Abstract- Cladistic analyses of data from allozyme polymorphisms in paper WASP SOCIAL parasites and THEIR HOSTS do not support the hypothesis that social parasites are most closely related to THEIR HOSTS. Electrophoretic data are adduced for nine species of Polistes, including all three known species of social parasites (Sulcopolistes) and THEIR HOSTS. Three different coding methods are investigated; in no case do the social parasites cluster most closely with THEIR HOSTS. Rather, there is limited evidence that they form a monophyletic group. However, formal taxonomic recognition of Sulcopolistes is not justified, as it renders Polistes sensu stricto paraphyletic. Although the social parasites are not most closely related to THEIR HOSTS, hosts and parasites belong in the same subgenus and share many characteristics that may have facilitated the exploitation and deception practised by the parasites on the hosts.
References
Baroni Urbani, C. 1989. Phylogeny and behavioural evolution in ants, with a discussion of the role of behaviour in evolutionary processes. Ethol. Ecol. Evol. 1: 137-168.
Beaumont, J. De. 1944. Les guêpes (Vespa L. s. l.) de la Suisse. Bull. Soc. Vaudoise Sci. Nat. 62: 329-362.
Beaumont, J. De. 1958. Le parasitisme social chez les Guêpes et les bourdons. Mitt. Schweiz. Ent. Ges. 31: 168-176.
Beaumont, J. DE and R., Matthey. 1945. Observations sur les Potistes parasites de la Suisse. Bull. Soc. Vaudoise Sci. Nat. 62: 441-454.
Bequaert, J. 1932. A tentative synopsis of the hornets and yellow-jackets (Vespinae; Hymenoptera) of America. Ent. Amer., n. s., 12: 71-138.
Bischoff, H. 1931. Zur Kenntnis der Gattung Pseudovespa. Sitzungsber. Ges. Naturforsch. Freunde, 16: 329-346.
Blüthgen, P. 1938. Systematisches Verzeichnis der Faltenwespen Mitteleuropas, Skandinaviens and Englands. Konowia, 16: 270-295.
Blüthgen, P. 1943. Die europäischen Polistinen (Hym. Vespidae Vespinae). Arch. Naturges., N.F., 12:94-129.
Bourke, A. and N., Franks. 1991. Alternative adaptations, sympatric speciation and the evolution of parasitic inquiline ants. Biol. J. Linn. Soc. 43: 157-178.
Buckup, P. 1991. Cladogram characters: predictions, not observations. Cladistics, 7: 191-195.
Buschinger, A. 1986. Evolution of social parasitism in ants. Trends Ecol. Evol. 1: 155-160.
Buschinger, A. 1990. Sympatric speciation and radiative evolution of socially parasitic ants- heretic hypotheses and their factual background. Z. Zool. Syst. Evolut.-forsch. 28: 241-260.
Buth, D. G. 1984. The application of electrophoretic data in systematic studies. Annu. Rev. Ecol. Syst. 15: 501-522.
Carlin, N. 1988. Species, kin and other forms of regcognition in the brood discrimination behavior of ants. In: J. C., Trager (ed). Advances in Myrmecology. E. J. Brill, Leiden , pp. 267-295.
Carpenter, J. M. 1981 1982. The phylogenetic relationships and natural classification of the Vespoidea (Hymenoptera). Syst. Ent. 7: 11-38.
Carpenter, J. M. 1987a. Phylogenetic relationships and classification of the Vespinae (Hymenoptera: Vespidae). Syst. Ent. 12: 413-431.
Carpenter, J. M. 1987b. On “The evolutionary genetics of social wasps” and the phylogeny of the Vespinae (Hymenoptera: Vespidae). Insectes Soc. 34: 58-64.
Carpenter, J. M. 1988. Choosing among multiple equally parsimonious cladograms. Cladislics, 4: 291-296.
Carpenter, J. M. 1990. Of genetic distances and social wasps. Syst. Zool. 39: 391-396.
Carpenter, J. M. 1991. Phylogenetic relationships and the origin of social behavior in the Vespidae. In: K. G. Ross and R. W., Matthews (eds). The Social Biology of Wasps. Cornell Univ. Press, Ithaca , pp. 7-32.
Carpenter, J. M. 1992. Random cladistics. Cladistics, 8: 147-153.
Cervo, R., M. C., Lorenzi and S., Turillazzi. 1990a. On the strategies of host nest invasion in three species of Sulcapolistes, social parasites of Polistes wasps. Actes. Coll. Insectes Soc. 6: 69-74.
Cervo, R., M. C., Lorenzi and S., Turillazzi. 1990b. Subcopolistes alrimandibularis, social parasite and predator of an alpine Polistes (Hymenoptera, Vespidae). Ethology 86: 71-78.
