Warnckandrena Pisanty

Pisanty, Gideon & Wood, Thomas James, 2024, The early-diverging subgenera of the bee genus Andrena (Hymenoptera: Andrenidae) in the Old World, Zootaxa 5474 (5), pp. 451-488 : 453-455

Subgenus Warnckandrena Pisanty subgen. nov.

Type species: Melitta curiosa Morawitz, 1877 , designated here.

Description. Medium to large-sized (10–18 mm), usually robust bees ( Figs. 8 , 10, 12, 14, 16 , 18, 21, 24, 27 , 30, 33). Integument dark, male clypeus yellow, male paraocular area dark. Head 1.0–1.2 times broader than long ( Figs. 9 , 11, 13, 15, 17 , 19, 22, 25, 28 , 31, 34). Mandible normally developed in both sexes, strongly bidentate. Condylar lamella of female mandible well-developed. Galea finely shagreened, matt to shiny, impunctate or very finely punctured, apex more or less rounded. Maxillary palpus extending as far as galea or beyond, six-segmented, segment 2 longest, 1.2–1.7 times longer than 1. Shape of glossa highly variable, 3–9 times longer than broad. Labial palpus shorter than or subequal to glossa, four-segmented, segment 1 longest. Subgenal coronet well-developed along hind and inner margin of paramandibular process. Labral process broader than long, trapezoidal, rectangular or arched, unnotched. Clypeus weakly to strongly protuberant, arched or medially flattened, 1.2–1.8 times broader than long. Malar space weakly to strongly developed, 0.2 to 0.6 times as long as width of mandible base. Inner margins of compound eyes parallel-sided to convergent above ( Figs. 9 , 11, 13, 15, 17 , 19, 22, 25, 28 , 31, 34). Facial foveae broad and long, sometimes narrowing at mid-length, sometimes separated from compound eye by raised linear area, fovea width equals 0.5–0.7 times antennocular distance. Distance of fovea from lateral ocellus less than 1 ocellar diameter ( Fig. 2 ). Flagellomere 1 in females 0.9–1.1 times as long as 2+3+4 ( Fig. 1 ), in males more variable, but always longer than 2 and shorter than 2+3+4. Vertex and gena more or less normally developed, genal area 0.7–1.3 times as broad as compound eye, ocelloccipital distance 1–2 times the diameter of the lateral ocellus ( Fig. 2 ). Preoccipital ridge dorsally weakly to moderately carinate, laterally rounded in females, slightly angled in males. Dorsolateral angle of pronotum slightly to moderately elevated, sometimes creating a transverse dorsal pronotal ridge that does not descend much laterally. Mesepisternum, propodeum and dorsal mesosomal surfaces finely shagreened, punctation very fine, shallow and often inconspicuous ( Figs. 2, 4, 5 ). Profile of propodeum weakly separated into narrow, curved basal region versus broad, vertical apical region, basal region about 0.9 times as wide as metanotum ( Fig. 4 ). Propodeal triangle very finely shagreened, without rugae ( Fig. 5 ). Propodeal corbicula incomplete, corbicular fringe thick, limited to dorsoposterior margin of corbicula, composed of long, minutely to moderately plumose hairs ( Fig. 4 ). Inner side of hind femur rounded. Inner hind tibial spur normally developed, not broadened or curved. Femoral and tibial scopae composed of simple hairs. Flocculus present, more or less complete ( Fig. 6 ). Hind pretarsal claw with distinct inner tooth. Forewing with three submarginal cells. Stigma slender, 4–5 times longer than broad, about as broad as prestigma. Submarginal crossvein 1 enters marginal cell 5–6 vein widths from stigma. Recurrent vein 1 enters submarginal cell 2 at its middle or proximal to it. Recurrent vein 2 enters submarginal cell 3 at 0.6 its length. Nervulus antefurcal to interstitial ( Fig. 7 ). Jugal lobe of hind wing 0.7–0.8 times shorter than vannal lobe. Profile of tergum 1 clearly separated into declivous basal region versus horizontal posterior region, basal region with medial longitudinal rim. Tergal discs shagreened, densely and finely punctured, discs 1–3 often with conspicuous hair cover. Tergal apical hair bands moderately developed ( Fig. 3 ). Female pygidial plate normally developed, without distinct elevated medial area; male pygidial plate undeveloped. Male sternum 7 normally developed, with two medioapical lobes which are fully separated to partly fused medially ( Figs. 42–43 ). Male sternum 8 more or less normally developed, columnar, without lateral extensions, pilosity of ventral side more or less uniform, apical process often broadened ( Figs. 36–41 ). Genital capsule variable, often highly modified. Dorsal gonocoxite lobe absent or developed, gonostylus blade broadened, penis valves elongate and narrow. Volsella often enlarged.

