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Effects of Cultivation and Proximity to Natural Habitat on Ground-Nesting Native Bees in California Sunflower Fields

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Effects of Cultivation and Proximity to Natural Habitat on Ground-Nesting Native Bees in California Sunflower Fields JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 79(4), 2006, pp. 309–320 Effects of Cultivation and Proximity to Natural Habitat on Ground-nesting Native Bees in California Sunflower Fields 1 2 3 JOHN KIM, NEAL WILLIAMS, AND CLAIRE KREMEN ABSTRACT: Agricultural conversion is one of the most prevalent anthropogenic uses on the terrestrial earth. Persistence of organisms in such landscapes is thought to be related to species- specific characteristics such as life history traits and dispersal distance. In an agricultural landscape in California, we examined local (farm-level) and landscape variables associated with nesting pref- erences of native ground-nesting bees. Compared to the known ground-nesting visitors to crops, bee community nesting on farms was depauperate. Further, more abundant and diverse communities of bees were found nesting at farms with patches of natural habitat near by than farms that were far away from natural habitat. Species responded differently to soil conditions created by farming practices, but the variability in nesting bee abundance was lower in farms near natural habitat than farms far from natural habitat. These findings suggest that most bee species are affected adversely but to varying degrees by agricultural intensification, and that natural habitats may buffer against the bee population variability in agricultural landscape. We present source/sink dynamics and resource limitations as possible explanations for the observed patterns. KEY WORDS: Ground-nesting bees, sunflower, nesting density, source-sink dynamics, agriculture, biodiversity, pollination Humans have appropriated an estimated 40% of the terrestrial surface for agriculture, greatly modifying the environment and ecosystem processes (Olson et al., 1983; Vitousek et al., 1997; Chapin et al., 2000; DeFries et al., 2004). Consequent loss of habitat and fragmentation are often cited as the greatest threats to biodiversity (Saunders et al., 1991; Wilcove et al., 1996), along with climate change and invasive species (Lodge, 1993; Sala et al., 2000). However, agricultural areas may also act as refuges (Becker et al., 1991; Ricketts et al., 2001; Soto-Pinto et al., 2001) and maintain biodiversity for some taxa (Mander et al., 1999; Noy and Kaplan, 2002; Luck and Daily, 2003). Response to agricultural conversion is thought to be highly species-specific and dependent on life history traits, vagility, and habitat specialization of the organisms (Tucker, 1997; Cane, 2001; Tworek, 2002). Bees represent a group of diverse and understudied organisms whose persistence in the agricultural landscape is poorly understood. In seasonally dry ecosystems, such as the Central Valley in northern California, irrigated agricultural areas may benefit bees by providing more floral resources at certain times of the year than natural habitat has historically offered (Banaszak, 1992; Corbet, 2000). Floral resources are inherently patchy in natural landscapes, and bees may have evolved to find patches of suitable habitats in a matrix of unsuitable areas (Kremen and Ricketts, 2000; Cane, 2001). However, agri- cultural intensification may deprive bees of other necessary resources, such as appropriate nest sites and nest construction materials. For example, ground-nesting bees may suffer from cultivation practices if tilling alters their preferred nesting conditions or destroys nests during larval development. 1 Nicholas School of the Environment and Earth Sciences, P.O. Box 90328, Duke University, Durham, North Carolina 27708. 2 Bryn Mawr College, Department of Biology, 101 N. Merion Ave. Bryn Mawr, Pennsylvania 19010-2899. 3 University of California, Dept. of Environmental Science, Policy and Management, 137 Mulford Hall #3114, Berkeley, California 94720. Accepted 20 September 2005; Revised 20 June 2006 Ó 2006 Kansas Entomological Society 310 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY Bees exhibit a wide range of nesting requirements. Some species have highly specific preferences for substrate (soil, twig, leaf, etc.), moisture, hardness, and amount of vege- tation cover (Potts and Willmer, 1997; Wuellner, 1999) and spend considerable time assessing edaphic conditions at different locations before initiating nests (Brockmann, 1979; Westrich, 1996). Nesting habitats may be limiting for such species partly because sites satisfying all their requirements are relatively rare (Evans, 1966; Michener, 1969; Potts and Willmer, 1998; Wuellner, 1999). Other species have more generalized nesting preferences, and soil and vegetation conditions do not explain the distribution of their nests (Michener et al., 1958; Wcislo, 1984; Mader, 