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Bombus Terrestris L Apidologie 39 (2008) 419–427 Available online at: c INRA/DIB-AGIB/ EDP Sciences, 2008 www.apidologie.org DOI: 10.1051/apido:2008020 Original article Foraging distance in Bombus terrestris L. (Hymenoptera: Apidae)* Stephan Wolf, Robin F.A. Moritz Institut für Biologie / Institutsbereich Zoologie, Martin-Luther-Universität Halle-Wittenberg, Germany Received 11 October 2007 – Revised 7 February 2008 – Accepted 25 February 2008 Abstract – A major determinant of bumblebees pollination efficiency is the distance of pollen dispersal, which depends on the foraging distance of workers. We employ a transect setting, controlling for both forage and nest location, to assess the foraging distance of Bombus terrestris workers and the influence of environmental factors on foraging frequency over distance. The mean foraging distance of B. terrestris workers was 267.2 m ± 180.3 m (max. 800 m). Nearly 40% of the workers foraged within 100 m around the nest. B. terrestris workers have thus rather moderate foraging ranges if rewarding forage is available within vicinity of the nests. We found the spatial distribution and the quality of forage plots to be the major determinants for the bees foraging decision-making, explaining over 80% of the foraging frequency. This low foraging range has implications for using B. terrestris colonies as pollinators in agriculture. Bumblebee / foraging / pollination / decision-making 1. INTRODUCTION efficiency (Gauld et al., 1990; Westerkamp, 1991; Wilson and Thomson, 1991; Goulson, Pollen dispersal through animal pollinators 2003). This is partly due to the more robust is essential for plant reproduction. The effi- handling of flowers by bumblebees and their ciency of pollinators depends on various fac- ability of buzz-pollination (e.g. in tomatoes) tors including the number of individuals that (Kevan et al., 1993; Morandin et al., 2001a,b; carry pollen from one flower to another, the Goulson, 2003) but also due to fundamen- ff number of pollen grains actually transferred on tal di erences in foraging strategies. Foraging the flower, and the distance over which pollen Apis workers take advantage of their very so- is transferred. The latter is of particular impor- phisticated communication system, which al- ffi tance with respect to rare and widely dispersed lows e cient recruitment of large numbers of plants (e.g. Kwak et al., 1998) but also for the foragers to highly rewarding sites to exploit pollination service in agriculture. these in short time. The communication in Bumblebees (Bombus spp.) are regarded as bumblebees, however, is much less advanced most efficient pollinators, not only for the pol- and only the type of resource but not its posi- lination of wild flowers but also for pollination tion is communicated (Dornhaus and Chittka, services, used in both outdoor and greenhouse 1999, 2001, 2004). Therefore, Bombus work- horticulture and orchards (e.g. Morandin et al., ers primarily forage based on individual expe- 2001a, b; Dasgan et al., 2004; Velthuis and rience and colonies have more scattered for- van Doorn, 2006). They have been shown age grounds (Westerkamp, 1991; Kearns and to be able to out-compete honeybees (Apis Thomson, 2001; Goulson, 2003). mellifera) in individual workers pollination In commercial use, the foraging range determines the optimal density of bumble- Corresponding author: S. Wolf, bee colonies for facilitating pollination ser- [email protected] vices. The foraging distance of workers has, * Manuscript editor: Jacqueline Pierre therefore, been the research focus of many Article published by EDP Sciences and available at http://www.apidologie.org or http://dx.doi.org/10.1051/apido:2008020 420 S. Wolf, R.F.A. Moritz studies. Walther-Hellwig and Frankl (2000a) outdoor boxes (80×30×22 cm) with about 100–130 estimated a maximum foraging distance for workers per colony provided by the supplier (Katz Bombus terrestris of up to 1750 m in a Biotech AG). The workers were briefly treated with mark and recapture study along a transect. carbon dioxide gas to colour-mark them with paint They found the majority of the workers (75%) pens (green, yellow, red, silver, white, blue, Edding foraging in distances of less then 1500 m XCXX ) according to their colony affiliation. The from the nest, and 43% of the foragers were day after marking, all colonies were placed in the found within a radius of 500 m. Other stud- field in the shelter of a hedge along an agricultur- ally used field track (nest position: 51◦ 23.53’ N, ies, also relying on transects but assigning ◦ B. terrestris workers genetically to common 11 42.40’ E; Fig. 1). colonies, obtained much smaller maximum foraging distances of 758 m (Knight et al., 2.2. Transect 2005) or even less than 625 m (Darvill et al., 2004). Estimates based on a range of assumed Bumblebee surveys were conducted at the end colony densities and genetic colony assign- of the season from October 4 to October 14 2006, ment (Chapman et al., 2003) resulted in a max- in the agricultural “desert” of the Querfurt plateau / imum foraging distances ranging from 870– 30 km in the west of Halle Saale, Germany. A 3900 m for B. terrestris workers. Obviously 3100 m long straight field track, flanked by inter- spersed flower rich plots was used as a transect estimates of foraging ranges have one aspect in (Fig. 1). Satellite image analysis and explorations common: they are highly variable. They range on foot ensured that the adjacent landscape (min from a few hundred meters to several kilo- 3 km from nest position) completely lacked any metres, which is not particularly satisfactory forage because the fields were ploughed and pre- given the high significance of bumblebees as pared for winter. The vegetation along both sides commercial and natural pollinators. It is un- of the track consisted predominantly of Loesels clear to what extent the high variance among rocket (Sisymbrium loeselii), several species of this- previous studies reflects differences among tles (Carduus crispus, Carduus nutans, Cirsium the studied colonies or resulted from different arvense, Cirsium acanthoides, Onopordum acan- experimental approaches. Also, variable cli- thicum), corn poppy (Papaver rhoeas), mugwort mate and weather conditions among studies (Arthemisia vulgare), dandelion (Taraxcacum of- are bound to generate additional variance for ficinalis), yarrow (Achillea millefolium), camomile foraging distance. Finally, plant density is one (Anthemis arvensis), and two species of bur (bur- of the most important factors for foraging dis- dock (Actium lappa)andActium tomentosum). The tances. If forage is sparse, bees must fly much thistles were the only rewarding forage for the bum- farther for rewarding food plants than in a set- blebees during the experiments Number of flowers ting with high plant density (Heinrich, 1976). and numbers of inflorescences within a 100 m in- Here we study foraging flight distances of terval along the track were used as determinants for B. terrestris workers by choosing an experi- patch quality. mental design that provided, for the first time, full control of the external factors by control- 2.3. Monitoring ling both colony positions as well as loca- tion and quality of the forage in a linear tran- Based on precursory observations the monitored sect setting. In addition to the flight distance, transect length was restricted to 1600 m. This tran- the role of these controlled external factors on sect was in addition to the 100 m intervals divided flight distance could be determined. into four monitoring sections. Daily flight entrance observations showed that foragers from all colonies left the nest exclusively in direction of the tran- 2. MATERIALS AND METHODS sect. Four observers monitored flight distances of marked foragers (three times a day, between 1000 2.1. Bombus terrestris and 1700 h) in a randomly assigned section by counting labelled foragers on flowers while slowly We used six commercially reared B. terrestris walking along the transect section (speed: 10 min colonies, three colonies in each of two Styrofoam per 100 m interval). Each section was covered twice Foraging distance in Bombus terrestris L. (Hymenoptera: Apidae) 421 Figure 1. Study site in the Querfurt Plateau in Saxony-Anhalt Germany. Labelled B. terrestris workers from six commercial colonies were observed foraging on patches of thistle flowers along a 1.6 km transect with 100 m intervals (arrows). No food plants other than those along the transect were available within 3 km distance (white: harvested fields, dotted: woody boundary ridges). per observation walk counting all individuals only unconstrained foraging with bright skies and once, which was possible due to low foraging fre- daytime ambient temperatures ranging from quencies (see results). Unmarked workers were nei- 11 ◦Cto23◦C. In total 126 marked foragers ther recorded nor analysed because we had no in- were observed in the 10 days observation pe- formation concerning the nest site of these workers. riod. Unmarked bumblebees (including B. ter- Monitoring was performed over a two-week period. restris, B. pascuorum) were observed in very low numbers only (< 3 per day), and there was 2.4. Data analysis no competition for resources during the exper- iment. Bumblebee workers were almost exclu- The forager counts per distance interval were sively (98.4%, n = 126) observed exploiting pooled over all days and all colonies to obtain an the thistle flowers along the transect and only overall distribution along the entire transect. The these data are included in the analyses. Pollen mean foraging distance was inferred from the mean distance between observation point and colony of loads of returning foragers were consistently all observed foragers. The mean maximum forag- pale yellow corresponding to the colour of the ing distance was calculated as the mean of the max- pollen of the thistle flowers available (Kirk, imum observed flight distance for a worker of each 1994), further indicating that workers exclu- colony.
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