nutrients in wasp honey

8/12/2019 Nutrients in Wasp Honey

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PHYSIOLOGY, BIOCHEMISTRY, AND TOXICOLOGY

Nutrients in Social Wasp (Hymenoptera: Vespidae, Polistinae) HoneyJAMES H. HUNT, ANTHONY M. ROSSI, 1 NELS J . HOLMBERG,2 SAMUEL R. SMITH,3

ANDWILLIAM R. SHERMAN4Department ofBiology,University of Missouri-St. Louis, St. Louis, MO 63121

Ann. Entomol. Soc. Am. 91(4): 466-472 (1998)ABSTRACT Previous investigators have questioned the temporal occurrence, biochemistry, andnutritional use of honey sometimes present in nests of some social polistine wasps. Honey of speciesin the generaPolistesandPolybiacontains diverse amino acids. Inositols (alicyclic polyalcohols) alsoare present in the honey of both genera; quercitolwasthe most abundant inositol in honey ofPolistesmetricus(Say) from Missouri, but it was not present in honeyofPolybia diguetanaR. du Buyssonfrom CostaRica.Honey ofP.metricusfrom Missouri is similar to that ofBrachygastramellifica (Say)because it contained more fructose than glucose. Honeyhasbeen seen inP.metricusnestsinMissouriat all phases of the colony cycle except late preemergence (1st pupae); scattered but suggestiveobservations indicate that swarm-founding trop ical polistine wasps store honey primarily during lowactivity phases of the colony cycle in locales with alternating wet and dry or subtropical seasons.Although the nutritional value of wasp honey seems clear, the nutritional role of honey in waspcolonies remains unknown.KEY WORDS Polistes, Polybia,Epiponini, wasp, honey, amino acid, inositol, quercitol, glucose,fructose

HONEY STORAGE IS well-known in highly social beessuchashoneybees (Apis.Apinae),stinglessbees (e.g.,Melipona, Trigona: Meliponinae), and bumble beesBombus:Bombinae) (all reviewed by Michener1974). Honey storage by social polistine wasps (Vespi-dae:Polistinae)is lesswell-known, althoughit was firstreported nearly 200 yrago.Azara (1809,pp. 171-172)drew the attention of European naturalists when hedescribed the collection in Paraguay of a social waspnest that contained honey. Although Walckenaer(footnote in Azara,1809,p. 172) and Latreille (1824)believed Azara had mistaken bees for wasps, St. Hi-laire's description (1825) of honey in a nest made ofpaperlikethose of European wasps confirmed Azara'sobservations, as reported by Latreille (1824) and re-confirmed by White (1841). Thesewaspswere swarm-founding species,nowplacedinEpiponini (C arpenter1982); they have large colonies and build enclosednests with multiple combs of cells. The biology ofEpiponini is reviewed by Jeanne (1991). Honey stor-age by the epiponineBrachygastra mellifica (Say) insouth Texas has been described by Schwarz (1929),Bequaert (1932), and Sugden and McAllen (1994).

Polistes(Polistini) are small-colony paper waspsthat found nests independently and that build single-1 Department of Biology, University of South Florida, Tampa, FL33620.2Me tabolism Division, Wa shington U niversity School of Med icine,St. Louis, M O 63110.3Department of Pathology, Washington University School of Med-icine, St. Louis, MO 63110.4Department of Psychiatry, Washington University School of Med-icine,St. Louis, M O 63110.

