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AMERICAN' MUSEUM Norntates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN' MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3188, 39 pp. December 31, 1996 Distributional Checklist of Species of the Genus Polistes (Hymenoptera: Vespidae; Polistinae, Polistini) JAMES M. CARPENTER' ABSTRACT A distributional checklist of the species of the de Beauvois, 1818, = tepidus malayanus Camer- paper wasp genus Polistes is presented. New syn- on, 1906; and arthuri vechti Petersen, 1990, non onyms are multipictus Smith, 1860, non Haliday, Das and Gupta, 1989, = arthuri Cameron, 1901. 1836, = stigma maculipennis Saussure, 1853; Two new replacement names are proposed: inter- stigma var. thoracica Buysson, 1913, non Fox, medius Kojima, for medius Kojima, 1988, non 1898, also = stigma maculipennis Saussure, 1853; Palisot de Beauvois, 1818; and torresae Silveira, tepidus var. minor Schulthess, 1932, non Palisot for occultus Silveira, 1994, non Kojima, 1988. INTRODUCTION Polistes is the only cosmopolitan genus of ful regional catalogs and revisions have social wasps, and, with 203 presently rec- been published in the intervening 90 years ognized extant species, one of the most spe- (Liu, 1937; Bequaert, 1938; Guiglia, 1972; cies-rich. The lack of an adequate catalog or Richards, 1978a, b; Krombein, 1979; Car- checklist for all the species of the genus Pol- dale, 1985; Das and Gupta, 1989; Starr, istes has led to a number of taxonomic errors 1992), but these works have not been inte- in recent time, as well the perpetuation of grated, and significant gaps remain. Yoshi- other problems. There have been no compre- kawa (1962) attempted a world checklist, hensive listings since the catalogs of Dalla but, as Richards (1973: 85) put it, he did so Torre (1894, 1904), which of course are now "somewhat imperfectly." His list had many too far out of date to be of general use. Use- omissions and misspellings, and the fact that Curator, Department of Enotomology, American Museum of Natural History. Copyright X American Museum of Natural History 1996 ISSN 0003-0082 / Price $4.00 2 AMERICAN MUSEUM NOVITATES NO. 3188 he often eschewed citations of original ref- can be diagnosed are species (Nelson and erences limited its taxonomic usefulness. Platnick, 1981; Nixon and Wheeler, 1990). The present listing redresses this situation. But this means that taxa currently recognized Nevertheless, it is a successor to Yoshikawa's as subspecies are either species or synonyms, work, in the sense that it is a distributional and moreover there must be a mixture. With- checklist. Following the citation of the orig- out careful study on a case by case basis, the inal description and synonymy for each spe- proper status of subspecific taxa cannot be cies, the distribution is given. The distribu- determined. I have applied this approach tion is listed using current political entities elsewhere to some groups of Vespidae (Car- (so far as they can be determined), because penter, 1988; Carpenter and van der Vecht, they are unambiguous compared to more 1991), Archer (1991, 1992, 1994, 1995) has generalized terms for areas, which may be done so for various species of Vespa, and idiosyncratic, and would require detailing the Snelling (1983) has followed this course for geographical definition. Where political di- a few species of Polistes. Until such studies vision could not be determined precisely can be undertaken for Polistes, I am unwill- from the literature, terms such as "New ing to proceed with wholesale changes in sta- Guinea" are used. Citations given at the end tus of subspecific taxa. Furthermore, I be- of the distribution are to the most recent lieve that such changes are best done as part summary. If there has been no report of a of a revision, rather than in a catalog or range extension since the description, no fur- checklist, so that adequate explanation of each ther reference is given. decision may be supplied. Therefore, I have The format of the synonymies is standard generally maintained subspecies, or treated in taxonomic catalogs, with each reference varieties as subspecies, if the most recent au- given in abbreviated form, to expedite track- thor did so-a total of 103 additional names. ing it down. Misspellings are indicated by The cases where I did not are as follows. exclamation points in brackets. The format Bequaert (1938) reduced many Ethiopian for dates lists the true date of publication, so species to varieties of a few species (he did far as can be determined. Where two dates so with many American species as well in are listed, the first is the true date of publi- other works). Bequaert's varieties of Ameri- cation, and the second, in parentheses, is that can species were first simply treated as sub- printed on the publication. The status given species (Bohart, 1951), and now many are each taxon generally follows the decision of treated as species (Richards, 1978a; Krom- the most recent author to treat that taxon. bein, 1979). I agree with Giordani Soika This authority may typically be