Cervo, R. and S., Turillazzi. 1985. Associative foundation and nesting sites in Polities nimpha (Hymenoptera: Vespidae). Naturwissenschaften 72: 48-49.
Clayton, J. W., and D. N., Tretiak. 1972. Amine-citrate buffers for pH control in starch gel electrophoresis. J. Fish. Res. Bd Canada, 29: 1169-1172.
Crother, B. I. 1990. Is “Some better than none” or do allele frequencies contain phylogenetically useful information Cladistics 6: 277-281.
Dani, F.R., and R., Cervo. 1992. Reproductive strategies following nest loss in Polistes gallicus (Hymenoptera, Vespidae). Ethol. Ecol. Evol., Special Issue 2, in press.
Day, M. C. 1979. The species of Hymenoptera described by Linnaeus in the genera Sphex, Chrysis, Vespa, Apis and Mutilla. Biol. J. Linn. Soc. 12: 45-84.
Eck, R. 1979. Biometrische Untersuchung zur Klarung der Artunterschiede bei sozialen Faltenwespen (Hymenoptera: Vespinae). Ent. Abh. Mus. Tierk. Dresden, 42: 315-344.
Emery, C. 1909. Über den Ursprung der dulotischen, parasitischen und myrmekophilen Ameisen. Biol. Centralbl. 29(11): 352-362.
Evans, H. E., and M. J., West-Eberhard. 1970. The Wasps. Univ. Michigan Press, Ann Arbor .
Farris, J. S. 1969. A successive approximations approach to character weighting. Syst. Zool. 18: 374-385.
Farris, J. S. 1981. Distance data in phylogenetic analysis. In: V. A. Funk and D. R., Brooks (eds). Advances in Cladistics: Proceedings of the First Meeting of the Willi Hennig Society. New York Bot, Garden , Bronx , pp. 3-23.
Farris, J. S. 1988. Hennig86, Version 1.5. Program and documentation. Port Jefferson Station, New York .
Farris, J. S. 1989. The retention index and the rescaled consistency index. Cladistics, 5: 417-419.
Feisenstein, J. 1985. Confidence limits on phylogenies: An approach using the bootstrap. Evolution, 39: 783-791.
Fitch, W. M. 1971. Toward defining the course of evolution: Minimum change for a specific tree topology. Syst. Zool. 20: 406-416.
Gamboa, G. 1978. Intraspecific defense: Advantage of social cooperation among paper wasp foundresses. Science, 199: 1463-1465.
Greene, A. 1991. Dolichovespula and Vespula. In: K. G. Ross and R. W., Matthews (eds). The Social Biology of Wasps. Cornell Univ. Press, Ithaca , pp. 263-305.
Guiglia, D. 1948. Le vespe d'Italia. Mem. Soc. Ent. Ital., Fasc. Suppl. 28: 5-84.
Guiglia, D. 1971. Les guêpes sociales (Hymenoptera Vespidae) d'Europe occidentale et septentrionale. Faune de l'Europe et du Bassin Mediterraneen 6. Masson et Cie, Paris .
Hauser, D. L., and W., Presch. 1991. The effect of ordered characters on phylogenetic analysis. Cladistics, 7: 243-265.
Heinze, J. 1991. Biochemical studies on the relationship between socially parasitic ants and their hosts, Biochem. Syst. Ecol. 19: 195-206.
Hölldobler, B. and E. O., Wilson. 1990. The Ants. Harvard Univ. Press, Cambridge. Massachusetts .
Hughes, C. R., and J. E., Strassmann. 1988. Foundress mortality after worker emergence in social wasps (Polistes). Ethology 79: 265-280.
Hunt, J. H., and G. J., Gamboa. 1978. Joint nest use by two paper wasp species. Insectes Soc. 25: 373-374.
Janet, C. 1903. Observations sur les Guêpes. C. Naud, Paris .
Kasuya, E. 1982. Take-over of nests in a Japanese paper wasp, Polistes chinensis antennalis (Hymenoptera, Vespidae). Appl. Ent. Zool. 17: 427-431.
Klahn, J. 1988. Intraspecific comb usurpation in the social wasp Polistes fuscatus. Behav. Ecol. Sociobiol. 23: 1-8.
Klahn, J. and G., Gamboa. 1983. Social wasps: Discrimination between kin and nonkin brood. Science, 221: 482-484.
Kojima, J. and K., Kojima. 1988. Three new species of Polistes Latreille (Hymenoptera: Vespidae) from Papua New Guinea, with notes on the taxonomic status of the subgenus Stenopolistes van der Vecht. J. Aust. Ent. Soc. 27: 69-80.