Diagnosis. Warnckandrena does not possess any unique single character which is fully diagnostic. However, the subgenus can be characterised by the combination of robust body shape, large and broad facial foveae, very long flagellomere 1 especially in females, finely sculptured mesosomal and metasomal surfaces, weakly differentiated and strongly sloping profile of propodeum, and rather modified genitalia. Other important characters, such as mouthparts, clypeal protrusion and malar space, are too variable among the included species to provide any meaningful diagnostic criterion.

Andrena melittoides , the sole representative of subgenus Melittoides s.str., is very easily distinguished from members of Warnckandrena by the basally orange-coloured terga 1–3 ( Figs. 65, 69 ), and the male’s yellowmarked paraocular areas ( Fig. 70 ) and highly unusual genitalia ( Fig. 71 ). Other important diagnostic characters include the very short glossa ( Figs. 66, 70 ; this trait highly variable in Warnckandrena ), more strongly plumose mesosomal hair (especially on the propodeum), very strongly postfurcal nervulus ( Fig. 68 ) (antefurcal to interstitial in Warnckandrena ), and narrower scopa ( Fig. 67 ).

Aside from Melittoides s.str., Warnckandrena are most easily confused with members of the species-rich subgenus Truncandrena . The best way to distinguish these two subgenera is by the sculpture of the mesosoma. In Warnckandrena , all dorsal and lateral mesosomal surfaces (mesepisternum, entire propodeum, mesonotum and scutellum) are very finely shagreened or alveolated and without strong punctation or any striations, and the propodeum in profile is strongly sloping almost in its entirety, without a clearly differentiated horizontal basal region ( Figs. 2, 4, 5 ). By comparison, the mesosomal sculpture of Truncandrena species is highly variable, but all of them have distinct punctation or striation on some mesosomal surfaces, and/or a strongly differentiated basal region of the propodeum. Similar characters can be used to distinguish the subgenus Nobandrena . In members of this subgenus, the propodeum is very strongly divided into basal versus apical regions with an almost 90 degree angle between them, and the basal part is very broad and almost horizontal. In addition, the males have yellow markings on the paraocular areas.

Biology. Warnckandrena are active in springtime on diverse plant families. Pollen collection data are available for four species, all of which appear to be pollen specialists. Three plant families are known pollen hosts ( Brassicaceae , Fabaceae , Papaveraceae ), and additional families are exploited as sources of nectar. The use of diverse flower hosts can explain the great variation in glossal length and head morphology among species of this group.

Distribution: North Africa, Aegean Islands and Turkey, the Levant, the Caucasus, and Iran. The centre of diversity is in Turkey with five of the six known species.

Etymology. The subgenus is named in honour of Klaus Warncke (1937–1993), the renowned German expert on Andrena (as well as many other bee groups), in combination with the latter genus name. The gender of the name is feminine.

Included species. Andrena coromanda Warncke, 1975 ( Turkey and Iran), A. curiosa ( Morawitz, 1877) ( Greece, Levant, Turkey, the Caucasus, and Iran), A. innesi Gribodo, 1894 (North Africa, Levant, and Turkey), A. otoolei Pisanty sp. nov. (Levant and Turkey), A. papio Pisanty sp. nov. ( Turkey), A. ramlehiana Pérez, 1903 ( Israel). Species belonging to Warnckandrena were originally assigned to the subgenus Melittoides Friese, 1921 ( Warncke 1968).

