Jerome G. Rozen

Eggs, Ovariole Numbers, and Modes of Parasitism of Cleptoparasitic Bees, with Emphasis on Neotropical Species (Hymenoptera: Apoidea)

Abstract The shapes, sizes, and chorionic ornamentation of mature oocytes/eggs are described along with ovariole and mature oocyte numbers of six lineages of primarily South American cleptoparasitic bees. This information is related to whether the eggs are introduced into brood chambers that are still open and being provisioned by the host female or whether the chambers have already been closed by the host females. The lineages, all in the Apidae, are as follows: (1) Kelita (Nomadinae: Brachynomadini), (2) Isepeolus and Melectoides (Apinae: Isepeolini), (3) Leiopodus (Apinae: Protepeolini), (4) Rhathymus (Apinae: Rhathymini), (5) Mesoplia and Epiclopus (Apinae: Ericrocidini), and (6) Exaerete (Apinae: Euglossini). A table in the section on Discussion and Analyses summarizes information on mature oocyte/egg size (egg index), total number of mature oocytes, mature oocytes per ovariole, and ovariole number (ovariole formula) for all taxa of cleptoparasites, worldwide, that have been studied to date. It shows that almost all of the Nomadinae have more than the plesiomorphic number of ovarioles, a feature also found in two of the three studied genera of the Ericrocidini. All other cleptoparasitic lineages lack extra ovarioles. The potential selective advantage of extra ovarioles is discussed. Also discussed is whether the large number of mature oocytes carried by cleptoparasites might result, in part, from the length of time required for chorion deposition after the oocytes reach maturity. The table shows not only that the mature oocytes/eggs of cleptoparasitic bees in general tend to be smaller than those of solitary bees, but that the mature oocytes/eggs of those cleptoparasites that hide their eggs in open host brood cells are significantly smaller than those that introduce their eggs into cells that have been closed by the host. The potential selective advantages of small egg size in cleptoparasitism are explored. Lastly, the unusual modified shapes of mature oocytes/eggs and the thick chorions of cleptoparasites that oviposit in open host cells are attributed to ways of protecting the eggs from discovery and damage by returning host females. Appended is a scanning electron micrograph of the micropyle of the North American Stelis elongativentris Parker (Megachilidae: Anthidiini), the ovariole and oocyte statistics of which have been published earlier. Also appended are a description and illustrations of the oocyte of Coelioxys novomexicana Cockerell (Megachilidae: Megachilini).

Nesting Biologies and Immature Stages of the Tapinotaspidine Bee Genera Monoeca and Lanthanomelissa and of Their Osirine Cleptoparasites Protosiris and Parepeolus (Hymenoptera: Apidae: Apinae)

Abstract The nesting biologies of Monoeca haemorrhoidalis (Smith) and Lanthanomelissa betinae Urban (Tapinotaspidini) are described from southeastern Brazil. Both are ground nesting; the nests of the former are attacked by the cleptoparasite Protosiris gigas Melo (Osirini), and those of the latter are attacked by Parepeolus minutus Roig-Alsina (Osirini). Egg eclosion, larval feeding behavior, and cocoon spinning of M. haemorrhoidalis are detailed. A female of P. gigas opens the closed cell of M. haemorrhoidalis by making a large opening in the cell cap (which is plugged after ovipositioning) through which she apparently extends her metasoma. Indirect evidence suggests that she uses her metasomal apex, and perhaps even the sting, to kill the host egg or early instar. Protosiris eggs are either attached to the cell-wall surface of the nearly vertical host cells or dropped onto the surface of the provisions. First instars of P. gigas, with strongly curved, sharply pointed mandibles, are also capable of killing host immatures or competing cleptoparasites. Cocoons of all four species are compared and contrasted. The egg, all larval instars, and pupa of Monoeca haemorrhoidalis are described, as are the egg and postdefecating larva of Lanthanomelissa betinae. The egg, all larval instars, and pupa of Protosiris gigas are described, as is the postdefecating larva of Parepeolus minutus. Both Monoeca haemorrhoidalis and Protosiris gigas have four ovarioles per ovary. The egg indices and other ovarian features of both species are identified and discussed. The possible phylogenetic relationship of the Tapinotaspidini with the Osirini is briefly explored on the basis of data from this study. Possible phylogenetic relationships of the Osirini with other cleptoparasitic apids are analyzed. In the appendix,, the identity of the species of Monoeca, whose nesting biology is presented in the main paper, is discussed. The species is M. haemorrhoidalis (Smith, 1854), a species closely related to M. schrottkyi (Friese, 1902) and M. xanthopyga Harter-Marques, Cunha, and Moure, 2001. An identification key for distinguishing these three species is presented. Tetrapedia piliventris Friese is placed as a junior synonym of M. haemorrhoidalis (new synonymy). A lectotype is designated for Pachycentris schrottkyi Friese. The species of Protosiris found attacking M. haemorrhoidalis is here described as new, P. gigas Melo sp. nov. It is structurally most similar to P. caligneus (Shanks), from which it differs by its abundant yellow marks, plumose pubescence on the lower paraocular area, protruding anterior mesoscutum, and sparser punctation on the metasomal terga. Appendix: Taxonomic Notes on Monoeca and Description of a New Species of Protosiris, by Gabriel A. R. Melo

