Floral visitation patterns of bees during spring in Constantine, Algeria
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Floral visitation patterns of bees during spring in Constantine, Algeria K. Louadi*, K. Benachour & S. Berchi Laboratoire de Biosystématique et Ecologie des Arthropodes, Département des Sciences de la Nature et de la Vie, Université Mentouri, Constantine 25.000, Algeria Although there have been studies on Algerian from 08:00 to 12:00 (GMT +1) from 26 February to Hymenoptera by Saunders (1908), Alfken (1914), 20 June 1996 and 1997 in fallow lands, grasslands Morice (1916), Schulthess (1924), Roth (1923, 1924, and along road verges. Small bees were captured 1930) and Benoist (1961), the bee fauna of some using a plastic aspirator 5 × 3 cm diameter contain- regions of Algeria, North Africa, is little known. ing filter paper moistened with 4–5 drops of ethyl Relationships of plants and bees have been exam- acetate (Guiglia 1972; Louadi & Doumandji 1998a; ined extensively in habitats having very different Louadi 1999). Large bees were netted. Samples of flora to those found in Algeria (Batra 1977; plants visited by insects were collected and depos- Michener 1979, 2000; Jacob-Remacle 1989; ited in the herbarium. Identification of plants was Jennerstein et al. 1991; Dafni 1984, 1991, 1992; done according to keys from Quezel & Santa (1962); Petanidou 1993; Petanidou & Smets 1995). Louadi Polunin & Huxley (1965) and Beniston (1984). The & Doumandji (1998a) examined only two species, software Microbanque fauna-flora (Rasmont et al. Apis mellifera L. and Bombus ruderatus siculus Dalla 1995) was used to manage the converted data. Torre in the Constantine area. The aim of this study The food niche of bees was quantified by diversity is to determine the floral choices of others species indices (Louadi & Doumandji 1998a; Jacob- of wild bees in natural habitats during spring. Remacle 1989). The floral visits were quantified by The Wilaya (Department) of Constantine Simpson’s concentration indices (Is) (1949). For (2287 km2) is situated in the eastern part of Algeria each taxon, the visits observed for each family or (36°36’N 06°62’E, 660 m a.s.l.). Vegetation is plant species corresponded to the percentage of composed of forest and scrub (9.2 %), steppe and individuals of the taxon that visited this family or brushwood (24.2 %) (Mebarki 1984), natural and these species of plants. Simpson’s index was cultivated grasslands (2.5 %) fields of cereals and calculated with regard to all floral visits made by other crops (63.8 %). Orchards occupy only 0.3 %. each bee species. Simpson’s index varies between Beekeeping in the region is little developed. We 0 and 1, the closer to unity, the greater the only noted 15 hives in an orchard at Hamma- specialization. It provides an evaluation of host Bouziane (36E41’N 06E59’E, 460 m a.s.l.). plant selection. q The Constantine area has a Mediterranean flora with a high number of flowering species ∑ n (n i i − 1) i= 1 (Petanidou & Vokou 1990; Dafni & O’Toole 1994). Is = , N( N − 1) There is a dominance of therophytes over perennial plants. Many annual plants live three to four where ni = number of visits observed on the ith months but some only live a few weeks (Beniston plant species, and N = total number of visits 1984). Their flowering cycle depends on the timing observed on the whole of the q plant species. and amount of rain (Mebarki 1984). Asteraceae is The width of the food niche is expressed by the dominant family including Silybum marianum Shannon’s index (H’) (Daget 1976). (H’) will increase L. (Greath), Scolymus hispanicus L., Senecio nebro- proportionately with higher numbers of plants densis L., Calendula arvensis L., C. suffruticosa Bat. B. visited. It also increases if bees are distributed and T. and Chrysanthemum palladium Poiret. evenly on the different plants. Brassicaceae, namely Sinapis arvensis L. and Bras- H' = − ∑ p i ln p i , sica fruticulosa Cyr, Lamiaceae, Rosmarinus sp. and i= 1 Lamium sp. and Malvaceae (Malva sp. and Lavatera where pi = the proportion of visits on the ith plant, sp) are also common. pi = ni /N, and ln = Neperian logarithm Bees visiting flowers were collected once per week Bees from four families were collected: Andreni- *To whom correspondence should be addressed. dae (six taxa), Megachilidae (11 taxa) Apidae and E-mail: l_louadi @ hotmail.com Halictidae (16 taxa each). We observed 15 families African Entomology 15(1): 209–213 (2007)