comb, unenclosed nests (biology reviewed by Evansand West Eberhard 1970, Reeve 1991). Honey storagebyPolisteswas reported by Lepeletier (1836, pp. 482-483),Rouget (1873,pp. 196-197), Brongniart (1895),and Marchal (1896). von Ihering (1904) reported ob-servations on honey storage by Brazilian wasps, in-cluding species ofMischocyttarus (Mischocyttarini),which resemblePolistesin most aspects of their biol-ogy (reviewed by Gadagkar 1991). Jeanne (1972) de-scribes honey storage byM. drewsenide Saussure.Wheeler (1908) and Rau (1928) provide extensivediscussions of honey storage by wasps, especiallyPolistes. Honey of epiponine wasps typically fills halfor more ofanest cell thathas no eggor larva, and the remay be hundreds to thousands of honey-containingcells in a nest at one time. Such honey is sometimescollected for human consumption (e.g., Vellard 1939,Crane 1990). Honeyinepiponine nests also may occuras hard crystalline (candied) spheres in nest cells oras small droplets on nest cell walls or, uncommonly, asdroplets in interstices of the nest envelope (e.g., Po -lybiaaequatorialis Zavattari, S. O'Donnell, personalcommunication). Honey in nests of PolistesandMischocyttarus occurs most often as droplets, eachusually = 5jui,orlessfrequentlyascandied blobs. Thehoney occurs on the walls of nest cells that are em ptyor that contain an egg or small larva (J.H.H., unpub-lished data).

In no case is the honey in wasp nests sited wherelarvae can feed on it. Only adults can feed directly onit, and it is not known if adult wasps, like honey bees(Winston 1987), regurgitate honey from their crops tolarvae, as Marchal (1896) believed to be the case.0013-8746/98/0466-0472$02.00/0 1998 Entomological Society of America


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July 1998 HUNT ET AL.: NUTRIENTS IN SOCIAL WASP HONEY 467Pathways of nutrient movement among individuals ofa social wasp colony can be as intertwined as those ofany social insect (Hunt etal.1987,figure12.2 in Hunt[1991]), and intracolony movement pathways andprimary consumers of honey in wasp colonies remainto be fully documented.Lepeletier (1836) observed honey inPolistesnestsduring the phase of the colony cycle in which malesand reproductive females are produced, and he spec-ulated that the honey must be essential for the devel-opment of these reproductives. Marchal (1896) wasskeptical, because he found honey in Polistes nestsonly in the preemergence phase. Rouget (1873) sawno natural honey storage at all inPolistes,but coloniesthat w ere fed sugar stored itashoney at the end of thecolony cycle, when the re were few ifanylarvae to befed, and Rouget saw no use in such end-of-seasonhoney storage. Strassmann (1979) addressed Rouget'sconcern when she described the use of honey in nestsofPolistes annularis(L.) in Texas as a source of mid-winter nourishment for gynes (inseminated femalesthat are potential queens of the coming nesting sea-son) that emerged from sheltered hibernacula onwarm midwinter days, and she proposed an adaptiverole for honey in enhanced gyne survivorship. Hunt(1982,1991) proposed honeytobe1of several sourcesof liquid proteinaceous nourishment that can under-gird the origin and elaboration of sociality in Hyme-noptera.

Lassaigne (1824) described the color, solubility, andcrystalization of the wasp honey collected by Saint-Hilaire (1825). Bertrand (1895) reported that honeycollected fromaPolistesnest did not turn the plane ofpolarized light to the left, as does honey bee honey.The disaccharide sucrose, which turns polarized lightto the right,isa principle carbohydrate of many flowernectars (Maurizio 1975). Honey bees mix nectar w ithsalivary enzymes (Maurizio 1975, White 1975) thatcleave sucrose into the monosaccharides glucose andfructose and, concomitantly, cause polarized light tobe turned to the left (i.e., the plane of polarized lightbecomes inverted). Bertrand (1895) concluded thatwasp honey must contain primarily sucrose and dex-trose (glucose) without levulose (fructose), and heproposed that wasp honey must therefore consist ofconcentrated nectar without alteration by invertase.However, Santolaya and Gentile (1953) reported 74%reducing sugars, which include glucose and fructose,and5%sucrose (sacarose) in honey ofPolybiascute-llaris(White) from Cordoba, Argentina. Sugden andMcAllen (1994) performedgaschromatographic anal-yses of major sugars in honey ofBrachgastra mellifica(Say) from southern Texas, and they reported theprobable identity (given here in rank order from mostto least) of fructose, other , glucose, sucrose, stachy-ose, raffinose, melibiose, and arabinose. Burgett(1974) gave evidence for the presence of glucoseoxidase, which can inhibit bacterial activity, in honeyof the epiponine Protonectarina sylveriae (de Saus-sure).Crane (1990) said, it appears likely that bothinvertase and glucose oxidase systems were developed.. . by all 'honey -storingHym enopterain the Apidae,

