J. HYM. RES. Vol. 12(2), 2003, pp. 220-237 of Andrena Foraging Strategy and Pollen Preferences vaga (Panzer) and Colletes ciinicularius (L.) (Hymenoptera: Apidae) Inge Bischoff, Kerstin Feltgen, and Doris Breckner Museum Alexander Adenauerallee 160, (IB) Zoologisches Forschungsinstitut und Koenig, D-53113 Bonn, Germany, email: [email protected]; (KF) Dottendorferstr. 29, D-53123 Bonn, Germany, email: [email protected]; (DB) Johann-Sebastian-Bach-Str. 4, D-77654 Offenbach, Germany vernal Abstract.—Andrena vaga (Panzer) and Colletes ciinicularius (L.), both ground nesting bees, western Both were studied in the years of 1996-1999 in a lowbush heath near Cologne, Germany. form with thousands species are solitarily but nest gregariously and sometimes large aggregations the Salix. We observed the of nests. They are reported to feed strictly oligolectic on genus daily ciinicularius foraging rhythms of both species and compared their foraging strategies. Colletes start- than A. and finished ed provisioning trips earlier in the morning, made more trips per day vaga, in the dark after 08.00 nest provisioning later in the evening. Colletes ciinicularius burrowed even p.m. Andrena vaga collected pollen and nectar on different days each. One pollen day included 1 the number of and the to 5 pollen trips. There was no clear correlation between pollen trips occurrence of a subsequent nectar day. We found also no correlation between the occurrence of nectar-provisioning trips and weather conditions. Pollen loads of both species were analyzed qualitatively and quantitatively with a cell counter and two different hand-counting systems. Andrena vaga collected nearly twice as much pollen as C. cuuiciilarius during one foraging trip. Cells and pollen loads of C. ciinicularius contained large portions of other pollen types, mostly Rosaceae such as Prunus, Sorbus and Pyrus or Acer, Quercus and Ilex. Thus, C. ciinicularius is not oligolectic as described in the literature. The percentage of pollen types other than Salix increased at the end of the flowering period of Salix, which indicates a resource restriction at the end of the season. The reproductive success of C. ciinicularius measured by nest provisionment exceeded that of A. vaga, because of longer activity per day and digging activity in the evening. Andrena vaga (Panzer 1799) and Colletes number of mostly Nearctic bee species (cf. ciinicularius (L.) are univoltine, vernal sol- Batra 1999, Lind 1968, Michener and Ret- itary bees which are distributed through- tenmeyer 1956, Schonitzer and Klinksik out the entire Paleartic Region. They pre- 1990, Stephen 1966), but precise data on fer to nest in sandy soils and often form the life cycle of European species of An- large aggregations (Friese 1923, Moeschler drena and Colletes are very scarce (Geb- 1938, Vleugel 1947, Westrich 1990). Both hardt and Rohr 1987, Malyshev 1927, Witt species are reported to be specialized on 1992). Salix as a resource pollen for their larvae By analyzing thoroughly the daily activ- and Schmidt (Westrich 1987). Being spe- ity patterns, niche differences between two cialized on the same host-plant, their sea- species may be shown. Levermann et al. sonal activity strongly overlaps (Westrich (2000) investigated the diurnal cycle and 1990). we the di- Therefore, investigated niche differentiation of Dasypoda hirtipes urnal activity pattern and foraging strate- (Fabricius) and Panurgus calcaratus (Scopo- of both Several studies have gy species. li). They demonstrated that apart from dealt with diurnal activity patterns and weather conditions, body mass and pollen- the of weather conditions impact on a collecting apparatus are important factors Volume 12, Number 2, 2003 221 determining the diurnal activity cycle. Re- and overlaying quicksand, soils are most- garding solitary Hymenoptera, female size ly sandy, permeable, and poor in nutri- appears to have a great influence on pro- ents. The climate is humid-oceanic with an visioning- and reproductive success (Al- annual mean temperature of 9.5 °C and an cock 1979). Colletes cunicularius is larger annual mean precipitation of 804 mm. than A. vaga and has more hairs, especially Due to drainage and loss of traditional ag- on the thorax. Therefore, we hypothesized ricultural use, grass and bushland domi- that better thermoregulatory abilities allow nate great areas of the heathland (Inter- C. cunicularius an earlier start to provision- kommunaler Arbeitskreis Wahner Heide ing activity in the morning. 