AMIERICANt MUSEUM Novitattes PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3224, 39 pp., 26 figures April 6, 1998

A Generic Key to the of , , and Paper Worldwide (: , )

JOHN W. WENZEL'

ABSTRACT The 31 genera of Vespinae and Polistinae tary with which they may be con- worldwide are identified in a key to struc- fused. Many characteristics are illustrated or de- ture. Fifty-nine couplets and more than 80 pho- scribed here for the first time, with notes on tographs and illustrations permit both special- both anomalous species and anomalous forms ists and amateurs to recognize these nests in the of nests of common species. Pertinent published field or museum collections. A brief overview figures and museum collections are cited to explains the distinction between nests of these assist the professional in finding reference ma- social wasps and those of other social or soli- terial.

INTRODUCTION All over the world, both entomologists and female (Wenzel, 1987) or millions (Zucchi et the lay public recognize and fear colonies of al., 1995). The aggressive, boldly striped social wasps. More than 900 species range adults advertise their unforgettable stings, from the Arctic to Tasmania, from prairie to and many moths, , and other defenseless rain forest to desert, from pristine habitats to have developed elaborate morpholog- industrial cities. Their sophisticated, all-fe- ical and behavioral mimicry to benefit from male societies provided the inspiration for a general desire among most to several of the major discoveries in be- avoid wasps. In addition to supporting these havior and sociobiology (Wilson, 1971: 7). mimicry complexes, their large numbers Mature colonies may consist of a single make wasps important predators of other in-

' Assistant Professor of Entomology, Department of Entomology, The Ohio State University, Columbus, Ohio.

Copyright C) American Museum of Natural History 1998 ISSN 0003-0082 / Price $6.30 2 AMERICAN MUSEUM NOVITATES NO. 3224 sects in many ecosystems, composing up to or plant resins. Soil or glandular secretion about 6% of total insect biomass (Fittkau and may be used to reinforce or repair nests, but Klinge, 1973). They are well known for sym- they rarely constitute the primary building bioses with each other (Windsor, 1972) and material for mature colonies. Brood combs with other organisms such as (Espelie are usually one sided, with an hexagonal ar- and Hermann, 1988), honeydew insects (Le- rangement of cells sharing thin (less than 0.5 tourneau and Choe, 1987), and birds (Hind- mm) walls. There are no specialized storage , 1955, Joyce, 1993, Dejean and Fotso, chambers, but one may find honey or prey 1995). This commensalism attracted artists ants and stored in ordinary brood of ancient civilizations (Coe, 1982) and cells or in spaces between envelope sheets. forms the foundation of the mythology and In Southeast Asia, nests of rituals of certain indigenous peoples (hover wasps) may be confused with those (Schoepf, 1990). The architecture of of Vespinae and Polistinae, but stenogastrine nests has been cited for its perfection as ev- nests can be separated by the following char- idence against modem evolutionary theory acters: cells more than 1.3 times as large as (Anon., 1986). Yet, despite this rich resource the head diameter of the wasp; cells usually wasps provide for behaviorists, ecologists, narrowed at their openings by a short collar anthropologists, and others, identifying and may be closed with carton by the adults wasps to has always required the care when the pupates; pupae naked and of a few overburdened specialists with ex- leaving no cocoon inside the cell; carton tensive libraries and reference collections. fragile and brittle; nest lacking a narrow, The present key to nests is offered to provide tough pedicel (although sometimes produc- an alternative that will be useful to specialists ing illusion of one by hanging from rootlets, and amateurs alike. fungal hyphae, or other such substrate); cells The wasps' industriousness as builders and sometimes connected to the substrate along cleverness as architects are appreciated even their side walls; and cells easily separated by those of us who dread unexpectedly dis- from the substrate (except when built of covering them making their home in the shel- clay). In contrast, Vespinae and Polistinae ter of our own. Their elegant paper nests build cells barely larger than the head di- sometimes attract attention as decorative cu- ameter of the wasp; cell openings not nar- rios in markets. These nests can be the size rowed by a collar nor closed over pupae by of a thimble or more than a meter long, as adults (occasionally closed in some Vespa) durable as hard felt or more fragile than egg but rather capped by a portion of the silk shells, more regular and uniform than the cocoons found in the cells; carton either frag- much-celebrated honeybee comb or wildly ile and brittle or tough and supple; nest with chaotic with an intricate mazelike interior. In- or without pedicel (mostly with pedicels in deed, variation in nest architecture between Southeast Asia); cells rarely connected to the distantly related species is far more striking substrate along their side walls; and, in living than is the morphological variation, and colonies without pedicels, cells usually not some species were initially sorted into certain easily separated from the substrate. genera primarily based on their nests. Recent Nests of a few more distant relatives of treatments demonstrate that architecture-re- Vespinae and Polistinae may be confused lated behaviors are particularly useful at trac- with them occasionally. The eumenine gen- ing phlyogenetic lineages in many animals era Zethus and Calligaster build clusters of (Wenzel, 1992a) and that architectural details cells of resin and plant material, often with of wasp nests reflect well the relationships pieces cut from , but each of these bar- derived from morphological study (Wenzel, rel shaped cells has its own thick walls rather 1993). than sharing thin, common walls with neigh- The nests of Vespinae (hornets and yello- boring cells, and rarely are all cells parallel. wjackets) and Polistinae (paper wasps) are Clay nests of the Neotropical sphecids Try- easy to recognize. Except for four Neotrop- poxlyon fabricator, T. maraballi, and some ical species that build nests of mud, the nest other species may be confused occasionally are all made of vegetable fiber without wax with those of paper wasps, but the former are 1998 WENZEL: NESTS OF SOCIAL WASPS 3 naked combs of closely adjacent, circular whatever means, be accompanied by a cells each with its own walls, built without a change in architecture in one or both descen- pedicel, and often plastered with mud on the dant lineages. Eberhard's (1990: 342) ap- sides of the comb. Social vespid nests built praisal that apparent species specificity of in cavities have an entrance that is a simple webs is an illusion due to lack of data hole or crack, never an entrance tube of resin probably applies to wasp nests as well. Intra- as found in stingless , for example, Tri- specific variation between habitats or popu- gona or Melipona (Apidae). Carton nests of lations can be great, and limited sampling ants are usually composed of a coarse uneven may lead to generalizations that are false or envelope only, and never include a hexago- hold for only one part of the species' range. nal comb. A few Neotropical polistines live The illustrations offered are only to assist inside the carton nests of ants or termites, the user in deciding if he or she has found and observers should take care not to confuse roughly the right description of the nest. the structures built by these latter with the Nests are far more variable than are the mor- parallel, naked, carton combs of the wasps phologies of the animals who build them, that will be found within a cavity. and some identifiable nests will not match Museum nest specimens are often de- closely the figure chosen to characterize the cayed, discolored, distorted, lacking parts, or genus. Problematic nests, such as those in- confused with parts and adults of other nests. cluded among the diverse architectural forms Therefore, one must be cautious in certain of Paleotropical Ropalidia, are sometimes decisions, such as whether a given specimen best determined by excluding the other pos- naturally lacks an envelope or has merely sibilities (i.e., the nest does not resemble Pol- lost it in handling. I have made use of bio- istes, Belonogaster, , or Parapo- geographic and other information that will lybia). In the neotropics the number of alter- help identification in the field or with prop- natives is much larger, and several genera erly labeled material, but some information, present problems due to their diversity of such as "in a cavity," may not be evident in nest forms or a superficial resemblance to museum specimens. Although some nests other taxa. Parachartergus includes forms have survived two centuries in collections similar to Leipomeles and Chartergellus. (Wenzel, 1992b), I do not recommend bor- Protopolybia and Agelaia both have forms rowing nest specimens from museums be- that may resemble other genera, depending cause they are bulky and travel poorly, and on the exact details of the site and colony there is generally little the novice will gain size. Pseudopolybia sometimes superficially from examining a specimen that could not resembles Vespinae (but fortunately the most have been learned from published accounts. similar forms are allopatric). Some nests of This is the first effort to include all 31 gen- can be separated from era of Vespinae and Polistinae worldwide. It only with difficulty and both of these from is not possible to identify all specimens to Protonectarina only by destructive exami- genus without error before investing a great nation. Neotropical, naked, pedicelate combs deal of study, and several genera may not be in exposed sites are usually or Mis- separable on the basis of very young nests. chocyttarus, but the key helps separate these Effort is rewarded, though, because an ex- from particular forms of other genera (usu- perienced observer can identify nearly all ally Agelaia or Protopolybia). Polybioides, nests to genus from a distance of a meter or Synoecoides, Marimbonda, Clypearia (in- more, and many to species. In my experi- cluding Occipitalia), and Asteloeca are poor- ence, indigenous people who are alert to in- ly known and may not be properly charac- sects can identify the majority of local spe- terized here. Among nests of the approxi- cies by nest structure alone. mately 350 species I have studied (appendix Despite much variation, I do not believe 1) are several anomalous ones that will run that every species has a distinctive nest ar- through this key to an incorrect genus (e.g., chitecture. As a premise, species specificity the long-fiber nests of car- requires an assumption about evolution that bonarius in and M. flavitarsis in the I am not willing to make: that speciation, by Rocky Mountains may key to Polistes; Ro- 4 AMERICAN MUSEUM NOVITATES NO. 3224

A

Fig. 1. Supple paper, rolled around a pencil without breaking into separate pieces. palidia loriana will key to ; the short-chip nests of Polistes stenopus in the C South Pacific or Neotropical Polistes bicolor resemble Mischocyttarus). I believe most of these exceptions are encountered rarely, but as this key is used and improved, perhaps these exceptions will be treated better. The generic nomenclature used here is that of Carpenter (1996) or essentially that of B Richards (1978) with the addition of Aste- loeca (Raw, 1985), synonymies of Protopo- lybia and Pseudochartergus (Carpenter and Wenzel, 1989) and Clypearia and Occipitalia Fig. 2. Suspension. A. Narrow pedicel. B. (Carpenter et al., 1996), and the replacement Buttressed sheet. C. Cell-marginal pedicel, be- of Stelopolybia with Agelaia (Carpenter and coming flat apically and serving as the wall be- tween the first two cells, typical of Polistes, Mis- Day, 1988). Dolichovespula is separated chocyttarus, and others. D. Cell-central pedicel, from Vespula, but Paravespula is not rec- becoming cone shaped apically and serving as the ognized as separate from Vespula. Although base of the first cell, unique to Ropalidia, Para- phylogenetic relationships of the genera of polybia, and Belonogaster. paper wasps are not yet definitively estab- lished, an interim cladogram is available from Wenzel and Carpenter (1994) and is modified and reproduced here as figure 14. IMLT Instituto Miguel Lillo, Tucumdn, Ar- gentina This cladogram may be useful for interpret- KUSM University of Kansas Snow Museum, ing information presented here and else- Lawrence, Kansas where, but it is to be considered preliminary MALS Museum am Lowentor, Stuttgart and is presented here only for reference and MNHN Museum National d'Histoire Naturelle, without discussion. A thorough treatment of Paris the phylogeny of social Vespidae is yet forth- MRAT Musee Royal de l'Afrique, Tervuren, coming. Belgium NMBS Naturistorische Museums, Bern INSTITUTIONAL ABBREVIATIONS NNML Nationaal Natuurhistorische Museum, Leiden AMNH American Museum of Natural History, WDH William D. Hamilton personal collec- New York tion, Oxford BMNH The Natural History Museum, London CMNH The Carnegie Museum, Pittsburgh, DEFINITIONS Pennsylvania CUIC Cornell University Insect Collection, Brittle: Carton that breaks into separate frag- Ithaca, New York ments when rolled around to the opposite side of FMNH Field Museum of Natural History, Chi- a pencil. Contrasts with "supple" (fig. 1). Dry cago museum specimens may become brittle despite HMCZ Harvard Museum of Comparative Zo- being supple in nature. ology, Cambridge, Massachusetts Carton: Material composed of masticated veg- HUMB Humbolt Universitat, Berlin etable matter or soil, bound in a matrix of dried 1998 WENZEL: NESTS OF SOCIAL WASPS 5

