Biological Observations on a Neotropical Parasocial Bee, Eulaema Nigrita, with a Review on the Biology of Title Euglossinae (Hymenoptera, Apidae)
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Biological Observations on a Neotropical Parasocial Bee, Eulaema nigrita, with a Review on the Biology of Title Euglossinae (Hymenoptera, Apidae). : A Comparative Study (With 21 Text-figures, 2 Plates and 9 Tables) Author(s) ZUCCHI, Ronaldo; SAKAGAMI, Shôichi F.; CAMARGO, João M. F. de Citation 北海道大學理學部紀要, 17(2), 271-380 Issue Date 1969-12 Doc URL http://hdl.handle.net/2115/27489 Type bulletin (article) File Information 17(2)_P271-380.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP Biolo~ical Observations on a Neotropical Parasocial Bee, Eulaema nigrita, with a Review on the Biolo~y of Eu~lossinae (Hymenoptera, Apidae). A Comparative Studyl) 2) By Ronaldo Zucchi, Shoichi F. Sakagami and Joao M.F. de Camargo Departamento de Genetica, Faculdade de Filosofia, Ciencias e Letras, Ribeirllo Preto, Zoological Institute, Hokkaido University, Sapporo and Departamento de Genetica, Faculdade de Medicina, Ribeirllo Preto (With 21 Text-figures, 2 Plates and 9 Tables) The female of the handsome golden and black Euglossa Surinamensis visits the caj u trees, and gathers with its hind legs a small quanti ty of the gum which exudes from their trunks. To this it adds the other material from the neighbouring bushes, and when laden flies off to its nest. H.W. Bates Naturalist on the River Amazons. Contents Introduction ... .. 271 6. Male behavior. .. 352 Materials and methods .............. 274 7. Associated animals .......... 366 Results and discussions .............. 282 8. Habitat preference and pheno- 1. Nests ........................ 282 logy ........................ 369 2. Immature stages .............. 305 Summary ........................ 370 3. Female behavior .............. 311 Acknowledgements ................ 372 4. Social organization ............ 331 References . .. 372 5. Flower visits .................. 344 Addenda. .. 379 Introduction The aim of the present paper is twofold, first to give the full account of observations made on nest architecture and behavior of an euglossine bee, Eulae- 1) Contribution No. 871 from the Zoological Institute, Faculty of Science, Hokkaido University, Sapporo, Japan 060 2) Dedicated to Prof. Theodosius Dobzhansky at his 70th birthday (Jan. 25 1970). Jour. Fac. Sci. Hokkaido Univ. Ser. VI, Zool. 17, 1969 271 272 R. Zucchi et al. ma (Apeulaema) nigrita Lepeletier (cf. Zucchi 1966, Zucchi MS), together with some notes on nests and flower visits of Euplusia auriceps (Friese) and Euglossa (Euglossa) cordata Linne, secondly to summarize the previous information published on the biology of the Subfamily Euglossinae. The euglossine bees form, together with bumblebees (Bombinae), stingless bees 1 (Meliponinae) and honeybees (Apinae), the Family Apidae ), which represents by its behavioral differentiation one of the summits in invertebrate evolution. Com paring the biological information so far obtained, however, we at once notice a conspicuous unbalance among these four groups. Although there are still much to be clarified as to the complicated life of honeybees, they are certainly biologically one of the best known insectB (Ribbands 1953, v. Frisch 1965, etc.). Our knowledge on bumblebees is still far less satisfactory than on honeybees, but the outline of their mode of life is already well recognized as compiled by Free and Butler (1959). Concerning stinglees bees, the previous information was summarized by Schwarz (1948), followed by Moure, Nogueira-Neto and Kerr (1958). There after some interesting facts on behavior have successively been recorded (Esch 1967, Lindauer and Kerr 1958, Kerr 1950, Nogueira-Neto 1954, Sakagami et al. 1963, Sakagami and Zucchi, 1966, 1968). But the biology of this Pantropic group is a rich but still insufficiently explored treasure, in spite of their highly evolved social organization, attaining a level comparable to honeybees. Finally the biology of euglossine bees is least known. Virtually no single species has biologically precisely been studied. Nevertheless they form an important link for the full understanding of apid evolution. Fortunately we could discover a nest of Eulaema nigrita, transfer it into an observation case and study their intranidal behavior, which has so far been concealed from any specialists of bee biology. The obtained results are still fragmentary in many aspects but involve some interesting novelties in comparison with the behavior of the other three groups. In preparing the descriptions of these results, we felt the necessity of compiling the previous information on the biology of Euglossinae. Obviously we need a number of studies for the future, specially planned and executed for this group. But such work