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Nordic Society Oikos Nordic Society Oikos The Socioecology of Body Size Variation in the Primitively Eusocial Sweat Bee, Halictus ligatus ( Hymenoptera: Halictidae) Author(s): Miriam H. Richards and Laurence Packer Source: Oikos, Vol. 77, No. 1 (Oct., 1996), pp. 68-76 Published by: Wiley on behalf of Nordic Society Oikos Stable URL: http://www.jstor.org/stable/3545586 Accessed: 14-02-2016 03:43 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/3545586?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Nordic Society Oikos and Wiley are collaborating with JSTOR to digitize, preserve and extend access to Oikos. http://www.jstor.org This content downloaded from 130.63.180.147 on Sun, 14 Feb 2016 03:43:15 UTC All use subject to JSTOR Terms and Conditions OlKOS 77: 68-76. Copenhagen1996 The socioecologyof bodysize variationin theprimitively eusocial sweatbee, Halictusligatus (Hymenoptera: Halictidae) Miriam H. Richardsand LaurencePacker Richards,M. H. and Packer,L. 1996. The socioecologyof body size variationin the primitivelyeusocial sweat bee, Halictus ligatus(Hymenoptera: Halictidae). - Oikos 77: 68-76. Patternsof demographicand social variationexhibited by the sweat bee Halictus ligatus,reveal that the expressionof eusocial colony organizationis associated with local environmentalconditions, harsher conditions promoting more classical eusocial- ityand gentlerconditions promoting diminished eusociality. We examineda variety of factorsaffecting body size variationof H. ligatusat a nestingaggregation near Victoriain southernOntario, Canada duringthe summersof 1984, 1990, and 1991. Body size variationwas associated withgender and reproductivecaste as expected, and therewere significant colony effects on size as well. An unexpectedresult was the dramaticeffect of local environmentalconditions on body size. Changes in weather patternsfrom year to yearresulted in significantbody size variationamong all classes of adult sweatbees, includingqueens, workers, gynes, and males. In 1990 cool, rainy weatherled to the productionof brood withrelatively small body sizes,while in 1984 and 1991, relativelydry, warm weatherhad the opposite effect.Weather probably influencedbrood body size by affectingthe abilityof adult femalebees to gather provisionsfor the larvae,and so indirectlyaffecting food availabilityto brood. Since queens are produceda yearbefore their own workers,environmental factors affecting theirrelative body sizes are uncorrelated.Nevertheless, the degreeof eusocial colony organizationis stronglyaffected by variation in the relativesizes of queens and workers,and this illustratesthe extentto whichstochastic environmental variation structuresopportunities for behavioural interactions in primitivelyeusocial bees. The patternof inversevariation between temperature and body size in thisbee contradicts the generalpattern of increasedbody size at highertemperatures generally seen in insectsand otherectotherms, and indicatesthat social insectscould be an exception to this rule. M. H. Richardsand L. Packer,Dept of Biology,York Univ., North York, ON, Canada M3J 1P3 (presentaddress of MHR: Dept of Biological Sciences, Brock Univ.,St. Catharines,ON, Canada L2S 3A1). For ectothermicorganisms, two major environmental adult body sizes (Ray 1960, Atkinson 1994, Berrigan influenceson body size are food availabilityand devel- and Charnov 1994). In specieswith parental care, adult opmental temperature.In insects,the more food an body size may also be influencedby factorsaffecting individualconsumes during the larval stages,the larger the ability of adults to provision the young. Many it is as an adult. This apparentlysimple relationship is insectsare mass provisioners,providing a larva withall complicatedby the effectsof temperature.At higher the food it will consume before developing into an temperatures,insects may develop faster,and so move adult. In such species,there exists the opportunityfor fromone larval stage to the next more quickly,spend provisionersto preciselycontrol the size of theiroff- less timeas feedingjuveniles, and end up withsmaller springin responseto ecological factorssuch as season Accepted20 February1996 Copyright?) OIKOS 1996 ISSN 0030-1299 Printedin Ireland - all rightsreserved 68 OIKOS 77:1 (1996) This content downloaded from 130.63.180.147 on Sun, 14 Feb 2016 03:43:15 UTC All use subject to JSTOR Terms and Conditions Table 1. The influenceof weatheron the demographyand social behaviourof VictoriaH. ligatusin 1984, 