Oocytes, Eggs, and Ovarioles of Some Long-Tongued Bees (Hymenoptera: Apoidea)

Abstract NOTE: Appendix: Parammobatodes rozeni, a New Bee Species from Israel, by Maximilian Schwarz3 We present here new information regarding the number of ovarioles and the number and size of mature oocytes of certain bee taxa collected on a field trip in Turkey in June and July 2001. This information is augmented with similar data concerning taxa related at the tribal level to those that we found in Turkey. The mature oocytes/eggs of all taxa listed below are described and most are illustrated by photographs, scanning electron micrographs, and/or line drawings, and comparisons are made with previously published descriptions. These taxa, arranged by family, subfamily, and tribe, are as follows: MEGACHILIDAE: Megachilinae: Dioxyini: Dioxys cincta (Jurine), Dioxys pacificus Cockerell. APIDAE: Xylocopinae: Xylocopini: Xylocopa (Proxylocopa) olivieri Lepeletier; Nomadinae: Ammobatoidini: Holcopasites insoletus (Linsley), Holcopasites tegularis Hurd and Linsley; Nomadinae: Biastini: Biastes brevicornis (Panzer); Nomadinae: Ammobatini: Ammobates carinatus Morawitz, Parammobatodes rozeni Schwarz, Oreopasites (Oreopasites) favreauae Rozen, Oreopasites (Oreopasites) vanduzeei Cockerell, O. (Perditopasites) barbarae Rozen, O. (P.) linsleyi Rozen, “Parammobatodes” orientana (Warnke), Pasites maculatus Jurine, Sphecodopsis (Pseudodichroa) capensis (Friese), S. (P.) fumipennis (Bischoff); Apinae: Melectini: Melecta albifrons albovaria Erichson, Melecta species, Thyreomelecta kirghisia Rightmyer and Engel, Thyreus lieftincki Rozen, Xeromelecta californica (Cresson). The mature oocyte of Xylocopa (Proxylocopa) olivieri, a ground-nesting species, is found to be large relative to the body size of the female but somewhat smaller than the “giant” eggs of wood-nesting Xylocopa, as classified by Iwata and Sakagami (1966. Gigantism and dwarfism in bee eggs in relation to the mode of life, with notes on the number of ovarioles. Japanese Journal of Ecology 16: 4–16). Egg deposition habits of some cleptoparasitic taxa are discussed. In addition to smaller size relative to body size, eggs of cleptoparasitic bees show great variation in micropylar structure, dimensions, and chorionic ornamentation, thickness, and patterning, compared with eggs of nonparasitic bees. An appendix by Maximilian Schwarz describes and names Parammobatodes rozeni, new species, from Israel.

Biology of the Cleptoparasitic Bee Mesoplia sapphirina (Ericrocidini) and Its Host Centris flavofasciata (Centridini) (Apidae: Apinae)

ABSTRACT This paper investigates the bionomics of the cleptoparasitic bee Mesoplia sapphirina Melo and Rocha-Filho, sp. nov. (described in the appendix), and of its ground-nesting host Centris flavofasciata Friese found along the Pacific coast of Guanacaste Province, Costa Rica. We explore the host-nest searching behavior, egg deposition, and hospicidal behavior of M. sapphirina. Anatomical accounts of its egg, first, second, and fifth larval instars are presented and compared with published descriptions of other ericrocidine taxa. Nests of the host bee as well as its egg and method of eclosion are also described.

Immature Stages of the Cleptoparasitic Bee Dioxys cincta (Apoidea: Megachilidae: Megachilinae: Dioxyini)

Abstract The fourth larval instar and pupa of Dioxys cincta are described and illustrated, and anatomical details of the first and last larval instar are interpreted from cast exuviae. The cocoon of this species is also described. The discovery of a hatched egg containing the first-instar exuviae indicates that the first instar remains mostly surrounded by the egg chorion. The second instar emerges through a hole in the thick dorsal surface of the chorion, presumably by chewing its way out. This discovery and the interpretation of the larval head anatomy of this species and that of D. pomonae Cockerell indicate that the genus has five larval instars. The second, third, and fourth instars are adapted to destroying the host egg or larva and any competing cleptoparasites. The fifth instar is not so adapted. As with Dioxys, other cleptoparasitic Megachilidae tend to have a sequential series of instars modified for attacking immatures of hosts and competing cleptoparasites, although which instars are so adapted varies. This contrasts with the cleptoparasitic lineages of the Apidae in which only a single instar, usually the first, is hospicidal.

