J. HYM. RES. Vol. 19(1), 2010, pp. 84-93

Chartergellus golfitensis West-Eberhard: a new species of Neotropical swarm-founding (: , ) with notes on the of Chartergellus zonatus Spinola

Mary Jane West-Eberhard, James M. Carpenter, Luiz Fernando F. Gelin, and Fernando B. Noll

(MJW-E) Smithsonian Tropical Research Institute, c/o Escuela de Biologia, Universidad de Costa Rica, Costa Rica; [email protected] (JMC) Department of Entomology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024, U.S.A.; [email protected] (LFFG) Departamento de Zoologia e Botanica, Instituto de Biociencias, Letras e Ciencias Exatas, Sao Paulo State University, Rua Cristovao Colombo, 2265 CEP 15054-000 Sao Jose do Rio Preto, SP, Brazil; [email protected] (FBN) Departamento de Zoologia e Botanica, histituto de Biociencias, Letras e Ciencias Exatas, Sao Paulo State University, Rua Cristovao Colombo, 2265 CEP 15054-000 Sao Jose do Rio Preto, SP, Brazil; [email protected]

Abstract.— Chartergellus golfitensis West-Eberhard new species, is described from Central An\erica and compared with C. zonatus Spinola, a species heretofore inadequately described.

Chartergellus is a genus of 1978, and C. sanctus Richards 1978. He (Hymenoptera, Vespidae, PoHstinae) rang- treated Chartergus zonatus Spinola 1851, as ing from southeastern Brazil to Costa Rica an unrecognized species and did not

(West-Eberhard et al. 2006). The taxonomic include it in his key, because the descrip- background of Chartergellus is somewhat tion ''does not agree fully with any speci- troublesome. It was described by Bequaert mens I have seen of Chartergellus'' (Rich- (1938) as a subgenus of Chartergus sensu ards 1978: 217). Most recently an eighth Bequaert. The use of the name Chartergus species has been added, C. afoveatus was due to overlooking the type designa- Cooper 1993. tion for that genus, and Bequaert (1943) With the exception of C. atectus (see pointed out that the name Parachartergus Richards 1978, and below), the species of von Ihering, 1904 was correct. Bequaert Chartergellus build nests consisting of mul- (1938) designated as type species Vespa tiple combs attached laterally to the sub- frontalis Fabricius, 1804, the only species strate by pedicels and covered by an included in Chartergellus. Richards (1978) envelope of a single sheet, with the raised Chartergellus to genus, and pointed entrance a short spout (Wenzel 1998). The out that Vespa frontalis Fabricius was envelope may be irregular, with the lines of preoccupied by Vespa frontalis Latreille construction evident and contributing to 1802. He proposed C. amazonicus as a camouflaging of the nest. Chartergellus is replacement name, and described five characterized by having a prominent new species: C. atectus Richards 1978, C. curved bristle on the third labial palpo- communis Richards 1978, C. nigerrimus mere, the maxillary palpi five-segmented Richards 1978, C. punctatior Richards and labial palpi three-segmented, lacking —

Volume 19, Number 1, 2010 85

Figs 1-8. Chartergellus golfitensis West-Eberhard new species. Holotype. 1. Habitus. Lateral view. 2. Dorsal view. 3. Head, frontal view. 4. Head, lateral view. 5. Head, dorsal view. 6. Mesosoma, dorsal view. 7. Mesosoma, lateral view. 8. Scutellum, metanotum and propodeum, dorsal view. Scale bars = 1 mm. an occipital carina, the mesepisternum wider than the diameter of the median lacking a dorsal groove, and the metano- ocellus. tum rounded (Carpenter 2004). Description. Female. Holotype fore wing Previously, only Chartergellus atectus was length 6.72 mm. recorded for the Costa Rican fauna (Rich- Color. Black. Reddish brown: clypeus, ards 1978). We describe here Chartergel- first antennal segment and flagellum be- lus golfitensis, a new species from Costa neath, malar space, lower 3/4 of the head, Rica. mandibles except teeth, fore tibia, tips of tarsi. Whitish anterior stripes on metano-

