 30.9833°N 34.8833°E, 8–20.v.2010, C. Drees, pitfall.

Observation records: 1♀ Golan Heights: Ya'ar Odem, Jubat el-Kabira, 24.iv.2011, Th. Assmann.

Distribution range: From Macaronesian Islands, Morocco and Portugal eastwards to India (incl. the Himalayas) and Central Asia, northwards to southern Scandinavia and southwards to the North and East Africa (Austin et al. 2008; Bruschi 2013).

Distribution range in southern Levant: Widespread, mostly in regions with the Mediterranean climate: Egypt (El-Akkad et al. 1998; HÄckel 2017), although Alfieri (1976) mentioned that the species was wrongly reported from Egypt; Israel (from Upper Galilee to northern Negev: Bodenheimer 1937; pers. obs., SMNHTAU); Jordan (Bruschi 2013: Zarqa; Madaba: pers. obs.); Lebanon (HÄckel 2017); and Syria (Bruschi 2013).

Taxonomic notes: Numerous taxa are now ranked as subspecies or older synonyms of maderae (HÄckel 2017). Particularly, C. m. auropunctatum (Herbst, 1748) has been treated very controversially by numerous authors: as a junior synonym of C. m. maderae (Bruschi 2013; Cavazzuti 2017; HÄckel 2017), as an independent species of its own (Jeannel 1940, 1941–1942; Casale et al. 1982; Forel & Leplat 2001; Arndt & Trautner 2006; Coulon et al. 2011), or as a subspecies (Breuning 1927; Lindroth 1985; Maguerre 2016) with some similarities to C. m. dsungaricum Gebler, 1833. Toussaint and Gillett (2018) reveal strong differences of mitochondrial and nuclear DNA sequences between the nominative subspecies and auropunctatum. The scale of variation at the given loci is comparable to some interspecific differences within the genus Calosoma. The molecular clock dates splitting of the two linages (maderae s.str. and auropunctatum) back to a few million years ago. Therefore, a revision (including molecular data) is needed to solve the taxonomic conundrum in this group. Genetic and phenotypic differences between maderae s.str. and auropunctatum prompt a separate discussion of the latter below. HÄckel (2017) lists both C. m. maderae and C. m. dsungaricum from Cyprus and Syria, but only maderae s.str. for Egypt, Israel, Jordan and Lebanon. In fact, beetles from the southern Levant show moderately convex distinctly transversely wrinkled elytral intervals. In maderae s.str., the elytral intervals are nearly completely flattened and wrinkles are indistinct; in auropunctatum, the intervals are flat, but distinctly transversely wrinkled; in dsungaricum, the elytral intervals are strongly convex and distinctly transversely wrinkled (Fig. 7). The beetles from the southern Levant show an intermediate phenotype between auropunctatum or maderae s.str., on one hand, and dsungaricum, on the other hand. Therefore, it is possible that the populations in the southern Levant represent a hybrid zone between two (or more?) subspecies. Bruschi (2013) indicates a contact zone between maderae s.str. and dsungaricum west of the southern Levant and classifies the beetles from the southern Levant as dsungaricum.