Vespidae, and Formicidae. She advocated use of theword honey for the stored carbohydrate food ofanysocial insect that has invertase and glucose oxidasesystems.Two studies strongly suggested that honey can beimportant to nutrient economy in wasps during theactive phases of the colony cycle. Rossi and Hunt(1988) placed 5-ix\drop lets of slightly dilute Apishoney into nests ofP. metricus beginning soon afternest founding and continuing twice weekly until off-spring emergence. Supplemented nests produced 1stoffspring earlier than did control colonies, and lst-emerged offspring of supplemented colonies had ahigher percen tage of fat than not only control off-spring but also foundresses. Dove (1994) repeated theexperiment and found that P. metricus colonies thatreceived honey supplementation in the preemergencephase attained larger nest sizes and produced moreoffspring by the end of the season, but the frequencyof workers among total offspring was lower than incontrols.It seems reasonable that honey can significantlyaffect both development and life history of otherhoney-storingwasps asit does forP.metricus.Nutrientanalyses of wasp honey can thus be of direct relevanceto broad issues in the biology of social wasps. ThecontentsofPolisteshoney have never been examined,nor are there analyses of amino acids in any wasphoney, although amino acids are well-known constit-uents ofApishoney (e.g., White 1975, Davies 1975)and, as free amino acids, of floral and extrafloral nec-tars (Baker and Baker 1973). Here we analyze aminoacids and inositols in honey of wasps in the generaPolistes (Polistini) andPolybia (Epiponini) plus majorsugars in honey of aPolistes species.

Materials and MethodsStudy and Collection Sites. Honey-containing nestcombs from an active colony ofPolybia occidentalis(Olivier) and anotherofPolybia diguetanaR.du Buys-

son were collected on31January1984at Hacienda LaPacifica nearCanas,Guanacaste Province, Costa Rica.Nest combs containing honey were taken from a col-ony ofP.diguetanaat Monteverde, Puntarenas Prov-ince, Costa Rica, on 9 January 1996. Observations ofhoney in epiponine nests have been made occasion-ally at other times at these and other Neotropicallocations.Three honey-containing preemergence nests ofPolistes humilissynoecus de Saussure were collectedfrom Decem ber 1988 to February 1989 at the CSIROBlack Mountain Site, Canberra, ACT, Australia. Twohoney-containing, preemergence nests ofP.metricuswere collected at Washington University's Tyson Re-search Cen ter near Eureka, St. Louis County, MO, on2 May 1988. Three similarP.metricusnests were col-lected on 17 May 1996. Three postemergence P. met-ricusnests were collected on7July 1997 and anotheron 30 July 1997 at the Missouri Botanical Garden'sShaw Arboretum near Gray Summit, Franklin County,MO. Sporadic observations of honey in P. metricus


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8 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 91, no. 4nests have been made over the past 20 yr at TysonResearch Center and Shaw Arboretum. Vouchersspecimens of thePolybiaspecies and P.metriciisare inthe Museum of Natural History of the University ofMissouri-St. Louis.

Amino Acids. The samples of P. occidentalisandP.diguetanahoney collected in 1984 were held in situ inthe nest comb under laboratory ambient conditionsuntil May 1988. At that time samples of honey weretaken from these combs and from combs ofPolisteshumilis collected a few months earlier, and from P.metricus combs, which were freshly collected. Foreach nest of each species, honey droplets were placedon the walls of a clean glass vial by using a spatula, andthe samples were frozen at80C until analysis in July1990. Sample preparation was preceded by suspensionof the honey droplets in deionized water. A 10-mgaliquot of each sample was hydrolyzed with 6 M HClat 110C for 24 h under a nitrogen atmosphere andthen freeze-dried. Samples were derivatized prior toamino acid analysis by adding 100 jLtl of2:2:1ethanol:triethanolamine (TEA):H2O, drying under a vacuum,and then adding 20 jal of 7:1:1:1 ethanol:TEA:H2O:phenylisothiocyanate (PITC) to each sample. The de-rivatizing mixture was allowed to react for 20 minunder a nitrogen atmosphere at 22C. Samples weredried and reconsitituted in a phosphate buffer con-taining 5 mM sodium phosphate with 6 acetonitrile.Samples and external amino acid standards were quan-tified with a Waters Associates high-performance liq-uid chromatography (HPLC) gradient system equippedwith a Pico*Tag column (Heinrickson and Meredith1984, Andersen et al. 1989).