.-.t.mg...... -

lom -Now= 14 - .. ..!"EM-W, x A C B

x Fig. 3. Lines of construction. A. Parallel lines of pulp, as in laminate envelope. B. Concentric arcs of pulp and imbricate pattern. C. Cross sec- Fig. 4. Composition of paper, low magnifica- tion showing superimposed structure of imbricate tion. A. Long fiber, typical of supple paper. B. envelope. Short chips, typical of brittle paper. salivary secretions. May have the consistency of mature nests (fig. 2). Synonymous with "petiole" paper, felt, or dried mud. or "peduncle" of other authors. Short connections Cell-central: Condition in which the apex of that do not function until they are several milli- the pedicel is gently flared into a cone that be- meters across are excluded from this definition, comes the base of the first cell of the comb (fig. although other authors sometimes include them. 2). Found in Ropalidia, Belonogaster, and Para- Secretion: Resinous, glossy material of glan- polybia. Contrasts with "cell-marginal." dular origin, added to pulp for strength and water Cell-marginal: Condition in which the apex of resistence. May be clear or any color; often re- the pedicel is flattened to become the side wall of sembles the basic color of the wasps' cuticle. the first cell (fig. 2), usually the common wall Short chips: Vegetable fragments less than between the first two cells. Found in Polistes, four times as long as wide, usually brittle. Con- Mischocyttarus, and other New World taxa. Con- trasts with "long fiber" (fig. 4). trasts with "cell-central." Supple: Carton that can be rolled around to the Envelope: A sheet of carton, mud, or secretion opposite side of a pencil without breaking apart protecting the combs and hiding them from the into separate fragments (fig. 1). Constrasts with environment. "brittle." Dry museum specimens may become Imbricate: Built or reinforced with shell-like brittle despite being supple in nature. structures usually less than 10 cm in breadth; lines of construction often tightly arced structures su- KEY TO NESTS OF VESPINAE perimposed upon each other like roof tiles or fish AND POLISTINAE scales (fig. 3). Laminate: Built or reinforced with sheetlike In case of error, the reader may trace de- structures usually more than 10 cm in breadth, cisions backwards by following the numerals lines of construction usually nearly linear and par- in parentheses to find the couplet at which allel, sheets mostly parallel and interconnected the alternative branch was more appropriate only intermittently (fig. 3). than the path chosen originally. An asterisk Lines of construction: A pattern visible in the means that the nest of this taxon is rarely carton due to the successive addition of separate seen or infrequently takes a form that will loads of building material to the margin of the key to the place given. envelope (fig. 3). Long fiber: Vegetable fiber that is slender and 1. Nest exposed ...... 2 more than four times as long as wide, usually sup- - Nest concealed in folded , hole in tree ple. Contrasts with "short chips" (fig. 4). or ground, hollow wall, attic, or other Pedicel: A tough, sometimes resinous, structure cavity ...... 48 that supports the nest from above or from the side, 2(1). Without envelope ...... 3 capable of function when diameter is 1 mm. It - With envelope ...... 9 may become broad, branched, or buttresslike in 3(2). Vacated pupal cells with bottoms lacking 6 AMERICAN MUSEUM NOVITATES NO. 3224

carton, either replaced by translucent secretion or completely open revealing A interior. No meconia in older cells (Old World only) ...... 4 - Vacated pupal cell bottoms of carton. -JL-- --L.- __iu_ With or without meconia (New World r Ir-73C -,Y--] only or cosmopolitan Polistes) .... 6 AQ .;Q::x ;IQ ir- 71 4(3). Cell bottoms open, revealing cocoons. k-4=46=36a Single comb, usually hammock or ball shaped and hung from one edge, some- times a linear string of cells (Madagas- car only). Sparse carton, silk cocoons providing most of the structure. Some- times all material removed from region of old cells and only reinforced comb margin remains; Africa to India ..... B ...... Belonogaster - Cell bottoms replaced by translucent se- cretion, not open to cocoons ...... 5 5(4). Combs single or multiple, elongate, and Fig. 5. Pedicellate combs. A. Horizontal, sus- hung from one edge. Mature combs pended from center. B. Inclined, suspended from may have single pedicel several centi- one edge. meters across. Carton sometimes trans- lucent, sometimes lacelike when new, composed of profuse secretion and yel- - Comb not deeply lobed, or, if so, then low or amber long fibers, rarely white with only one or few pedicels. Cell or brown; Iran to New Guinea, north to walls usually higher in older region of Korea and Japan (fig. 5) ...... nest and gradually shorter toward new- ...... Parapolybia er marginal regions ...... 8 - Combs single or multiple, shape variable, 8(7). Carton composed mostly of short vege- hung from one edge or supported cen- table chips, gray or brown in color. trally. Carton generally opaque (except Cells not larger than 3 mm wide, or, if for cell bottoms), usually composed of so, then with single pedicel more than short vegetable chips and profuse se- 20 mm long (except in USA); South cretion, inorganic matter, or, rarely, America, Carribean, southern USA, fiber. Pedicles narrow but possibly and Colorado, and occasionally farther long North along Great Basin (fig. 5) ..... numerous. Envelope sometimes pres- ...... Mischocyttarus ent; Africa, Asia, and Australia (fig. 5) - Carton composed mostly of long, woody ...... Ropalidia fibers, usually gray in color, cells larger 6(3). Pedicel absent. Comb single, horizontal, than 3 mm wide in Americas; cosmo- broadly attached to branch or limb; up- politan (fig. 5) ...... Polistes per surface composed of soft felt 3 mm 9(2). Nest broadly appressed to trunk or limb, or more thick, forming a hemispherical longest axis along line of attachment, or slightly peaked cap that conceals not on a leaf (figs. 6, 7); south cell bases and usually encompasses to central and Trinidad supporting branch; Central and South ...... 10 America ...... - Nest clearly pendant from leaves, branch- - Pedicel single or multiple. Combs single es or limbs. Shape variable (figs. 8, 9, or multiple, orientation variable, at- 10, 11, 13) ...... 18 tached to leaves, human domicile, or 10(9). Nest shaped like inverted flask, remod- other substrate. Without thick felt cap elled mostly downward. Envelope car- ...... 7 ton gray, usually streaked with other 7(6). Comb beneath Heliconia leaf, deeply lobed. colors. Entrance in lowermost region of Pedicels multiple and numerous. Cell envelope (fig. 6) ...... 11 walls uniform in height; Central and - Nest dome shaped, remodelled mostly South America ...... laterally or upward. Envelope not pa- ...... Protopolybia scutellaris pery, often very brittle, brown, gray, or 1998 WENZEL: NESTS OF SOCIAL WASPS 7

Fig. 7. Dome appressed to broad surface; en- Fig. 6. Inverted flask appressed to broad sur- trance as short upward spout; cut away to show face or leaf (rarely among twigs); entrance as sessile comb; and Metapolybia; Clypear- downward spout; cut away to show pedicelate ia and Asteloeca with entrance variable. combs; Parachartergus, Chartergellus, and Lei- pomeles. Nectarinella and Marimbonda with ses- sile comb. rough, and variable, sometimes brittle. Lines of construction strong. Entrance short collar; Central and tropical South tan, often spotted or mottled. Entrance America .... Chartergellus* variable (fig. 7) ...... 14 14(10). Envelope more than 4 cm deep, often 11(10). Entrance distinct, gently narrowed, corrugated with orderly ridges and fur- downward spout ...... 12 rows converging on a central longitu- - Entrance simple, not a spout ...... 13 dinal line. Carton coarse, with little ev- 12(11). Envelope bearing numerous fine carton pil- ident secretion. Entrance usually in up- lars 1-2 mm high and tipped with a per part of nest, separate from final gap sticky secretion. Carton color closely re- in construction, ringed by short collar

sembling substrate. Comb sessile upon ...... Synoeca substrate; and South - Envelope less than 4 cm deep, no corru- America .. .. Nectarinella* gation. Carton with profuse secretion (possibly Marimbonda*, and short vegetable chips, including but without pillars) small windows or layers of pure, - Envelope without sticky pillars, some- glossy, transparent secretion ..... 15 times corrugated with orderly ridges 15(14). Nest projecting beyond margin of narrow and furrows. Carton color not closely support ...... 16 resembling substrate. Combs pedicel- - Nest wrapping around narrow support, or late, oval to kidney shaped; Central and on broad support ...... 17 tropical South America ...... 16(15). Comb back showing cell bases, comb

...... Parachartergus sides showing cell contours. Carton 13(11). Envelope without carton pillars or floss, smooth, glossy. Lines of construction funnel shaped, very supple and soft. long, parallel, clear ..... Asteloeca * Lines of construction only weakly vis- - Comb back thick, obscuring cell bases, ible. Entrance simple hole. Combs ped- comb sides generally not showing cell icellate, oval to kidney shaped; Central contours. Carton rough, many fine par- and tropical South America ...... ticles. Lines of construction short, ir-

...... Parachartergus regular, or obscured ..... Clypearia* - Envelope with fine carton pillars or floss 17(15). Envelope shallow, may be flat, some- at envelope margin, especially on inner times sharply angled and resembling side of upper margin; envelope flat, eaves where side walls meet dome. En- 8 AMERICAN MUSEUM NOVITATES NO. 3224

Fig. 10. Chaotic nest; cut away to show mul- Fig. 8. Inverted flask pendant below leaves; tiple combs sessile on multiple envelopes; possi- entrance as downward, curved spout; cut away to bly multiple entrances; usually incorportating show pedicellate combs suspended from those nearby vegetation; Protopolybia, Brachygastra, above; Angiopolybia. and Ropalidia.

trance separate from final gap in con- - Envelope variably shaped, widest below midline, sometimes angled abruptly struction, upward spout or short collar. where rounded bottom meets sides, or, Cells 2.5 mm in diameter or less .... if ball shaped, then with rough and ir- ...... Metapolybia regular exterior. Entrance variable, but, - Envelope dome arching gradually. Entrance if spoutlike, then not at lowest region at final gap in construction, simple hole of the nest. Combs' position often or short collar ...... Clypearia* strongly marked on envelope exterior 18(9). New World south from California, Ari- (figs. 9, 10, 11) ...... 27 zona, Texas ...... 19 20(19). Nest supported primarily by single broad - Old World or North America to southern leaf, sometimes a few leaves, or a few USA ...... 43 twigs that terminate in leaves touching 19(18). Envelope simple sheet or shaped like in- or very close to nest ...... 21 verted flask, smoothly curved and gent- - Nest suspended from limb or twigs sev- ly narrowed below, widest above mid- eral centimeters from leaves, or at- line, or, if ball shaped, then with tached to a broad vertical surface such smooth and uniform exterior. Entrance lowest structure, often spoutlike. Combs' position not reflected on en- velope exterior (figs. 6, 8) ...... 20

Fig. 11. Pendant nest; widest below midline; Fig. 9. Inverted mushroom pendant below cut away to show combs sessile on previous en- leaves; central pedicel or distinct short stalk; cut velopes (envelope sometimes imbricate in upper away to show primary envelope from margin of region, not shown); tiers of combs strongly comb; Protopolybia and Charterginus (some Po- marked on exterior of side wall; Brachygastra, lybia similar). Polybia, Protonectarina, and Synoecoides. 1998 WENZEL: NESTS OF SOCIAL WASPS 9