Lorenzi, M. C., and R., Cervo. 1992. Intraspecific usurpation in Polistes biglumis bimaculatus (Hymenoptera, Vespidae). Ethol. Ecol. Evol., in Press.
Lorenzi, M. C., and S., Turillazzi. 1986. Behavioural and ecological adaptations to the high mountain environment of Polistes Uglumis bimaculatus. Ecol. Ent. 11: 199-124.
MacDonald, J. F., and R. W., Matthews. 1984. Nesting biology of the southern yellowjacket, Vespula squamosa (Hymenoptera: Vespidae): social parasitism and independent founding. J. Kansas Ent. Soc. 57: 134-151.
Makino, S. 1985. Foundress-replacement on nests of the monogynic paper wasp Polities biglumis in Japan (Hymenoptera: Vespidae). Kontyü 53:143-149.
Makino, S. 1989. Usurpation and nest rebuilding in Polistes riparius: Two ways to reproduce after loss of the original nest (Hymenoptera: Vespidae). Insectes Soc. 36: 116-128.
Makino, S. and K. Sayama. 1991. Comparison of intraspecific nest usurpation between two haplometrotic paper wasp species (Hymenoptera: Polistes). J. Ethol. 9: 121-128.
Matsuura, M. 1991. Vespa and Provespa. In: K. G. Ross and R. W., Matthews (eds). The Social Biology of Wasps. Cornell Univ. Press, Ithaca , pp. 232-262.
Matthews, R. W. 1982. Social parasitism inyellowjackets (Vespula). In: P. Jaisson (ed.). Social Insects in the Tropics, Vol. 1. Universite Paris-Nord, Paris , pp. 193-202.
Michener, C. 1974. The Social Behavior of the Bees: A Comparative Study. Harvard Univ. Press, Cambridge , Massachusetts .
Mickevich, M. F. 1982. Transformation series analysis. Syst. Zool. 31: 461-478.
Mickevich, M. F., and M. S., Johnson. 1976. Congruence between morphological and allozyme data in evolutionary inference and character evolution. Syst. Zool. 25: 260-270.
Mickevich, M. F., and C., Mitter. 1981. Treating polymorphic characters in systematics: A phylogenedc treatment of electrophoretic data. In: V. A. Funk and D. R., Brooks (eds). Advances in Cladistics: Proceedings of the First Meeting of the Willi Hennig Society. New York Botanical Garden, Bronx , pp. 45-58.
Mickevich, M. F., and C., Mitter. 1983. Evolutionary patterns in allozyme data: A systematic approach. In: N. I. Platnick and V. A., Funk (eds). Advances in Cladistics 2: Proceedings of the Second Meeting of the Willi Hennig Society. Columbia Univ. Press, New York , pp. 169-176.
Mickevich, M. F., and S. J., Weller. 1990. Evolutionary character analysis: tracing character change on a cladogram. Cladistics, 6: 137-170.
Murphy, R. W., J. W., Sites, Jr., D. G., Buth and C. H., Haufler. 1990. Proteins I: Isozyme electrophoresis. In: D. M. Hillis and C., Moritz (eds). Molecular Systematics. Sinauer Associates, Sunderland , pp. 45-126.
Nixox, K. C. 1991. Clados, Version 1.0. Program and documentation. Trumansburg, New York .
O'Donnell, S. and R. L., Jeanne. 1991. Interspecific occupation of a tropical social wasp colony (Hymenoptera: Vespidae: Polistes). J. Ins. Behav. 4: 397-400.
Ondricek-Fallscheer, R. L. 1992. A morphological comparison of the sting apparatuses of socially parasitic and nonparasitic species of yellowjackets (Hymenoptera: Vespidae). Sociobiol. 20: 245-300.
Pardi, L. 1942. Richerche sui Polistini, V. La poliginia iniziale di Polistes gallicus. Boll. Inst. Ent. Univ. Bologna, 14: 1-104.
Platnick, N.I. 1989. Cladistics and phylogenetic analysis today. In: B. Fernholm, K. Bremer and H. Jörnvall (eds). The Hierarchy of Life. Etsevier, Amsterdam , pp. 17-24.
Reed, H. C. and R. D., Akre. 1982. Morphological comparisons between the obligate social parasite, Vespula austriaca (Panzer), and its host, Vespula acadzca (Sladen) (Hymenoptera: Vespidae). Psyche 89: 183-195.
Reeve, H. K. 1991. Polities. In: K. G. Ross and R. W., Matthews (eds). The Social Biology of Wasps. Cornell Univ. Press, Ithaca , pp. 99-148.
Richards, O. W. 1971. The biology of the social wasps (Hymenoptera, Vespidae). Biol. Rev. 46: 483-528.