Key to the species of Warnckandrena

Females

1- Mid and hind basitarsi orange. Disc of tergum 4 with black hair. Basal half of clypeus shagreened, apical half smooth. Malar area developed, about 0.4 times as long as broad. Glossa short, 3 times longer than broad. Body length 13–15 mm ( Turkey, Iran).................................................................................... A. coromanda

- All basitarsi dark ( Figs. 8 , 10, 12, 14, 16 ). Disc of tergum 4 with mostly light- to reddish-coloured hair ( Figs. 3, 8 , 10, 12, 16 ) and/or clypeus entirely shagreened ( Fig. 15 ). Malar area, glossa and body length variable............................ 2

2- Clypeus largely shiny, smooth to very superficially shagreened, relatively sparsely punctured, distance between punctures 1–2 puncture diameters ( Fig. 9 ). Glossa 6 times longer than broad ( Fig. 8 ). Body length 15–17 mm (Aegean Islands, Levant, Turkey, Caucasus, Iran)................................................................................ A. curiosa

- Clypeus strongly shagreened at least in basal half ( Figs. 11, 13, 15, 17 ). Glossa and body length variable................ 3

3- Clypeus densely punctured except for shiny impunctate medioapical area (distance between punctures 0–1 puncture diameters), and usually with slightly elevated midline ( Fig. 11 ). Malar area linear, 0.2 times as long as broad ( Figs. 10–11 ). Glossa short, 3.5 times longer than broad ( Fig. 10 ). Body length 13–15 mm (North Africa, Levant, Turkey)................... A. innesi

- Clypeus uniformly shagreened and matt at least in basal ¾, more sparsely punctured (distance between punctures 0.5–2 puncture diameters), without shiny impunctate medioapical area, without elevated midline ( Figs. 13, 15, 17 ). Malar area longer, at least 0.3 times as long as broad ( Figs. 12–17 ). Glossa longer, 6–9 times longer than broad ( Figs. 12, 14, 16 ). Body length variable. ................................................................................................... 4

4- Malar area almost half as long as broad ( Figs. 12–13 ). Clypeus slightly flattened centrally. Tergum 4 with light-coloured hair. Glossa 8 times longer than broad. Body length 16–18 mm ( Turkey)................................ A. papio sp. nov.

- Malar area 0.3 times as long as broad ( Figs. 14–17 ). Clypeus evenly domed. Tergum 4 with light- or dark-coloured hair. Glossa 6–9 times longer than broad. Body length 11–15 mm ......................................................... 5

5- Tergum 4 with dark brown hair, in sharp contrast to the light-coloured terga 1–3 ( Fig. 14 ). Clypeus fully shagreened up to apex ( Fig. 15 ). Glossa 6 times longer than broad. Body length 11–13 mm ( Israel)............................ A. ramlehiana

- Terga 1–4 all with white or yellowish hair ( Fig. 16 ). Apical margin of clypeus often smooth to weakly shagreened ( Fig. 17 ). Glossa 9 times longer than broad. Body length 13–15 mm (Levant and Turkey)...................... A. otoolei sp. nov.

Males

1- Flagellomere 1 1.2 times as long as 2. Hind basitarsi and often also mid basitarsi orange ( Fig. 18 ). Dorsal lobe of gonocoxite large, extending more than half the distance to the gonostylus blade ( Fig. 20 ). Glossa short, 3 times longer than broad ( Fig. 18 ). Body Length 10.5–12 mm ( Turkey, Iran)........................................................ A. coromanda

- Flagellomere 1 much longer than 2, at least 0.9 times as long as 2+3 ( Figs. 25, 28 , 31). All basitarsi dark ( Figs. 21, 24, 27 , 30, 33). Dorsal lobe of gonocoxite absent or smaller ( Figs. 23, 26, 29 , 32, 35). Glossa and body length variable.............. 2

2- Gonocoxites curved medioapically, without dorsal lobes ( Figs. 23, 26 ). Glossa long, 8–9 times longer than broad ( Fig. 24 ). Body length 13–14.5 mm ............................................................................... 3

- Gonocoxites with distinct dorsal lobes ( Figs. 29 , 32, 35). Glossa shorter, 3.5–6.5 times longer than broad. Body length 10–15 mm ................................................................................................ 4

3- Malar area 0.5–0.6 times as long as broad ( Figs. 21–22 ). Penis valves uniformly broad, digiti very large. Inner margins of gonocoxites strongly diverging apically ( Fig. 23 ) ( Turkey)........................................ A. papio sp. nov.