210 African Entomology Vol. 15, No. 1, 2007 Table 1. Total number, floral visit rate and number of native plant-visitor species (all Apoidea, 1996–1997). Plant families Plant species visited Total number of % of floral visits Number of visitor visits species Lamiaceae Rosmarinus officinalis 2087 55.91 22 Malvaceae Malva sylvestris 413 11.06 21 Crepis vesicaria 274 7.34 9 Carthamus sp. 237 6.35 16 Calendula suffruticosa 193 5.17 10 Asteraceae Senecio nebrodensis 136 3.64 13 Chrysanthemum paludosum 32 0.86 9 Silybum marianum 30 0.80 9 Scolymus hispanicus 19 0.51 6 Anthemis chia 12 0.32 2 Papaveraceae Papaver rhoeas 182 4.88 16 Boraginaceae Borago officinalis 44 1.18 5 Brassicaceae Sinapis arvensis 30 0.80 8 Brassica fruticulosa 8 0.21 5 Convolvulaceae Convolvulus tricolor 24 0.64 3 Primulaceae Anagallis monelli 4 0.11 1 Fumariaceae Fumaria officinalis 3 0.08 2 Fumaria capreolata 2 0.05 1 Resedaceae Reseda alba 2 0.05 2 Fabaceae Hedysarum corronarium 1 0.03 2 Total 3733 100 162 of the most abundant native plants in our region. visited by all of Apoidea families. The full set of Only 11 plant families were visited by all of the observations are recorded in Tables 1 and 2. bees (Table 1); the Lamiaceae (56 %) and the The long-tongued bees visited plants with deep Asteraceae (25 %) accounted for 81 % of the visits. corollas, such as Rosmarinus officinalis L. The The Fumariaceae, Resedaceae and Fabaceae were predominant plant family for the long-tongued of little interest to Algerian bees. Bees never visited bees such as Apidae and Megachilidae was the the Euphorbiaceae. Lamiaceae. The short-tongued bees, Andrenidae More than half of all bee species observed visited and Halictidae, concentrated their visits on the Asteraceae. Bees focussed on relatively few plants, Asteraceae. These plants were characterized by some of which were relatively common. Of the flowers with particularly accessible nectar. three species of the Lamiaceae family only one The quantification of the degree of feeding species, Rosmarinus officinalis L, attracted the most specialization of most observed wild bees was bee species and the most visits. Many bee species carried out using Simpson’s floral visit indices (Isf) but fewer numbers of individuals also visited for the plant families and (Isp) for the visited plant Malva sylvestris L. (Malvaceae). Both these plant species. Values calculated for these indices are species are common throughout the Mediterranean listed in Table 3 for 15 bee species. Eucera obliterata region. Perez, 1896, was the species with the highest indi- Tables 1 and 2 show that of all plant families, the ces (Isf). The other species showed some very ele- Lamiaceae received the highest number of visits, vated indices relative to the remainder of the particularly from the Apidae and Megachilidae. Apoidea. They concentrated their visits on only The Andrenidae and the Halictidae especially one or two families of plants: the Lamiaceae for visited the Asteraceae. The Papaveraceae were E. obliterata and the Asteraceae for Andrena albo-
Short communications 211 Table 2. Analysis of visitation activity according to bee family. Apoidea families Andrenidae Megachilidae Apidae Halictidae Apoidea Number of visits 143 659 2178 753 3733 % of visits 3.83 17.65 58.34 20.17 100 Number of taxa 6 11 16 16 49 Number of plant families visited 4 7 9 6 14 Number of plant species visited 10 12 17 13 18 punctata Rossi, 1792, Andrena flavipes Panzer, 1798, slightly lower than that one of Evylaeus mediterra- and Anthophora plumipes Pallas, 1772. Species that neum. It would seem to indicate that the Halictidae presented a mid-level index were Eucera notata were more eclectic in their choices of plants than Lepeletier, 1841, Eucera oraniensis Lepeletier, 1841, the Andrenidae. The Apoidea Andrena florentina and Evylaeus malachurum Kirby, 1802, showing a had the highest Isf/Isp ratio. This species visited preference for several plant families, Lamiaceae, several plants in its favourite family, Asteraceae. Malvaceae and Papaveraceae for the first two; Osmia signata Erichson, 1835, possessed the lowest Lamiaceae and Asteraceae for the third. Andrena ratio; it was limited to some species therefore florentina Magretti, 1883, Bombus terrestris L., 1758, within the preferentially exploited families. Evylaeus subhirtum Lepeletier, 1841, and Lasio- The feeding niche is expressed by Shannon’s glossum clavipes Dour, 1872, had