Four new Palaearctic Brassicogethes (Coleoptera, Nitidulidae, Meligethinae), and phylogenetic inference on the B. coracinus group

Abstract

Combined morphological and ecological data, together with molecular evidence from a companion paper, are presented to clarify the taxonomic position of four problematic species, members of the Brassicogethes coracinus group, from north-western Spain, eastern Turkey, and Tajikistan (Coleoptera, Nitidulidae, Meligethinae). This species-group was represented by a dozen anthophagous species associated with Brassicaceae, and distributed from North Africa to Middle Asia and southern Siberia. The analysis was mainly focused on the specific distinction and formal description of two new species from E Turkey, B. gloriae sp. nov., and B. coracimimus sp. nov., one species from Tajikistan, B. capannai sp. nov., and one additional new species, B. cantabricus sp. nov., from Cantabrian Mts in NW Spain. B. gloriae sp. nov., strictly associated with Neotchihatchewia isatidea (Boissier) Rauschert (Brassicaceae, Lunarieae), is morphologically related to but molecularly distinct from the rare and sympatric B. longulus (Schilsky, 1894). B. coracimimus sp. nov., whose larval host-plant is unknown, is morphologically and molecularly related to the amply sympatric and common B. coracinus (Sturm, 1845). B. cantabricus sp. nov., strictly associated with Sisymbrium irio L. (Brassicaceae, Sisymbrieae), is morphologically related to but molecularly markedly distinct from the parapatric B. subaeneus (Sturm, 1845). B. capannai sp. nov., whose larval host-plant is unknown, is morphologically related to the rare and allopatric B. longulus (Schilsky, 1894). A short discussion of the state of art of the taxonomy and phylogeny of members of the B. coracinus group, a key to the identification of the 16 known species, and an updated synoptic table including larval host-plant relationship and geographical distribution of each included species, are finally provided.

Appendix

1.1 Key to the species of the Brassicogethes coracinus complex

In the following key, all species so far recognized on morphological and ecological bases within the B. coracinus species-complex, are included. Refer to Table 2 and Fig. 76 for abbreviations of measurements used in the key.

• 1. (2) Pronotum markedly rounded at sides, with markedly obtuse posterior angles (Fig. 16). Head and pronotum with spaces between punctures more or less dull, contrasting with elytra, usually more shiny and slightly more coarsely punctured. Body brown to blackish-brown with usually yellowish legs and antennae. Male genitalia peculiarly shaped (Figs. 40, 41). Female ovipositor as in Fig. 54. Length 2.4–2.6 mm. Southern Turkey. ……………………………..………….………1. cristofaroi (Audisio and De Biase, 2005)

• 2. (1) Pronotum not so markedly rounded at sides, with always clearly distinct and less obtuse posterior angles (Figs. 1–9, 10–15). Male genitalia more or less differently shaped (Figs. 24–39, 42–53). Female ovipositors as in Figs. 55–64. …..………..……………………………………….3

• 3. (6) Elytra and especially pronotum rather strongly depressed, with peculiarly widely flattened lateral sides of pronotum (Figs. 9, 15). Body brown to blackish-brown with generally yellowish legs and antennae. Male genitalia with apex of the aedeagus bearing a more or less markedly distinct excision (Figs. 44–47). Length 1.7–2.7 mm. Species from central Asia and the Syrian Desert. ……………………….………….……………….………………….…….4

• 4. (5) Anterior margin of clypeus feebly arcuately emarginated (Fig. 9). Body less elongate (Fig. 9; ratio LELY/WELY = 1.03–1.10). Male genitalia as in Figs. 44, 45, with only minutely incised apex of the aedeagus. Suberemic areas of Tajikistan. Larval host-plant unknown. …………………..……………………………………………2. accentus (Kirejtshuk 1978 )