25(24). Envelope a broad funnel. Entrance a sim- ple hole at bottom. Combs never hung one from another, often highly re- curved, overlapping but rarely fusing with others. Cocoons domed above cell walls by more than cell diameter ... .. Leipomeles - Envelope more bottle shaped. Entrance a long downward spout with flared open- ing directed sideways. Combs may be suspended from those above by central pedicel, neighboring combs may fuse. Fig. 12. Nest between leaves or in other cav- Cocoons domed barely, if at all, above ity; envelope as simple sheets across gaps (rarely from margin of comb); combs variable; Agelaia, cell walls ...... Angiopolybia 26(20). Envelope laminate, layers either supple, Protopolybia, and some Ropalidia. pale gray, and joining smoothly (com- pressa, difficilis), or with horizontal as a trunk; usually white or gray, rarely stripes of more brittle brown carton and yellow or brown, sometimes streaked long vertical lines where two layers with other colors ...... 26 meet (vespiceps). Combs round or 21(20). Envelope not domed more than 2 cm oval, secondary combs suspended from from leaf surface, color variable . . 22 those above by central pedicel. Cocoon - Envelope domed more than 2 cm from caps simple. Entrance simple at lowest leaf surface, usually amber or brown point . Pseudopolybia 24 ...... - Envelope mostly single layer or a few 22(21). Envelope color variable, usually yellow layers skewed and not closely parallel, or amber, rarely brown, green, or gray, usually gray or beige, with orderly sometimes nearly transparent with pro- ridges and furrows or sometimes fuse secretion, often with ridges or lumpy and irregular. Combs kidney streaks of color suggesting leaf veins. shaped and attached to substrate, rarely Entrance usually 5 mm wide or less. (Central America) secondary combs Sometimes with inconspicous, fine oval and suspended from those above. sticky carton pillars 1-2 mm high around leaf petiole. Combs often highly Cocoon caps often crossed with walls recurved, often overlapping but rarely of pulp. Entrance usually at least a fusing with others. Cocoons extending short spout ...... Parachartergus above cell mouths by more than cell 27(19). Primary comb suspended from leaf by a central pedicel or short distinct stalk diameter ...... Leipomeles (possibly Marimbonda*, and bearing from its margin a domed but with sessile combs) primary envelope not more than about - Envelope gray or white, uniform in tex- 2 cm deep, sometimes resembling an ture and color, or if streaked then not inverted mushroom (fig. 9) ...... 28 so as to suggest veins of a leaf. En- - Primary comb not suspended from a ped- trance usually more than 5 mm wide, icel or short distinct stalk, or, if so, then with or without spout. Combs planar to envelope much more than 2 cm high slightly recurved, sometimes fusing. and not resembling a mushroom (fig. Cocoons barely if at all domed above 11) ...... 30 cell walls ...... 23 28(27). Envelope same color as pulp of carton. 23(22). Fine carton pillars or floss at envelope Nest shape variable. Entrance lateral or margin, especially on inner side of up- ventrolateral. Secondary comb distin- per margin ...... Chartergellus* guishable from primary. Contours of - No such pillars or floss cells visible around much of external ...... Parachartergus smithii, colobopterus, amazonensis comb margin ...... 29 24(21). Envelope shaped like funnel or inverted - Envelope painted partly or completely bottle. Entrance at bottom of narrow white externally, remaining yellow, spout ...... 25 amber, or brown internally. Nest mush- - Envelope as simple sheet. Entrance mere- room shaped, sometimes star shaped ly a gap between envelope and sub- with six points, with single, stout, cen-

strate (fig. 12) . . Agelaia cajennensis tral pedicel. Entrance dorsolateral or 10 AMERICAN MUSEUM NOVITATES NO. 3224

dorsal through comb. Secondary comb 35(33). Mexico; if elsewhere, then with new en- continuous with primary, indistinguish- velope and comb added chaotically at able from it when mature. Contours of several places at the bottom and side cells rarely visible around external walls of nest (fig. 10). May have mul- margin of comb ...... Charterginus tiple entrances. Sometimes contains 29(28). Carton composed of long fiber, often very honey .. Brachygastra azteca, fine, usually pale. Pedicels often mul- mellifica, lecheguana tiple, sometimes very fine. Secondary - Not Mexico; if in Mexico, then with combs either continuous with primary combs added in orderly fashion one be- comb, pedicellate, and with single lat- low the other (fig.11) ...... 36 eral entrance, or secondary combs ses- 36(35). Cell walls easily peel away from backing sile upon primary envelope and nest sheet to yield an intact mat of hollow with separate lateral entrances for each hexagons. Entrance in center of lowest comb. Cells less than 2 mm in diameter envelope. Upper combs spherical, so ...... Protopolybia that small nests are ball shaped; lower - Carton composed of short dark chips. Sin- combs more planar (but curvature and gle stout stalk never resembling a fine spacing irregular), so large nests cylin- drical or sausage shaped. Seams mark- pedicel. Secondary combs sessile upon ing successive envelopes sinuous and primary envelope and nest with single irregular. Carton sometimes glossy. ventrolateral entrance and internal pas- Nest sometimes contains honey; south- sage connecting separate levels of ern Brazil, Paraguay, northern Argen- combs. Envelope sometimes bears long tina .Protonectarina extensions of carton at margins. Cells - Cell walls firmly joined to backing, can- over 2 mm in diameter ...... Polybia not be peeled off without destroying bistriata, platycephala, bicyttarella, the hexagons. Combs curved similarly. catillifex, incerta, scrobalis sometimes Seams marking successive envelopes occidentalis (Central America) straight or gently curving. Carton dull

30(27). Carton ...... composed of fine, soft, durable, ...... 37 white plant hairs resembling cotton, often 37(36). Entrance lateral or ventrolateral, some- with blots of dark material painted on times with a short lateral or upward surface. Entrance ventrally in center of spout, no honey ... Polybia ruficeps, bottom envelope; South America only sometimes occidentalis, Chartergus some other Polybia - Not as above .31 - Entrance dorsal. Nest sometimes with 31(30). Envelope abruptly angled where bottom honey .. Brachygastra moebiana meets side wall. Carton brittle, dark 38(32). Carton thin and fragile. Lines of con- brown, composed of plant chips; portions struction obvious in upper region. Of- of side walls and most of bottom enve- ten imbricate in upper region .... 39 lope painted white. Entrance lateral, - Carton dense and firm, sometimes reso- nating when struck with a finger or may be long slit . Brachygastra bilineolata, scutellaris, smithii stick. Lines of construction not obvi- ous. or - Not as above .32 Imbrication weak absent in up- per region ...... 40 32(31). Carton supple, resilient ...... 33 39(38). Entrance lateral, long vertical slit at least - Carton brittle, stiff ...... 38 three times as high as wide ...... 33(32). Carton yellow or amber ...... 34 ...... Brachygastra augusti - Carton gray ...... 35 - Entrance circular or oval ..... Polybia 34(33). Comb continuous spiral or concentric 40(38). Mud or clay ...... Polybia emaciata, spheres, forming a ball growing in all spinifex, singularis, furnaria directions. Cell diameter more than 2 - Vegetable fiber ...... 41 mm .... Agelaia areata, flavipennis 41(40). Bottom of nest convex, roughly radially - Comb many separate blocks, or, if contin- symmetrical. Entrance lateral .... 42 uous spiral, then expanding downward. - Bottom flat or slightly concave, distinctly Cell diameter 2 mm or less. Nest may asymmetrical, sometimes with one side be chaotic in structure (fig. 10) ...... drooping lower than other. Entrance .... Protopolybia sedula, acutiscutis, clearly ventral and opposite drooping eximia, weyrauchi lower side ...... Epipona 1998 WENZEL: NESTS OF SOCIAL WASPS I1I

sometimes sessile marginally on sub- strate or basally on envelope; southern A India to northern Australia, New Guin- ea, and Philippines ...... Ropalidia 46(44). Envelope often of independent arcs 10 cm wide to completely circumferential. Internal surface of envelope smoothed and almost continuous. Combs slightly B recurved and connected by broad flat- tened butresses. Cell walls slightly curved; Holarctic .... Dolichovespula - Envelope arcs usually less than 10 cm wide and overlapping. Internal surface of envelope usually rough and reveal- ing many pockets between separate im- bricate arcs. Combs flat and connected Fig. 13. A. Nest pendant (sometimes subter- by stout posts especially in lower levels ranean); upper region imbricate, lower region of nest, or by several buttresses not more laminate; pedicellate combs suspended from continuous across back of comb. Nests those above; entrance simple; abrupt switch to sometimes more than 1 m in diameter; queen cells. B. Sometimes with conical roof; Dol- northern Europe, introduced into USA, ichovespula, Vespula, Vespa, and Provwspa. Pseu- Australia, New Zealand, Chile, South dopolybia arboreal only, without queen cells, no conical roof. Africa, and Argentina ...... * (rarely nests in the open, usually in cavities) 47(43). Carton entirely of coarse, short chips. 42(41). Envelope extensively painted white ...... Synoecoides* Lines of construction strong. Primary - Envelope brown or gray, not painted comb supported by one or more flat- white ...... Polybia tened pedicels spread across comb 43(18). Carton flexible, supple (fig. 1) ..... 44 back. Face of combs sometimes cov- - Carton stiff, brittle ...... 47 ered over with carton to block cells. 44(43). Not ball or flask shaped, or, if so, then Nest sometimes with conical cap above new carton is yellow or tan and not point of support ...... Vespa striped. Envelope either of simple sep- - Carton at least partly of long amber plant arate sheets or many irregular, fused hairs. Lines of construction sometimes sheets ...... 45 not visible on upper envelope. Primary - Nest ball or flask shaped, sometimes with comb supported by parallel, rodlike a tall peaked cap above point of sup- pedicels from the substrate, or by a port. Envelope striped blue gray, lower broad butress formed by fusion of these region consisting of many complete or pedicels. Nest without conical cap. partly circumferential arcs. Combs hor- Back of primary comb sometimes izontal, hanging from those above (fig. glossy, sometimes with furrows be- 13) ...... 46 tween cell bases partly filled with car- 45(44). Envelope resembling bivalve shell, bilat- ton; Southeast Asia ...... Provespa erally symmetrical with sheets hanging 48(1). Nest of multiple combs (except in early on each side of combs. Entrance ven- stage) surrounded by an envelope ... tral, horizontal slit where envelope ...... 49 sheets fail to fuse. Combs vertical, - Single naked comb without an envelope hanging from one edge without a nar- ...... 55 row pedicel and rarely in contact with each other. Often hanging over water; 49(48). Envelope mostly of many interconnected central and western Africa ...... layers. Cell diameter varying within or ...... Polybioides between combs, never less than 3 mm. - Envelope not like a bivalve, variable. En- If envelope encloses combs, then en- trance usually a simple slit or hole, trance ventral or ventrolateral .... 50 sometimes multiple. Comb orientation - Envelope mostly a simple sheet, sometimes variable, sometimes growing spirally, a few layers, sometimes transparent and 12 AMERICAN MUSEUM NOVITATES NO. 3224

celophanelike. Cell diameter uniform 56(55). Comb sessile inside sheatfing leaf of a bro- throughout nest, 3 mm or less. Entrance meliad; South America ......

