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Of a Solitary Stem-Nesting Bee, Osmia Rufa (L.)

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Of a Solitary Stem-Nesting Bee, Osmia Rufa (L.) Original article The function of the vestibulum in nests of a solitary stem-nesting bee, Osmia rufa (L.) Karsten Seidelmann Institut für Zoologie, Martin-Luther-Universität Halle-Wittenberg, Domplatz 4, 06099 Halle (Saale), Germany (Received 2 April 1998; accepted 21 November 1998) Abstract - Nests of the stem- or hole-nesting megachilid bee, Osmia rufa, were analysed to help clar- ify the function of the outermost empty chamber of the nest, the vestibulum. Only nests in an exposed nesting environment had a long vestibulum, whereas nests protected from sun and temperature fluc- tuations (within a honey bee hive body) had short vestibuli or none at all. The rate of parasitism of the first cell from nests with a vestibulum did not differ from nests without a vestibulum in both nesting environments. The short vestibulum built by O. rufa in protected nests probably represents the remaining space of a nest hole that is too short for an additional cell. Nests in exposed environments suffered much higher mortality in all brood cells owing to parasites attacking open cells during nest construction. Additionally, mortality caused by the cleptoparasitic drosophilid, Cacoxenus indaga- tor, one of the main parasites of the Red Mason bee, increased greatly from the rear to the entrance of nests. When nesting bees detect the cleptoparasitic fly, they do not provision the outermost space of nesting holes, thus constructing a vestibulum, to avoid misinvestment due to the high risk of par- asitism. In nests exposed to normal weather factors, vestibular cells probably shelter the brood also from high fluctuating temperatures. &copy; Inra/DIB/AGIB/Elsevier, Paris Osmia rufa / vestibulum / nest architecture / avoidance of parasitism 1. INTRODUCTION the nest structures are a defence against par- asites, predators and nest destroyers. By models et al. Nests of many Aculeata have empty cells probability Tepedino [21] were in variable numbers and positions [9, 10, able to demonstrate that empty, yet sealed 19]. To date, the function of empty cells has cells can reduce the success of parasites of remained unclear. Usually it is thought that mud-nesting Hymenoptera. These species E-mail: [email protected] attach several cells on a surface or construct &times; 160 mm in length) in each block. To test for subterranean nests with cells branching off possible influences of the nesting environment on the of a these from a main tunnel in the soil. production vestibulum, nesting blocks were placed either in an intact (’protected Empty cells are also found in twig- or environment’) or dismantled (’exposed environ- hole-nesting, xylophilous Aculeata, espe- ment’) honey bee hive body (box with no combs cially megachilid bees that construct their or bees). The protected nest blocks could be reached via the entrance of brood cells in a linear order one behind the only regular flight the hive. Thus, nests in the intact hive were pro- other. In such linear nests an empty cell is tected from direct weather influences. In the usually found between the final provisioned exposed environment, the entire front side of the cell and the nest plug. These so called hive box was removed. Nesting blocks in such vestibular cells are believed to be a protec- dismantled hives were sheltered from rain but tion against parasitism of the nest [7, 11, exposed to normal temperature fluctuations. 12, 22]. Krombein [9] assumed that the par- A total of 1 712 complete nests of Osmia rufa asites that inserted their ovipositor through from within 22 nest blocks were evaluated. the nest plug no longer exist. Therefore the A nest was considered complete when the nest- tube contained several brood had a vestibulum has become a behavioural relict ing cells, reg- ular nest and, in cases where the nest had a without current value. plug, adaptive vestibulum, if the outermost brood cell was com- According to Rust [15], a vestibulum is pletely sealed off by a cell partition. These restric- typical for all Osmia species. However, dur- tions were necessary because at the end of the season old females also seal incomplete the analysis of Osmia rufa nests around flight ing nests [18]. In this case, it is difficult to interpret vestibuli were observed to Halle, Germany, the empty space between the nest plug and the be facultative structures. This was not to be first provisioned cell. under their assumed function in expected In all nests, the thickness of the nest defense. the lack of complete parasite Furthermore, plug (except protected environment in 1992) and a vestibulum did not result in a marked vestibulum length were measured. The contents increase in parasitism of outermost brood of the outermost cell (= first