• 5. (4) Anterior margin of clypeus distinctly truncate (Fig. 15). Body more elongate (Fig. 15; ratio LELY/WELY = 1.13–1.17). Male genitalia as in Figs. 46, 47, with much more distinctly incised apex of the aedeagus. Subdesertic areas of Uzbekistan, SE Kazakhistan, Turkmenistan and Syria. Larval host-plants: Crambe spp. ….. 3. explanatus (Reitter 1900 )

• 6. (3) Elytra and especially pronotum never so strongly depressed. Male genitalia with apex of the aedeagus always without excisions, or only obtusely and shallowly emarginated (Figs. 24–39, 42, 43, 48–53)……………………………………………………………………………………………..…..7

• 7. (8) Metasternum in males with deep and long longitudinal median impression. Male genitalia with very peculiar tegmen, very narrowly and deeply excised distad (Figs. 42, 43; angular measurement ANGT = 18°–20°). Female ovipositor with peculiarly shaped apex (Fig. 55). Habitus as in Fig. 14. Length 1.7–2.7 mm. Western Europe, western North Africa. Rocky outcrops, xeric meadows, sunny hill slopes, coastal sand dunes, cultivated fields. Larvae on Biscutella cichoriifolia Loisel., Brassica spp. and Sinapis spp. …………………………………………………4. fulvipes (C. Brisout de Barneville, 1863)

• 8. (7) Metasternum in males with much shallower longitudinal median impression, or almost not impressed. Male genitalia with tegmen always with normal, more or less widely V-shaped excision (Figs. 24–39, 48–53; angular measurement ANGT = 40°–80°). Female ovipositors with differently shaped apex (Figs. 56–64). ……………………………………………………9

• 9. (10) Front tarsi in males as wide as or hardly narrower than antennal club (ratio WFTA/W10J = 1.00–0.94; ratio LFTA/WFTA = 3.23–3.50; Figs. 1, 17, 18). Male genitalia (Figs. 48, 49) with median lobe of aedeagus showing a peculiarly shaped and sclerotized proximal edge of the ejaculatory ostium, this being very widely and transversely U-shaped (ratio V/H = 1.44–1.53; Fig. 49). Female ovipositor with blunt and usually markedly darkened apex (Fig. 56). Habitus as in Fig. 1. Length 1.5–2.8 mm. Europe, northern Near East, central Asia, and Siberia towards the Russian Far East. Cultivated fields, meadows, wetlands, shaded hill slopes. Larval stages mostly on Barbarea spp., Sisymbrium spp., Brassica spp., and Sinapis spp. ………………………………………………..………………9. coracinus (Sturm, 1845)

• 10. (9) Front tarsi in males more distinctly to markedly narrower than antennal club (ratio WFTA/W10J = 0.65–0.40; ratio LFTA/WFTA = 3.55–4.83; Figs. 2–8, 10–13, 19, 20). Male genitalia with median lobe of aedeagus showing the proximal edge of the ejaculatory ostium regularly arc-like shaped, widely U-shaped or widely V-shaped (Figs. 24–39, 50–53), never widely transversely U-shaped as in the preceding species (ratio V/H = 0.89–1.33). Female ovipositor with usually less blunt apex (Figs. 57–64). Body outline more elongate (Figs. 2–8, 10–13). ……………………………………………………………………………………………………………….….11

• 11. (18) Pronotum showing its maximum width at the posterior angles, from there slightly but regularly arcuately narrowed towards the anterior pronotal margin (Figs. 10–13; ratio WPR1/WPR2 > 1). Body usually dark brown with faint bronze reflections …………….…12

• 12. (13) Dorsal punctures dense on pronotum and elytra, each puncture separated by its closest neighbor by one diameter or less, combined with surface between pronotal punctures moderately dull, microsculpture reticulate. Median lobe of aedeagus more pointed at apex and elongate (ratio LEAE/WIAE = 2.10), maximum width at distal three-fifths (Fig. 29), distinctly narrowed both proximad and distad; tegmen (Fig. 28) usually more pigmented and darker than median lobe. Ovipositor not darkened at apex (Fig. 60), apical suture of coxites narrowly darkened, and “central point” at distal three-fifths. Length: 2.2–2.4 mm (Fig. 13). Larval stages on Cardamine (s.str.) glauca Sprengel (Brassicaceae). Probably a Balkan endemic, known from NW Greece. .……….……… 5. epeirosi Audisio, Mancini and De Biase, 2006

