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Novel Blend of Life History Traits in an Inquiline Ant, Temnothorax Minutissimus, with Description of the Male (Hymenoptera: Formicidae)

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Novel Blend of Life History Traits in an Inquiline Ant, Temnothorax Minutissimus, with Description of the Male (Hymenoptera: Formicidae) Myrmecologische Nachrichten 6 67 - 76 Wien, Dezember 2004 Novel blend of life history traits in an inquiline ant, Temnothorax minutissimus, with description of the male (Hymenoptera: Formicidae) Alfred BUSCHINGER & Timothy A. LINKSVAYER Abstract Forty-five colonies of Temnothorax curvispinosus (MAYR, 1866) containing the rare inquiline ant T. minu- tissimus (SMITH, 1942) have been collected near Bloomington, Indiana, USA. The colonies were censused and some were kept in laboratory culture. Dealate and alate females of T. curvispinosus and T. minutis- simus were dissected. The parasite females have a total of six ovarioles. Both, T. minutissimus and the host species, are facultatively polygynous, and the parasite is host-queen tolerant. Alate males were found in August and September. High numbers of alate and dealate, mated but not yet reproductive, young females of T. minutissimus were present in colonies collected in early spring, a feature which had been known as "Intranidal Mated Offspring Hibernation" (IMOH). Mated young queens of T. minutissimus seem to dis- perse in spring to invade host colonies. Apparently they are accepted quite easily by host colonies. Rearing of colonies collected in the early spring, or hibernated in the laboratory, yielded first a brood of sexuals of T. curvispinosus, and subsequently considerable numbers of gyne pupae of T. minutissimus appeared. Only very few males were produced (sex ratio about 0.1 ♂/♀). Intranidal mating attempts were observed, and newly mated young females were detected in colonies having reared gynes and males of T. minutissimus. Life history of the species thus is a novel combination of traits found in different other parasitic ant species: Intranidal mating and IMOH as in a few European "degenerate slavemakers" of the genus Myrmoxenus, but the parasite is host-queen tolerant, as is the case in two of the three European inquiline species of Lepto- thorax (former Doronomyrmex). The development of the parasites after the host species sexuals is a novel trait. The male of T. minutissimus is described. It is characterized by a reduced number of antennomeres (9-11 instead of 12), and a certain "morphological feminization". Key words: Temnothorax minutissimus, Temnothorax curvispinosus, inquilinism, intranidal mating, female-biased sex ratio, IMOH, morphological feminization of male Prof. Dr. Alfred Buschinger, Rossbergring 18, D-64354 Reinheim, Germany. E-mail: [email protected] (contact author) Timothy A. Linksvayer, Department of Biology, Indiana University, Bloomington, IN 47405, USA. Introduction "Social parasitism" occurs when a parasitic social in- In temporary social parasitism, a young parasite sect species depends on another, usually closely re- queen penetrates a host nest, replaces the host queen, lated social insect species for at least one stage in and rears her first brood with the aid of the remain- colony development. A parasite-host relationship be- ing host workers. Later, after the natural death of tween two closely related species may frequently the host workers, such colonies have numerous wor- originate through intraspecific parasitism (e.g., BU- kers of the parasitic species, living similarly to in- SCHINGER 1990, BOURKE & FRANKS 1991, SAVO- dependent species. A second group of social para- LAINEN & VEPSÄLÄINEN 2003). However, social para- sites practice dulosis, and are the slavemakers. Again, sitism in the broad sense also includes the "guest the young queen replaces the host colony queen(s), ants" that live together with only distantly related host usually by force, and in some species the queen also species (e.g., the myrmicine genus Formicoxenus with kills or ejects all host workers, taking over only the the other myrmicine genera Myrmica and Manica or host colony brood from which her first "slaves" with the formicine genus Formica, respectively). Sev- emerge. Slavemaker workers also develop, but they eral major types of social parasitism in the strict are specialized on slave raiding and cannot forage or sense have been described, including temporary so- rear the slavemaker's broods. Instead, they attack cial parasitism, dulosis, and inquilinism. neighboring nests of the host species from where they retrieve worker pupae, thus replenishing the stock of slaves in the slavemaker colony. The third group of social parasites, the "inqui- lines", are species that usually have lost their own worker caste. Mated young queens, by one or the other technique, join the host colonies in which they typically coexist with the host species queen(s). A few inquiline species are dubbed "host queen intol- erant" because they are never found in queenright host nests. A surprising wealth of morphological and be- havioral particularities has been found in such