Chartergellus golfitensis West-Eberhard, tum curved at margin of propodeum, new species pronotal carina, and broadly along entire (Figs 1-9) length of gena. Wings hyaline, venation brownish. Head. Eyes with short sparse Diagnosis.—Chartergellus golfitensis is dis- hairs, touching clypeus for short distance. tinguished by the presence of a raised Clypeus wider than long (median length x black-lipped arc at the base of the man- width at middle = 0.74). Entire surface of dible; clypeus in short contact with eye; head pubescent except for malar space and pale markings on the scutellum; frons with gena posterior to it, and clypeus below three-pointed reddish brown mark with level of eyes, where there are long bristles. the central point nearly reaching the Frons with sparse and shallow punctures. median ocellus and in the intraocular space Malar space present, about as wide as to the center of the ocular sinus; and a length of fourth antennomere. Gena with broad pale stripe along the entire length of width equal to width of compound eye. the gena, usually without red or black Widest part of eye seen from front only region below, which when present is no half length of first antennomere. Vertex 86 Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling

Figs 1-8. Continued.

Figs 1-8. Continued. Volume 19, Number 1, 2010 87

Figs 1-8. Continued. hairs present, sparser and shorter than arc with a black basal lip, dorsal third those of frons which are nearly as long as covered by lateral clypeal lobe. Mandible bristles of cl3^eus below level of eyes. Base smooth, outer surface with a few long of mandible with raised rim, forming an bristles. Mesosoma. Pronotum, scutum and

Figs 1-8. Continued. Journal of Hymenoptera Research: Festschrift Honore^g Roy Snelling

Figs 1-8. Continued.

Figs 1-8. Continued. Volume 19, Number 1, 2010 89

Figs 1-8. Continued. scutellum with fine, dense punctation and fovea present. Propodeum without poster- covered with straight hairs. Mesepister- ior concavity, with hairs on entire surface, num with scrobal furrow beginning at propodeal valve narrow throughout. margin beneath tegula and curving poste- Male. riorly to form a gentle arc about half as Color. As in female but metasoma (gas- long as the mesepisternum, which is ter) and legs dark brown. Reddish-brown rounded anterior to the furrow. Dorsal of face fills entire ocular sinus and lacks pronotal carina well developed, ending three medial peaks on frons. Whitish genal close to the posterior margin. Pronotal stripe touching eye. Additional white:

Fig. 9. Male genitalia of Chartergellus golfitensis. From left to right, aedeagus in ventral view, aedeagus in lateral view, digitus, cuspis and paramere. Scale bar = 1 mm. 90 Journal of H^-mexopter-a Research: Festschrift Honoring Roy Snelling

