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A Biochemical Approach Toward the Systematics of the Leptothorax "Muscorum" Group in North America (Hymenoptera" Formicidae)

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A Biochemical Approach Toward the Systematics of the Leptothorax Biochemical Systematics and Ecology, Vol. 17, No. 7/8, pp. 595-601, 19~9. 0305-1978/89 $3.00 + 0.00 Printed in Great Britain. © 1989 Pergamon Press plc. A Biochemical Approach Toward the Systematics of the Leptothorax "muscorum" Group in North America (Hymenoptera" Formicidae) JORGEN HEINZE Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, U.S.A.* Key Word Index--Leptothorax "muscorum"-complex; Leptothoracini; Formicidae; Hymenoptera; genetic variability; electro- phoresis. Abstract--Enzyme patterns have been used to distinguish among a number of morphologically very similar ant species belonging to the Leptothorax "muscorum" group in North America. Not counting the already described species L. retractus, L. sphagnicolus and L. crassipilis, the complex apparently consists of at least three or four more different taxa. Introduction As part of a taxonomic revision of this com- Ever since Provancher described the myrmicine plex, two species have recently been described: ant Leptothorax canadensis about 100 years ago, L. retractus, a common ant throughout boreal taxonomists have disagreed on the systematic North America, and L. sphagnicolus, known only position of this and related taxa from North from some spruce-bogs in Quebec [5]. In most America. Ants belonging to the myrmicine areas of North America at least two additional genus Leptothorax, subgenus Leptothorax (s.str.) taxa can be distinguished, which have been M. R. Smith (=Myrothorax Ruzsky) are rather referred to as "small brown" and "large black" inconspicuous and small in size. They nest in Leptothorax "muscorum" in Alberta [3, 8] or preformed cavities in decaying wood, in bark, Leptothorax spp. A and B in Quebec and New under stones, and in moss in boreal and alpine, England [6, 7]. It is still unknown whether large mostly coniferous forests. Because intraspecific black Leptothorax from Quebec are conspecific variablity is often high, making distinctions with the large black ants from Alberta and between species based on morphological Colorado, or whether small brown specimens characteristics is frequently difficult. from Alaska belong to the same taxon as small In Eurasia, four non-parasitic species are brown L. "muscorum" from New Hampshire. known: Leptothorax acervorum (Fabricius), L. Material has almost exclusively been collected muscorum Nylander, L. gredleri Mayr and L. in the Rocky Mountains and in Northeast scamni (Ruzsky). Specimens from North America, and records from the connecting America have been treated as subspecies of areas, e.g. the Canadian Northwest Territories or palaearctic L. acervorum and L. muscorum by northern Ontario, are rare [1, 2]. some authors, as varieties of L. canadensis, or as Though the characterizations "large black" or additional species by others (for a detailed "small brown" suggest reliable differences in review see [1]). Brown's synonymization of all size and color, more intensive morphological North American taxa with the palaearctic L. and ethological studies indicate that some large muscorum [2] certainly does not reflect the black ants actually are "small brown" L. actual situation. In recent publications several muscorum or that bicolored, small black, and authors have agreed that the subgenus Lepto- large brown "Leptothorax" (s.str.) from thorax (s.str.) comprises a number of different Northeast America usually can be sorted into species in North America [3-7]. one of the two taxa. In a number of ant genera biochemical markers have been used for the definition of *Present address: Zool. Inst. I1., D8700 WLirzburg, F.R.G. species in sibling complexes such as Irido- (Received 13 June 1989) myrmex "humilis" [9], Aphaenogaster "rudis" 595 596 JORGEN HEINZE [10], and Rhytidoponera "irnpressa" [11 ]. Heinze colonies could be collected, there is a chance and Buschinger [7] have shown that esterase that some principally detectable variation has stains of crude Leptothorax homogenates, been overlooked. The probability (P) of missing separated by IEF, can help to distinguish a rare allele with the frequency a in a small between morphologically similar taxa such as L. sample of n diploid individuals is muscorum and L. gredlerL The esterase "finger- P= (l--a) 2n [13]. prints" of species belonging to the North Ameri- can L. "muscorum "/L. canadensis complex, However, if n whole colonies rather than n however, are very similar and differences are individuals are used to collect data, the expo- difficult to interpret. In addition, reliable results nent becomes 3n. The probability of an allozyme could be obtained only with white pupae, which with a frequency of 0.2 to remain undetected if a are not always available. sufficient