1980). Such diversity among bee nesting requirements implies that those with broader requirements may find farms to be as suitable for nesting as undisturbed areas, while other species suffer from widespread loss of nesting habitats upon agricultural conversion. Bees are of special concern for biodiversity conservation because they are important contributors to agricultural production (McGregor, 1976; Parker et al., 1987; Southwick and Southwick, 1992; Morse and Calderone, 2000). Honeybees, Apis mellifera, form a cornerstone of agricultural pollination. Recent declines in the number of managed honeybee colonies (Watanabe, 1994; Ingram et al., 1996), coupled with a growing recognition that naturally occurring native bees also are important contributors to crop pollination (Kremen et al., 2002a; Klein et al., 2003; Ricketts, 2004; Ricketts et al., 2004), suggest that it is important to understand not only how agricultural landscapes benefit from wild bees but also to what extent they support wild bee populations. Recent studies in an agricultural landscape in California have documented declines in abundance and diversity of native bees with increasing agricultural intensification (Kremen et al., 2002a, b; Kremen et al., 2004; Larsen et al., 2005; Greenleaf, 2005). Some species were relatively insensitive to the amount of natural habitat near the farm, but many were absent on farms with less natural habitat nearby (Kremen, 2004; Greenleaf, 2005). One possible mechanism contributing to this pattern is loss of nesting habitat on farms. Data on nesting patterns within such areas are absent, and the literature on species-specific nesting requirements of these native species is woefully incomplete, although basic nesting guilds of bees are well known. If tilled grounds at farms are unsuitable nesting habitat for ground- nesting bees, nesting sites may be limited in predominantly agricultural landscapes. In such cases, neighboring natural habitat may act as nesting resources for native bees on farms, and farms adjacent to patches of natural habitat could support higher population densities due to surplus bees dispersing from local natural habitat. Some species, however, may persist at low densities on farms if they can accept surrogate nesting habitats such as fallow field borders. Assessing the quality of farms as nesting habitats in a landscape context can reveal how farms and adjacent lands should be managed to maintain a high diversity and density of native bees. We assessed the effect of habitat conversion on bee nesting densities at farm and landscape scales in the same landscape in California studied by Kremen et al. (2002a, 2004). We determined which species of ground-nesting native bees nest on farms, whether proximity to natural habitats affects their nesting densities, and what soil conditions they prefer for nesting. Materials and Methods Site Description We studied bee nesting densities at 10 conventional sunflower farms located in the Capay Valley in the Central Valley of California (35840–539N 122815–509W). Sunflowers VOLUME 79, ISSUE 4 311 (Helianthus annuus L. (Asteraceae)) are native to the area and cultivated forms attract more than 30 species of native bees (Kremen et al., 2002b; Greenleaf, 2005). Farms were classified by their proximity to natural or semi-natural chaparral, mixed oak, woodland, and riparian habitats. ‘‘Near’’ farms included .25% of such natural habitat within a 2 km radius of the farms and ‘‘Far’’ farms had ,2% of natural habitat within a 2 km radius. The designations were based on a previous study showing that bee com- munities responded to spatial scales between 1.2 km and 2.4 km (Kremen et al., 2004). The proportion of natural habitat surrounding each farm (henceforth proximity gradient) was calculated using Arcview GIS, 3.3 by ESRI. Farmers in this region till sunflower fields to 0.3 meter and apply herbicide a few days before the seeds are planted. In a given season, about 1/3 of the field sites are sprayed for the main insect pest, sunflower head-moth, Homoeosoma electellum Hulst (Lepidoptera: Pyralidae) (Carl Hjerpe, Eureka Seeds, pers. comm.). Planting dates vary from farm to farm. Fields are flood irrigated three to four times during the four to five months growing season. Field borders, usually next to dirt roads or irrigation ditches, are left weedy and untilled. Farms were sampled in a random order between July and August of 2003. In our system, number of days between planting and sampling dates and pesticide application did not differ significantly between ‘‘Near’’ and ‘‘Far’’ farm types (number of days between planting and sampling dates: d.f. 5, t-statistics 0.54, P . 0.61; pesticide application: v2 0.62, P . 0.42). All farms had blossoming at the time of the sampling. The surveyed farms were under contract by two different seed companies, but standard method of cultivation was used for all farms (Carl Hjerpe, Eureka Seeds, pers.
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