Stripe anterior to scrobal furrow, and seum of Zoolog}' of the Uni\ersit\' of Sao Paulo, coxae. Head. Mandibles with silvery ap- Sao Paulo, Brazil. Paratypes. In Museo de pressed hairs on basal half of outer surface, Zoologia, Escuela de Biologia, University of Costa Rica: Puntarenas Prov. 40 females, 17 apical half with a few long bristles, under- males, aU with the same locality and collector as surface covered with long hairs. Ch^eus as the holot\^e. Golfo Duke, 24 km W. Piedras long as wide (L:W = 1.06), in extensive Blancas, alt. 200m, iii 1992, I9 (Paul Hanson); contact with eyes except for a short 3 km SW Rincon, VI 1991, 2 99 (Paul Hanson); distance dorsally, and covered with silver}^ III-IV 1992, 19 (Paul Hanson). In Instituto pubescence, ventrally forming a dense Nacional de Biodiversidad (INBio), Costa Rica: band of long appressed bristles which Puntarenas Prov. Est. Agujas, Sendero Zamia, gleam in direct light as do similar hairs 300m. 22-30 AGO 1996. L_S_276750_526550 on lateral portions of frons, white portions ^8318, 19 (E. Retes, A. Azofeifa, M. Lobo); of coxae, and a thin line on cur\^ed distal Albergue Cerro de Oro, 200m., 5-9 MAY 1995, L margin of last metasomal sternum. Malar N 279650 ^4745, I9 (B. Gamboa); 4-14 May space about half as wide as length of fourth 1995, L_S_280450_517500 #5919, I9 (E. Alfaro); antennomere. Gena only about half width Rancho Quemado, 200m., Die 1992, L-S 292500_511000, I9 (F. Quesada); Abr 1992, I9 of eye. Widest part of eye seen from front (K. Hores); 299 (D. Brenes); lun 1992, I9 (F. as long as first antennomere; saber-like Quesada); Est. Esquinas, Peninsula de Osa, ventral mandibular tooth longer and sharp- 200m. Abr 1993, L S 301400.542200 #2076, I9 er-edged than in female. Q.F. Quesada); Est. Sirena, 0-lOOm, P.N. Cor- Male genitalia (Fig. 9). Aedeagus serrate covado, Jun 1991, L-S270500_508300, I9 (G. beneath. Ventral process (at base of serrate Fonseca); P.N. Manuel Antonio, Quepos, 80m. portions of aedeagus) forming a "U" with Abr 1993, L S 370900_448800 #2140, !£ (G. outer end sharply pointed, as in C. commu- Vaiela); Oct 1992, L-S 370900_448800, I9 (G. nis (Richards 1978, fig.88b). Serrations: Varela). proximal nine very fine and triangular Etymology.—The name is given after the (pointed), more distal 25 broader, curved type locality, the Golfito region in Costa on the posterior edge and pointing for- Rica, Central America. ward. Serrate edge slightly longer than Remarks.—INBio collecting stations Est. distance from end of serrations to tip (Estacion) Agujas, Est. (Estacion) Esquinas, of aedeagus. Parameral spine hairless, Albergue Cerro de Oro, and Rancho slightly curved laterad, with elongate Quemado are all in the Osa Peninsula of lateral opening near tip. Inner surface of southwestern Costa Rica. digitus with long fine hairs, longer than The holot}''pe and paratypes were com- width of digitus seen in profile, and shorter pared with paratypes of three other species toward tip, which is bare. of Chartergellus: C. punctatior Richards, C. Distribution. —Known only from Pacific communis, and C. zonatus Spinola. The lowlands of Costa Rica. parat}^pe of C punctatior Richards, labeled Habitat.—The tv\^o obser\'ed living colo- as such by Richards, is from the same nest nies of C. golfitensis (the colony of the as a specimen (COLOMBIA: Valle, Anchi- holotype and another from the same caya, 27.vii.73, M.J.W. Eberhard) mista- locality), and all other collected specimens, kenly listed by Richards (1978: 222) as a were found in areas of lowland humid paratype of C. nigerrimus. Richards (1978) forest. Nests were built on a vertical tree listed no specimens of C. punctatior from trunk and the wall of a building. Colombia in the description, but included Colombia in the distribution of this species Types.—Holotype, 9, COSTA RICA, Punta- renas Prov., Refugio de Vida Silvestre Golfito, in his key, further indicating his intention 83 11'55"W, 8-39'25"N, 500m from the sea level. to consider the Colombian specimen 2.i.2006 (L. Chavarria), deposited in the Mu- (which fits the description of punctatior) a VoLLAiE 19, Nl-mber 1, 2010 91 paratype of this species. Besides this speci- Color variation.—^The colony of the t\^pe men, two other Colombian (Anchicaya) specimen contained three queens and two specimens of C. punctatior (28.ix.76, M. J. readily distinguished age cohorts: young West-Eberhard) were examined by Rich- individuals (16 females and 17 males), ards in London. One was retained in the distinguished by wide clear metasomal Natural History Museum, London, and the apodeme margins with only a very thin other, from the same colony, is in West- black marginal line, as is characteristic of Eberhard's collection, labeled by Richards recently emerged vespid adults (West- as a paratype of C. atectus. But it is clearly a Eberhard 1975), and 22 older females specimen of C. punctatior that conforms in (including the holotype), with wide black structure (mandibles with strong basal rim, apodeme margins. The young females and first metasomal tergum with broad resembled the queens in being lighter in yellow apical band) and nest type (un- color: meso- and metasoma and some parts exceptional for the genus: combs on a rock of legs that are black in the older females rather than leaves, covered by a full are dark to light brown in the young envelope) to the published description of females and queens, suggesting that the punctatior, not to those of the holotype of C. recently emerged females, although lack- atectus collected at the same locality and ing sperm in the spermatheca and with described as having a nest unusual for the undeveloped ovaries, are young queens. genus in being constructed betw^een leaves Queens have light brown first antenno- and lacking a full envelope (Richards 1978). mere and nearly white mandibles. White The paratype of C. communis examined here lines at the posterior margin of the prono- (BRAZIL: MT, Ilha de Bananal, Santa tum and the scrobal furrow are present in Isabella [recte: do Bananal, Santa Isabel], some females, absent in some (including 25.viii.68, W. D. Hamilton) was deposited in the holotype), and a brownish spot in the West-Eberhard collection by Richards. some, variation not correlated with age or Along with other West-Eberhard voucher caste. A white stripe on the distal margin of specimens, these paratypes will eventually the first metasomal tergum is complete or be deposited in the US National Museum, absent in some, or at the lateral margins Smithsonian Institution, Washington DC. only in most females. A detailed morpho- We also compared a paratype female of C. metric analysis of both sexes and castes golfitensis wilh three female syntypes of the will be published elsewhere. Spinola type series of C. zonatus in the The Sendero Zamia, Rancho Quemado, collection of the Museo di Zoologia Siste- and Manuel Antonio specimens have dark matica delTUniversita di Torino, now brown rather than reddish brown facial housed in the Museo Regionale di Scienze markings, and light markings are brown or Naturali in Turin. yellowish brown rather than whitish. Richards (1978) described three Charter- NOTES CHARTERGELLUS gellus species (amazonicus, punctatior, and ON ZONATUS SPINOLA communis) as having a ''raised rim" at the base of the manidible. In the communis Spinola (1851) referred to four female paratype this is a rim-like ridge that rises specimens collected by M. Ghiliani in Para, abruptly forming a dark-brown-edged arc with the male unknowm, but only three near the inner margin of the mandible, specimens are present in the Turin collec- continued as a straight black line to the tion and there is no record of the location overlapping comer of the clypeus. In the of the missing female. Examination of the punctatior parat\^es and in C. golfitejtsis the remaining specimens indicate that Rich- arc arises less abruptly and is not contin- ards' (1978: 218) doubts regarding the ued as a line. Spinola (1851) description of C. zonatus. 92 Journal of Hymenoptera Research: Festschrift Honoring Roy Snelling