variable (fig. 12) ...... 52 ...... Metapolybiabromelicola 50(49). Envelope blue gray, supple; Holarctic, - Comb pedicellate ...... 57 historically south through montane 57(56). Older cell bottoms replaced with win- Mexico and ..... Vespula dows of clear secretion; Africa, India rufa group (Palearctic), V. germanica, to Australia, New Guinea, and Japan

pensylvanica, squamosa (New World), ...... Ropalidia V. penslyvanica (introduced) Hawaii, - Older cell bottoms retaining carton; Central V. germanica (introduced) Australia, and South America .... Mischocyttarus New Zealand, Chile, South Africa, 58(55). Carton and pedicels fibrous, brown, am- and Argentina ber, or yellow, with very little secre- - Envelope tan or brown, brittle ..... 51 tion. Combs often orderly parallel or 51(50). Envelope usually completely enclosing nest; concentrically arched, interconnected most imbricate patches closed completely by pedicels; Central and South Ameri- to the outside; often many pillars con- ca .Agelaia necting envelope to cavity ceiling. Older - Carton brown or gray. Usually a single cells mostly less than 5 mm in diameter; continuous comb; if multiple, then sep- Holarctic .... Vespula vulgaris group arate and not as above ...... 59 except germanica and pensylvanica 59(58). Pedicels fibrous, easily broken or de- - Envelope often broadly open below combs, tached; cells less than 3 mm in diam- or reduced to sheets closing access to eter; Central and tropical South Amer- cavity. Older cells mostly more than 6 ica ...... Protopolybia mm in diameter. Palearctic and Southeast - Pedicels resinous, tough, and well an- Asia, eastern USA ...... Vespa chored; cells more than 3 mm in di- 52(49). Older cell bottoms removed and without ameter in Central and South America, variable elsewhere; cosmopolitan .... carton ...... 53 - Older cell bottoms retaining carton, or ...... Polistes sessile ...... 54 53(52). Comb carton composed of short chips. ILLUSTRATIONS Older cell bottoms replaced with win- In addition to the photographs presented dows of clear secretion. Comb, if spiral here, previously published photographs and shaped, separate from envelope; southern detailed illustrations are cited for the reader India to northern Australia, New Guin- ea, and Philippines ...... Ropalidia to consult. Some of these are obscure, but - Comb carton composed of long plant fiber. they are included so that thorough students Older cell bottoms open, not replaced will not overlook certain noteworthy works. with secretion. Comb, if spiral shaped, Boxed arrow in figures indicates upward. All continuous with envelope; Southeast photographs are by the author unless other- Asia and Indonesia ..... Polybioides wise stated. Scale bar equals 25 mm unless 54(52). Envelope supported by walls of cavity otherwise noted. Nomenclature is cited par- and composed of yellow or amber long enthetically where modem nomenclature dif- fiber. Comb usually over 50 cm2 in area fers from that used by earlier authors. Read- when mature. Nest with or without ers who wish to consult a few texts that cover pedicel ...... Agelaia cajennensis the diversity well should examine Berland - Envelope supported by comb margin and and Grasse (1951), Jeanne (1975), Wenzel variable in color, or supported by walls (1991), or scattered photographs in the Ross of cavity and transparent with little or and Matthews (1991) volume. Many fine il- no vegetable fiber. Comb usually less in the classical than 50 cm2 in area when mature, ped- lustrations are available icellate ...... Protopolybia works by Saussure (1853-1858) and Mobius 55(48). Carton composed of short chips; some- (1856), though these are not commonly times glossy with profuse secretion, available and their nomenclature is archaic. sometimes with patches of pure, trans- Institutions housing certain specimens are parent secretion ...... 56 abbreviated on page 4, and localities are of- - Carton of long fiber; duller, glossy only fered for photographs from the field for near pedicel ...... 58 which no specimen was retained. A cladistic 1998 WENZEL: NESTS OF SOCIAL WASPS 13 matrix of architectural characters for the gen- tical pedicel. Carton supple, long fiber, felt- era of Polistinae is offered in Wenzel (1993). like, amber or yellow, rarely brown. Comb An interim cladogram of genera of social single, growing gradually off substrate at Vespidae is offered in figure 14. Ranges of margins, facing downward, sometimes re- the genera are approximate. sembling an inverted bowl or plate. Foun- dation and back of comb thickened second- DESCRIPTIONS OF NESTS arily by addition of felt, obscuring cell bases, often including supporting branch into comb Agelaia: Variable. Swarm-founded. In apex. Cell walls straight and parallel. No en- cavities or subterranean, sometimes exposed velope. Mexico to Argentina. See figures arboreal. Nests can grow to millions of cells 17A, 21B; Buysson (1906); Ducke (1910); (A. vicina). Pedicels multiple, fibrous, cell- Evans and West Eberhard (1970); Vecht marginal, vertical or horizontal, from cavity (1972); Richards (1978); and Schremmer walls; sometimes sessile initiation. Carton (1986). supple, long fiber, amber or yellow. Combs Asteloeca: Poorly known. I have seen one growing gradually at margins, may be sus- complete specimen and a fragment of a sec- pended from each other, fusing, double sided ond. Swarm-founded. Arboreal. Sessile ini- (A. lobipleura), planar, conical, or spiralling tiation on a narrow branch. Carton brittle, of outward. Cell walls straight, parallel to di- short chips, very glossy. Comb single, grow- vergent. Cavity species without envelope or ing off narrow substrate, expansion probably sometimes with a reduced envelope restrict- suddenly in blocks. Cell bases visible on ing access to cavity (A. cajennensis); ex- back of comb, cell walls straight and parallel. posed arboreal nests (A. areata and flavipen- Envelope a single sheet, from comb margin, nis) with single or multiple ball shaped en- abruptly angled where walls meet flattened velope supported by combs within or by ped- dome, about one cell length distant from icels arising from combs, not removed but comb face, initial sheet apparently covered rather built over as nest expands in all direc- with clear secretion, may be partly removed tions. Mexico to Argentina. See figures 5, 12, to allow contiguous expansion. Entrance hole 17C, D, 23C; Vesey-Fitzgerald (1938: fig. a short collar, apparently at center of dome, 12, as Gymnopolybia); Evans and West Eber- at final remaining gap in envelope construc- hard (1970: fig. 91, as Stelopolybia); Jeanne tion. Amazonian and , Brazil, (1970: as Stelopolybia, 1973); Richards and French Guyana. See figures 7, 19D, 24B (1978: figs. 28-30, as Stelopolybia); Wenzel and Garcia (1978, as Polybia trailii). (1991, 1992b); and Zucchi et al. (1995). Belonogaster: Independent-founded. Ar- Angiopolybia: Swarm-founded. Arboreal, boreal. Pedicel resinous, cell-central, vertical. often on leaves. Pedicel initially single, Carton supple, long fiber but sometimes very sometimes later multiple, fibrous, cell-mar- spare with cocoons or re-used silk providing ginal, vertical. Carton supple, long fiber, am- structure. Comb single, growing gradually at ber or yellow, rarely brown. Combs growing margins, usually hammock shaped due to di- gradually at margins, adjacent and fusing or vergent cell walls (sometimes vertical string suspended from each other, slightly recurved, of adjacent cells-Madagascar only). Cell not in contact with envelope. Cell walls bottoms removed but not replaced, cocoons straight, subparallel. Envelope from sub- slit open, no meconium in old cells, some- strate, single sheet, flask shaped, partly re- times old cells near top completely removed moved to allow downward expansion. En- and only reinforced comb margin supplies trance long downward spout with hole facing support in mature nests. No envelope. Africa, horizontally. to Brazil. See figures 8, Arabian Peninsula, and perhaps India. See 17B, 23F; Ducke (1910: fig. 12, as Stelopo- figure 16F; Iwata (1966); Marino Piccioli lybia); Vesey-Fitzgerald (1938: fig. 3, as Ste- and Pardi (1970, 1978); Pardi and Marino lopolybia); and Jeanne (1975). Piccioli (1970); Keeping and Crewe (1983); Apoica: Swarm-founded. Arboreal, usual- and Gadagkar (1991). ly on narrow branch. Sessile initiation, some- Brachygastra: Variable. Swarm-founded. times resembling fibrous, short, broad, ver- Arboreal. Sessile initiation. Carton either 14 AMERICAN MUSEUM NOVITATES NO. 3224

