Factors Affecting Offspring Body Size in the Solitary Bee Osmia Bicornis (Hymenoptera, Megachilidae) Sabine Radmacher, Erhard Strohm
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Factors affecting offspring body size in the solitary bee Osmia bicornis (Hymenoptera, Megachilidae) Sabine Radmacher, Erhard Strohm To cite this version: Sabine Radmacher, Erhard Strohm. Factors affecting offspring body size in the solitary bee Osmia bicornis (Hymenoptera, Megachilidae). Apidologie, Springer Verlag, 2010, 41 (2), 10.1051/apido/2009064. hal-00892048 HAL Id: hal-00892048 https://hal.archives-ouvertes.fr/hal-00892048 Submitted on 1 Jan 2010 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie 41 (2010) 169–177 Available online at: c INRA/DIB-AGIB/EDP Sciences, 2009 www.apidologie.org DOI: 10.1051/apido/2009064 Original article Factors affecting offspring body size in the solitary bee Osmia bicornis (Hymenoptera, Megachilidae)* Sabine Radmacher,ErhardStrohm Institute of Zoology, University of Regensburg, 93040 Regensburg, Germany Received 12 February 2009 – Revised 11 August 2009 – Accepted 15 August 2009 Abstract – Body size is related to fitness in many insects. In solitary bees offspring body size is largely determined by maternal provisions and microclimate. We studied the effect of quantity and quality of pro- visions and rearing temperatures (20, 25 and 30 ◦C) on body size in the Red Mason bee, Osmia bicornis. Cocoon weight increased with provision weight and decreased with increasing temperature. High temper- ature (30 ◦C) led to high mortality of larvae and smaller body size, partly due to incomplete consumption of the provisions. Diversity of pollen in the provisions was low, and most provisions primarily consisted of mainly oak and maple pollen. Pollen diversity did not affect cocoon weight and males provided with pure oak or pure maple pollen showed similar adult size. Our results suggest that the quantity of provisions and temperature, but not pollen quality influence the development of O. bicornis progeny and might affect population dynamics of this abundant European pollinator. Osmia bicornis / body size / pollen diversity / temperature 1. INTRODUCTION Maeta, 1989;Kim,1997), nest usurpation (Tepedino and Torchio, 1994;Kim,1997), and Solitary bees are important pollinators of overwintering survival (Tepedino and Torchio, both wild and crop plants (Batra, 1995; Freitas 1982; Bosch and Kemp, 2004). Thus, fac- and Pereira, 2004). Due to the worldwide tors affecting offspring body size in soli- pollinator decline (Kearns et al., 1998), the tary bees could play an important role in re- economic importance of pollination services productive success and population dynamics. (Klein et al., 2007), and increasing knowledge In solitary bees body size might be influ- about the role of the so-called “wild bees” in enced by the amount of larval provisions, the (crop) pollination (e.g. Winfree et al., 2007), quality of provisions, and temperature during the interest in conservation and management development. Several studies have shown a of wild bees has considerably increased. Thus, strong correlation between the amount of pro- detailed knowledge is needed about the re- visions consumed by larvae and adult body quirements for successful reproduction and size (e.g. Klostermeyer et al., 1973; Bosch and population maintenance for at least in some Vicens, 2002), whereas heritability for body model species. size seems to be low (Tepedino et al., 1984). Body size is related to fitness in many in- Larval body size and performance might sects (Honek,ˇ 1993). In solitary bees, sev- be influenced by the quality of pollen-nectar eral studies have shown advantages for larger provisions as well. Pollen of different plant individuals in foraging efficiency (Torchio species differs in nutritional value (Roulston and Tepedino, 1980), fecundity (Sugiura and and Cane, 2000). Several studies have shown that different pollen types differentially influ- Corresponding author: S. Radmacher, ence larval growth and development in honey [email protected] bees (e.g. Standifer, 1967), bumble bees (e.g. * Manuscript editor: Jean-Noël Tasei Tasei and Aupinel, 2008) and solitary bees Article published by EDP Sciences 170 S. Radmacher, E. Strohm (e.g. Levin and Haydak, 1957;Prazetal., pupate and undergo metamorphosis in late summer. 