- Malar area 0.3–0.4 times as long as broad ( Figs. 24–25 ). Penis valves slightly swollen medially, digiti small, hidden behind gonocoxite in dorsal view. Inner margins of gonocoxites parallel almost to apex ( Fig. 26 ) (Levant, Turkey).................................................................................................... A. otoolei sp. nov.

4- Dorsal gonocoxite lobes well-developed, parallel-sided, relatively broad, rounded apically. Digiti short, hidden under gonocoxite in dorsal view. ( Fig. 29 ). Head as long as broad. Clypeus smooth and shiny at least in apical ½ ( Fig. 28 ). Body length 13.5–15 mm (Aegean Islands, Levant, Turkey, Caucasus, Iran)................................................. A. curiosa

- Dorsal gonocoxite lobes short, converging apically, narrow and pointed. Digiti well-developed, knob-shaped, clearly visible above gonocoxites ( Figs. 32, 35 ). Head 1.1–1.2 times broader than long. Clypeus fully shagreened to apically smooth ( Figs. 31, 34 ). Body length 10–14 mm ............................................................................. 5

5- Gonostyli strongly curved towards each other, with strong inward bulge at mid length ( Fig. 32 ). Flagellomere 1 as long as 2+3 or slightly shorter. Glossa short, 3.5 times longer than broad. Apical ¼–½ of clypeus shiny, more or less smooth ( Fig. 31 ) (North Africa, Levant, Turkey)......................................................................... A. innesi

- Gonostyli extended apically, with very weak inward bulge at mid length ( Fig. 35 ). Flagellomere 1 longer than 2+3. Glossa longer, 6–6.5 times longer than broad. Clypeus fully shagreened ( Fig. 34 ) ( Israel)....................... A. ramlehiana

Figures

FIGURES 1–9. Andrena (Warnckandrena) curiosa (Morawitz), female: 1, flagellum; 2, vertex and mesosoma; 3, metasoma; 4, profile of propodeum; 5, propodeal triangle; 6, flocculus and scopa; 7, forewing venation; 8, habitus; 9, head.

FIGURES 10–17. Andrena (Warnckandrena) female habitus (10, 12, 14, 16) and heads (11, 13, 15, 17): 10–11, A. innesi Gribodo; 12–13, A. papio sp. nov.; 14–15, A. ramlehiana Pérez; 16–17, A. otoolei sp. nov.

FIGURES 18–29. Andrena (Warnckandrena) male habitus (18, 21, 24, 27), heads (19, 22, 25, 28) and genitalia (20, 23, 26, 29): 18–20, A. coromanda Warncke; 21–23, A. papio sp. nov.; 24–26, A. otoolei sp. nov.; 27–29, A. curiosa (Morawitz).

FIGURES 30–43. Andrena (Warnckandrena) male habitus (30, 33), heads (31, 34) genitalia (32, 35), eighth sterna (36–41) and seventh sterna (42–43): 30–32, 38, A. innesi Gribodo; 33–35, 41, A. ramlehiana Pérez; 36, A. coromanda Warncke; 37, 42, A. curiosa (Morawitz); 39, A. otoolei sp. nov.; 40, 43, A. papio sp. nov.

FIGURES 65–73. Andrena (Melittoides) melittoides Friese, female (65–68) and male (69–73): 65, 69, habitus; 66, 70, head; 67, flocculus and scopa; 68, forewing venation; 71, genitalia; 72, eighth sternum; 73, seventh sternum.