brood cell behind the cells, although all major parasites of O. rufa vestibulum) was recorded for 1 413 nests, and and contents of all were present. The vestibulum could there- for 1 073 of these, the length brood cells were also recorded. From the fore represent remaining space which is too length of the provisioned cells of each nest, the mean small to construct an additional brood cell. length for male and female cells were calculated. To test these and thus to hypotheses, clarify In O. rufa nests, sons occupy smaller cells than the function of a vestibulum, its frequency daughters, and they are located towards the and length were recorded in nests of O. rufa entrance [14, 16, 17]. Therefore, the space which from different nesting environments, and would have been needed for an additional male cell in that nest was to the the outermost cells were exam- particular compared provisioned of the if it existed. To ined for parasites. To identify possible influ- length vestibulum, compute mean vestibulum length for nest blocks, vestibu- ences of the construction of a vestibulum lum length was assumed as zero for nests lacking on parasitism of inner brood cells, the total this empty space. nest contents of random were exam- samples To ascertain the distribution of ined. parasites within nests, every nest was equally divided into 300 sections by length and the contents of each section registered. A distribution curve of para- 2. MATERIALS AND METHODS sites could be constructed by totalling the num- ber of parasitized cells in every section over all nests [17]. The study was carried out on nests of Osmia rufa from the Botanical Garden of the Univer- Differences in vestibulum length and entrance sity of Halle (Germany). In 1992 and 1993, plug thickness between exposed and protected females of this species were offered wooden nest nesting environments were statistically analysed blocks with 100 nesting holes (8 mm in diameter by a two-way model I ANOVA with year and nest environment as fixed effects followed by a factor NE (F = 413.62, P < 0.001). In 1993, Scheffé test for a posteriori comparison of means. the vestibuli were slightly shorter than in The existence of a vestibulum and the parasitism 1992. of the outermost cell were added as fixed effects in a model I ANOVA to analyse influences of Only 15 % (1992) and 5 % (1993) of nest parasitism rate on outermost cell parasitism. vestibuli in protected nests were large Parasitism rate (PR) was computed as number enough to accommodate an average-sized of cells parasitized per provisioned cells. A log- male cell. In exposed nests, 63 % (1992) linear analysis according to Warnstorff and Dör- and 50 % (1993) of vestibuli would have fel [23] of a four-dimensional contingency table been to accommodate another was used to detect influences on the occurrence large enough male cell of a vestibulum and a dependence of the outer- (figure 1). most cell’s contents on the existence of a vestibu- lum. In the contingency table, year (Y), nesting environment ( NE), and existence of a vestibulum 3.2. Thickness of the nest plug (V) were considered design variables, and the content of outermost brood cell (BC) was the Osmia nests had nest from response variable. All calculations were com- rufa plugs puted with the Statistica&reg; package (StatSoft Inc.). 1.3 to 14.5 mm in thickness (table II). The thickness of the plug depended mainly on the occurrence of a vestibulum (F = 149.71, 3. RESULTS P < 0.001), but was not correlated with the vestibulum length (all R2 < 0.01). In nests 3.1. Frequency and length without a vestibulum, the plug was signifi- of the vestibulum cantly thicker than in nests with one (table II, all P < 0.001). An influence of the The occurrence of a vestibulum was nest environment on plug thickness was not related to the type of nesting environment: found (F = 0.64, n.s.). (NE x V 2I = 274.04, P < 0.001). A vestibu- lum was found in only half of the protected nests compared to ca. 90 % of exposed nests 3.3. Contents (table I). Also the mean vestibulum length of the outermost brood cell differed between the types of nesting envi- ronments. In both years, vestibuli from Parasites found in the outermost brood exposed nests were twice as long as vestibuli cells were Cacoxenus indagator (Diptera: from protected nests (table I; Scheffé-Test, Drosophilidae) in 10.4 % of nests, Anthrax all P < 0.001). Most of the variance in anthrax (Diptera: Bombyliidae) in 4.6 % of vestibulum length was associated with the nests, and Chaetodactylus osmiae (Acari: in 0.3 % of nests. These Chaetodactylidae) protected nests) was the mortality in nests species attack open cells during provision- without a vestibulum higher than in nests ing. In none of the 1 712 investigated nests with one ([V x BC]/Y1 NE2: 2I = 6.387, nest had a parasite broken through an intact P = 0.041).
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