1990,and 1991 (table slightlymodified from Richards and Packer 1995). Year 1984 1990 1991 Temperature warm cool hot Rainfall below average heavy moderate Nest initiationand synchronous synchronous asynchronous firstbrood emergence Nest and brood survival good poor good No. firstbrood per nest 6.65 5.83 >9.0 Firstbrood sex ratio 14.6 11.5 5.5 (% male) No. workersper nest 5.65 5.16 8.5 Second brood emergence protandrous not protandrous protandrous Ovipositionpatterns dominatedby queens large contributionby workers Colony organization more eusocial more eusocial less eusocial and food availability,and social factorssuch as the survival,produced few workers, and queens dominated population-widesex ratio (Wrenschand Ebbert 1993). oviposition.Under more favourableconditions due to In social insects,body size is also an importantcorollary unusuallywarm temperatures,colony and brood sur- of reproductivecaste (Breed 1976, Packer and Knerer vival were high, large numbersof workerswere pro- 1985, Richards and Packer 1994), so environmental duced, and workerslaid a large proportionof eggs in influenceson body size are inextricablyentwined with the reproductivebrood. Such effectsmay also underlie those affectingdemographic and social variation. continent-wide,north-south dines of increasingcolony A seriesof recentstudies points to the role of envi- size and decreasingeusociality in H. ligatus.In southern ronmentalvariation in organizingthe behaviour of areas with long nestingseasons, there are very large primitivelysocial sweat bees (Wcislo in press). In sev- numbersof workersper nest,making queen controlof eral species,there is markedgeographic variation in the workerbehaviour less effectivethan in northernareas expressionof eusociality,with populations in harsh withshort nesting seasons wherethere are many fewer climates (e.g. high altitudes)reverting to solitarybe- workersper nest. haviour (Sakagami and Munakata 1972, Packer 1990, In H. ligatus and other primitivelyeusocial sweat 1991, Richards 1994a, b, Eickwort et al. in press). bees,queens controlworker behaviour through physical Variationin social behaviourand colony organization aggressionso large numbersof workersmake queen is also observed in sweat bees with less extremebe- controlof workerbehaviour less effective(Kukuk and haviouralflexibility. One exampleis Halictus ligatus,a May 1991). However,adult body size must also be an primitivelyeusocial species that has been studied importantfactor governing the outcomeof social inter- throughoutits geographicalrange, and in which soli- actions. Halictus ligatus queens have been observed tary populations have never been observed. Halictus pummellingworkers into the sides of nest tunnelsor ligatusexhibits strong variation in the degreeof euso- forcingthem out the entranceto forage (Packer and cial colony organization:harsher environmental condi- Richards unpubl.), and the larger size of the queens tionspromote more classicallyeusocial behaviour,and clearly confersan advantage in such confrontations. gentlerconditions lead to diminishedeusociality, trends Variationin body size would thereforebe expectedto evidentat both local and continentalscales (Kirkton contributesubstantially to variationin the expression 1968,Michener and Bennett1977, Richards and Packer of eusocial behaviour.Halictus ligatusexhibits consid- 1995, Richards et al. 1995). By diminishedeusociality, erable geographic variation in body size (Kirkton we mean that workers'reproductive efforts shift from 1968), bees fromnorthern populations being generally assistingthe queen to raise the queen's reproductive smallerthan bees fromsouthern populations. The dine offspring,to raisingtheir own reproductiveoffspring, in body size is very likelyrelated to the north-south includinggynes. dines of increasingcolony size and decreasingeusocial- Our studies of a population of H. ligatus near ity,and likewise,could also mirrorlocal patternsof size Toronto in southernOntario (Table 1; Richards and variation. Packer 1995, Richardset al. 1995) indicatethat annual In this study,we investigatevariation in adult body variationin local weatherconditions directly influences size and its social and demographiccorrelates and the demographyof sweat bee colonies, and indirectly consequencesas part of our overall studyof environ- influencesthe natureof social interactionsamong adult mentallyorganized social behaviourof H. ligatus.We females.Under adverseconditions caused by excessive foundthat
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