is, or has been, not always easy to realize, because of their tropically limited range, their scarcity and secretive mode of life. Consequently the majority of previous information is incidental and fragmentary. Attempts to summarize such records were tentatively made by Schrottky (1907) and Friese (1930, 1941). Maa (1953) gave a synoptic table of both morphological and biological features of four groups of Apidae. But none of them tried to gather all available records nor to analyse them sufficiently. Fortunately, several papers on the biology of Euglossinae, much more precise than those in earlier times, have recently been published, encouraging for the advance of our knowledge on these bees. In such circumstance, it 1) The classification system differs among specialists. Michener (1944) : The Subfamily Apinae with four tribes, Euglossini, Bombini, Meliponini and Apini; Moure, Nogueira·Neto and Kerr (1958) : The Family Apidae with four subfamilies, Euglossinae, Bombinae, Meliponinae and Apinae; and Michener (1965a) : The Family Apidae with three subfamilies, Euglossinae, Bombinae and Apinae (with Meliponini and Apini). Here we follow the system by Moure et al., simply for the convenience in discussions. Biological Observations on Eulaema nigrita 273 may be appropriate to give a compilation of previous records, to facilitate further studies of this fascinating, but so far ignored group. Probably because of its rather singular external appearance, the Subfamily Euglossinae has occasionally been placed outside of Apidae (for instance, Hand lirsch 1925), but it shares several good characters with other apid groups, though some of them such as the possession of developed corbiculae are secondarily modified in parasitic genera. Within Apidae Euglossinae is (PI. IV-B) distinguished by a number of conspicuous features, including brilliantly metallic integument (Plate IV), extraordinary prolongation of glossa, peculiarily concave male sternum VIII, highly differentiated male legs, etc. All four groups of Apidae are quite distant one another, but Euglossinae may stand relatively closer to Bombinae than the other two highly social groups. Maa (1953) considers Euglossinae as the most primitive among four groups, without giving sufficient explanation. But it is still premature to give any definite conclusion from the morphological point of view, because beside some apparently primitive features, this group possesses some highly specialized ones. Sociologically it is certainly most primitive in the sense that many species are seemingly solitary or possessing only weakly developed social system (cJ. Section 4). For the time being, however, it may be wise to avoid a careless amalgamation of biological and morphological interpretations. Since a monograph by Friese (1899) a good number of species had been described but no attempt to establish the taxonomic framework had been made until Moure (1950) gave a key to the genera. Thereafter he described a number of additional species (1963",1967a) and gave a check list of so far known species (1967b). According to his system, the present status of the subfamily is summa rized as follows: Genera and main characters Subgenera and main characters Number of species in Moure (1967 b) Euglossa Latreille 1802 Euglossa 8. str. (PI. IV-A, 1,2,4) 39 Nonparasitic, small to Small, male mandibles medium sized, brilliantly bidentate, glossa long metallic with poor pubescence Euglossella Moure 1967 15 (PI. IV-A, 3,5). Small, male mandibles tridentate, glossa long Glossura Cockerell 1917 12 (PI. IV-A, 6~ 9, B). Medium, glossa very long Euplusia Moure 1943 (Pl.IV-A, 12~ 23) 45+ (=Plusia Hoffmannsegg 1817) Nonparasitic, medium to large, moderately metallic and haired Eufriesea Cockerell 1908 (PI. IV-A, 11) 1 Ditto, with large plate like mesoscutellum 274 R. Zucchi et al. Eulaema Lepeletier 1841 Eulaema s.str. (Pl.IV-A, 27,30-32) 14 (=Eulema auct.) Metasoma. extensively metallic, Large to very large, less male face without white marks metallic and more hairy Apeulaema Moure 1950 (Pl.IV-A, 26,28,29) 6 Metasoma less metallic, male face with white marks Aglae Lepeletier et Serville 1825 1 (Pl.IV-A, 25) Parasitic, brilliantly metallic, body fiat, mesoscutellum plate like Exaerete Hoffmannsegg 1817 5 (Pl.IV-A, 24) (=Chrysantheda Perty 1833). Parasitic, brilliantly metallic, mesoscutellum tuber culate In total 137 Certain biological differences among these genera will subsequently be mentioned. Concerning the nonparasitic genera, three items are specially noteworthy: 1) Absence of typical subterranean nidification by self-excavation. 2) Abundant use of resin for nidification and 3) Appearance of an incipient social complexity in some species. Furthermore, males of many species have a special preference for the flowers of certain, often quite definite orchid species, accompanied with