Glucose and Fructose. Nests collected in 1997 werefrozen with honey droplets in situ at20C within 5 hof collection. For analysis, honey droplets were re-moved from nests, weighed, and dissolved in deion-ized water. Honey droplets from the 3 nests collectedon7July were pooled for1sample; droplets from thenest collected on30July were pooled fora2nd sample.Both samples were analyzed enzymatically by themethod of Passonneau and Lowry (1993). The reagentconsisted of50mM Tris buffer at pH 8.1, 2 mM mag-nesium chloride, 0.5 mM ATP, 0.5 mM NADP, and0.02 BSA. To measure glucose, 1.0 unit per ml hex-okinase and 0.1 unit per ml glucose-6-phosphate de-hydrogenase were added to amixof 0.025 ml of samplein 3.0 ml of reagent, and the increase in NADPH wasmonitored inaPerkin-Elmer Lambda3Bdouble-beamspectrophotometer. At the completion of the glucosereaction, fructose was measured by addition of 0.4units per ml phosphoglucoisomerase and spectopho-tometric monitoring. The millimolar extinction coef-ficient of NADPH was taken to be 6.22 at 340 nm.

Inositols. Inositols are alicyclic polyalcohols (poly-ols) with hydroxyl groups directly attached to thecarbons of the ring. Samples collected in 1996 wereprepared for analysis byaprocedure that removes99of the reducing sugars and that retains polyols (Os-tlund et al. 1993).Awater-diluted sample of honey wasshaken with 250y\ of washed Amberlite IR-120(H+)for 45 min; the supernatant was then transferred to a

tube containing 350 \x\of Amberlite IRA-440c(OH)and shaken for2.5h. Next, the supernatant was passedover a C-18 Sep-Pak (Waters Associates, Milford, MA)that had been washed with methanol and water, andfinally the samples were vacuum-dried before deriva-tization.

Samples were derivatized with pentafluoropropio-nyl (PFP) groups (Ostlund et al. 1993) by reactingwith 40 jLtl of an acetonitrile solution containing 10pentafluoropropionic anhydride (PFPA) for 30 min at65C. An aliquot was then dried and redissolved in asolution of 0.35 PFPA in acetonitrile prior to chro-matography. For derivatization with trimethylsilyl(TMS) groups (Sherman et al. 1970), dried sampleswere treated with 50 bis(trimethylsilyl)trifluoro-acetamide (Regis) in dry pyridine for 1 h at 65C.

The derivatized samples were analyzed by gas chro-matography/mass spectrometry (GC-MS). PFP-deri-vatized samples were run on a Hewlett-Packard 5988AGC-MS in the negative ion chemical ionization(NICI) mode; TMS-derivatized samples were run ona Hewlett-Packard 5970 GC-MS with electron ioniza-tion in the positive ion detection mode. Separation ofthe PFP-derivatized samples was carried out on aChirasil-Val fused silica capillary column (25mby 0.32mm i.d., Alltech Associates, Deerfield, EL) with tem-perature programming as follows: 92C for1min, 12/min to 112, 217 min to 180C, followed by a bake-outof 70C/min to 205C. For the TMS derivatives, sep-aration was performed on a J&W DB-210 column(Alltech) (15 m by 0.32 mm i.d.) with temperatureprogramming of 90C for 0.5 min followed by 20C/min change to 200C where it was held for 7 min forbake-out.