Richards, O. W. 1973. The subgenera of Polistes Latreille (Hymenoptera, Vespidae). Rev. Bras. Ent. 17:85-104.
Richards, O. W. 1978. The Social Wasps of the Americas, Excluding the Vespinae. British Museum (Natural History), London .
Röseler, P.-F. 1991. Reproductive competition during colony establishment. In: K. G. Ross and R. W., Matthews (eds). The Social Biology of Wasps. Cornell Univ. Press, Ithaca , pp. 309-335.
Snelling, R. R. 1952. Notes on nesting and hibernation of Polistes (Hymenoptera, Vespidae). PanPacif. Ent. 28:177.
Snelling, R. R. 1954. Wasps of the genus Polistes in California and Arizona (Hymenoptera, Vespidae). J. Kans. Ent. Soc. 27: 151-155.
Spradbery, J. P. 1991. Evolution of queen number and queen control. In: K G. Ross and R. W., Matthews (eds). The Social Biology of Wasps. Cornell Univ. Press, Ithaca , pp. 337-388.
Strassmann, J. E. 1981. Evolutionary implications of early male and satellite nest production in Polistes exdamans colony cycles. Behav. Ecol. Sociobiol. 8: 55-64.
Swofforo, D. L., and S. H., Berlocher. 1987. Inferring evolutionary trees from gene frequency data under the principle of maximum parsimony. Syst. Zool. 36: 293-1525.
Swofford, D. L., and G. J., Olsen. 1990. Phylogeny reconstruction, in: D. M. Hillis and C., Moritz (eds). Molecular Systematics. Sinauer Associates, Sunderland , pp. 411-501.
Taylor, L. H. 1939. Observations on social parasitism in the genus Vespula Thomson. Ann. Enl. Soc. Amer. 32: 304-315.
Turillazzi, S. 1991. The Stenogastrinae. In: K. G. Ross and R. W., Matthews (eds). The Social Biology of Wasps. Cornell Univ. Press, Ithaca , pp. 74-98.
Turillazzi, S. 1992. Nest usurpation and social parasitism in Polistes wasps: New acquisitions and current problems. In: Bitten, J. (ed.). Biology and Evolution of Social Insects, in Press.
Turillazzi, S. and A. Ugolini. 1978. Osservazioni su colonie miste sperimentali di Polistes gallicus (L.) e Polistes foederatus (Kohl) (Hymenoptera, Vespidae). Redia 61: 233-249.
Varvio-Aho, S. I., P., Pamilo and A., Pekkarinen. 1984. Evolutionary genetics of social wasps (Hymenoptera, Vespidae, Vespula). Insectes Soc. 31: 375-386.
Vecht, J. van der and J. M., Carpenter 1990. A catalog of the genera of the Vespidae (Hymenoptera). Zool. Verhand. Leiden, 260: 3-62.
Werdelin, L. 1989. We are not out of the woods yet-A report from a Nobel Symposium. Cladistics, 5: 192-200.
West-Eberhard, M. J. 1986. Alternative adaptations, speciation and phylogeny (a review). Proc. Natl. Acad. Sci. USA, 83: 1388-1392.
Weyrauch, W. 1937. Zur Systematik und Biologie der Kuckuckswespen Pseudovespa, Pseudo-vespula und Psmtdopolistes. Zool. Jahrbüch. 70: 243-290.
Weyrauch, W. 1938. Zur Systematik und Biologic der palearktischen Polistinen. Arb. Physiol. Angewand. Ent. 5: 273-278.
Wheeler, W. M. 1928. The Social Insects. Harcourt, Brace and Co., New York .
Wheeler, W. M., and L. H., Taylor. 1921. Vespa arctica Rohwer, a parasite of Vespa diabolica de Saussure. Psyche, 28: 135-144.
Wilson, E. O. 1971. The Insect Societies. Harvard Univ. Press, Cambridge , Massachusetts .
Wolf, H. 1991. Körperbau und Lebensweise unserer Sozialen Wespen als Ergebniss der Evolution. Verh. West. Ent. Tag. 1990: 45-54.
Yoshikawa, K. 1955. A polistinae colony usurped by foreign queen. Ecological studies of Polistes wasps, II. Insectes Soc. 2: 255-260.
Zikan, J. F. 1949. O gênero Mischocyttarus Saussure, (Hymenoptera, Vespidae), com a descriçâo de 82 espécies novas. Bol. Parque Nac. Itatiaia, 1:1-251.
Zimmermann, K. 1930. Zur Systematik der palearktischen Polistes (Hym. Vesp.). Mitt. Zool. Mus. Berlin, 15: 609-621.