Russian Entomol. J. 20(3): 265271 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2011 Morphotypes of chromosome sets and pathways of karyotype evolution of parasitic Hymenoptera Ìîðôîëîãè÷åñêèå òèïû õðîìîñîìíûõ íàáîðîâ è íàïðàâëåíèÿ ýâîëþöèè êàðèîòèïà ïàðàçèòè÷åñêèõ ïåðåïîí÷àòîêðûëûõ (Hymenoptera) V.E. Gokhman Â.Å. Ãîõìàí Botanical Garden, Moscow State University, Moscow 119991, Russia. E-mail: [email protected] Áîòàíè÷åñêèé ñàä Ìîñêîâñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà, Ìîñêâà 119991, Ðîññèÿ. KEY WORDS: chromosomes, karyotype evolution, morphotypes, parasitic Hymenoptera. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: õðîìîñîìû, ýâîëþöèÿ êàðèîòèïà, ìîðôîëîãè÷åñêèå òèïû, ïàðàçèòè÷åñêèå ïåðåïîí÷àòîêðûëûå. ABSTRACT. Chromosomal diversity of parasitic ly ants [Imai et al., 1988; Hoshiba, Imai, 1993]. Fur- Hymenoptera has been analyzed with the help of the thermore, karyotypes, or rather karyomes (term intro- logical possibility space approach. Using three param- duced by Smirnov [1991]), usually evolve as holistic eters (haploid chromosome number, length ratios of objects [Lukhtanov, 1999] (see also Rasnitsyn, 1987), chromosomes within the haploid set, and the degree of and construction of the so-called chromosomal alter- karyotypic metacentricity), 18 classes, or morpho- ation networks [Imai, 1991, 1993], an apparent tool logical types, of chromosome sets have been delimited, proposed for analysis of karyotype evolution in many of which only 13 contain at least one karyotype. Possi- groups of living organisms including Hymenoptera, is ble major pathways of karyotypic transformation in probably unable to describe this process in an adequate parasitic wasps have been outlined. manner. The main aim of the present paper is therefore a thorough analysis of the major morphotypes and path- ÐÅÇÞÌÅ. Õðîìîñîìíîå ðàçíîîáðàçèå ïàðàçè- ways of karyotypic transformation in parasitic Hy- òè÷åñêèõ Hymenoptera ïðîàíàëèçèðîâàíî ñ ïîìî- menoptera. ùüþ ïîäõîäà, ïðåäóñìàòðèâàþùåãî ñîçäàíèå ïðî- ñòðàíñòâà ëîãè÷åñêèõ âîçìîæíîñòåé. Ñ èñïîëüçî- Materaial and methods âàíèåì òðåõ ïàðàìåòðîâ (ãàïëîèäíîå ÷èñëî õðîìî- ñîì, ñîîòíîøåíèå äëèí õðîìîñîì â ãàïëîèäíîì To cover the whole range of karyotypic variation in íàáîðå è ñòåïåíü «ìåòàöåíòðè÷íîñòè» êàðèîòèïà) parasitic wasps, their chromosome sets have been ana- óñòàíîâëåíî 18 êëàññîâ (ìîðôîëîãè÷åñêèõ òèïîâ) lyzed by means of the technique that is widely used in õðîìîñîìíûõ íàáîðîâ, èç êîòîðûõ òîëüêî 13 âêëþ- evolutionary morphology, namely, by delimitation of ÷àþò õîòÿ áû îäèí êàðèîòèï. Îïðåäåëåíû îñíîâ- the so-called logical possibility space (see Meyen, íûå ïðåäïîëàãàåìûå íàïðàâëåíèÿ ïðåîáðàçîâàíèé 1975). To adequately describe it, three main parame- êàðèîòèïà ïàðàçèòè÷åñêèõ ïåðåïîí÷àòîêðûëûõ. ters have been selected to characterize more or less substantial groups of parasitic Hymenoptera: (a) hap- Introduction loid chromosome number (n = 3 to 7, 8 to 13 and 14 to 23); (b) length ratios of chromosomes within the hap- Parasitic Hymenoptera belong to the very large, loid set (one or two chromosomes are not less than 1.5 taxonomically complicated and economically impor- times shorter/longer than the others, or they more or tant group of insects [Rasnitsyn, 1980]. Nevertheless, less gradually decrease in size; if chromosomes are the detailed picture of karyotype evolution of parasitic obviously uneven by size in any other way, those kary- wasps is far from being complete, although a few suc- otypes are conditionally included into the latter class); cessful attempts of its reconstruction have been made (c) degree of karyotypic metacentricity [not less than (see Gokhman, 2009). Moreover, research of that kind one-half of all chromosomes are metacentric in a broad usually requires a detailed chromosomal study of a sense (including submetacentrics), or acrocentrics (again certain group, but comparatively few karyotypes of in a broad sense, including subtelocentrics) predomi- parasitic Hymenoptera have been studied to that extent nate within the given karyotype]. Karyotypic data from up to now, apart from aculeate Hymenoptera, especial- Gokhman [2009] and a number of recent papers [Gokh- 266 V.E. Gokhman man, Gumovsky, 2009; Gokhman, 2010b; Gokhman et doidea: many species of the families Chalcididae, Tri- al., 2010] together with a few unpublished observa- chogrammatidae and some other groups. tions made by the author were used in the analysis; (8) Ichneumonoidea: Ichneumonidae: a few Pim- however, information on chromosome sets of certain pliformes, most Ichneumoniformes and a number of groups was also taken from a few extra papers [Drey- Ophioniformes; Braconidae: species of the genus Bas- fus, Breuer, 1944; Hung, 1982; Sanderson, 1988; Hoshi- sus Fabricius, 1804 (Agathidinae) and a few other ba, Imai, 