Chemical standards of inositols were obtained asfollows: D-c/uro-inositol from Calbiochem; authentic(+)-quercitol and L-c/iiro-inositol as gifts of LaurensAnderson (University of Wisconsin, Madison); andduterium-labeled (l&S^S^-dg-D^-c/iiro-inositol fromKen Sasaki (Connaught Centre for Biotechnology Re-search, Toronto). The preparation of the latter com-poundisdescribed in Sasaki et al. (1987). This materialalso contained small amounts of d6-mi/o-inositol, d6-neo-inositol, and d6-sq///o-inositol.

ResultsAmino Acids. In the honey of 4 wasp species the

number of detected amino acids ranged from 12 (P.diguetana to 17 (P.metricus1) (Table 1). All of thedetected amino acids are nutritive. Three of the sam-ples contained 6 of the 10 essential amino acids; 2samples contained 9, and 2 samples contained all 10.The proportional abundances of amino acids in wasphoney are less equitable than those of typical proteinreported by King and Jukes (1969). Wasp honeys hadfrom 2 to 4 amino acids with proportions higher thanthose of the most common amino acid in typical pro-tein and, concomitantly, from 3 to 9 amino acids withproportions lower than the least abundant amino acidin typical protein.


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July 1998 HUNT ET AL.: NUTRIENTS IN SOCIAL WASP HONEY 469Tulilc I. Concentrations (mol ) of total aniino acids in the honey of 4 wasps species

AsparagineClutamineSerineGlycineHistidine*Arginine*Threonine*AlunineProlineTyrosine*Valine*Methionine*CysteineIsoleucine*Leucine*Phenylalanine*Lysine*

P liA11.126.42.74.64.14.414.38.51.41.11.617.52.4

P.h.27.69.03.94.67.217.617.43.313.62.51.51.81.73.22.23.1

P.h.321.316.9

4.46.73.02.39.614.510.11.81.91.41.01.91.71.7

P.m.l33.125.2

2.61.33.54.67.28.76.41.41.70.90.40.71.01.00.4

P.m.24.140.02.51.11.715.54.35.38.72.15.00.62.51.41.04.5

P.d.23.221.31.71.36.624.310.45.21.11.51.12.3

P O

15.517.42.44.21.33.19.819.312.23.91.71.70.92.81.72.0

K-J10.89.2

8.58.03.04.46.57.85.23.57.11.94.08.04.26.0P.h.,P. humilis (3 nests);P.m.,P.metricus (2 nests);P.d., P.diguetana;P.O., P. occidentalis.Also given is the com position of average pro tein,as calculated by King and Jukes (1969). Essential amino acids are marked with an *.

Glucose and Fructose. Table 2 gives the mean of 2assays for each pooled honey sample as weight, mi-cromoles, and percentage of total sample weight forboth glucose and fructose. In both samples fructoseconstitutes20%by weight of the total honey. Glucoseis much less abundant, at 1-3%.Inositols. InP.metricushoney, the principal inositolof both PFP and TMS derivatives was identified asquercitol (lL-l,2,4/2/5-cyclohexanepentol) by thefollowing criteria. The PFP-derivatized honey sampleshad 2 principal GC peaks with retention times of135.25 and 138.00sand of equal intensity; the authen-tic + )-quercitol as the PFP derivative had a singlemajor component GC peak at134.71s.The mass spec-trum of both of the major GC peaks of the PFP-derivatized honey had majorionsatm/z894(M~) andm/z874(M~-HF;ion intensity ofm/z874>894);thesame majorionsand relative intensities were observedin the spectrum of the PFP derivative of authenticquercitol. We therefore ascribe the presence of2GCpeaks identified as quercitol in the honey sample, incontrast to only 1 analogous peak in the authentic + )-quercitol, to the chiral separatory properties ofthe Chirasil-Val GC column (Leavitt and Sherman1982). We suggest that the P .metricushoney samples

Tnltlc 2. Glucose and fructose by weight, micronioles, and aspercent of total sample weight for 2 p ooled samp les of P. metricushoney7 July nests 30 July nest

Total honeyWeight, mgGlucoseWeight, mgMicroniolesPercentFructoseWeight, mgMicroniolesPercent

