AMERICAN MUSEUM Notltates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET NEW YORK, N.Y
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AMERICAN MUSEUM Notltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2630 AUGUST 22, 1977 JEROME G. ROZEN, JR. Biology and Immature Stages of the Bee Genus Meganomia (Hymenoptera, Melittidae) AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2630, pp. 1-14, figs. 1-27 August 22, 1977 Biology and Imnmature Stages of the Bee Genus Meganomia (Hymenoptera, Melittidae) JEROME G. ROZEN, JR.' ABSTRACT The hibernating, postdefecating larva and the Melittidae cannot be evaluated on the basis pupa of Meganomia binghami (Cockerell) are of larvae because the numerous similarities described and compared with similar stages of shared by Meganomia, Melitta, and Macropis are other melittid bees. Biological information on symplesiomorphic. However, there are no larval this species and on an undescribed species of characteristics that would exclude Meganomia Meganomia includes the following information: from the family. Cladistic analysis of features of nest ecology, nest configuration and structure, the mature larva indicates that Meganomia is a provisioning, development, number of genera- sister group to all other melittids whose imma- tions a year, mating, sound production and daily tures are known. activity of adults. The placement of the genus in INTRODUCTION Meganomia is a southern African genus of families except for Ctenoplectrinae, the imma- large-bodied, conspicuously marked bees that had tures ofMeganomia were then unknown. been placed in the halictid subfamily Nomiinae On a trip of South West Africa in 1976, my until Stage (1971) demonstrated that it belonged wife and I made a special effort to find and to the Melittidae on the basis of numerous struc- recover larvae of this genus, partly to confirm tural features of adults. Transferring it to that Stage's taxonomic placement of the genus and family, he stated that "it fit nicely" in the partly to see how the larvae would fit Rozen Melittinae although he did not enumerate what and McGinley's hypothesized phylogenetic rela- characters he used for this decision. More re- tionships within the family. Not only did we cently Rozen and McGinley (1974) investigated collect larvae and pupae ofMeganomia binghami the phylogeny and systematics of the Melittidae (Cockerell) we also obtained considerable bio- using mature larvae as the data source. Although logical information on the same species and a larvae of a number of taxa were described and limited amount of such information on another, analyzed, including representatives of all sub- distinctive species ofMeganomia yet to be named 'Deputy Director for Research and Curator of Hymenoptera, the American Museum of Natural History. Copright O The American Museum of Natural History 1977 ISSN 0003-0082 / Price $1.15 2 AMERICAN MUSEUM NOVITATES NO. 2630 and described. The present paper describes the vanced than others. At 17 km. east of Usakos, immature stages of M. binghami and reports on we saw no pollen-bearing females entering nests; all biological information that was gathered; con- at 62 km. west of Omaruru, only a few pollen clusions regarding relationships and placement of carrying females were observed and no fresh the genus are offered. cells were uncovered. However, at 52 and 58 km. I thank my wife, Barbara, who helped in all southwest of Omaruru pollen-laden females were aspects of the field investigations. Drs. Mary- abundant, and provisioned cells were encoun- Louise and Michael Penrith and Director C. G. tered although none was occupied by last instar Coetzee of the State Museum, Windhoek, South larva. Meganomia species B apparently flies West Africa, extended many courtesies and somewhat earlier than M. binghami, for we often advice while we were in southern Africa, for which saw pollen-laden females with frayed wings, we are grateful. Most of the manuscript was dic- whereas females of M. binghami, all fresh, were tated on tape in noisy hotel rooms or still noisier just beginning to appear. airplanes. My thanks go to Ms. Phyllis Browne Abundant rains had fallen in South West who painstakingly deciphered these tapes as she Africa during the late summer so that vegetation has done for many other similar studies. Dr. at all sites was green and flowers plentiful. Late Vincent Whitehead, South African Museum, summer rains also had been moderately heavy Capetown, and Dr. L. Vari, of the Transvaal for several preceding years. Museum, Pretoria, kindly permitted me to ex- Description of Sites. All nesting sites of amine their respective collections in connection M. binghami were in nearly horizontal ground with this study. This investigation was partly where the soil was sandy, moist below the sur- supported by National