Stenogastrinae Vespa Provespa Dolichovespula Vespula Polistes I Mischocyttarus I _{ , Ropalidia I Parapolybia I Polybioides I ______Belonogaster I Apoica AgelaiaI Lu..Angiopolybia I Pseudopolybia Parachartergus Chartergellus Nectarinella Leipomeles Marimbonda CZ Synoeca I . Clypearia Metapolybia Asteloeca I Protopolybia Charterginus Chartergus Brachygastra Protonectarina Polybia Synoecoides Epipona Fig. 14. A preliminary cladogram for social wasps. Provided for basic reference only, this tree is modified from Wenzel and Carpenter (1994) and is not definitive. 1998 WENZEL: NESTS OF SOCIAL WASPS 15 brown, brittle, and of short chips or gray, Envelope from comb margin, a single sheet, supple, and of long fiber; without obvious dome, sometimes deep with acute ridges and secretion reinforcement. Primary comb built broad furrows or flattened with hollow points rapidly to full size, entirely sessile beneath protruding hexagonally. Lines of construc- broad surface or growing off narrow or up- tion obscured externally by pulp. Surface of right surface; may be planar or spherical. nest sometimes spotted or entirely covered Secondary combs strictly sessile on preced- with white substance externally. Envelope ing envelope, sometimes chaotic and starting margin partly removed to allow contiguous at several places in gray nests; orderly in expansion. Entrance hole simple, peripheral brown nests. Cell walls straight and subpar- or dorsal. Panama to Brazil. See figures 9, allel. Primary envelope from comb margin or 17F, 26F; Ducke (1905: pl. 1); Bequaert substrate, usually a deep dome. Secondary (1938); Evans and West Eberhard (1970); envelope built from previous envelope and Schremmer (1978). (brown nests), sometimes partly from sub- Chartergus: Swarm-founded. Arboreal, strate (gray nests). Brown nest side wall of- often high in tree. Sessile initiation, point of ten reinforced by imbrication or surface blots contact later encompassing branch. Carton of peripherally, but not on area destined to sup- fine, long white fiber resembling cotton, felt- port secondary combs. Brown nests may be like, supple when thin, later stiff and easily painted white on sides and below. Entrance withstanding strong pressure of fingers. simple, sometimes aligned to form an inter- Combs multiple, sessile on envelope, grow- nal passage between stories, sometimes pe- ing by rapid construction of several layers. ripheral, long, vertical slit (brown nests), or Cell walls straight and subparallel. Primary multiple and chaotic (gray nests only). May envelope from substrate or comb margin, contain honey. Arizona and Texas to north- secondary envelopes from preceding enve- ern Argentina. See figures 10, 11, 21D, 25F; lope, initially a single sheet but rapidly re- Buysson (1905); Ducke (1910: fig. 4, as Nec- inforced to become thick, heavy, and durable tarina); Richards (1978); and Starr (1991). dome, usually angled abruptly where side Chartergellus: Swarm-founded. Arboreal, walls meet bottom cell-bearing region of Pedicel fibrous, cell-marginal, horizontal to dome. Lines of construction obscured by margin of downward-facing comb. Carton of pulp, surface of nest sometimes spotted with long, gray fiber, either supple, smooth, and dark pulp externally. Entrance hole simple, uniform or brittle and irregular with promi- in ventral center of dome which may become nent lines of construction. Combs multiple, funnel shaped, little more than one head- growing gradually at margins, suspended width in diameter, aligned to form internal from substrate one below the other, not in passage between stories. to Brazil. contact with envelope. Cell walls straight, See figure 26C; Ducke (1910); Berland and subparallel. Envelope from substrate, a single Grasse (1951); Evans and West Eberhard sheet, flask shaped, sometimes much flat- (1970); Wilson (1971); Spradbery (1973); tened when substrate is broad, partly re- and Richards (1978). moved to allow downward expansion. Fine Clypearia (= Occipitalia): Poorly known. paper pillars or floss between substrate and Swarm-founded. Arboreal. Sessile initiation envelope margin, particularly on inside of on trunk or branch. Carton brittle, of short upper region. Entrance short downward chips. Comb single, entirely sessile on broad spout. to Brazil. See figures 6, surface or growing off narrow substrate; in 18C, 23A, B and Richards (1978). latter case back of comb reinforced with pulp Charterginus: Swarm-founded. Arboreal, obscuring cell bases. Cell walls straight and often on broad leaf. Sessile initiation, resem- parallel or subparallel. Envelope single sheet, bling a stout pedicel. Carton of fine, long fi- from substrate or comb margin, dome ber, feltlike but brittle. Comb single, resem- shaped, sometimes less than 1 cm distant bling an inverted mushroom, only a few ini- from comb face. Initial sheet apparently cov- tial cells reaching substrate, nest growing by ered with clear secretion (except in C. sul- sudden blocks contiguous with existing cata) and distinctive, nonpapery particles ap- comb. Cell walls straight and subparallel. plied to the secretion separately. Envelope 16 AMERICAN MUSEUM NOVITATES NO. 3224 margin may be partly removed to allow con- of construction obscured by pulp added ex- tiguous expansion. Entrance hole simple or a ternally. Cross section of nest sometimes in- short collar, corresponding to final remaining creasing for several layers then gradually de- gap in envelope construction at top, periph- creasing. Entrance hole simple, very near ery, or center of dome. C. sulcata often sit- shorter side wall of dome, aligned to form an uated near arboreal nests, such as those internal passage between stories. Mexico to of Azteca. Panama to southern Brazil, and Brazil. See figures 20D, 26E; Vesey-Fitzger- Bolivia. See figures 7, 24C, D, F; Ducke ald (1938: fig. 7, as Tatua); Berland and (1910); Araujo (1955: fig. 8); Evans and Grasse (1951: fig. 1026); Wilson (1971); West Eberhard (1970: fig. 92); and Jeanne Jeanne (1975); Richards (1978); and Wenzel (1979). See Carpenter et al. (1996) for syn- (1991, 1992b). onymy. Leipomeles: Swarm-founded. Arboreal, Dolichovespula: Independent-founded. beneath single or among few leaves, some- Arboreal, occasionally subterranean (western times with leaf petiole bearing pillars 1-2 plains of USA and Canada). Hanging sheet mm long tipped with sticky secretion. Pedi- of fibrous paper from substrate precedes ped- cel fibrous, cell-marginal, vertical or oblique icel. Pedicel single, twisted, and reinforced from leaf veins, single or multiple. Carton with secretion in new nests, fibrous in mature supple, long fiber, thin and often translucent. nests, cell-marginal, vertical. Carton supple, Combs adjacent, growing gradually at mar- gray. Combs recurved, suspended from each gins, rarely fusing, planar to highly recurved, other by buttresses, cell walls curved, diver- not in contact with envelope. Cell walls gent, with abrupt switch to large queen cells, straight, subparallel to divergent. Cocoons not in contact with envelope. Envelope from domed 1-2 mm above cell walls. Envelope hanging sheet or substrate, mostly laminar, from leaf, single sheet, flattened dome to fun- sometimes with imbricate conical roof, re- nel shape, may be any color including green, moved and smoothed internally as nest sometimes with ridges or streaks mimicking grows. Lines of construction obvious. En- leaf veination, sometimes with a central, im- trance hole long downward spout in young pressed, longitudinal furrow composed of nests, simple in mature nests. Holarctic, Asia tight arches of pulp. Envelope margin partly excluding India and Indochina. See figures removed to allow downward or peripheral 13, 26A; Hungerford (1930); Duncan (1939); expansion. Entrance toward tip of leaf, some- Berland and Grasse (1951); Spradbery times with downward spout. Brazil to Costa (1973); Akre et al. (1980); Edwards (1980); Rica. See figures 6, 18E, 23E, 24A; Ducke Mastuura and Yamane (1984); and Greene (1910); Richards (1978); Schremmer (1983, (1991). 1986); and Wenzel and Carpenter (1994). Epipona: Swarm-founded. Arboreal, usu- Marimbonda: Poorly known. I have seen ally high in tree. Sessile initiation, point of only two specimens that I believe are of this contact encompassing branch. Carton of genus. Swarm-founded. Arboreal on vertical short chips and coarse matter, gray or brown, branch or leaf. Sessile intiation. Carton sup- brittle, very dense, without obvious secre- ple, long fiber. Comb probably expanding tion, easily withstanding pressure and reso- gradually at margins and mostly downward, nating if struck with a finger. Combs multi- restricted to substrate. Envelope flattened, ple, sessile on envelope, growing by rapid rather similar to some Leipomeles but with- construction of several layers. Cell walls out leaf veination, sometimes with central, straight and subparallel. Primary envelope longitudinal crease composed of tight arches from substrate or comb margin. Secondary of pulp. Brazil, probably Amazonian Peru, envelopes from preceding envelope, single , and Colombia. See figures 6 (but sheet but rapidly reinforced to become thick, sessile), 23D. usually abruptly angled where side walls Metapolybia: Swarm-founded. Arboreal. meet bottom cell-bearing region of dome, pe- Sessile initiation on broad surface. Carton of ripheral walls drooping lower on side oppo- short chips, supple when new, brittle later. site entrance than elsewhere and sometimes Single comb expanded suddenly in blocks on unsuitable for comb in lower stories. Lines substrate adjacent to earlier comb and on any 199818WENZEL: NESTS OF SOCIAL WASPS 17 side of it when nest is on horizontal surface, Carton supple, long fiber. Combs adjacent growing mostly upward when on vertical (sometimes fusing), or one below the other, surface, growing around a curved substrate or suspended from each other (Central Amer- rather than off it. Cell walls straight and usu- ica only), growing gradually at margins. Cell ally parallel (except where nest grows around walls straight, subparallel, sometimes pulp curve). Envelope (lacking in M. bromelicola) aligned with neighboring walls extending from substrate or from fully elongated cell across pupal caps. Envelope from substrate, walls (M. docilis), single sheet, not papery, single sheet, flat dome or flask shaped, some- generous secretion often forming clear win- times contacting combs. Envelope margin dows. Short envelope walls perpendicular to partly removed to allow expansion in any di- substrate, straight or curving outward, eaves rection but primarily downward, sometimes often forming an acute angle with the flat- with regular corrugations converging on a tened, rough, and uneven central region of weak keel. Entrance usually a downward dome. Envelope margin partly removed and spout. Mexico to Argentina. See figures 6, expanded to cover contiguous combs togeth- 17E, 18A, B; Ducke (1910: fig. 16); Vesey- er. Entrance spout peripheral, curved upward, Fitzgerald (1938); Berland and Grasse (1951: built separately from final remaining gap in fig. 1025); Spradbery (1973); Richards dnvelope construction. Mexico to Paraguay. (1978); Schremmer (1978, 1986); Strass- See figures 7, 19A, 24E; Vesey-Fitzgerald mann et al. (1990); and Wenzel (1992b). (1938); Richards (1978); and Schremmer Parapolybia: Independent-founded. Arbo- (1984). real. Pedicel resinous, cell-central, vertical to Mischocyttarus: Independent-founded. Ar- oblique, sometimes absent in secondary boreal, sometimes in cavities. Pedicel resin- combs, may be several centimeters wide in ous, cell-marginal, vertical or oblique. Car- mature combs. Carton supple, long fiber with ton supple or brittle, short chips (except M. conspicuous glossy secretion, usually yellow carbonarius in eastern Brazil), sometimes or light brown, sometimes very pale, some- with conspicuous glossy secretion. Comb times translucent or lacelike when new. may be any shape, may be multiple. No en- Combs long and slender, hung from one velope. Southwestern Canada to northern Ar- edge, may be multiple and closely packed. gentina. See figures 5, 15D, E, F, 16A, B, C, Cell bottoms and base of cocoons removed 22A; Ducke (1914); Richards (1945); ZikaLn and replaced with transparent secretion, no (1949); Jeanne (1975); Herre et al. (1986); meconium in old cells. No envelope. Iran to Carpenter and Wenzel (1988); Gadagkar New Guinea, north to Korea and Japan. See (1991); and Starr (1991). figures 5, 22C; Vecht (1966); Sekijima et al. Nectarinella: Swarm-founded. Arboreal (1980); Yamane (1984); and Gadagkar on broad vertical or slanting surface. Sessile (1991). initiation. Carton supple, long, gray fiber. Polistes: Independent-founded. Arboreal, Single oval comb expanding gradually at sometimes in cavities. Pedicel resinous, cell- margins but primarily downward. Envelope marginal, vertical or oblique. Carton supple, from substrate, flattened dome with irregular of long fiber (except P. stenopus in South small ridges and furrows, externally bearing Pacific or Neotropical P. bicolor), usually paper pillars 1-2 mm long and tipped with glossy only near pedicel but sometimes ex- sticky secretion, often blotted with lichens tensively over comb back. Comb planar or and fragments of bark as camouflage, some- recurved, rarely as a long string of cells (P. times many fine paper pillars or floss be- goeldi, South America and P. stenopus, tween substrate and envelope margin or South Pacific); single, rarely multiple. Indo- downward entrance spout. Costa Rica to Co- pacific species sometimes with two brood per lombia, also Mato Grosso, Brazil. See figures cell in tandem. No envelope. Cosmopolitan. 6 (but sessile), 18D and Schremmer (1977, See figures 5, 15C; Rau (1928); Yoshikawa 1986). (1964); Evans and West Eberhard (1970); Parachartergus: Variable. Swarm-found- Yamane and Okazawa (1977); Schremmer ed. Arboreal. Pedicel fibrous, cell-marginal, (1986); and Starr (1991). vertical to horizontal, sometimes multiple. Polybia: Variable. Swarm-founded. Arbo- 18 AMERICAN MUSEUM NOVITATES NO. 3224 real. Sessile initiation or suspended by but- low or amber, sometimes brown. Combs tressed sheet from substrate (subgenus Tri- multiple, long, hung from one edge, and par- chinothorax). Carton brittle, of short chips or allel, facing outward from orginal back-to- mud (P. emaciata, furnaria, singularis, and back pair and growing gradually at the mar- spinifex), and without obvious secretion re- gins (Africa), or single, horizontal, and as a inforcement. Primary comb entirely sessile downward spiral (Asia). Cell walls straight and built rapidly to full size beneath broad and subparallel, cell bottoms and base of co- and horizontal surface, or growing gradually coons removed and not replaced, no meco- at margins off narrow or upright surface. nium in old cells. Envelope of separate, in- Secondary combs strictly sessile on preced- completely fused sheets perpendicular to ing envelope, often growing by rapid con- combs, open below, resembling bivalve shell struction of several layers. Cell walls straight (Africa), or formed by marginal fusion of and subparallel. Primary envelope from spiral comb with subtle imbricate reinforce- comb margin or substrate, usually a deep ment above (Asia). West and Central Africa dome, rarely resembling an inverted mush- and Southeast Asia. See figures 21A, B, 22F; room (few one-level nests of P. bicyttarella, Bequaert (1918: pl. 19); Pagden (1958); Wal- bistriata, catillifex, incerta, platycephala, recht (1963); Vecht (1966); Richards (1969); and scrobalis), sometimes with projections and Turillazzi and Francescato (1994). extending radially outward (particularly P. Protonectarina: Swarm-founded. Arbore- catillifex). Secondary envelopes built from al. Sessile initiation. Carton gray, supple, of previous envelope, each envelope as a single long fiber, slightly glossy. Primary comb sheet, built to full size rapidly, side wall of- built rapidly to full size, spherical. Secondary ten quickly reinforced by imbrication or sur- combs strictly sessile on preceding envelope face blots peripherally but not on area des- and successively less curved, but curvature tined to support secondary combs. Upper re- and spacing of combs irregular. Nest devel- gion of nest sometimes slightly glossy on ex- oping from ball to sausage shaped, often terior (subgenus Trichinothorax). Entrance growing by rapid orderly construction of sev- simple, peripheral or in lower region of eral layers. Cell walls straight and subparal- dome, aligned to form internal passage be- lel. Comb backing (internalized envelope) is tween stories, modified in mud nesters into a easily peeled away from hexagonal pattern of small ventral hole (P. furnaria), projecting cell walls leaving hollow cell walls behind. snout (P. emaciata), or vertical slit continued Primary envelope from substrate, usually a across all combs (P. singularis, and spinifex). deep dome. Secondary envelope built from Polybia richardsi (Cooper, 1993) builds an previous envelope or partly from substrate, initial nest of thick walls (AMNH) that is seams marking successive envelopes sinuous expanded to peculiar chaotic shape with mul- and irregular. Entrance simple, in ventral sur- tiple entrances and extraneous flanges or face of nest, aligned to form internal passage blisters of carton. The interior may be sec- between stories. Nest sometimes contains ondarily smoothed with carton, unique honey. Southern Brazil, Paraguay, and north- among Polybia. Arizona and Texas to Ar- eastern Argentina. See figures 11, 21E, 25D; gentina. See figures 9, 11, 20A, B, C, E, F, Shima et al. (1996). 21F, 25E; Ducke (1910); Vesey-Fitzgerald Protopolybia (= Pseudochartergus): (1938); Berland and Grasse (1951); Windsor Highly variable, including within species. (1972); Spradbery (1973); Jeanne (1975, Swarm-founded. Arboreal, frequently on 1986); Garcia (1978); Richards (1978); Ho- leaves or in cavities. Pedicel fibrous, cell- fling and Machado (1985); Schremmer marginal, vertical, multiple, or sessile initia- (1986); and Cooper (1993). tion on surface (P. acutiscutis, and sedula). Polybioides: Asian species poorly known. Carton supple to brittle, long fiber. Comb and Swarm-founded. Arboreal or in cavities cell walls may be constructed rapidly to large (Asia). Sessile initiation, but supporting mar- or final size. Cell walls straight and subpar- gin of comb may become reinforced with allel. Secondary combs may be built rapidly dark secretion to resemble a fat pedicel (Af- to large or final size from margin of primary rica). Carton supple, long fiber, usually yel- comb and coplanar with it, or sessile on the 1998 WENZEL: NESTS OF SOCIAL WASPS 19 primary envelope, sometimes spiraling 13-15, as Parachartergus; 1914); and Rich- downward (P. acutiscutis, and sedula). En- ards (1978). velope, when arising from margin of comb Ropalida: Highly variable. Independent- or cell walls, then partly removed and ex- or swarm-founded. Arboreal or in cavities. panded to cover contiguous, pedicelate Pedicel resinous, cell-central, vertical to hor- combs together; when from primary enve- izontal, or initiation sessile. Carton supple or lope or substrate, then covering successive, brittle, usually short chips with conspicuous sessile combs separately. Envelope as a sin- glossy secretion. Comb may be any shape, gle sheet, dome, or absent; or as a dry, clear may be multiple; large single comb may be sheet of secretion without wood pulp, arising cut to produce separate daughter combs. from substrate and restricting access to the Cells bottoms and base of cocoons removed cavity (usually folded leaves). Envelope not and replaced with transparent secretion, no reinforced by imbrication or lamination, rare- meconium in old cells. If envelope present, ly thickened by blots of pulp. Entrance hole then as simple sheets, often joined chaotical- simple, usually peripheral, often multiple ly or closing gap between leaves or nearby providing separate access to each layer, not objects, may be pure secretion without pulp. aligned to form internal passage between sto- Entrances simple, may be multiple, but do ries. Honduras to Brazil. See figures 9, 10, not align to form internal passage. Africa, In- 12, 19E, F, 25A, B, C; Ducke (1910: figs. 2, dia to Australia and New Guinea, and north 3; 1914, as Pseudochartergus); Jeanne to Japan. See figures 5, 10, 12, 16D, E, 21C, (1970, as Pseudochartergus); and Schrem- 22D, E; Carl (1934); Vecht (1962); Yoshi- mer (1984, as Pseudochartergus). For syn- kawa (1964); Hook and Evans (1982); Ko- onymy, see Carpenter and Wenzel (1989). jima (1982, 1984, 1988); Ito (1986); Kojima Provespa: Swarm-founded. Arboreal or in and Jeanne (1986); Ito et al. (1988); Sprad- tree cavities. Pedicel fibrous, cell-marginal, bery and Kojima (1989); Maschwitz et al. vertical, multiple but later fused to form sin- (1990); Gadagkar (1991); Starr (1991); Ya- gle broad pedicel. Carton supple to brittle, of mane et al. (1991); and Makino et al. (1994). short chips to long fiber. Combs suspended Synoeca: Swarm-founded. Arboreal, usu- from each other, usually with one central ally on broad slanting surface, often near ar- pedicel between combs, growing gradually at boreal ant nest. Sessile initiation. Carton brit- margins, queen cells mixed with tle, short chips, often coarse. Each comb ex- worker panding gradually at margins but mostly cells, not in contact with envelope. Back of downward. Secondary combs built contigu- comb sometimes with pulp placed in furrows ous with primary but mostly higher on sub- between cell bases. Cell walls straight, sub- strate, may be built partly on envelope when parallel. Envelope from substrate or base of substrate is narrow. Cell walls straight, par- pedicel, laminate to imbricate. Entrance hole allel to divergent. Envelope from substrate or simple. Southeast Asia. See figures 13, 15B; margin of comb, each a single sheet, deep Matsuura and Yamane (1984: figs 7.3, 7.4); dome sometimes with regular transverse cor- Maschwitz and Hanel (1988); and Matsuura rugations, sometimes chaotically supported (1991). from previous envelope. Entrance as a short Pseudopolybia: Swarm-founded. Arboreal. upward collar near top of envelope. Lower, Pedicel fibrous, cell-marginal, vertical, sin- older levels of nest may be abandonned gle, sometimes becoming broad. Carton sup- while upper levels are active. Mexico to Ar- ple. Combs stacked downward, occasionally gentina. See figures 7, 19B; Buysson (1906); multipedicellate, growing gradually at mar- Ducke (1910); Vesey-Fitzgerald (1938); gins, slightly recurved, not fused with en- Schremmer (1973, 1986); Jeanne (1975); velope. Cell walls straight. Envelope from Richards (1978); and Castell6n (1980). substrate, laminar or loosely imbricate, Synoecoides: Poorly known. I have seen spherical or tapering below, removed inter- one specimen. Swarm-founded. Arboreal. nally and smoothed as nest grows. Entrance Sessile initiation, point of contact encom- hole simple at lowest point. Nicaragua to passing branch. Carton brittle, of short chips, Brazil. See figure 18F; Ducke (1910: figs. without obvious secretion. Primary comb 20 AMERICAN MUSEUM NOVITATES NO. 3224 sessile on substrate, growing off it gradually ripherally or in lower combs. Cell walls at margins. Secondary comb sessile on pre- straight to slightly curved, subparallel. En- vious envelope. Cell walls straight and sub- velope from hanging sheet or from previous parallel. Primary envelope from substrate. sheet of envelope, imbricate in upper region, Secondary envelopes from preceding enve- but more laminar below, highly imbricate lope, single sheet, lines of construction ob- when nest is mature. Envelope removed in- scured by long fiber pulp added externally, ternally as nest grows and internal surface sometimes pale or white. Colombia, Ecuador, left rough, revealing many pockets between Peru, Bolivia, and Brazil. See figures 11, separate imbricate arcs, often very brittle and 26D and Araujo (1944). tan in color. Entrance hole simple. Vespula Vespa: Independent-founded. Arboreal, in rufa group. As V. vulgaris but carton always cavities, or subterranean. Pedicel fibrous, supple and gray. Usually with butressed cell-marginal, vertical, sometimes multiple. sheets between combs. Comb planar. Cell Carton brittle, of short chips. Cell walls walls curved, divergent. Envelope laminar or straight or slightly curved, subparallel. Sec- loosely interconnected, imbrication vague or ondary combs with single or multiple pedi- absent. Entrance hole simple. Holarctic, with cels, stacked downward. Cell size increasing some extension into southern Asia and Cen- gradually to queen size, some cells may have tral America, introduced into Australia, New open ends closed with paper. Envelope from Zealand, Chile, South Africa, and Hawaii. substrate or base of pedicel, laminate to im- See figures 13, 26B; Duncan (1939); Sprad- bricate, removed internally as nest grows and bery (1973); Edwards (1980); Akre et al. internal surface left rough, revealing many (1981); Yamane et al. (1981); Matsuura and pockets between separate imbricate arcs. En- Yamane (1984); Schremmer (1986); and velope sometimes reduced to restrict natural Vasic (1986, as Vespa). access to a cavity, or absent in cavity nesters. Nest may develop imbricate conical roof in ACKNOWLEDGMENTS exposed sites. Entrance often as long down- I thank C. D. Michener (KUSM), who en- ward spout when nest is young, simple hole couraged me to undertake this project and in mature nests. Southeast Asia, Palearctic, helped me through an early effort. J. M. Car- and introduced to eastern USA. See figures penter (AMNH) lent considerable financial 13, 15A; Vecht (1957); Matsuura (1971, and scientific aid over several years. W. D. 1984, 1991); Edwards (1980); Seeley and Hamilton (Oxford University) and R. L. Seeley (1980); Starr and Jacobson (1990); Jeanne (University of Wisconsin, Madison) and Starr (1991). unselfishly offered their field notes and pho- Vespula (= Paravespula): Vespula vulgar- tographs. C. K. Starr (University of the West is group. Independent-founded. In cavities or Indies, Trinidad) and J. M. Carpenter im- subterranean. Hanging sheet of fibrous paper proved the manuscript. J. Kojima (Ibaraki from substrate precedes pedicel. Pedicel fi- University, Japan) reviewed my characteriza- brous, cell-marginal, vertical, twisted, single. tion of Asian taxa. This work was supported Multiple pedicels occuring between second- in part by NSF grants BSR-8817608 and ary combs, peripheral pedicels from combs BSR-9006102 to J. M. Carpenter, by a Poste to envelope, sometimes between envelope d'Acceuil of the MNHN, Paris, and the Re- and cavity wall, butresses sometimes built search Foundation of The Ohio State Uni- secondarily above primary comb. Carton versity. Photographic reproduction was made supple to brittle. Combs stacked downward, possible by the Edward Hoffman Fund, abruptly switching to large queen cells pe- AMNH.