1993; Abe, 1998; Baldanza et al., 1999; Fusu, groups. Diaprioidea: Diapriidae: all karyotypically 2008]. Using this technique, I have focused on rear- known species. Cynipoidea: Cynipidae: species of the rangements that cause substantial changes between cer- genus Diplolepis Geoffroy, 1762; Figitidae: a few spe- tain types of chromosome sets of parasitic wasps. With- cies. Platygastroidea: Scelionidae: Telenomus fariai in-type rearrangements are therefore not considered Costa Lima, 1927. Ceraphronoidea: Megaspilidae: Den- in the present paper, although they may well be respon- drocerus carpenteri (Curtis, 1829). Chalcidoidea: My- sible for additional karyotypic diversity (see for details maridae: Anaphes iole Girault, 1911; Aphelinidae: a Gokhman, 2009). The diversity of morphotypes of chro- few species; Eurytomidae: a few species; Eulophidae: mosome sets within all karyotypically studied super- Elachertus sp. families of parasitic Hymenoptera has been analyzed (9) Evanioidea: Gasteruptiidae: Gasteruption jacu- using corresponding phylogenetic reconstructions (taken lator (Linnaeus, 1758). Ichneumonoidea: Ichneu- from Gokhman [2009] and a few other sources) to monidae: most Pimpliformes and a few Ichneumoni- reveal main transformation pathways of karyotype evo- formes; Braconidae: a number of species mostly be- lution in the group studied. The principal phylogeny of longing to the cyclostome group of subfamilies (Do- the parasitic Hymenoptera used in the present paper is ryctinae, Opiinae, Alysiinae etc.). based on the study by Ronquist et al. [1999], which, in (10) No parasitic wasps having chromosome sets of turn, is a computerized reanalysis of the dataset pre- that kind are currently known. sented in the pioneering work by Rasnitsyn [1988]. In (11) Ichneumonoidea: Ichneumonidae: many Ophio- the end, possible mechanisms of the above mentioned niformes and a few Ichneumoniformes; Braconidae: a transformations have been suggested. few species, e.g. Alysia manducator (Panzer, 1799) (Alysiinae). Cynipoidea: Cynipidae: most species. Chal- Results and discussion cidoidea: Encyrtidae: a number of species; Eurytomi- dae: a number of species; Aphelinidae: some species of Morphotypes of chromosome sets of parasitic the subfamily Coccophaginae. Hymenoptera (12) Ichneumonoidea: Ichneumonidae: a few Pim- Eighteen possible classes (morphological types) of pliformes (Orthocentrinae) and Ichneumoniformes chromosome sets of parasitic wasps have been delimit- (Cryptinae and Ichneumoninae). ed. General distribution of those morphotypes by su- (13) Chalcidoidea: Aphelinidae: a few species of perfamily is listed in the Table. This information is the genera Encarsia Förster, 1878 and Aphelinus Dal- also given below in more detail. man, 1820. (1) Ichneumonoidea: Ichneumonidae: Ophioni- (14) Ichneumonoidea: Ichneumonidae: a few Ophio- formes (Mesochorinae): Mesochorus sp.; Braconidae: niformes and Ichneumoniformes (Ichneumoninae). Aphidius matricariae Haliday, 1834 (Aphidiinae) and (15) No parasitic wasps having karyotypes of that most Cheloninae. Cynipoidea: Cynipidae: Andricus kind are currently known. mukaigawae (Mukaigawa, 1913). Chalcidoidea: many (16) Chalcidoidea: Aphelinidae: Encarsia tricolor species of the families Eulophidae, Torymidae, Eu- Förster, 1878; Eupelmidae: Eupelmus linearis Förster, pelmidae and some other groups. 1860. (2) Ichneumonoidea: Ichneumonidae: Ichneumoni- (17) Ichneumonoidea: Ichneumonidae: Ichneumon- formes (Ichneumoninae): Dirophanes fulvitarsis (Wes- iformes: a few Ichneumoninae. Cynipoidea: Figitidae: mael, 1845) and Vulgichneumon saturatorius (Linnae- Ganaspis xanthopoda (Ashmead, 1896) and Leptopili- us, 1758); Braconidae (Meteorinae): Meteorus versi- na boulardi (Barbotin, Carton et Kellner-Pillault, 1979). color (Wesmael, 1835). Chalcidoidea: Mymaridae: Ana- Chalcidoidea: Eupelmidae: Arachnophaga picardi (Ber- phes listronoti Huber, 1997. nard, 1936); Encyrtidae: Ageniaspis fuscicollis (Dal- (3) Ichneumonoidea: Ichneumonidae: Ichneumoni- man, 1820). formes (Ichneumoninae): Virgichneumon digrammus (18) Evanioidea: Gasteruptiidae: Gasteruption as- (Gravenhorst, 1820). sectator (Linnaeus, 1758). Ichneumonoidea: Ichneu- (4) (6). No parasitic wasps having karyotypes of monidae: Ophioniformes (Orthopelmatinae): Orthopel- that kind are currently known. ma mediator (Thunberg, 1824). (7) Ichneumonoidea: Ichneumonidae: Ophioni- formes (Tryphoninae): Netelia latungula (Thomson, Main pathways of karyotype evolution of para- 1888); Braconidae: Charmon cruentatus Haliday, 1833 sitic Hymenoptera (Charmontinae) and a few Aphidiinae. Cynipoidea: It is difficult at present to precisely identify the Cynipidae: Andricus kashiwaphilus Abe, 1998. Chalci- ancestral karyotype for the parasitic Hymenoptera. Spe- Morphotypes of chromosome sets and karyotype