• 13. (12) Dorsal punctures less dense, on pronotum and elytra punctures separated by 1–2 diameters, or, if more dense, surface between pronotal punctures almost smooth and shining. Median lobe of aedeagus more deeply sinuate and wider at apex (ratio LEAE/WIAE = 1.68–2.13), and more parallel-sided (Figs. 24–27); tegmen never more pigmented and darker than median lobe. Ovipositor with “central point” at distal four-ninths (Figs. 58, 59). …………..…14

• 14. (15) Elytral punctures finer and shallower, with more or less dull spaces between punctures. Ovipositor at “central point” usually with apparently fused proximal portions of the main sclerites of coxites and valvifers (Fig. 58), and apex never darkened. Median lobe of aedeagus relatively short and wide (ratio LEAE/WIAE = 1.66–1.78; Fig. 25), and tegmen slightly wider (Fig. 24). Body moderately elongate (ratio LELY/WELY = 1.05–1.09); habitus as in Fig. 11. Length 1.7–2.8 mm. Europe, from the Pyrenean Mts. to the northern Caucasus. Mesophilous forests, edges of mesophilous bushy areas. Larval stages oligophagous on Cardamine spp. and Cardaminopsis spp. ……6. subaeneus (Sturm, 1845)

• 15. (14) Elytral punctures coarser and deeper, and spaces between punctures more or less shining. Ovipositors unicolorous or slightly darkened distad, with normal, not apparently fused, sclerites at “central point” (Fig. 59). ……………………………………………………………………..….16

• 16. (17) Body slightly more elongate (ratio LELY/WELY = 1.07–1.11); habitus as in Fig. 10. Dorsal punctures relatively less coarse and deep. Median lobe of aedeagus distinctly more elongate (ratio LEAE/WIAE = 1.95–2.13; Fig. 27), and tegmen slightly narrower (Fig. 26). Ovipositor as in Fig. 59, with apex usually slightly darkened. Body usually brown to dark brown, with distinct bronze reflections, and with yellowish-brown to brown legs and antennae. Length 2.2–3.1 mm. Europe, Caucasian countries, northern Turkey. Mesophilous forests, edges of mesophilous bushy areas. Larval stages monophagous on Hesperis matronalis L. or (in Caucasian areas) on the very closely related H. transcaucasica Tzvelev ………………………………………………8. matronalis (Audisio and Spornraft 1990 )

• 17. (16) Body slightly shorter (ratio LELY/WELY = 0.99–1.05); habitus as in Fig. 12. Dorsal punctures relatively coarser and deeper. Aedeagus distinctly less elongate (ratio LEAE/WIAE = 1.64–1.75; as in M. subaeneus, Fig. 25), and tegmen slightly wider (as in Fig. 24). Ovipositor as in M. subaeneus (Fig. 58), but at its “central point” with no apparently fused proximal portions of the main sclerites of coxites and valvifers, and with apex never darkened. Body usually blackish, with scarce metallic hues and with black to dark brown legs and antennae. Length 1.7–2.7 mm. Cantabrian Mts in NW Spain (Fig. 78). Meso-xerophilous and shady rocky habitats. Larval host-plant Sisymbrium irio L. ………………………………………………….7. cantabricus Audisio and Cline, sp. nov.