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Spradbery, J. P. Walrecht, B. J. J. R. 1973. Wasps: an account of the biology and 1963. Bijdrage tot de kennis van de nestbouw natural history of solitary and social van Polybioides naar aanleiding van het wasps. Seattle: Univ. Washington in de Rijkslandbouwhogeschool te Press. Gent aanwezige nest. Biol. Jaarb. Do- Spradbery, J. P., and J. Kojima donaea. 31: 244-258. 1989. Nest descriptions and colony popula- Wenzel, J. W. tion of eleven species of Ropalidia 1987. Ropalidia formosa, a nearly solitary (Hymenoptera, Vespidae) in New paper wasp from Madagascar (Hyme- Guinea. Japanese J. Entomol. 57: 632- noptera: Vespidae). J. Kansas Entomol. 653. Soc. 60: 549-556. Starr, C. K. 1991. Evolution of nest architecture. In K. G. 1991. The nest as the locus of social life. In Ross and R. W. Matthews (eds.), The K. G. Ross and R. W. Matthews (eds.), social biology of wasps, pp. 480-519. The social biology of wasps, pp. 520- Ithaca, NY: Cornell Univ. Press. 539. Ithaca, NY: Cornell Univ. Press. 1992a. Behavioral homology and phylogeny. Starr, C. K., and R. S. Jacobson Annu. Rev. Ecol. Syst. 23: 361-381. 1990. Nest structure in Philippine hornets 1992b. Les nids de guepes sociales d'interet (Hymenoptera, Vespidae, Vespa spp.). historique conserves au MNHN, a Paris Japanese J. Entomol. 58: 125-143. [Hymenoptera, Vespidae]. Rev. Fr. En- Strassmann, J. E., C. R. Hughes, and D. C. Quell- tomol., n. ser., 14: 1-11. [English trans- er lation available from the author]. 1990. Colony defense in the social wasp Par- 1993. Application of the biogenetic law to be- achartergus colobopterus. Biotropica havioral ontogeny: a test using nest ar- 22: 324-327. chitecture in paper wasps. J. Evol. Biol. Turillazzi, S., and E. Francescato 6: 229-247. 1994. Notes on the nest architecture and Wenzel, J. W., and J. M. Carpenter brood rearing of Polybioides raphigas- 1994. Comparing methods, adaptive traits and tra (Vespidae, Polistinae). Rend. Fis. tests of adaptation. In P. Eggleton and Acc. Lincei, ser. 9, 5: 367-375. R. Vane-Wright, (eds.), Phylogenetics Vasic, Z. and ecology: 79-101. London: Aca- 1968. Biological investigations on the Ger- demic Press. man wasp (Vespa germanica F.). Bull. Wilson, E. 0. Nat. Hist. Mus. Belgrade, ser. B, 23: 1971. The insect societies. Cambridge, Mass: 211-224. Belknap Press. Vecht, J. v. d. Windsor, D. M. 1957. The Vespinae of the Indo-Malayan and 1972. Nesting association between two neo- Papuan areas (Hymenoptera, Vespi- tropical polybiine wasps (Hymenop- dae). Zool. Verh. Leiden. 34: 1-83. tera, Vespidae). Biotropica 4: 1-3. 1962. The Indo-Australian specied of the ge- Yamane, S. nus Ropalidia (Icaria) (Hymenoptera, 1984. Nest architecture of two oriental paper Vespidae) (second part). Zool. Verhand. wasps, Parapolybia varia and P. no- Rijksmus. Nat. Hist. Leiden. 57: 1-71. dosa, with notes on its adaptive signif- 1966. The east Asiatic and Indo-Australian icance (Vespidae, Polistinae). Zool. species of Polybioides Buysson and Jahrb. Syst. 111: 119-141. Parapolybia Saussure (Hym., Vespi- Yamane, S., Y. Ito, and J. P. Spradbery dae). Zool. Verhand. (Leiden) 82: 1- 1991. Comb cutting in Ropalidia plebeiana: a 42. new process of colony fission in social 1972. The social wasps (Vespidae) collected wasps (Hymenoptera: Vespidae). Insec- in French Guyana by the mission du tes Sociaux 38: 105-110. Museum national d'Histoire naturelle Yamane, S., S. Makino, and R. P. MacFarlane with notes on the genus Apoica. Ann. 1981. Embryo nest architecture in three Ves- Soc. Entomol. Fr., n.ser. 8: 735-743. pula species (Hymenoptera, Vespidae). Vesey-Fitzgerald, D. Kontyuf 49: 491-497. 1938. Social wasps (Hym. Vespidae) from Yamane, S., and T. Okazawa Trinidad, with a note on the genus Try- 1977. Some biological observations on a pa- poxyolon Latreille. Trans. R. Entomol. per wasp Polistes (Megapolistes) tepi- Soc. London 87: 181-191. dus malayanus Cameron (Hymenopter, 1998 WENZEL: NESTS OF SOCIAL WASPS 25

