Apidologie 34 (2003) 321–331 © INRA/DIB-AGIB/ EDP Sciences, 2003 321 DOI: 10.1051/apido:2003035 Original article Nest construction and architecture of the Amazonian bumble bee (Hymenoptera: Apidae) Olivia Mariko TAYLORa, Sydney A. CAMERONb* a Department of Biology, University of Victoria, PO Box 1700 STN CSC Victoria, BC, V8W 2Y2 Canada b Department of Entomology, University of Illinois, Urbana-Champaign, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801, USA (Received 8 February 2002; revised and accepted 1 November 2002) Abstract – The Amazonian bumble bee, Bombus transversalis, is mostly restricted to tropical rain forest of the Amazon Basin. Little is known of its biology, in part because its surface nests are cryptic and hard to find. Here we examine nest site characteristics, nest architecture and construction behavior from 16 colonies observed in different regions of Amazonia. We quantify structural features of the nest habitat and external and internal characteristics of the nests. We ascertain that nests are constructed on terra firme, on the surface of the ground and incorporate elements of growing vegetation as physical support. Nests consist of a thatched canopy of tightly woven leaves and rootlets, beneath which lies the brood and food storage pots. To build the nest, workers cut and transport leaves from the surrounding forest floor. Nests are dry inside, despite torrential rainfall and external relative humidity levels near 100%. Nests may be reused although a colony appears to persist for only one season. corbiculate bees / Bombus transversalis / neotropics / leaf-cutting behavior / adaptation / colony life cycle 1. INTRODUCTION (Michener, 1974). However, the Amazonian bumble bee, Bombus transversalis (Olivier) In the family Apidae (sensu Roig-Alsina (subgenus Fervidobombus), is unusual in and Michener, 1993), the bumble bees being mostly restricted to tropical lowland rain (Bombus Latreille) comprise a group forests of the Amazon Basin (Lievano et al., of approximately 240 eusocial species 1991; Cameron et al., 1999), where perma- (Williams, 1998). They are most diversified nently humid climate, appropriate nesting sites and abundant in the temperate higher eleva- and intense predation present major challenges tions and cool, mesic regions of the Northern to most ground nesting bees, including bumble Hemisphere (Plowright and Laverty, 1984; bees (Janzen, 1971; Michener, 2000; Cameron Labougle, 1990), where they commonly nest et al., 1999; Ramírez and Cameron, in press). below ground in abandoned rodent burrows As is true of most Fervidobombus (Cameron (Sakagami, 1976). Within South America, and Williams, 2003), B. transversalis nests on bumble bees are more numerous at moderate the surface of the ground, yet this species to high altitudes in the mountains and in the found a way to survive the exigencies of living subtropical and temperate southern portions on the forest floor in tropical rain forest, of the continent, and are taxonomically which is often partially flooded. What features inconspicuous among tropical lowland fauna might have been selected or enhanced as * Correspondence and reprints E-mail: [email protected] 322 O.M. Taylor, S.A. Cameron mention the unique composition of the nest, which resembles a pile of thatched leaves (Fig. 1). Thus nesting biology seems a promising avenue from which to begin an examination of potential adaptations enabling B. transversalis to survive in the wet tropics. Nests have been described for a number of Neotropical Fervidobombus species (Tab. I), but detailed descriptions of nest site character- istics and nest architecture are wanting for B. transversalis (and many of the other Fervi- dobombus species). Four published accounts have reported on only three nests (Dias, 1958; Olesen, 1989; Cameron and Whitfield, 1996; Cameron et al., 1999), precluding comparative Figure 1. Bombus transversalis nest illustrating the conical shape, canopy characteristics and physical insights and a full understanding of their nest- support protruding through the nest (modified from ing biology. In this report we describe charac- Cameron et al., 1999). teristics of the nest sites and nest architecture from 16 colonies of B. transversalis studied at five sites within Amazonian Colombia, Brazil, B transversalis moved into rain forest habitat Ecuador and Peru. We focus on physiographic from surrounding temperate areas? Published and topological features of nesting sites and reports on colonies discovered in the wild all provide a detailed description of the nest Table I. Nesting sites described for tropical species of Bombus within the subgenus Fervidobombus (Burks, 1951; Krombein, 1967; Sakagami, 1976). Regions where the species are thought to occur (documented by Williams, 1998) are shown in italics. Species of Areas observed Habitat Nest site preference Author(s) Bombus atratus South America Tropical rain Principally surface Dias, 1960; Sakagami Franklin (Brazil) forest – highlands nesters (arboreal and et al., 1967; Zucchi, 1973; regions: W, E, S and lowlands below ground nests Lievano et al., 