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• W438512627 abstract "Nesting soils of 32 species of fossorial bees were sampled from diverse habitats across the continental United States. These soils classified texturally as sands or loams. Broadly diverse values for bee size, cell depth, soil moisture and texture, and average annual precipitation and temperature were represented in the sample, but each varied independently from the others, and was not predictable from a bee species' familial or tribal affiliation. species secreting markedly different Dufour's gland lipids, which are used to waterproof the walls of their subterranean nest cells, could be found nesting under similar edaphic and climatic conditions. Most species of bees nest in the soil. Females excavate subterranean tunnels terminated by enlarged chambers or cells which they provision with pollen and nectar masses. Fossorial nesting is characteristic of the Andrenidae, Fideliidae, Halictidae, Melittidae, Oxaeidae and Stenotritidae. The majority of the Colletidae and Anthophoridae also nest underground. Nesting above-ground only predom inates among the Megachilidae, Apidae, Hylaeinae and Xylocopinae. Although bees are best known as adults that appear briefly each year, individuals of most species spend the greater part of their lives underground as immatures (Linsley, 1958; Malyshev, 1935; Michener, 1974; Roubik, 1989). Bees nest in diverse earthen habitats, ranging from weathered sandstone (Custer, 1928) and the mortar of prehistoric adobe walls (pers. obs.) to embankments of dense clay, deep alluvial silts, and sand dunes of deserts and beaches (Malyshev, 1935; Stephen et al., 1969; Roubik, 1989). Horizontal or vertical nesting sites are typically considered well-drained (Linsley, 1958), but species of neotropical Epicharis (Roubik and Michener, 1980), palearctic Dasypoda (J. Teng?, pers. comm.) and nearctic Nomia (P. Torchio, pers. comm.) are known to nest suc cessfully at sites that are periodically submerged. Many solitary bee species nest in dense aggregations, which suggests that females select sites with particular surface or edaphic attributes. The alkali bee, Nomia melanderi, for instance, may aggregate in the tens of thousands in soils charac terized by less than 8% clay and slow surface seepage (Stephen, 1960). Nesting females of this species respond to hot ambient temperatures by excavating nest burrows more deeply in the soil (W. P. Stephen, pers. comm.). In the blueberry barrens of Maine, Osgood (1972) found that species of Collet?s, Andrena and halictine bees nested more frequently in soils whose surface organic layers were thinner than neighboring locations. In contrast, a comparative study of Brazilian bank-nesting species (Michener et al., 1958) concluded that female philopatry was primarily responsible for the aggregation of nests, as neighboring, seemingly equiv alent sites in the same embankments remained unused. Few other comparative or quantitative studies have been made for the nesting soils of bees. Accepted for publication 4 March 1991. This content downloaded from 157.55.39.45 on Thu, 01 Sep 2016 05:08:35 UTC All use subject to http://about.jstor.org/terms VOLUME 64, NUMBER 4 407 My objectives here are to 1) survey the nesting soils of taxonomically diverse bee species across a broad range of soil moistures, textures, habitats, and biomes of North America, and 2) reveal any predictive correlations of these edaphic measures with bee size, climate, taxonomic affiliation or to each other. Materials and Methods Soil surrounding the shallowest provisioned cell of an active nest was collected into an airtight jar. The fresh 30-100 g sample was weighed, dried for 3-5 d in an oven at 30? to 40?C, and reweighed (?0.1 g). Soil moisture content was cal culated gravimetrically as: Weight loss/dry weight x 100 (Brady, 1984) Cell depth was measured vertically to the soil surface. The provisioning female bee was collected for identification. Her size was estimated by the shortest distance between her wing tegulae as measured using a dissecting stereomicroscope and ocular micrometer (Cane, 1987). Particle-size distribution of the dried soil was analyzed by standard sedimen tation techniques. Briefly, a sample was suspended in solution with the aid of a blender and a detergent dispersing agent. Specific gravity, measured using a hy drometer as the suspension settled, was used to calculate the proportion of silt and clay fractions, with sand as the remainder. In later analyses, particles >2 mm diam (gravel) were first removed by sieving. Textural classification of the soils follows the system of the U.S. Dep't. of Agriculture (Brady, 1984). Average annual temperature and precipitation were taken from the nearest reporting weather station (NOAA, 1983; Brown, 1982; Ting and Jennings, 1976) or more local sources (US Nat'l. Weather Service, Auburn, Ala.; Bodega Bay Marine Laboratories, Bodega Bay, Calif.; S'west. Research Station of the Amer. Museum Nat. Hist., Portal, Ariz.). Relationships among the variables were explored using several multivariate approaches available through NTSYS Ver. 1.50 (Rohlf, 1988). Following default standardization of the cases, a correlation matrix was calculated using all of the variables. The species were then clustered by UPGMA using the correlation val ues. These correlation values were also used to construct a matrix of cophenetic similarity values to assess the goodness of fit of the above clustering to the data. A second analysis, principal components, was performed with the standardized data for soil textures, moisture and nesting depth. Eigenvectors and eigenvalues were calculated from the correlation matrix. Data obtained for each bee species were then projected onto the first two or three eigenvectors and plotted against each other. In all, nests of 32 species of ground-nesting bees were analyzed, representing 22 genera and seven families of the North American fauna (Table 1). Results and Discussion Soil moistures for this sampling of bee species varied over a 15 x range, which reflected (at least in part) the 19 x range in annual rainfall. Soil textures ranged from sands to silt loams and clay loams. No bees were found nesting in clay or silt soils. All of the soils examined were at least %, and as much as 94% sand. The finer particle fractions, being silts and clays, were less abundant, ranging from absence to Vi of the soil dry weight (Table 1, Fig. 1). species nesting in This content downloaded from 157.55.39.45 on Thu, 01 Sep 2016 05:08:35 UTC All use subject to http://about.jstor.org/terms Table 1. Species of ground-nesting bees sampled in this survey. Size is intertegular span in mm, annual precipitation and nest cell depth are in cm, and annual temperature is degrees C. The Bee no. corresponds to those used in the figures. o 00 Family Subfamily/tribe Depth Moisture %clay %silt %sand %gravel Precip. Temp. Classification Macropodinae MEGACHILI DAE" @default.
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• W438512627 title "Soils of ground-nesting bees (Hymenoptera: Apoidea): texture, moisture, cell depth and climate" @default.
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