Vespidae) in New Guinea. Kontyfu 45: criqo de 82 especies novas. Bol. Parq. 283-299. Nac. ItatiMia (Rio de Janeiro) 1: 1-251. Yoshikawa, K. Zucchi, R., S. F Sakagami, F B. Noll, M. R. Me- 1964. Predatory hunting wasps as the natural chi, S. Mateus, M. V. Baio, and S. Shi- enemies of insect pests in Thailand. In ma T. Kira and T. Umesao (eds.): Nature and life in Southeast Asia. Kyoto: Fau- 1995. Agelaia vicina, a swarm-founding pol- na and Flora Research Society. istine with the largest colony size Zikin, J. F among wasps and bees (Hymenoptera: 1949. 0 ge-nero Mischocyttarus Saussure Vespidae). J. New York Entomol. Soc. (Hymenoptera, Vespidae), com a des- 103: 129-137.

Appendix 1 SPECIES FOR WHICH NESTS HAVE BEEN EXAMINED Genera are presented in phylogenetic order (see fig. 14). Nomenclature for Polistes follows Carpenter (1996), but Richards (1978) subgenera are listed in parentheses under Aphanilopterus, with which Car- penter synonymized them. Following Kojima and Carpenter (1996) for Ropalidia, no subgenera are listed. A list of institutions where these specimens may be found is available at http:Hiris.biosci.ohio- state.edu/people/jww/specieslist.html.

VESPINAE cavapyta stenopus Genus Vespa cinctus tenebricosus affinis comanchus tepidus analis crinitus Subgenus Polistella cincta cubensis (including Stenopolistes) crabro dominica albocalcaratus mongolica erythrocephalus aquilinus parallela exclamans bernardii tropica goeldii defectivus velutina infuscatus fastidiosus instabilis flavobilineatus Genus Provespa versicolor anomala haugi (Epicnemius) humilis Genus Dolichovespula bicolor madecassus arenaria billardieri mandarinus maculata cinerascens mandiburens media pacificus manillensis norvegica testaceicolor nigrifrons stotheri sylvestris (Fuscopolistes) sagittarius Genus Vespula apachus saussurei germanica carolina sikorae maculifrons dorsalis smithii pensylvanica fuscatus smithii neavei rufa metricus stigma squamosa poeyi takasagonus vulgaris (Onerarius) carnifex Subgenus Polistes POLISTINAE (Palisotius) (including Sulcopolistes) major africanus Genus Polistes associus Subgenus Aphanilopterus Subgenus Gyrostoma badius annularis (including Nygmopolistes) dominulus apicalis diabolicus gallicus aterrimus jadwigae marginalis buyssoni olivaceus nimpha canadensis minor tenellus 26 AMERICAN MUSEUM NOVITATES NO. 3224

Genus Mischocyttarus Subgenus Monacanthocnemis granulata Subgenus Clypeopolybia buyssoni gregaria carbonarius punctatus honkongensis piger vaqueroi horni Subgenus Monogynoecus ignobilis Subgenus Haplometrobius alienus jacobsoni alboniger fraudulentus lacustris artifex insolitus latebalteata cerberus metoecus maculiventris cooperi moralesi madecassa decimus pelor malayana dimorphus marginata iheringi Subgenus Phi mathematica illusorius alfleni melania mirificus araujoi minor nigropygialis barbatulus morosa oecothrix barbatus nitidula ornatus basimacula nobilis prominulus cassununga opifex silvicola cubensis phalansterica stenoecus flavicornis picta surinamensis flavoniger plebeiana sylvestris hirsutus prasina tenius imeldai ranavali undulatus lilae revolutionalis vredeni lules romandi weyrauchi mexicanus rufoplagiata xanthocerus oreophilus schultessi ypiranguensis pallidipectus shestakovi Subgenus Kappa paraguaensis stigma atramentaris phthisicus sumatrae bertoni tarmensis timida comunalis wagneri tomentosa injuncundus Genus Ropalidia turneri latior ambigua variabilis latissimus anarchica variegata metathoracicus antennata vitripennis pseudomimeticus aristocratica xanthura shunkei artifex Genus Parapolybia socialis bambusae nodosa tolensis bicincta orientalis bilineata varia Subgenus Megacanthopus brazzai collarellus cabeti Genus Polybioides collaris capensis melainus melanopygus carinata raphigastra parallelogrammus cincta tabidus saturatus constitutionalis Genus Belonogaster superus cyathiformis abyssinica Subgenus Mischocyttarus distigma brachycera cinerascens fasciata brevipetiolata drewseni ferruginea brunnea gynandormorphus flavopicta clypeata labiatus flavoviridis discifera matograssoensis formosa dubia melananus galimatia eumenoides rotundicollis guttatipennis filiventris tomentosus grandidieri fulvipennis 1998 WENZEL: NESTS OF SOCIAL WASPS 27

grisea cyanea Genus Polybia guerini septentrionalis Subgenus Alpha juncea surinama bifasciata rufipennis virginea Subgenus Apopolybia vassae Genus Clypearia jurinei Genus Apoica duckei Subgenus Cylindroeca flavissima sulcata dimidiata gelida weyrauchi Subgenus Formicicola pallens Genus Metapolybia rejecta pallida acincta thoracica Subgenus Furnariana azteca funmaria Genus Agelaia aztecoides richardsi angulata cingulata areata docilis Subgenus Myrapetra suffusa belemensis cajennensis bicyttarella flavipennis Genus Asteloeca bistriata fulvofasciata traili lobipleura catillifex myrmecophila Genus Protopolybia diguetana pallipes (including Pseudochartergus) dimorpha vicina acutiscutis erythrothorax xanthopus biguttata flavifrons chanchamayensis juruana Genus Angiopolybia chartergoides occidentalis pallens exigua parvulina paraensis fuscatus paulista Genus Pseudopolybia holoxantha platycephala compressa minutissima ruficeps difficilis pallidibaleatus scrobalis vespiceps panamensis scutellaris picteti Genus Parachartergus Subgenus Pedotheca sedula emaciata apicalis scutellaris aztecus singularis weyrauchi spinifex colobopterus wheeleri fraternus Subgenus Polybia fulgidipennis Genus Charterginus liliacea richardsi aberrans striata smithii fulvus Subgenus Platypolybia weyrauchi nevermanni incerta Genus Chartergellus Genus Chartergus procellosa amazonicus chartarius Subgenus Trichinothorax communis globiventris affinis punctatior metanotalis chrysothorax Genus Nectarinella Genus Brachygastra flavitincta championi augusti rufitarsis azteca sericea Genus Leipomeles bilineolata tinctipennis dorsata lecheguana velutina nana mellifica Genus Synoecoides Genus Marimbonda scutellaris depressus albogrisea smithii Genus Epipona Genus Synoeca Genus Protonectarina guerini chalibea sylveirae tatua 28 AMERICAN MUSEUM NOVITATES NO. 3224

E

Fig. 15. A. . Cells papered over (FMNH). B. Provespa anomala. Broad pedicel formed of secondarily fused, rod-like pedicels (BMNH). C. Polistes olivaceous. Tandem brood, eggs (arrows) distal to pupae (Toamasina, Madagascar). D. Mischocyttarus fraudulentus, from below. Multiple pedi- cels; comb fusion (Costa Rica; photo by C. K. Starr). E. Mischocyttarus pelor, from below. Comb two- sided, expanding outward from central row (between arrows) (Turrialba, Costa Rica; USNM). F. Mis- chocyttarus decimus, two nests from side, natural orientation (left) and inverted (right). Nest on midrib of leaf; paper cone smooth, extending beyond cell walls (arrows), not composed of separate cell bases (left) (Mazaruni Settlement, Guyana; BMNH). 1998 WENZEL: NESTS OF SOCIAL WASPS 29

Fr1.7 I vker- f

Fig. 16. A. Mischocyttarus weyrauchi (or possibly variant of nigropygialis), from above. Pedicel short; comb single row supported centrally; cells face horizontally; furrows between cells reduced; some cells rectangular in cross section (arrow) (Chapada, Mato Grosso, Brazil, CMNH). B. Mischocyttarus new species near punctatus. Cells round in cross section, from lip of preceeding cell (arrows). Terminal dark spots are glue fastening nest to card (BMNH). C. Mischocyttarus near alienus. Pedicel long; comb single verticle row of cells (Puerto Viejo, Costa Rica). D. Ropalidia bicincta. Comb pinnate; central rib built by foundresses, lateral branches added by workers (Antseranana, Madagascar; KUSM). E. Ropal- idia montana. Envelope sheets multiple; entrances variable and multiple (Mudumulai Preserve, Tamil Nadu, India). F. Belonogaster sp. Comb removed as pupae emerge, leaving hole. Back of comb visible through hole; comb supported by marginal strips of reinforced carton bordering hole (margin retouched for clarity) (HUMB). 30 AMERICAN MUSEUM NOVITATES NO. 3224

I

a C X . > -. \ i F Fig. 17. Comb expansions. A. Apoica sp. Broad attachment to branch; back of comb felt-like, showing no cell bases (HMCZ). B. Angiopolybia pallens, combs removed. Repeated expansion of en- velope corresponding to lateral growth of combs and addition of new combs suspended from those above; old entrances plugged (arrows) but not removed in this anomalous nest, current entrance open (KUSM). See typical form, figure 23F C. Agelaia testacea, comb back. Radial comb growth from point of intiation (arrows, pedicels removed); comb fusion, hexagonal pattern becoming continuous across both combs (upper region) (Santarem, Para, Brazil; HMCZ). D. Agelaia areata, nest bisected. Spiral envelope expanding over itself; entrances multiple; some cells destined to hold brood, others (arrow) papered over to provide support (KUSM). E. Parachartergus (apicalis group), comb, from below. Secondary,F'tcrosswalls> onX,top,+of pupal caps (Suriname; CUIC). F. Charterginus fulvus, incipient nest, from below. Lateral margins drawn across face of completed comb, eventually fusing to create six- pointed nest; entrance simple hole dorsally or at apex of one point (Suriname; CUIC). 1998 WENZEL: NESTS OF SOCIAL WASPS 31