Last Larval Instar and Mature Oocytes of the Old World Cleptoparasitic Bee Stelis murina, Including a Review of Stelis Biology (Apoidea: Megachilidae: Megachilinae: Anthidiini)

Abstract Herein we describe the mature oocyte and last larval instar of Stelis (Stelis) murina Pérez, a cleptoparasite associated with Osmia (Pyrosmia) submicans Morawitz near Ismailia, Egypt. The mature oocyte is compared with that of Stelis (Stelis) elongativentris Parker and found to be approximately equal in size. The mature oocyte of S. murina is also very close in size to that of its host, an unusual phenomenon in host-cleptoparasite relationships in bees. A review and analysis based on literature accounts of what is known about the mode of cleptoparasitism of Stelis is offered. Added are observations on the biology of Stelis murina resulting from our fieldwork. The mature larva of Stelis murina is described and found similar but not identical to those of other known Stelis larvae. We also include a preliminary key to the genera of cleptoparasitic megachilids based on known mature larvae and also a summary describing the modes of cleptoparasitism by these taxa.

Hospicidal Behavior of the Cleptoparasitic Wasp Sapyga luteomaculata And Investigation into Ontogenetic Changes in Its Larval Anatomy (Hymenoptera: Vespoidea: Sapygidae)

Abstract This study describes the nesting biology of the Egyptian cleptoparasitic wasp Sapyga luteomaculata Pic, which attacks the nests of two species of bees, Osmia submicans Morawitz and Megachile minutissima Radoszkowski, both belonging to the Megachilidae. We include descriptions of the egg/mature oocyte and of the first and last larval instars. We also identify the anatomical changes in the larva that take place as it transforms through intermediate instars from a host-killing first instar to a form adapted to feeding on the stored provisions supplied by the host. Biological subjects treated are egg deposition, ovariole statistics, eclosion, larval behavior including cocoon construction and defecation, and competition with other cleptoparasites. Comparisons are made throughout with accounts of other Sapyginae. Many biological and larval anatomical similarities between Sapyga and cleptoparasitic bees are recognized, and only a few possible cleptoparasitic novelties are identified.

Notes on the Egg and Egg Deposition of the Cleptoparasite Thyreus ramosus (Hymenoptera: Apidae: Melectini)

Abstract Evidence is presented that females of the cleptoparasite Thyreus ramosus (Lepeletier) insert their eggs into the closed host cells of an Anthophora (Dasymegilla), tentatively identified as A. muscaria Fedtschenko, just as other members of the Melectini whose egg deposition habits are known insert their eggs into host cells. The eggs of both host and cleptoparasite are described and illustrated. Comparison of the eggs of T. ramosus with those of other melectines strongly suggests that the melectines are monophyletic.

Interspecific Variation in Immature Larvae of the Cleptoparasitic Bee Genus Coelioxys (Hymenoptera: Megachilidae)

Abstract We describe the immature larvae of Coelioxys (Liothyrapis) decipiens Spinola and compare them with those of Coelioxys (Boreocoelioxys) octodentata Say and C. (B.) sayi Robertson. All three species have five larval instars, and, in each case, the third instar is primarily responsible for assassinating the host immature. However, the anatomy of the third instar of C. decipiens appears to be plesiomorphic compared with highly modified features of the third instars of the other two species. This suggests that a study of the third instars of other taxa in this diverse genus might play a significant role in sorting out the phylogenetic relationships within the genus, which contains more than 320 species worldwide.

Biology of the Bee Hoplitis (Hoplitis) monstrabilis Tkalců and Descriptions of Its Egg and Larva (Megachilidae: Megachilinae: Osmiini)

Abstract Herein we describe the nesting biology of the solitary ground-nesting bee Hoplitis (Hoplitis) monstrabilis Tkalců from eastern Turkey. Its shallow nests in the ground differ from the known nests of members of subgenus Hoplitis, most of which make mortar and pebble nests either on the exposed surfaces of rocks or within stems or other cavities. Cells are not lined with flower petals or other vegetative tissue, as expected for subgenus Hoplitis, but unlike other ground-nesting species of Hoplitis belonging to other subgenera such as Anthocopa. The egg of this bee is also described and illustrated, as is the fifth (last) larval instar.