1991; Neotropical found) Cameron and Jost, 1998 brasiliensis South America Tropical rain Semi-subterranean Moure and Sakagami, Lepeletier regions: E forest – lowlands 1962 Neotropical medius North America Tropical evergreen Principally surface Michener and LaBerge, Cresson (Mexico) forest nesters (below ground 1954; Sakagami et al., regions: S Nearctic nests found) 1967 and N Neotropical morio South America Tropical rain Principally surface Milliron, 1961; Zucchi, (Swederus) (Brazil) forest – mid-lands nesters (below ground 1973 regions: W, E nests found) Neotropical pullatus Costa Rica Tropical rain Principally surface Janzen, 1971; Lievano Franklin regions: N, W forest – wet lowlands nesters (arboreal nests et al., 1991 Neotropical and highlands found) transversalis South America Tropical rain forest – Surface nesters Dias, 1958; Olesen, 1989; (Olivier) (Brazil) wet lowlands Lievano et al., 1991; regions: W, E Cameron et al., 1999 Neotropical Nest construction and architecture of Bombus transversalis 323 architecture summarized from the collective colonies. We also describe the means by which workers construct nests and modify the forest floor in the nest vicinity by clearing patches bare of leaf litter. We conclude that the unusual nest structure is a critical factor in the successful survival of B. transversalis in the rain forest. 2. MATERIALS AND METHODS Field colonies of B. transversalis were examined at five locations: Colombia (Amacayacu National Park, along the Amazon river [03°49’S, 70°15’W], May 17-19, May 25-June 12, 2000); Brazil (Palmarí Figure 2. A graphical representation of the Reserve along the Javarí River [04°50’S, measurement of ground slope. Two poles placed k70°01’47”W], May 20-24, 2000); Ecuador vertically into ground on up and down slopes are (Tiputini Biodiversity Station [University of San separated by a taught 310 cm string. Vertical Francisco, Quito] along the Tiputini River distance from horizontal string to ground was [00°38’18”S, 76°09’W], June 17-26, 2000 and measured at both poles (A, B) and the difference Yasuní Scientific Research Station, near the between them was used as the vertical Tiputini River, Orellana [00°05’S, 75°47’W], June measurement (C) for the angle. String length gave 26-July 7, 2000); and, Peru (Tambopata-Candamo the horizontal measurement (D). The degree of Reserve in the state of Madre de Dios [12°49’S, slope was calculated as the tangent. Slope direction 69°24’W], August 9-22, 2000). All sites were loca- (aspect) was taken as the down-slope direction. ted in rain forest of the Amazon Basin. We collected habitat measurements and nest architectural data (canopy) encasing the brood comb, the materials from 12 active nests (Tab. II). Nine of the 12 active used to construct the canopy, including its internal nests were dissected (see below) and left intact in layers. Colony construction activities and the the field, one was excavated and taken to an indoor clearing of patches on the forest floor were laboratory, and two were left undissected in the observed and recorded with video camera (Canon field. Limited data were collected from an additio- GL-1) and focal observations of marked individuals nal four nests that were found vacant, without during 20-min periods each hour through the day. brood. Colony size (adult and brood population) was Five features of the nest site were measured: estimated by counting the number of adult bees (1) relative amounts of vegetation overhanging the collected at the nest (excluding absent foragers) and nest; (2) degree of shade; (3) directional aspect of the number of different brood cells (pupae, larval the nest; (4) ground slope; and (5) presence or clusters and egg cells). Each larval cluster was absence of plant structures (roots, saplings, tree estimated to contain an average of six larvae per buttresses) providing physical support for the nest. cluster and each egg cell was estimated to contain The overhanging vegetation included small shrubs five eggs per cell. These estimations were based on and saplings, understory trees, the general forest multiple observations of brood from colonies in canopy layer, and emergent trees. Shade was different locations. classified into five ranks based on pre-established categories of the number of vegetation layers Colony dissections were performed by teasing overhanging the nest: 1= complete sun exposure, no open the nest canopy from the top center after the shade; 2= high sun exposure (limited shade bees were collected with aerial nets and kept cool in provided throughout the day); 3= even mixture of an iced container. Dissections were conducted so as shade cover and sun exposure; 4= mostly shaded to minimize time of disturbance of the nest structure (limited sun exposure throughout the day); and and brood. To complete a dissection of large nests 5= complete shade cover, no direct exposure to swiftly, with minimum exposure of brood to the sun due to thick canopy coverage. The direction of humid ambient air, only half the brood comb was the nest entrance was taken with a compass bearing. measured for size and half the cells tallied.