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The Social Organisation of Halictus Ligatus (Hymenoptera; Halictidae) in Southern Ontario

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The Social Organisation of Halictus Ligatus (Hymenoptera; Halictidae) in Southern Ontario The social organisation of Halictus ligatus (Hymenoptera; Halictidae) in southern Ontario LAURENCEPACKER' Department of Zoology, University of Toronto, Toronto, Ont., Canada M5S 1Al Received November 14, 1985 PACKER,L. 1986. The social organisation of Halictus ligatus (Hymenoptera; Halictidae) in southern Ontario. Can. J. Zool. 64: 2317-2324. The social organisation of Halictus ligatus was studied at Victoria, southern Ontario. At this locality, the one worker brood has a protracted period of emergence; this results in small colony populations throughout the summer activity phase. Workers average 12.7% smaller than their queens, 60% of them have some ovarian development, and 42% of them mate. More males are produced towards the very end of the first brood than earlier in the spring provisioning phase. These late first brood males probably survive to mate with reproductive brood females. In orphaned nests, one worker dominates the others to become a replacement queen. Most replacement queens are mated and orphaned colonies produce reproductives of both sexes. Data from this population are compared with those of other studies of this, and other, halictine species. PACKER,L. 1986. The social organisation of Halictus ligatus (Hymenoptera; Halictidae) in southern Ontario. Can. J. Zool. 64: 23 17-2324. L'organisation sociale d'Halictus ligatus a fait l'objet d'une Ctude a Victoria dans le sud de I'Ontario. A cet endroit, l'unique cohorte d'ouvrikres a une pCriode d'kmergence prolongCe; les populations de cette colonie sont donc petites au cours de la phase d'activitk de 1'CtC. Les ouvrikes sont en moyenne de 12,7% plus petites que les reines, 60% d'entre elles subissent un certain dCveloppement ovarien et 42% d'entre elles s'accouplent. Plus de males sont produits a la toute fin de 1'Cmergence que plus t8t au printemps durant la phase d'approvisionnement de la colonie. Ces males qui apparaissent tardivement au cours de la premike reproduction survivent probablement pour s'accoupler aux femelles reproductrices de la cohorte. Dans les nids oh il n'y a plus de reine, une ouvrikre domine les autres et sert de reine rempla~ante.La plupart des reines de remplacement s'accouplent et les colonies orphelines produisent des individus reproducteurs des deux sexes. Ces donnCes sont comparCes a celles qui ont CtC obtenues au cours d'autres Ctudes sur cette espkce et sur d'autres espkces d'halictinCs. [Traduit par la revue] Introduction Halictus ligatus is an example of a species that has the Halictine bees exhibit a remarkable diversity of social second type of colony cycle. Although haplometrotic nests may organisation, with solitary and eusocial species often being have a period of inactivity during the development of the first found within the same subgenus. The colony cycles of the more brood, pleometrotic nests do not, and there is continuous advanced eusocial species may be divided into two broad activity after worker emergence until the departure of the last groups. In the first type, periods of complete inactivity separate reproductive brood individual. Despite there having been more differentFor personal use only. brood provisioning phases. This colony cycle is found field studies of this species than any other social halictine in the "carinate" members of the Lasioglossurn subgenus (Chandler 1955; Kirkton 1968; Roberts 1973; Raw 1975; Litte Evylaeus (Packer and Knerer 1985). In the second, nests are 1977; Michener and Bennett 1977; Knerer 1980; Eickwort 1985; active from the emergence of the first workers until their demise Packer and Knerer 1986a), much remains to be discovered and the emergence of the last reproductives in late summer or about its basic biology. autumn. This type of colony cycle is found in many species of The study reported here was carried out near Victoria, Lasioglossurn (Dialictus) (Breed 1976), Halictus (Knerer 1980), Ontario. Unless stated otherwise, only data from 1984 will be and in some "non-carinate" Lasioglossurn (Evylaeus) (Knerer presented here. In 1984, 130 nests were excavated; additional 1981). An additional feature, i. e. , pleometrotic founding of observations at the nest site were made in 1983 and 1985. some colonies, cuts across this classification. Pleometrotic Analyses of variation in nest architecture of this population of nests of the first colony type will often remain open between H. ligatus have been published elsewhere (Packer and Knerer spring and summer provisioning phases while the subordinates 19866) and data on pleometrosis are presented in an accom- panying article (Packer 1986). A detailed analysis of female continueCan. J. Zool. Downloaded from www.nrcresearchpress.com by YORK UNIV on 02/13/16 foraging. Pleometrotic nests of the second colony type often remain active from initiation in spring until their demise in reproductive success and the sex ratio is in