1989). The investigated aggregations of C. Apart from the different sizes of the two cunicularius and A. vaga lie on sandy in- species, the specific pollen-collecting ap- land dunes, paratus suggests differences in their pol- both METHODS len collecting efficiency. Although — species have trochanter-femur-baskets, Field work. To study their diurnal ac- floccus and thoracic pollen basket of A. tivity, individual female bees were vaga is more strongly developed than in marked with opalith-plates. A total of 238 C. ciniicularius. In a number of bee genera, females of C. cunicularius in 1996 and 174 the size and the type of pollen-collecting females of A. vaga in 1997 were marked, apparatus results in different amounts of Corresponding nests were identified by a pollen transported (Braue 1916, Friese colored nail of the same number as on the 1923, Michener et al. 1978, Pasteels and opalith-plate of the female. To record the Pasteels 1979, Westerkamp 1987, 1996, exact departure and return times of the fe- Westrich 1990). Thus, we examined the males, several nests were covered with number of collected pollen grains in both preserving jars (cf. Witt 1992). species. Climatic parameters were measured Although both species are regarded as with data loggers (Orion Tiny Logger strictly oligolectic on Salix (Vleugel 1947, Manager OTLM Tinytalk©). Soil temper- Westrich and Schmidt 1987, Westrich atures of all years were recorded 20 cm 1990), anecdotal observations reviewed by below the surface, and air temperature Mader (1999) suggest that C. cunicularius and atmospheric humidity in a portable visits also flowers other than Salix. A clear weather station were measured at a height evaluation about oligolecty in a bee spe- of 2 m in 1997-1999. Data from the Co- des can only be made by a quantitative logne/Bonn airport weather station were analysis of pollen loads or cell provisions, also used. The data on the activity and the The aims of this study were to analyze nectar trip frequency of the bees were test- niche differentiation of two synchronous ed for possible correlation against various and syntopic bee species on the basis of climatic parameters (daily hours of sun- activity patterns and the use of host shine, mean daily temperature, mean dai- plants, to asses the degree of oligolecty, ly atmospheric humidity, humidity of soil and to compare their pollen collecting ef- and minimal daily soil temperature), ficiency. Since the activity of most of the bees is The study area is part of the so-called extremely influenced by weather condi- "Wahner Heide", a large 5000 ha heath- tions (Larsson 1991, Lind 1968) we created land, east of the river Rhine near Cologne, a measure for the bees activity indepen- Since 1961 the heathland has been a mili- dent from weather conditions (cf. Midler tary training area for NATO and was des- 1994), which we referred to a so-called ignated a Nature Reserve in 1968. Due to "bee day" (= BD). Such a measure makes the presence of Quaternary gravel, sand it also easier to compare different years. 222 Journal of Hymenoptera Research = at the entrance = = bee in the nest provisioning trip waiting yyfa digging C. cunicularius n10 n423 n465 l l l I II first departure 8:35 last return 17:31 (n35) (n.20) A.vaga ("nectar-day") n19 n39 tirst departure 11:13 last return 14:29 (n25) (n39) A.vaga ("pollen-day") n19 n90 n82 last return 16:40 first departure 10:18 (n20) (n29) H 1 1 h H 1 1 1 1 h H 1 1 1 1 1 1 1 h H 1 1 1 h H 1 1 1 1 Time (MET) Fig. 1. "Ideal ethogram". Time schedule of provisioning behavior of A. vaga and C. cunicularius (dotted line = end of burrowing unknown). Further information's see text. = One bee day corresponds to a day with 0.75 BD flying activity more than Vi optimal weather conditions, which fe- of one BD = males could use completely for provision- 0.25 BD flying activity less than Vi of ing activities. A bee day for C. cunicularius one BD = had between 8-9 hours. The measure for BD no flying conditions the these 8-9 hours was the observed flying whole day activity at such an optimal day. Other — Nectar- and pollen days of A. vaga. To days with less conditions optimal flying assess a between A. nectar- were defined as follows: pattern vaga's and pollen-provisioning trips, we ana- = 1.0 BD flying activity of 8-9 hours lyzed the number of pollen days as well 0.75 = BD flying activity more than Vi as the number of pollen trips between nec- of one tar BD trips. One nectar trip corresponds to 0.25 = BD flying activity less than Vi of one nectar day and the patterns of pollen one BD trips between nectar trips occurs over a = BD no flying conditions the period days since pollen and nectar are whole day collected on different days. We obtained complete observations of pollen trips be- The classification of the bee day of A. tween nectar of 7 different females. vaga was more difficult because of the trips Pollen —For we variance in analysis. pollen analysis greater activity time, in spite excavated 10 cells of C. cunicularius in 1996 of good weather conditions: and 6 cells in 1998. The cells could not be 1.0 BD = of 8-11 flying activity hours attributed to a specific nest or time of com- Volume 12, Number 2, 2003 223 pletion.