• 18. (11) Pronotum showing its maximum width at the posterior third or fourth, from there slightly but regularly arcuately narrowed towards the anterior pronotal margin, and almost parallel-sided or distinctly narrowed also in posterior fourth or third, towards its posterior angles (Figs. 2–8; ratio WPR1/WPR2 < 1). ………………..………………………….………….……19

• 19. (22) Antennal club smaller (Figs. 2, 3, 65–66; ratio W10J/W03J = 3.70–4.20). Habitus as in Figs. 2, 3, with legs slightly wider and relatively shorter. Male genitalia as figured (Figs. 50–53). Ovipositors with relatively pointed apex (Figs. 57, 61). ……..………………………..…..…20

• 20. (21) Male genitalia as in Figs. 50, 51; median lobe of aedeagus distinctly more narrowed in its distal fifth, showing the proximal edge of the ejaculatory ostium widely V-shaped (Fig. 51). Male metasternum with usually shallower longitudinal impression. Ovipositor peculiarly pointed (Fig. 61). Habitus as in Fig. 3. Length 1.8–2.2 mm. Eastern Turkey (Fig. 77). Edges of meso-xerophilus bushy areas. Adults collected mostly on Arabis spp. and Erysimum spp.; larval host-plants unknown. …………………10. coracimimus Audisio and Cline, sp. nov.

• 21. (20) Male genitalia as in Figs. 52, 53; median lobe of aedeagus showing the proximal edge of the ejaculatory ostium regularly arc-like shaped (Fig. 53). Male metasternum with usually deeper longitudinal impression. Male genitalia slightly clinally variable in proportional length (ratio LEAE/WIAE = 1.92–2.32), more elongate in Balkan and Anatolian populations, shorter in SW European and central European ones. Ovipositor as in Fig. 57, moderately pointed. Habitus as in Fig. 2. Length 1.8–2.9 mm. Southern Europe from Iberian Peninsula to Turkey, Caucasus and southern Iran, northwards to the southern Czech Republic. Sunny rocky outcrops, xeric hillsides, sandy and gravel slopes, xeric meadows, mostly on mountain areas. Larval host-plants: Erysimum spp. ….11. erysimicola (Audisio and De Biase, 2001)

• 22. (19) Antennal club distinctly larger (Figs. 4–8; ratio W10J/W03J = 4.60–6.33). Habitus as in Figs. 4–8, with legs distinctly longer and weaker. Male genitalia as in Figs. 30–39. Ovipositors as in Figs. 62–64. …..…….………………………………………………..………………….…………23

• 23. (24) Antennal club comparatively smaller (Figs. 5, 71; ratio W10J/W03J = 4.60–5.12). Male genitalia as in Figs. 36, 37. Ovipositor as in Fig. 62. Body more or less elongate (ratio LELY/WELY = 1.03–1.17); habitus as in Fig. 5. Length 2.1–2.9 mm. Coasts of western central Italy, southern Italy and western Sicily, and inner areas of northern Spain and of central Sicily. Coastal sand dunes and coastal rocky promontories, but also rocky outcrops, xeric parasteppic meadows, and gravel hillsides in inner areas. Larval host-plants: Matthiola sinuata (L.) R.Br., M. incana (L.) R.Br., M. fruticulosa (L.) Maire, and M. perennis Conti. ………….….……………….…………….15. thalassophilus (Audisio and De Biase, 2005)

• 24. (23) Antennal club comparatively larger (Figs. 4, 6–8, 69–70; ratio W10J/W03J = 5.50–6.33). Male genitalia as in Figs. 32–35, 38, 39. Ovipositors as in Figs. 63, 64 (ovipositor of M. arankae nearly as in M. thalassophilus: Fig. 62). Habitus as in Figs. 4, 6–8. Species from central and eastern Mediterranean areas, Anatolia, Caucasus and Middle Asia. ….………25