8.90.10.61.02.011.122.0

36 2

1.16.13.07.340.520.0

Each value is the mean of 2 analyses.

contained both (+ ) -quercitol and()-quercitol, withthe (+ ) form being the 1st of the pairtoelute from theGC column. TheP.diguetanahoney sample containedno quercitol.The TMS-derivatized P.metricushoney revealed asingle major peak with retention time of 207.6 s; au-thentic +)-quercitol had an analogous single peakeluting at 207.4s.The presence ofasingle GC peak asthe TMS derivative in both cases is attributed to theachiral nature of the separation on the DB-210 GCcolumn. The mass spectra of both the P. metricushoney and the authentic quercitol contained the fol-lowing ions in decreasing order of intensity:m/z 318,m/z 305,m /z 344,m /z 419,m/z 331,andm /z 329. Oftheseions,m/z344can be related to the molecular ionm/z524 (not seen) by the loss of2TMSOH moieties,and m /z 419 can be th e result of the loss of1TMSOHand1methyl radical from M+. Ions atm/z318and 305are common to the TMS inositols.Although +)-quercitol appears to be the majorinositol in the P.metricus honey samples, retentiontimes and the presence of appropriate mass spectralions during GC-MS of the PFP and TMS derivativesindicate th e presence of small amounts of myo-inosi-tol, L-c/iiro-inositol, D-c/iiro-inositol, neo-inositol,sct/Zfo-inositol, and pinitol (Table 3).Because the mass spectrum of viburnitol (1L-1,2,4/3,5-cyclohexanepentol) would probably be indistin-guishable from that of quercitol, and because we hadno viburnitol for use as a gas chromatographic reten-tion time standard, we cannot exclude the possibilitythat our samples are, or contained, viburnitol. How-ever, this is unlikely considering the retention timeidentities to quercitol that we observed with 2 differ-ent derivatives and 2 different columns.

DiscussionBecause our amino acid analyses were of hydro-lyzed samples, we cannot conclude whether ouramino acid data represent protein , free aminoacids,or


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47 ANNALSO F THEENTOMOLOGICAL SOCIETY OFAMERICA Vol. 91, no. 4Table 3 . Inositols found in honey samples of P. metricus(3nests)and P.diguetanaandidentified as PFP or TMSderivativesWaspDerivative

P. met.1 P. met.2 P.met. 3 diguetanaPFP TMS PFP TMS PFP TMS PFP TMS

Inositolchiro-neo-nnjo-PinitolQuercitolsajllo-+, Indicates achromatographic retention time identity with anauthentic sampleaswellas thepresenceofmass spectral ions thatarerepresentativeof theinositolderivative.,Denotes thattheinositolwasabsentorbelowthedetection limitsof theanalysis.

both. Ingested pollenhasbeen foundinseveralPolistesspecies and inB.mellifica(Hunt et al. 1991), but wedid not examine wasps or honeys in the current studyfor the presence of pollen. Pollen could contributesignificantly to the amino acid composition of honey,and future investigations of wasp honeys should in-clude attention to this possibility. Differences in theranked proportional abundances of amino acidsamong the3samples ofP.humilishoney and betweenthe 2 samples ofP.metricushoney from the same siteand date suggest use of diverse, possibly nonfloralnectar or nonnectar, sources by these small-colonywasps. Similarities of ranked proportional abundancesof the more common amino acids in honey of the 2large-colonyPolybiaspecies may reflect use ofasim-ilar, possibly floral resource, even though the samplesrepresent different dates and sites separated by ^50km.The honey of P. metricusis similar to that ofB.melliferaas reported by Sugden and Me Allen (1994)because it contains more fructose than glucose. Thesmall available quantities of P.metricus honey pre-cluded analyses for sucrose or other major sugars. Atthis time, we cannot address the sources of the P.metricushoney. We strongly agree with Crane (1975)that the question of whether or not wasps do, indeed,have enzymatic cleavage (inversion) of sucrose meritscareful reexamination.Inositols are known from a wide range of plant andanimal tissues, are implicated inawide range ofphys-iological activities, and are apparently universal asphospho-inositides in phospholipids (Posternak1965). Plants produce inositols, and nearly all plantscontain them (Posternak1965). +)-Quercitol occursin various parts, including leaves and twigs, of severalspecies of oak Quercus) (Posternak 1965); variousoaks are abundant at Shaw Arboretum where thehoney samples we analyzed for inositols were col-lected. In insects, inositols have been shown to beinvolved in melanization inS chistocercagregaria (For-skal) (Dadd 1961), and they are essential for thegrowth ofPeriplaneta americanaL.,variousplant-feed-ing Orthoptera, Coleoptera, Homoptera, and severalLepidoptera, although not all insects require them(references in Wigglesworth 1972, Dadd 1973, House