2008). Mixing of different pollen species could They hibernate as adults inside their cocoons and be advantageous, because it results in a better emerge the next spring when temperature rises. balance of nutritive substances and dilution of Brood cells were obtained from a population of toxins (Bernays and Minkenberg, 1997; Tasei O. bicornis nesting in trap nests in two observa- and Aupinel, 2008; Unsicker et al., 2008). tion huts near the Biocenter of the University of In addition to larval food, characteristics of Würzburg, Germany. The vegetation surrounding the nesting site can affect offspring body size. the huts consisted of set-aside orchards (mainly ap- Temperature is probably the most important ple trees and grassland), landscaped grounds (sev- abiotic ecological factor and the “temperature- eral tree species and lawn), and farmland (rape and cereal crops). Female bees readily accepted size rule” for ectotherms states that lower tem- trap nests that were made of dense styrofoam peratures during development result in larger covered with transparent polycarbonate lids that adult body size (e.g. Atkinson, 1994). Since allowed continuous observation of nesting activ- nest sites of solitary bees probably vary in tem- ity and brood development (for more details see perature regimes (due to variable solar irra- Strohm et al., 2002). Between 8 May and 14 June ff diation and insulation), o spring body size is 2006 we selected 26 completed nests with a total probably influenced by these differences. of 205 brood cells that contained eggs or newly In this study we investigated possible ef- hatched larvae which had not yet started feeding on fects of (1) quantity and (2) quality of larval the provisions. We brought them into the laboratory food as well as (3) temperature during devel- to determine the mass (quantity) and composition opment on body size of the Red Mason bee Os- (quality) of pollen and to subject them to different mia bicornis (Linnaeus 1758, Hymenoptera, temperature treatments (see Sect. 2.4). Megachilidae, formerly O. rufa L.). Based on empirical knowledge about other bees and her- bivores, we predicted that the size of offspring 2.2. Pollen quantity bees will be positively correlated with both the amount and the diversity of pollen provisions Polycarbonate covers were carefully removed (nectar was not included in the analysis due to from the trap nests and brood cells were separated. The pollen loaf with the attached egg was removed its low amount in O. bicornis provisions). In with a custom made scoop of known weight. After brood cells provisioned with only one pollen weighing to the nearest 0.1 mg (Mettler AE 160) species, larval performance might differ be- / ff the provision with egg larva was carefully put back tween pollen types due to di erences in nu- into the brood cell. Weighing the pollen loaf with- tritional value. Furthermore, we tested the pre- out the egg was not possible because detaching it diction that lower rearing temperatures result from the provision would have harmed the egg or in larger body size of the bees. young larva. However, egg weight is largely the same for all brood cells; thus, the inclusion of egg 2. MATERIALS AND METHODS weight should not have biased the results for pro- vision weight. Brood cells were re-sealed with pa- 2.1. Bees per on the side and transparent foil on the top (fixed with adhesive tape) and subjected to different rear- O. bicornis is one of the most abundant solitary ing temperature treatments (see Sect. 2.4). At the bees in Central Europe (Westrich, 1989). The fe- end of development, pollen remnants were weighed males of this univoltine, polylectic species fly from and the difference between the initial and final pro- early April to mid-June and they accept a large vision weight was considered to be the mass of range of pre-existing cavities where they build their pollen actually consumed. (mostly) linear nests. Each nest consists of several brood cells that are provisioned with pollen and some nectar (< 4% of total provisions) and sep- 2.3. Pollen quality arated by loam partitions (Maddocks and Paulus, 1987; Strohm et al., 2002). Females attach a single After removal of nest lids (see Sect. 2.2), sam- egg onto the provisions of each brood cell. The lar- ples of pollen were taken using a moistened nee- vae hatch, consume the provisions, spin a cocoon, dle that was used to puncture the provisions at Effects on Osmia bicornis body size 171 five spots spread over the pollen loaf. After each different rearing temperatures but minimised fur- puncture, pollen grains adhering to the needle were ther mortality for the 30 ◦C group. On 5 Septem- wiped onto a slide where they were immersed in ber 2006, all brood cells were transferred to 10 ◦C Polyvinyllactophenol (PVL) and covered with a to initiate diapause. The cocoons were weighed to coverslip. To identify the bee-collected pollen in the the nearest 0.1 mg at the same individual age (119– provisions, a reference collection of pollen was pre- 122 days after oviposition) as a measure of adult pared from all plants that flowered during the nest- body size. Then mortality and sex were determined ing activity of the bees within a radius of 500 m by inspection of the bee after opening the apical end around the huts (107 plant species).