the same index. index (H’). By analogy with the previous index, we These bees visited several plant families as did calculated H’f for plant families and H’p for plant bees having the lowest indices between 0.3 and species to enable us to evaluate the feeding niche 0.4. and to distinguish whether the Apoidea species Concerning the indices of plant species visits distributed themselves evenly on plant groups (Isp), it appeared that species having the highest and if the number of plants visited increased. indices were those that had a high Isf index. Eucera obliterata presented the widest feeding Species having the lowest index were Evylaeus niche and the lowest H’f. Andrena albopunctata, mediterraneum Blüthgen, 1926, and Andrena Andrena flavipes, Anthophora plumipes and Andrena florentina. The index of Andrena florentina was florentina followed it. The twelve other Apoidea Table 3. Floral visit indices (Is) and the feeding niche (H’) of 15 bee species. (Isp = Index of plant species visits; Isf = Index of plant family visits; H’ = Shannon’s index; np = number of plant species visited; nf = number of plant families visited) Apoidea species np nf H’p H’f I sp I sf Isf/Isp Andrenidae Andrena albopunctata Rossi, 1792 3 2 0.773 0.378 0.747 0.861 1.153 Andrena florentina Magretti, 1883 6 3 2.368 0.97 0.145 0.467 3.221 Andrena flavipes Panzer, 1798 6 3 0.968 0.48 0.708 0.846 1.195 Megachilidae Osmia signata Erichson, 1835 11 8 2.006 1.917 0.318 0.316 0.994 Halictidae Lasioglossum clavipes Dour, 1872 10 6 2.019 1.338 0.374 0.46 1.23 Evylaeus malachurum Kirby, 1802 10 7 2.01 1.17 0.417 0.61 1.463 Evylaeus mediterraneum Blüthgen, 1926 9 6 2.675 1.702 0.179 0.385 2.151 Evylaeus subhirtum Lepeletier, 1841 6 4 2.012 1.378 0.3 0.467 1.557 Apidae Anthophora plumipes Pallas, 1772 7 4 1.364 0.824 0.58 0.734 1.266 Eucera punctatissima Panzer, 1895 4 3 2.063 1.673 0.234 0.324 1.385 Eucera oraniensis Lepeletier, 1841 10 7 1.24 1.15 0.61 0.625 1.025 Eucera obliterata Perez, 1896 2 2 0.709 0.098 0.974 0.974 1 Eucera notata Lepeletier, 1841 15 8 1.216 1.115 0.649 0.65 1.002 Tetralonia alternans Brulé, 1832 3 3 1.806 1.473 0.288 0.353 1.226 Bombus terrestris L., 1758 8 4 2.016 1.241 0.338 0.467 1.382
212 African Entomology Vol. 15, No. 1, 2007 had wide feeding niches. Evylaeus malachurum, (Lamiaceae) and had little interest in the native Eucera oraniensis and Eucera notata, had the plants, possibly due to the ability of honeybees narrowest feeding niche; Osmia signata had the to communicate information about good food largest. resources. Nevertheless, Lamiaceae have by far Concerning plant species, the feeding niche the most rewarding nectar with regard to volume of Bombus terrestris was large. Andrena flavipes and sugar content (Dafni 1991; Petanidou & Smets presented an index closer than 1. Among the 1995). According to Dafni (1984, 1992), morphology fifteen other Apoidea, Evylaeus mediterraneum and of flowers, size and length, colour and odour of the Andrena florentina showed the greatest feeding aromatic plant can cause deception in foraging. diversity. Eucera punctatissima Panzer, 1895, Lasio- Jacob-Remacle (1989), studying Anthophora glossum clavipes, Evylaeus subhirtum and Osmia plumipes, found a family visitation index of 0.158. signata also had large niches. The other Apoidea In our case, the visitation index of this species was had a narrower feeding niche (H’ 4 mm) may produce more nectar than shorter species on this family while the Lamiaceae attracted flowers (Petanidou & Smets 1995) and larger 13 species. In the present work 23 bee species were flowers with more display may offer higher calo- recorded visiting Lamiaceae. The results are not rific reward per flower (Dafni 1991). The nectar strictly comparable as both the plant and bee sugar concentration preferred by large bees species differed from those of our study with the ranges from 20 to 50 % (Petanidou 1993). exception of Anthophora plumipes (Apidae). Results The wild bees did not visit the Euphorbia could also depend on the time of flowering in the helioscopia perhaps on account of its toxicity year (Dafni 1991). (Maurizio 1968; Louadi & Doumandji 1998a). The It also appeared that species where H’f indices Halictidae and the Andrenidae especially observed were higher or equal to 2 visited plants of several had gathered on the Asteraceae. This family’s different families. Those of which H’p
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