resulting in its status as incertae sedis Other characters: eyes with sparse short (Carpenter and van der Vecht 1991; Cooper hairs; mandible with four teeth, base with 1993), are largely unjustified, though a few raised basal arc, scutal punctures separated inaccuracies were evident (see below). by one diameter or more, scrobal furrow rises Richards stated that ''the description of to gentle ridge at anterior side, coarsely and

. . .Chartergus zonatus does not agree fully irregularly punctuate, with appressed hairs with any specimens I have seen of Charter- on posterior part and striate anterior ridge. gellus" but the Spinola specimens accord As suspected by Richards regarding a with Richards' (1978) generic description strongly yellow-marked species, the man- for Chartergellus in having the propodeal dibles are not black; they are brown, with valve linear rather than in-curving, prono- only the teeth dark brown or black, in- tal fovea present, widely separated from cluding in the darkest of the three speci- carina; hind-wing vein cu-a much longer mens seen. The scutal margins are not than 1 Cui; labial palpi with three seg- yellow, as stated by Spinola; instead, it is ments (the maxillary palpi could not be the margin of the adjacent sclerites that are seen); scrobal furrow strong; margin of yellow; and only about half of the metano- strong; mesoscutum opposite tegula scu- tum is yellow, not the entire metanotum. tellum rounded in profile; metanotum not Spinola (1851) cited Ghiliani's notes as vertical; and occipital carina lacking. stating that the nest of this species, in Species level characters are as follows: contrast to that of Chartergus nitidulans