• 25. (26) Male genitalia with paramera distinctly blunter at apex (Fig. 32); aedeagus showing the proximal edge of the ejaculatory ostium regularly arc-like shaped and placed more distally than in all other species of the complex (ratio LEOS/LEAE = 0.08–0.10; Fig. 33). Female ovipositor as in Fig. 64. Body long and narrow (ratio LELY/WELY = 1.14–1.22), brown with entirely yellowish legs and antennae, elytra frequently with a large discal orange spot (Fig. 4). Length 2.7–3.0 mm. Eastern Turkey, but probably present also in other circum-Pontic and SW Asiatic areas. Steep sandy and terrigenous hillsides and slopes. Larval host-plant: Matthiola odoratissima (Bieb.) R.Br. ……………………………..…12. longulus (Schilsky 1894 )

• 26. (25) Male genitalia with paramera distinctly more pointed at apex (Figs. 30, 34, 38); aedeagus showing the proximal edge of the ejaculatory ostium regularly arc-like or U-shaped, but placed less distally than in the preceding species (LEOS/LEAE = 0.15–0.20; Figs. 31, 35, 39). Female ovipositors as in Figs. 62, 63, or unknown. ……………………………………………………….27

• 27. (28) Tarsi peculiarly long and narrow, especially the middle and posterior ones, metatarsal ungueal articles more than 6× longer than wide (Fig. 72). Body relatively shorter and wider, ratio LELY/WELY = 1.04 (Fig. 7), dark brown with brown legs and antennae, elytra vaguely paler. Male genitalia as in Figs. 30, 31. Female ovipositor unknown. Front tibiae weak and narrow, with larger and more pointed teeth on their outer edge (Fig. 23). Length 2.7 mm. Tajikistan, but probably present also in other Middle-Asiatic areas. Suberemic mountain areas. Larval host-plant unknown, but likely Matthiola sp. ………………………………..……………………..….…13. capannai Audisio and Cline, sp. nov.

• 28. (27) Tarsi never so long and narrow, especially the middle and posterior ones, metatarsal ungueal articles less than 5× longer than wide. Body more elongate, ratio LELY/WELY = 1.14–1.19 (Figs. 6, 8), dark brown with usually yellowish legs and antennae. Male genitalia as in Figs. 34, 35, 38–39. Female ovipositors as in Figs. 62, 63. Front tibiae with smaller and less pointed teeth on their outer edge (Figs. 6, 8). ……………………………………………………………….29

• 29. (30) Pronotum usually less regularly arcuated at sides (Fig. 8). Elytra with spaces between punctures slightly less shining and always without metallic reflections. Body dark brown to orange-brown, legs and antennae always entirely yellowish; elytra frequently with a small discal orange spot. Male genitalia as in Figs. 34, 35, with paramera more shortly pubescent at apex, and median lobe of the aedeagus more truncate at apex. Ovipositor as in Fig. 63. Length 2.6–2.8 mm. Eastern Turkey (Fig. 77). Eroded mountain slopes and ravines, wind-shaped inner sand dune systems on xeric highlands. Larval host-plant: Neotchihatchewia isatidea (Boiss.) Rauschert …………….………….….14. gloriae Audisio and Cline, sp. nov.

• 30. (29) Pronotum usually more regularly arcuated at sides (Fig. 6). Elytra with spaces between punctures slightly more shining and frequently with weak faint metallic reflections. Body uniformly dark brown to orange-brown, legs and antennae pale brown to orange-yellowish, with antennal club usually darker. Male genitalia as in Figs. 38, 39, with paramera more pubescent at apex, and median lobe of the aedeagus more sinuate at apex. Ovipositor nearly as in M. thalassophilus (Fig. 62). Length 2.4–3.0 mm. Mountain areas of central and southern peninsular Italy, Ponto-Pannonian areas, Balkans, Turkey, Caucasus. Sunny rocky outcrops, xeric hillsides, gravel slopes, xeric parasteppic meadows, mostly on mountain areas. Larval host-plants: Hesperis spp. (H. laciniata All., H. tristis L., H. bicuspidata (Willd.) Poiret, H. pendula DC., and related species). …………………………………………………………..……16. arankae (Audisio and De Biase, 2005)