Science Foundation grant face, and partly or completely barren on the no. GB32193. surface. At 17 km. east of Usakos (fig. 1) and Specimens of adults, immatures, and cocoons 62 km. west of Omaruru (fig. 2), the sites were in described here are deposited in the collections of a partly denuded area a few meters from a graded the American Museum of Natural History. dirt roadway, whereas at the two localities south- west of Omaruru (fig. 3) most nests were actually in the margin of a dirt roadway although some BIOLOGY nests occurred on adjacent ground. We excavated Description of Area. We commonly encoun- nests only at 62 km. west and 52 km. southwest tered Meganomia binghami in central South West of Omaruru. At the former locality the soil was Africa in thorn scrub savanna where its pollen compact, moderately fine, and easily excavated plant, Crotalaria podocarpa (Leguminosae), oc- sand; from here we gathered information regard- cured in sandy situations. We found the bees in ing cocoons and postdefecating larvae. At the large numbers at 17 km. east of Usakos, S.W.A.; latter, the surface soil was drier and harder, but 8 and 26 km. north of Karibib, S.W.A.; 52 and the subsurface soil was fairly easy to excavate. 58 km. southwest of Omaruru, S.W.A.; and 62 The soil texture differed primarily by being km. west of Omaruru. Nesting sites were discov- coarser. This locality yielded data on eggs, pollen ered at all localities except the two north of masses, and early larval instars. Karabib and were studied between March 18 and Nesting Activity. Females constructed nests in 27, 1976. open horizontal ground, under low plants (in- At 17 km. east of Usakos, another species of cluding Indigofera) growing on flat surfaces, and Meganomia flew in the same vicinity as M. bing- in the sloping surfaces of low banks along the hami, and collected pollen from Indigofera, also edges of roadways. Where abundant, nests were a legume; currently unnamed, I call it Megano- in large, loose aggregations, which at 58 km. mia species B. Because its nests were not found, southwest of Omaruru (fig. 3), the largest nesting most of the following biological information site, extended along the roadway and its margins refers to M. binghami for approximately 150 meters. Within aggrega- The nesting season was early for M. binghami tions, nests were irregularly distributed, and, although some nesting sites were farther ad- when dense, were often less than 10 cm. apart. 1977 ROZEN: MEGANOMIA 3 FIGS. 1-3. Nesting sites of Meganomia binghami in South West Africa. 1. At 17 km. east of Usakos. 2. At 62 km. west of Omaruru. 3. At 58 km. southwest of Omaruru. In figures 1 and 2, burrows were most common in middle foreground; in figure 3, dark tumuli of burrows can be seen along edge of road. At 58 km. southwest of Omaruru, the adjacent Our casual observations indicated that most tumuli formed a striking, continuous row of nests were occupied by only one female, but two freshly excavated soil along the edge of the road, nests at 62 km. west of Omaruru had two somewhat darker than the roadway (fig. 3). females each. Although these two nests may have 4 AMERICAN MUSEUM NOVITATES NO. 2630 been entered by females searching for places to begin nests, both females from one nest had somewhat frayed wings, suggesting that neither had just begun nesting. All entrances were sur- AULUS rounded by copious tumuli, measuring approxi- mately 10 cm. in diameter and 5 to 8 cm. high in cases of horizontal surfaces. The tumulus was loose, except that most incorporated a curved entrance turret (fig. 5) of consolidated sand buried in the soft excavated material. The turret rose from the nest, curved, then extended hori- 4 zontally as much as 5 cm. so that the entrance was near the edge of the tumulus. Nest entrances with or without turrets appeared open most of the time although several were found plugged, perhaps by females bringing soft soil to the sur- 6 face. Holes without tumuli, encountered at some 100 cm nest sites, were emergence openings. Main tunnels and branches, circular in cross FIGS. 4-6. Nest components of Meganomia section and about 10 mm. in diameter, possessed binghami. 4. General configuration of main burrow at 62 km. west of Omaruru. 5. Nest smooth, seemingly uncoated walls. Several main entrance. 6. Diagram of cell showing orientation, burrows excavated carefully at 62 km. west of orientation of provisions, and position of egg. Omaruru descended not quite vertically in a Scale refers to figure 5 only. slightly irregular path to depths of as much as 114 cm. (fig. 4) but as yet lacked cells. At this locality, numerous cells from previous genera- had applied a waterproof lining. The cell closure tions were encountered at depths commonly was a deeply concave spiral on the inside. That it between 70 and 120 cm.