1974). Inositols stimulate feeding in most (herbivo-rous?) insects (Hanson 1983). In Hymenoptera, ino-sitols are essential for larval development in honeybees (Winston 1987), and a red ant had an inositolcontent of 2,200 y/g (Posternak 1965). The nutritionalrole of inositols in the wasps we studied is unknown,but a significant role is probable. Two propositionsthat can be drawn from the presence of inositols inwasphoney are that the honeymay have come, at leastin part, from nonfloral sources and tha t the honey canserve the wasps as a reserve ofthispotentially impor-tant class of nutrients.Several of the pioneering authors cited in the in-troduction queried the temporal occurrence of honeyin wasp nests. Rau (1928) studiedPolistesin the sameregion of Missouri, bu t no tinthe same sites used in thecurrent study, and he noted occurrence of honeydroplets inPolistesnests throughout the nesting sea-son but in greater abundance toward the end of thesummer. Our observations in Missouri have revealedthat honey storage in early preemergenceP.metricusnests (lacking pupae) in May is commonplace. Inter-estingly, honey seems to be absent from late preem er-gence nests (with pupae) in June, but itisoften found(although sometimes difficult to observe) in nests inJuly and August from which offspring have emerged.Although the largestP.metricusnest studied by Dove(1994) had more abundant honey droplets in earlyOctober than any other Polistes nest we observed(J.H.H., unpublished data), other active nests at thesame site and date had none.Honey storage by tropical epiponines seems toreach highest levels in th e dry season of strongly sea-sonal locales (Hunt et al. 1987, Kojima 1996) and athigh elevation (e.g., Cooper 1993) and latitudinal ex-tremes (Texas and Paraguay) of their distribution. Itseems likely that honey storage by epiponines occursprimarily in seasonal sites during times when larvaeare few in number.Investigations describing2honey-storing species ofthe epiponinegenusBrachygastraatthe margins of thegenus' distribution in Texas B.mellifica:Sugden andMcAllen 1994) and Paraguay B. lecheguana: Azara1809) have noted that the wasps are not very aggres-sive (see also Vazquez de Espinosa1942 [1628]). Thiscontrasts markedly to experience with B.mellifica intropical dry forest regions of Costa Rica and savannasof Venezuela, where it is among the more aggressiveepiponine wasps (J.H.H., unpublished data,i eR. L.Jeanne), except when a nest is in decline and nearlydevoid of both brood and honey (J.H.H., unpublisheddata).

AcknowledgmentsWe thankJ.Philip Spradbery, who collected theP.humilisnests; Sean O'Donnell, who collected the 1996 P.diguetananest; Richard Coles, who welcomed and facilitated our fieldstudies at Tyson Research Center; James Trager, who wel-comed and facilitated our field studies at the Shaw Arbore-tum; Jeanne M organ Zarucchi, who assisted with the trans-lation from French to English of several key passages; and


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July 1998 H U N T E T A L . : N U T R I E N T S I N S O C I A L W A S P H O N E Y 471David W. Roubik and James M. Carpenter, who broughtreferences to our atten tion. C ollection of samples in 1984 wassupported in part by a grant to J.H.H. from The AmericanPhilosophical Society and by a Research Leave granted bythe University of Missouri-St. Louis.

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Received for publication 15 October 1997; accepted 1December 1997

nutrients in wasp honey

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