Russian Entomol. J. 20(3): 261264 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2011 Gynandromorphs of Megachile picicornis (Morawitz, 1877) and M. deceptoria (Peréz, 1890) (Hymenoptera: Megachilidae) and their evolutionary interpretation Ãèíàíäðîìîðôû Megachile picicornis (Morawitz, 1877) è M. deceptoria (Peréz, 1890) (Hymenoptera: Megachilidae) è èõ ýâîëþöèîííàÿ èíòåðïðåòàöèÿ A.V. Fateryga1,2, S.P. Ivanov2, M.A. Filatov3 À.Â. Ôàòåðûãà1,2, Ñ.Ï. Èâàíîâ2, Ì.À. Ôèëàòîâ3 1 Nikitskiy Botanical Garden National Scientific Center, Nikita, Yalta 98648, Ukraine. E-mail: [email protected] 2 V.I. Vernadskiy Taurida National University, 4 Academician Vernadskiy Avenue, Simferopol 95007, Ukraine. E-mail: [email protected] 3 V.V. Dokuchaev Kharkov National Agrarian University, P.O. Communist-1, bdg. 4, KhNAU Campus, Kharkov Province 62483, Ukraine. E-mail: [email protected] 1 Íèêèòñêèé áîòàíè÷åñêèé ñàä Íàöèîíàëüíûé íàó÷íûé öåíòð, Íèêèòà, ßëòà 98648, Óêðàèíà. 2 Òàâðè÷åñêèé íàöèîíàëüíûé óíèâåðñèòåò èì. Â.È. Âåðíàäñêîãî, ïðîñïåêò Àêàäåìèêà Âåðíàäñêîãî, 4, Ñèìôåðîïîëü 95007, Óêðàèíà. 3 Õàðüêîâñêèé íàöèîíàëüíûé àãðàðíûé óíèâåðñèòåò èì. Â.Â. Äîêó÷àåâà, ï/î Êîììóíèñò-1, ó÷åáíûé ãîðîäîê ÕÍÀÓ, êîðïóñ 4, Õàðüêîâñêàÿ îáëàñòü 62483, Óêðàèíà. KEY WORDS: bees, Megachilidae, Megachile, gynandromorphs, evolution of cleptoparasitism. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: ï÷åëû, Megachilidae, Megachile, ãèíàíäðîìîðôû, ýâîëþöèÿ êëåïòîïàðàçèòèçìà. ABSTRACT. Seven gynandromorphic specimens [Bischof, Ulrich, 1929; Tums, 1970]), but most of of Megachile picicornis and M. deceptoria collected in them were mosaic and frontal gynandromorphs that the Crimea are described. All specimens of both spe- belonged to the genus Megachile Latreille, 1802 [Mitch- cies have similar morphology, i.e. the male-like head ell, 1929; Wcislo et al., 2004]. (except for the 12-segmented antennae) and legs as More than one hundred years ago, Cockerell de- well as the female-like metasoma that totally lacks scribed a female of the new species, Megachile detersa scopa. Arguments in favor of the hypothesis that onto- Cockerell, 1910, with certain male characters, like 13- genetic anomalies of that kind could lead to the origin segmented antennae, dense pubescence of clypeus, of cleptoparasitic species of bees from the nest-build- spines on the fore coxae and bifurcate claws. In this ing ones are given. female, ventral surface of the metasoma was bare and had no trace of the scopa [Cockerell, 1910]. Later on, ÐÅÇÞÌÅ. Îïèñàíî ñåìü ãèíàíäðîìîðôíûõ ýê- the same author described the genus Androgynella çåìïëÿðîâ Megachile picicornis è M. deceptoria, ñî- Cockerell, 1911 on the basis of the series of 14 females áðàííûõ â Êðûìó. Âñå ýêçåìïëÿðû îáîèõ âèäîâ and one male of the species mentioned above [Cocker- èìåþò ñõîäíóþ ìîðôîëîãèþ: ãîëîâà (çà èñêëþ÷å- ell, 1911]. Since normal females of M. detersa were íèåì 12-÷ëåíèêîâûõ àíòåíí) è íîãè êàê ó ñàìöîâ, à unknown, he suggested that the genus Androgynella ìåòàñîìà êàê ó ñàìîê, íî ïîëíîñòüþ ëèøåííàÿ had been an early stage of the evolution of cleptopara- ñêîïû. Ïðèâåäåíû àðãóìåíòû â ïîëüçó ãèïîòåçû î sitic bees. Michener [1965] considered the name An- òîì, ÷òî ïîäîáíûå îíòîãåíåòè÷åñêèå àíîìàëèè ìîã- drogynella invalid because the corresponding genus ëè ïðèâîäèòü ê âîçíèêíîâåíèþ êëåïòîïàðàçèòè÷åñ- had been described on the basis of a group of sexually êèõ âèäîâ ï÷åë èç ãíåçäîñòðîÿùèõ âèäîâ. anomalous specimens, and gave this name the status of the junior synonym of the subgenus Eutricharaea Th- Introduction omson, 1872 of the genus Megachile. In 20022005, we collected seven gynandromor- Various sex-related anomalies are relatively more phic specimens of Megachile in the Crimea and found frequent in Megachilidae than in the other families of that they were generally very similar to Cockerells Apoidea as well as in the other insects in general [Mitch- Androgynella. These specimens belong to two dif- ell, 1929; Popov, 1953; Engel, 2007]. Only few of the ferent species, namely, Megachile (Eutricharaea) pici- anomalies described in this family were true (lateral) cornis (Morawitz, 1877) and Megachile (Eutricharaea) gynandromorphs belonging to a number of genera (e.g. deceptoria (Peréz, 1890). 