preparation. autumn. Methods These two basic types of social organisation present different The study site was visited several times each week from the problems to the student of halictine societies. Bees with bursts beginning of the prenesting phase until the emergence of the reproduc- of activity separated by periods of nest closure are easier to tive brood. The positions of nests were marked with bamboo skewers study. This is because brood development is comparatively with a reference number written on tape at the top. Bees were caught, synchronized and nest excavation towards the end of the marked, measured, and released, and nests were excavated as inactive phases will often give a complete picture of brood sex described by Packer and Knerer (1986~).The only methodological ratio and offspring size. Those species which are continuously difference was that Johnson's baby powder, rather than paprika, was active are more difficult to study: a picture of colony develop- blown down the nest entrances to facilitate excavation. ment has to be pieced together from data from many nests Traditionally, students of halictine biology count the number of excavated throughout the colony cycle. developed ovarioles to gain an estimate of relative fecundity of females. Previous studies of H. ligatus (Litte 1977; Michener and Bennett 1977) have distinguished four categories of ovarian develop- 'present address: Department of Biology, University College of ment: class A with five or six developed ovarioles, B with three or four, Cape Breton, P.O. Box 5300, Sydney, N.S., Canada BlP 6L2. C with one or two, and D with no ovarian development. This procedure Printed In Canada I Imprime au Canada 2318 CAN. J. ZOOL. VOL. 64, 1986 was followed here. Another technique used to judge ovarian develop- ment is to measure the width of the most developed oocyte. These methods have been improved upon here by separating developing and pollen resorbing ovariole contents (distinguished by shape and colouration balls differences) and by estimating the fraction of a whole oocyte present in each ovariole. Fractions of 0, 114, 112, 314, and 1 were estimated by eye. Thus, five scores are available for each female's ovarian development: (i) the traditional A to D categorization, (ii) the sum total of fractions of whole oocytes that were developing, (iii) the sum of resorbing portions of oocytes, (iv) the total of ii and iii, (v) the number of fully developed oocytes that were ready to be laid. The shape of the pollen balls constructed by H. ligatus females depends upon the sex and caste that the provision mass will produce -- larvae (Chandler 1955; Michener and Bennett 1977). Thus, male-producing T m - r prepupae pollen balls are small, slightly flattened spheres. Gyne-producing pollen balls are elongate and saddle shaped, and have a shallow depression on the dorsal surface to receive the egg. Worker pollen balls are like those of males, but slightly larger. In practice, it is not possible to distinguish male- and worker-producing provision masses on shape and size alone; however, they are produced at different times of year. Estimates of nest age were made by counting the 1st day that pollen collection was observed in any nest as day 1 and numbering from there. Data from nest excavations were lumped together into 10-day periods for analysis. Categories range from 1 to 9, although an additional male sample was excavated on day 116. r -a *male The number of nests excavated in each time period is shown at the bottom of Fig. 1. Nests were excavated from at least three different portions of the study aggregation within each 10-day time period. Several adjacent nests were excavated from each portion. These methods avoided bias in the choice of nests excavated within individual time periods. Results Description of colony cycle The bees awake from hibernal diapause in late April or during May. However, they do not leave their overwintering burrows Head width until late May or early June. During warm, sunny days in late T (mm) May, the gynes that have overwintered may fly to flowers and For personal use only. feed upon nectar and pollen. They establish new nests at this time or return to their natal burrows and refurbish them. Few bees are active on any given day during this period and, as a result, few observations were made. The first pollen-foraging trips were observed on June 6 in 1983 and June 3 in 1984. Haplometrotic nests were active daily for a period of about 2 weeks and then opened only intermit- tently until the emergence of the first workers. However, some gynes provisioned one or more cells just before the emergence queen survival -b -a queen head width of the first workers. This resulted in a wide range of develop- - mental stages in some nests excavated just before first worker emergence. Three nests excavated on July 19, when most nests Can. J. Zool. Downloaded from www.nrcresearchpress.com by YORK UNIV on 02/13/16 had active workers, contained only the queen, some cells with I width mouldy contents, and two, three, and four recently provisioned I I (mm) pollen balls with eggs.
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