Color. Black. Dark brown: antennae Fabricius, is found on the trunks of trees, (lighter at tip beneath), legs brown and and that the sting is very painful. Spinola and wing venation. Reddish- brown: cly- was sent a sample of the nest carton, which three-pointed area of frons to peus and 1/3 he described as "teme et grisatre," (dull distance antennal to from insertions me- and grayish) but this has evidently been dian ocellus. Yellow: intraocular space to lost. There is no holotype labeled, hence all ocular sinus; genal stripe half broad (about three specimens examined are syntypes. width of gena) from mandible nearly to dorsal margin of eye; propodeal carina; ACKNOWLEDGMENTS stripe anterior to scrobal furrow and about The authors thank the Ministerio de Ambiente, Va its length; anterior margins of scutellum, Energia y Telecomunicaciones de Costa Rica (MIN- metanotum; thin stripes on margins of all AET) for the collecting license, all the personnel of the sclerites bordering mesoscutum; posterior Refugio de Vida Silvestre Golfito for collaboration, the margins of terga 1, 2 (one female) or terga Universidad de Costa Rica for hospitality in Golfito, and Chaves Jimenez for help during the collection 1, 2, 3 (two females). Other variation: one Jose of colonies made by L. Chavarria in Costa Rica. We female, labeled with a small green tag also thank J. Diniz for helping with the dissection of marked "9," is darker in color than the male genitalia. MJWE thanks Prof. Maria Cristina other two: all of the markings that are Lorenzi, Guido Pagliano, and Dr. Stefania Fucini for yellow in the other two specimens and in help and hospitality during work on the Spinola the Spinola description, are brown, and all specimens in Turin; and Dr. Paul Hanson of the University of Costa Rica and Manuel Solis of the Costa brown markings are darker brown in this Rican National Biodiversity Institute (INBio) for help specimen, which is one of those with three with use of collections at their institutions. metasomal stripes. A female labeled "Pun- gono fortemente" (stung strongly) was LITERATURE CITED affected by mold, so the remaining speci- Bequaert, C. 1938. A new Charterginus from Costa men, with no special label, and having J. Rica, with notes on Charterginus, Pseudochartergus, yellow coloration like that described by Pseudopolybia, Epipona, and Tatua (Hymenoptera, Spinola, was used for the description Vespidae). Revista Entomologia Rio de Janeiro 9: reported here. 99-117, 1 pi., Ifig. Volume 19, Number 1, 2010 93

. 1943. The generic names Chartergus, Parachar- Richards, O. W. 1978. The Social of the Americas, tergus, Epipona and Tatua. A correction (Vespidae, Excluding the Vespinae. British Museum (Natural Hymenoptera). Bulletin Brooklyn Entomological History), London. 580 pp + 4 pp plates. Society 38: 4. Spinola, M. 1851. Compte rendu des Hymenopteres

Carpenter, J. M. 2004. Synonymy of the genus Mar- inedits provenant du voyage entomologique de imbonda Richards, 1978, with Leipomeles Mobius, Ghiliani dans le Para en 1846, 3-78. [Preprint of 1856 (Hymenoptera: Vespidae; Polistinae), and a Memorie della Reale Accademia della Scienze di

new key to the genera of paper wasps of the New Torino (2)13: 19-94,1853 [Spinola (1853) is avail- World. American Museum Novitates 3465: 1-16. able at http://ripley.si.edu/ent/nmnhtypedb/

and J. van der Vecht. 1991. A study of the images/pdfs/2746.pdf]

Vespidae described by William J. Fox (Insecta: Wenzel, J. W. 1998. A generic key to the nests of Hymenoptera), with assessment of taxonomic hornets, yellowjackets, and paper wasps world- implications. Annals of the Carnegie Museum 60: wide (Vespidae: Vespinae, Polistinae). American 211-241. Museum Novitates 3224: 1-39.

Cooper, M. 1993. A new species of Chartergellus (Hym., West-Eberhard, M. J. 1975. Estudios de las avispas Vespidae, Polistinae, Polybiini) from Bolivia. En- sociales (Himenoptera, Vespidae) del Valle del

tomologist's Monthly Magazine 129: 165-166. Cauca. I. Objetivos, metodos y notas para facilitar

Fabricius, J. C. F. 1804. Systema Piezatorum. 439 pp. la identificacion de especies comunes. Cespedesia 4 Brunsvigae. (16): 245-267.

Ihering, H. von. 1904. As vespas sociaes do Brasil. , J. M. Carpenter, and P. E. Hanson. 2006. Revista do Museu Paulista 6: 97-309, 5pls, 3 figs. Familia Vespidae. 617-644 in: Hanson, P. E., and

Latreille, P. A. 1802. Historic naturalle, generale et I. D. Gauld eds., Hymenoptera de la Region particulere des Crustacees es des Insectes. (Sonnini's Neotropical. Memoirs of the American Entomological suites a Buffon), Paris. 3 xxi + 468 pp. Institute 77: 1-994.