determined (1991) that, likewise, many of the Ethiopian from the citations given for the distribution. taxa treated as varieties by Bequaert (1938) However, there are several cases where I merit species status. Thus I have followed a have deviated from this practice. Articles classification of the Ethiopian species more 31(b) and 34(b) of the International Code of like that of Schulthess (1921) with respect to Zoological Nomenclature have been fol- the rank given. lowed with respect to agreement in gender of I have also not recognized any subspecies generic and specific names, necessitating in the Palearctic species biglumis (Linnaeus), correction of Vespa nimpha Christ to nim- dominulus (Christ) or gallicus (Linnaeus), phus, to agree with Polistes, which is mascu- synonymizing all, for two reasons. Firstly, line. However, I agree with Bohart and Menke some taxa have even recently been recog- (1974) and Krombein (1979) that Vespa car- nized as "forms" or "varieties" of subspe- olina Linnaeus should be treated as a noun in cies (e.g., Polistes biglumis bimaculatus var. apposition, hence it does not have to agree in arduinoi in Guiglia, 1972, with bimaculatus gender, and so Richards' (1978a) use of car- var. nigrinotum a synonym), but formal rec- olinus is incorrect (I here treat Vespa stigma ognition of any taxon below subspecies has Fabricius likewise, contra Carpenter, 1996). not been permitted through three revisions of With respect to the use of the subspecies the International Code of Zoological Nomen- category, I agree that it has no place in a clature, since 1961. To the extent that such phylogenetic system: species-group taxa that taxa are used to divide subspecies, they are 1996 CARPENTER: CHECKLIST OF POLISTES 3 nothing more than synonyms anyway. Sec- Guinean subspecies of tepidus (see Petersen, ondly, there has been considerable confusion 1990), malayanus Cameron, 1906. NEW in the application of the proper names to STATUS for the subspecies arthuri vechti is dominulus and gallicus for most of this cen- as a synonym of nominotypical arthuri. tury, encompassing the time when most of The only other taxonomic changes are the the subspecies were described. As shown by proposal of two replacement names. The first Day (1979), Kohl (1898) misidentified gal- new name is for medius Kojima, 1988, pre- licus, with the result that he described it occupied in Polistes by media Palisot de anew as foederata. Subspecies described un- Beauvois, 1818. The latter name is a syn- der foederata must consequently now be onym of crinitus (Felton); it was recorded as transferred to gallicus, but those described a synonym of americanus (Fabricius) by under gallicus must be transferred to domi- Saussure (1853), but unfortunately the name nulus. I propose to obviate any attendant was not mentioned in the revision by Rich- confusion by simply synonymizing these ards (1978a). Dr. Kojima has supplied a re- subspecies; those that I have examined ap- placement name for medius Kojima: inter- pear to be neither constant nor distinct in the medius Kojima, NEW NAME. The second color characters supposed to distinguish new name is for for occultus Silveira, 1994, them. This also applies to the commonly rec- preoccupied in Polistes by occultus Kojima, ognized subspecies biglumis bimaculatus, 1988. Sr. Silveira has supplied a replacement which does not differ from typical biglumis name for occultus Silveira: torresae Silveira, (contra Bluthgen, 1955: 398, who separated NEW NAME. the nominotypical Scandanavian- form from Finally, regarding higher classification, the supposedly more richly yellow-marked Polistes is the sole genus in the tribe Polistini bimaculatus; see also Guiglia, 1972: 59). (Carpenter, 1993), and the subgeneric system The last four cases where I have not rec- followed is that presented in Carpenter ognized subspecies despite previous treat- (1996). That system recognizes only four ment are the varieties stigma var. thoracica subgenera: nominotypical Polistes, Gyrosto- Buysson, 1913; and tepidus var. minor ma, Aphanilopterus, and Polistella. These Schulthess, 1932; multipictus Smith, 1860 four subgenera are monophyletic groups; the (all treated as subspecies of stigma by Peter- system of Richards (1973), with two genera, sen, 1987); and arthuri vechti Petersen, 1990. Polistes and Sulcopolistes, and 11 subgenera All four names are preoccupied in Polistes: in Polistes, included several paraphyletic stigma var. thoracica by thoracicus Fox, groups. But Richards' system also recog- 1898 (and cavapyta var. thoracica Buysson, nized some monophyletic groups not given 1906), tepidus var. minor by minor Palisot formal names in my system (see Carpenter, de Beauvois, 1818, multipictus by multipic- 1996: fig. 17), and to assist future investi- tus Haliday, 1836 (now placed in the genus gations of phylogenetic relationships among Agelaia), and vechti by rothneyi vechti Das species, I indicate the subgenera into which and Gupta, 1989. Rather than propose re- Richards placed species since transferred.
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