262 A.V. Fateryga, S.P. Ivanov, M.A. Filatov Fig. 12. Gynandromorph of Megachile picicornis: 1 frontal view; 2 lateral view. Ðèñ. 12. Ãèíàíäðîìîðô Megachile picicornis: 1 âèä ñïåðåäè; 2 âèä ñáîêó. Material and methods The gynandromorphs were found among the speci- mens collected by the authors and deposited in the collection of Vernadskiy Taurida National University (Simferopol). The collections of the Zoological Muse- um of M.V. Lomonosov Moscow State University, Zoological Institute of the Russian Academy of Sci- ences (St. Petersburg), Institute of Zoology of the Na- tional Academy of Sciences of Ukraine (Kiev) and Kharkov Entomological Society were also examined but any gynandromorphic specimens of Megachile from the Crimea were not found. In total, 109 specimens of M. picicornis and 57 specimens of M. deceptoria col- lected in the Crimea have been studied. Results Six gynandromorphic individuals belonging to M. picicornis were collected, specifically: 1 specimen, Crimea, Karadag Nature Reserve, 01.08.2002 (S.P. Iva- nov); 2 specimens, Crimea, Opuk Nature Reserve, Fig. 35. Megachile picicornis: 3 antennae of gynandro- 11.08.2003, on Centaurea diffusa (M.A. Filatov); 3 morph; 4 ditto, normal female; 5 ditto, normal male. Scale specimens, ibid., 06, 08 and 12.08.2004 (M.A. Fila- bar 0.5 mm. Ðèñ. 35. Megachile picicornis: 3 àíòåííû ãèíàíäðîìîðôà; tov). In addition, another specimen belonging to 4 òî æå, íîðìàëüíîé ñàìêè; 5 òî æå, íîðìàëüíîãî ñàìöà. M. deceptoria was also collected: Crimea, Tarhankut Ìàñøòàá 0,5 ìì. Peninsula, Kipchak Gully, 23.06.2005 (A.V. Fateryga). All specimens of the both species have similar mor- phology. Specifically, they have a typical male-like scopa (Fig. 2). Second to fifth sterna bear only one to head with dense whitish pubescence of the clypeus three hairs per sternum; moreover, the pubescence is (Fig. 1). Their mandibles are tridentate with the broad developed only on the first sternum and apical margin third teeth that is characteristic of the male (normal of the sixth sternum. Examination of the genitalia con- females have four-toothed mandibles). Nevertheless, firms that they are indistinguishable from those of the the antennae combine both male and female features normal female. (Fig. 3): they are 12-segmented as in the female (Fig. We have found deutonymphs of two species of 4), but long enough almost like those of the male (Fig. symbiotic mites of the genus Vidia Oudemans, 1905 5). The legs are characteristic of the normal male, with (Acari, Winterschmidtiidae) on the apical sterna of spines on the fore coxae and bifurcate claws. The meta- M. deceptoria and on the propodeum of the only speci- soma is generally female-like except for lack of the men of M. picicornis. Gynandromorphs of Megachile picicornis and M. deceptoria 263 Discussion parasitic Coelioxys and Radoszkowskiana Popov, 1955 [Michener, 2007]. We have examined females of 15 Gynandromorphs of M. picicornis and Palaearctic Coelioxys species of the subgenera Allo- M. deceptoria are described here for the first time, but coelioxys Tkalcù, 1974, Boreocoelioxys Mitchell, 1973 similar sexually anomalous specimens are already and Coelioxys Latreille, 1809, and found some charac- known in a number of other members of the genus, i.e. ters which were identical to the male traits of the de- in M. detersa [Cockerell, 1910]; M. (Ptilosarus) bertony scribed gynandromorphs of M. picicornis and M. decep- Schrottky, 1980 [Mitchell, 1929]; M. (Eutricharaea) toria, i.e. relatively long antennae, tridentate mandi- pilidens Alfken, 1924 [Wolf, 1998a] and M. (Eutri- bles with the broad third tooth and an incision between charaea) leachella Curtis, 1828 [Wolf, 1998b]. How- the third and the second teeth. In addition, females of ever, all previously described gynandromorphs can be one of the examined species, Coelioxys (Allocoelioxys) distinguished from the present ones by their 13-seg- emarginata Förster, 1853 had entirely bare metasomal mented antennae. Nevertheless, one of those gynandro- sterna. The latter character state has never been found morphs of Megachile (Eutricharaea) subrixator Cock- either in the females or males of nest-building species erell, 1915 had 12-segmented antennae ([Cockerell, (even in those using specialized structures of their legs 1918], quoted in [Mitchell, 1929]). Moreover, Cocker- to transport the collected pollen), but was characteris- ell described two specimens with different numbers of tic of the described gynandromorphs. Thus cleptopara- antennal segments (12 and 13). In addition, Mitchell sitic taxa in Megachilini could originate from the nest- [1941] found a similar gynandromorph of Megachile building species of Megachile through expression of (Chelostomoides) angelarum Cockerell, 1902 with 12- the male traits in the females via the apparent gynan- segmented antennae but with the well-developed sco- dromorphic stage. pa. Sex-related anomalies of that kind, i.e. combina- From the formal point of view, the described gy- tions of the male-like head and thorax with the female- nandromorphs of M. picicornis and M. deceptoria are like metasoma that lacks the scopa, are therefore quite females, because they have normal female genitalia widespread in Megachile and especially in the subge- and the functional sting as well as six metasomal seg- nus Eutricharaea. It is also noteworthy