Fig. 18. A. Parachartergus apicalis, sectioned. Envelope generally single sheet with large regular corrugation, remodelled to allow comb expansion; pedicelate combs supported independently from ver- tical or inclined branch; new combs added either above or below older combs (Suriname, NNML). B. Parachartergus amazonensis. Envelope simple or flattened dome; entrance simple gap in lower region (between arrows) (near Iquitos, Peru, AMNH). C. Chartergellus punctatior. Margin of envelope bearing flossy, paper pillars internally, particularly in top region. carcass from museum pest (Barbacoas; Colombia, BMNH). D. Nectarinella championi. Envelope with fine irregular corrugations, highly cryptic on broad trunk; primary envelope (between black arrows) expanded by adding secondary envelope below and contiguous with it (between white arrows); entrance simple or spout at lowest region (white arrow) (Atena, Costa Rica; AMNH). E. Leipomeles dorsata, corrugated form. Envelope with fine, ir- regular corrugations laterally, smooth central region bordered by dark bands of pulp (arrows); entrance simple gap at leaf apex (Barcellos, Brazil; NMBS). F. Pseudopolybia difficilis. Envelope round or ovoid, multiple, laminae closely spaced; entrance simple hole in lower region (Suriname; CUIC). 32 AMERICAN MUSEUM NOVITATES NO. 3224

's -

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i 3 - !!1 .S ;!E :::--A fse'-

Fig. 19. A. Metpolybia cingulata, incipient nest, from below. Small swarm rebuilding sessile comb on site of destroyed nest (cement ceiling); entrance (arrow) distinct, away from last gap in envelope, which will be closed completely (Tena, Ecuador; AMNH). B. , section through lower uninhabited levels of large, active nest. Comb sessile; old envelopes (arrows) not removed as nest expands upward; entrance (not shown) at top (San Vito; Costa Rica). C. Clypearia duckei, damaged nest. Sessile beneath branch distal to ant nest; envelope rough, reinforced with blots, rounded, not showing cell walls; entrance short collar, lateral or upward (near Manaus, Brazil; WDH, photo by W. D. Hamilton). D. Asteloeca traili, from below. Sessile beneath branch; envelope smooth, not concealing marginal cell walls or bases; entrance central, short collar. This nest was expanded once (near Iquitos, Peru; AMNH). E. Protopolybia near holoxantha, incipient expansion of mature nest. Pedicellate comb expanded contiguously with older comb; envelope ultimately arising from marginal cell walls and closed with old envelope; entrance simple, dorsal or dorsolateral, sometimes distinct from last gap in incipient envelope (KUSM). F. Protopolybia acutiscutis, dry season nest, bisected. Envelope multiple, sometimes chaotic; entrances multiple, lateral or ventrolateral; later combs sessile upon preceding envelope (Canal Zone, Panama; KUSM). 1998 WENZEL: NESTS OF SOCIAL WASPS 33

00 i I Fig. 20. A. Polybia scutellaris, sectioned. Envelope reinforced initially by fine, imbricate carton, exterior ultimately thickened by blots; stout spines are typical of this species but rare in the genus. Genera Chartergus and Epipona are similarly heavily reinforced, but without spines (half of nest at MNHN, other half in Geneva; see Wenzel 1992b). B. Polybia bistriata, bisected. Initial cells are sessile on substrate, but comb grows off it, creating illusion of having a pedicel. Some Apoica, Brachygastra, Chartergus, and Charterginus are similar (Suriname, NNML). C. Polybia affinis, sectioned. Primary comb suspended by butressed sheet (arrows), typical of subgenus Trichinothorax (NNML). D. Epipona sp., sectioned and damaged. Nest of 13 levels completed before the hatching larvae in the top (oldest) level required cell walls to be elongated, hence construction in as rapidly as 10-15 days. Some nests of this genus are not expanded beyond the effort of the original swarm for several years (but see fig. 26E). The hole in the lower center and the broken lowest level are the result of damage during collecting (Costa Rica?; KUSM). E. Polybia diguetana (?), bisected. Envelope with imbricate reinforcement; lower combs with successively greater radius of curvature (KUSM). F. Polybia dimidiata, sectioned, and M. Hennigan. Nest surrounds vertical support; envelope with thick imbrication in upper portion (NNML). 34 AMERICAN MUSEUM NOVITATES NO. 3224

Fig. 21. Comb expansions. A. Polybioides melaina and the author. This species achieves the largest colony size of any African social wasp (MRAT, photo by J. M. Carpenter). B. Polybioides raphigastra, from below. Built in cavities, this comb spirals outward and downward. Imbricate reinforcement of the margins of the comb provide added support and form a protective envelope around more centrally located cells (BMNH). C. Ropalidia montana, comb. Protected inside a separate envelope, continuous combs growing in a downward spiral (= klimakatocyttarus sensu Carl, 1934) allow this and some other Ropalidia to create huge nests without requiring many distinct structures for support or modifications thereof to allow expansion (see also fig. 16E) (AMNH). D. Brachygastra mellifica, from below. Chaotic expansion of comb and envelope in mature nests produces many entrances and loosely interconnected regions of cells. Young nests are more orderly and form discrete levels with a single entrance (not shown) (USNM). E. Protonectarina sylveirae, comb, from above. The supple paper of the comb back (a former envelope) can be easily peeled away to leave the hexagonal walls of brood comb behind (see also fig. 25D) (Brazil; HMCZ). F. Polybia ruficeps. Many Polybia have the entrance as a lateral or dorsolateral hole or short spout above the level of the lowest brood brood comb. This species exaggerates the form to resemble a tea pot with the spout pressed back against the pot. The entrance (center) is well above the level of the lowest comb (arrow), with which it communicates by a curved passage. Subse- quent envelopes must incorporate the old entrance, becoming increasingly asymmetrical and protruding on one side (Chapada, Mato Grosso, Brazil; CMNH). 1998 WENZEL: NESTS OF SOCIAL WASPS 35

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Fig. 22. A. Mischocyttarus sp., dead nest. Multiple pedicels and comb fusion can produce linear nests of dozens of cells. Such Mischocyttarus nests are usually built on leaves guarded basally by dolichoderine ants (San Vito, Costa Rica). B. Apoica flavissima. Nocturnal habits of Apoica allow the entire workforce to remain on the nest in the day, where they may be so numerous as to completely cover the face of the nest and hang from the bodies of their nestmates. Pale-bodied species make an impressive display (Iguazu, Argentina; AMNH). C. Parapolybia varia, comb back. Removal of meco- nium through the back of the cell leaves a hole that is closed by transparent oral secretion, appearing dark here. Such "windows" in the back of the comb are unique to Ropalidia and Parapolybia (AMNH). D. , from below. Many Ropalidia begin nests with multiple combs, which generally proliferate separately but may fuse (Bangalore, India). E. Ropalidia romandi. Ropalidia that build en- velopes tend to make amorphous structures of multiple layers, but the exterior is generally smooth in appearance (Queensland, Australia; AMNH; photo by J.M. Carpenter). F. Polybioides tabidus, one envelope sheet folded up. The origin of the nest is revealed by the first pair of back-to-back combs, which also correspond to the oldest region of the bivalve-like envelope. The first cells of each comb are sessile by their walls on the branch, but are chewed down and reinforced with secretion to produce a narrow support resembling a pedicel (between arrows) for the comb that hangs from it. This species has two simple envelope sheets, one on either side, but P. melaina is complex (see fig. 20A) (MNHN). 36 AMERICAN MUSEUM NOVITATES NO. 3224

NW

Fig. 23. A. Chartergellus punctatior. Envelope is a flatenned dome (black arrows), rough, uneven, and cryptic against a broad trunk; entrance (white arrow) is a short spout easily concealed by loitering wasps (Tena, Ecuador; AMNH). B. Same as A, with envelope removed, revealing combs. Former margin of envelope denoted by arrows; dark area above is a wound in the tree, white area just below combs is a spider egg sac. C. Agelaia xanthopus, in a tree cavity. Combs stacked, conical, and partly spiral and fused. See also cross-section photo in Wenzel (1992b) (Mexico, MNHN). D. Marimbonda (?). Envelope simple on the underside of a narrow leaf; central furrow (arrow) like Leipomeles (see E) (BMNH). E. Leipomeles dorsata, furrow form. Central furrow (black arrow) mimics leaf midrib, ridges (white arrows) mimic veins; entrance at apex of leaf (Santarem, Para, Brazil; photo by R. L. Jeanne). F. Angiopolybia pallens. Typical form; smooth envelope with elegant flared spout. Boundary between carton of different colors (arrow) marks beginning of new lower construction to accommodate growing combs (see also fig. 17B) (Misahualli, Ecuador; photo by J. M. Carpenter). 1998 WENZEL: NESTS OF SOCIAL WASPS 37

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Fig. 24. A. Leipomeles dorsata, ant guard on leaf petiole. (Belem, Brazil; photo by R. L. Jeanne). B. Asteloeca traili. Smooth carton, margin of nest showing lines of construction, cell bases, and walls (near Iquitos, Peru; AMNH). C. Clypearia (= Occipitalia) sulcata, broken. Rough, thick carton, rein- forced to obscure lines of construction, cell bases, and walls (near Iquitos, Peru; AMNH). D. Clypearia sulcata, incipient, from below. Nest (between black arrows) in close proximity and apical to Azteca ant nest (white arrow) (Iquitos, Peru). E. Metapolybia cingulata. Brittle envelope damaged slightly, revealing comb within. Envelope rough, sometimes glossy, rarely showing lines of construction. Entrance at top concealed by sharp eaves around envelope margin (Tena, Ecuador; AMNH). F. Clypearia weyrauchi, oblique side view. Envelope glossy, smooth, lines of construction obvious (IMLT). 38 AMERICAN MUSEUM NOVITATES NO. 3224

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Fig. 25. A. . Multilevel form among leaves (near Santa Cruz, Bolivia; AMNH). B. Protopolybia chartergoides, in curled leaf, from below. Transparent form, envelope consists of sal- ivary matrix only, no pulp, restricting access to cavity; entrance is simple gap on right side where new membrane appears_M4+t.comb visible' as brown carton rough; ordinary beyond transparent envelope (near confluence of Rio Napo and Rio Amazon, Loreto, Peru; AMNH). C. Protopolybia exigua binominata, from below. Planar form (in situ), envelope exterior green (fading in collections), acute margins of envelope remain natural pale color of carton (near Tena, Ecuador; AMNH). D. Protonectarina sylveirae, lower region, sectioned. Supple, gray wavy carton; seams marking successive combs wiggly and irreg- ular (see also fig. 21E) (MALS, photo by T Osten). E. Polybia sp., from below. Defense display: wasps cover nest and nearby branches with wings upraised, next response is to attack moving or alien objects (Iquitos, Peru). F. Brachygastra scutellaris. Nest is sessile despite image of narrow pedicel (cf. P. bistriata, fig. 20B). Carton is naturally brown, but lower region of envelope is later colored white below (see also fig. 26D, F). This form found in B. scutellaris and B. smithii (near Iquitos, Peru; AMNH). 1998 WENZEL: NESTS OF SOCIAL WASPS 39

Fig. 26. A. Dolichovespula maculata, sectioned. Laminae of concentric envelope sheets (Islip, New York). B. Vespula maculifrons, mature envelope surface. Tight imbricate reinforcement, lines of con- struction evident (Lawrence, Kansas; KUSM). C. Chartergus chartarius, broken, and P J. DeVries. Cylindrical white nest, entrance in center of combs (left arrow), top of nest encompassing branch (right arrow) (Sucumbios, Ecuador; photo by J. Clark). D. Synoecoides depressus. Carton is naturally brown, but lower region of envelope is later colored white (see also figs. 23F, 26F) (BMNH). E. Epipona niger, expanded nest. Original nest was tapered below, second construction effort represented by broader lower region; comb becomes increasingly sloping, entrance near comb margin on higher side of incline (arrow). Cotton stuffed in entrance and area damaged during collection (Gamboa, Panama). F. Charterginus nevermanni, broken. Comb reveals natural amber color of carton, exterior of envelope later colored white (see also figs. 25F, 26D) (La Selva, Costa Rica; photo by R. M. Timm). Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates and Bulletin published during the last five years are available at World Wide Web site http://nimidi.amnh.org. Or address mail orders to: American Museum of Natural History Library, Department D, Central Park West at 79th St., New York, NY 10024. TEL: (212) 769-5545. FAX: (212) 769-5009. E-MAIL: [email protected]

8 This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).