so- cial parasites, and nearly each species, or at least each genus of parasitic ants exhibits its own blend of techniques to ensure its survival and to exploit Fig. 1: Part of a Temnothorax curvispinosus colony with the social characteristics of its respective host spe- workers and a large queen, and a few alate gynes of the cies. Some species, instead of mating during a nup- inquiline, Temnothorax minutissimus. tial flight, mate inside the mother nests and thus are adelphogamous. Hibernation of dealate, mated, young T. minutissimus have also been collected in Michi- queens is observed in some of these species making gan (ALLOWAY undated), West Virginia (S. Foitzik, the colonies seemingly polygynous during parts of the pers. comm.), New York (COVER 1996) and Ohio annual cycle. Wingless, workerlike females may oc- (HERBERS 2004). Apparently, multiple gynes of T. cur, and the sex ratio may be considerably female- minutissimus can be found in a host species colony, biased due to reduced male production. Both host and but only the "socially dominant" female lays eggs, inquiline species may be monogynous or facultative- and this individual spends most of its time riding on ly polygynous, etc.; for details, particularly referring the back of the host queen (ALLOWAY undated). Males to the various hypotheses on the evolution of social of T. minutissimus apparently are produced in low parasitism, see BOURKE & FRANKS (1991), BUSCHIN- numbers (HERBERS 2004). In this paper, we report GER (1986a, 1990, 2002) and HÖLLDOBLER & WIL- collection data and further observations from 45 colo- SON (1990). nies of T. curvispinosus with T. minutissimus, and In this paper, we will investigate a poorly known we provide an as yet lacking description of the male inquiline species of which we recently were able to of T. minutissimus. gather a number of colonies from the field, Temno- thorax (= Leptothorax, subgenus Myrafant) minutis- Material and methods simus (SMITH, 1942). Our experiments and observa- A total of 45 T. curvispinosus colonies with T. minu- tions suggest that this species exhibits a novel blend tissimus were collected from June 2002 to August of life history traits. 2004 (Tab. 1) from the Griffy Nature Preserve and Temnothorax minutissimus has been described by the adjacent Indiana University Griffy Woods Re- SMITH (1942) after four females from the collec- search and Teaching Preserve, near Bloomington, In- tions of the United States National Museum of Nat- diana, USA. For laboratory rearing, proven tech- ural History, Washington DC, USA (USNM). Be- niques were applied as described by BUSCHINGER cause the females were in the same series with sever- (1995). Dissections of females were made according al workers of T. curvispinosus (MAYR, 1866), the au- to BUSCHINGER (1968) and ALLOWAY & al. (1982). thor suggested that the new species is an inquiline Laboratory colonies were checked once a week, ex- of the latter. cept for the time of hibernation. Since we lacked T. Until recently, practically nothing new has been minutissimus colonies of similar size and composi- published on the species' life history. HEINZE (1989) tion in large numbers, the laboratory studies were mentions preliminary enzyme-electrophoretic results designed in a way that a maximum of information suggesting a close relationship of T. minutissimus, T. could be gained from each single colony. Therefore duloticus (WESSON, 1937) and Protomognathus ame- the fate of a number of colonies is described in de- ricanus (EMERY, 1895) to their hosts. Temnothorax tail (Appendix). duloticus and P. americanus are slavemaker species that both enslave T. curvispinosus and in addition a Results few other species of Temnothorax. A couple of web- Temnothorax minutissimus is a rare workerless para- sites refer to new records and provide a few more bio- site of T. curvispinosus. For example, among 556 logical observations of T. minutissimus. and 610 colonies of T. curvispinosus collected for Besides the original collection from Washington different studies in 2003 and 2004, only 18 (3.3 %) D.C. (SMITH 1942), T. curvispinosus colonies with and 14 (2.3 %), respectively, were parasitized by T. 68 Tab. 1: Details of colonies of Temnothorax minutissimus when collected, and production in laboratory culture. NA is entered when initial collection data is not available. w = workers; Lv = larvae; p = pupae; P. am. = Protomognathus americanus. Coll. date Colony # T. curvisp. T. minut. ♀♀ T. minut. Notes T. minut. ♀♀/ w/ Lv Dealate/ alate ♂♂ Production (♀/♂) 26 Jun 02 251 1/ 41/ 45 ca 3 / 0 0 See Appendix 18 / 1 26 Jun 02 267 2/ 41/ 67 ca 2 / 0 0 30 Sep 02 302 1/ 18/ 10 1/ 1 1 30 Sep 02 308 0/ 17/ 10 5/ 0 0 30 Sep 02 317 1/ 76/ 11 0/ 2 0 30 Sep 02 321 1/ 35/ 7 0/ 6 0 30 Sep 02 326 0/ 32/ 7 3 / 4 0 See Appendix 0 / 0 15 Mar 03 519 1/ 228/ 56 11/ 8 0 See Appendix 65 / 6 15 Mar 03 521 1/ 76/ 57 3/ 3 0 See Appendix 6 / 1 10 Apr 03 883 1/ 34/ 33 19/ 9 0 See Appendix 5 / 1 10 Apr 03 884 1/ 30/ 20 42/ 45 0 See Appendix 4 / 0 10 Apr 03 891 1/ 65/ 11 8/ 0 0 10 Apr 03 898 4/ 161/ 121 1/ 0 0 10 Apr 03 900 0/ 75/ 40 1/ 0 0 + 1 w P.
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