number of workers from five colonies To find enzyme systems that can be visualized is studied, therefore, is about 3.5%, and the per- in tissue-homogenates from adult workers and centage is even smaller if the colonies contain that can be used for the distinction between several queens (polygyny). Though additional different taxa of L. "muscorum", samples of allozymes might have been missed in some Leptothorax (s.str.) from various populations in populations in this study, it is improbable that North America have been studied with electro- the observed specific differences of IDH (isocitric phoresis in cellulose acetate plates. dehydrogenase, EC 1.1.1.42), MDH-1, and PGD (6- phosphogluconic dehydrogenase, EC 1.1.1.44) in Results and Discussion the various taxa are accidental. In a sample of 20 The results of this study demonstrate that colonies the chance that rare allozymes with a enzyme data can be used to define taxa of Lep- frequency of less than 0.05 are not found is tothorax (s. str.); they also show, however, that below 5%. AIIozymes that occur only with fre- this subgenus is far more complex in North quencies less than 0.05 have often been America than in Europe. neglected in comparative studies and have been Of about ten loci that could be reliably stained defined as "absent" [14]. in homogenates of individual adult Leptothorax If data from all populations are compared, it workers, only ME (malic enzyme, EC 1.1.1.40) becomes likely that in spp. A and B, and prob- and MDH-2 (malic dehydrogenase, EC 1.1.1.37) ably also in additional species, MDH-1, IDH and showed considerable variation within species. In PGD are almost monomorphic within species, as a sample of about 20 colonies of large black L. they are in L. acervorum from southern Sweden "muscorum" (sp. B) from Mount Monadnock, (Douwes, personal communication). PGI (phos- New Hampshire, the most extensively studied phoglucose isomerase, EC 5.3.1.9) is known to population, two different electromorphs of be polymorphic in L. acervorum [15]; however, MDH-2 and three, perhaps four of ME could be in various taxa from North America, variability in separated by electrophoresis in cellulose acetate this enzyme could not be found. plates. Similar variability was found in small The low gene diversity in social Hymenoptera brown L. "muscorum" (sp. A) from Mount was explained by the influence of sociality on Monadnock and from Tadoussac, Quebec. Addi- the effective population size (e.g. [16]). In some tional data indicate that these two enzymes are species, such as the sweat-bee Lasioglossum polymorphic also in other taxa of the subgenus zephyrum [17] or the parasitic ant Epirnyrma Leptothorax (s.str.). In Harpagoxenus sublaevis, a kraussei (Jessen and Klinkicht, in preparation), palaearctic slave-making ant closely related to no variation at all could be detected by electro- this subgenus, MDH-2 and ME have also been phoresis, and in numerous species belonging to found to be variable [12], and at least MDH-2 is the subgenus Leptothorax (Myrafant) most of the variable in L. acervorum (Douwes, personal com- five examined loci were found to be mono- munication). morphic within species [18]. It thus is not Other enzyme systems show little or no surprising that most of the studied enzymes are intraspecific variability in Leptothorax (s.str.). more-or-less constant either within taxa (MDH-1, Since in a number of populations, only few PGD, IDH and, perhaps, PGI), or within the SYSTEMATICS OF THE LEPTOTHORAX "MUSCORUM" GROUP IN NORTH AMERICA 597 whole subgenus (SOD, superoxide dismutase, Quebec (B), Alberta (D), Colorado and New EC 1.15.1.1; LDH, lactic dehydrogenase EC Mexico (E), nor between small brown Lepto- 1.1.1.28). thorax A from Northeast America, C from The low variability of MDH-1, IDH and PGD in Alberta and H from Colorado. Identity of enzyme Leptothorax (s.str.) from North America makes it patterns, especially when based on a small possible to define taxa by species-specific sets amount of data, does not necessarily imply that of enzymes. The results of isoenzyme studies two taxa are conspecific. It still remains to be strongly support provisional distinctions based settled whether large black B from Northeast on morphological, ethological and karyological America is the same species as large black D characters. and E from the Rocky Mountains. The three taxa As can be seen in Table 1, taxa are charac- are very similar in morphology and biology too. terized by one of four different combinations of Both Leptothorax A and B can be found in coni- fast (f), slow (s), and very slow (v) migrating ferous forests throughout Northeast America. MDH-1, IDH and PGD electromorphs. Though Whereas in New Hampshire and Massachusetts each taxon does not have its own specific com- they are common only in the mountains (Mount bination, the following distinctions become Greylock, Mount Monadnock), in Quebec or possible: northern Maine they nest at lower elevations or (1) L. retractus (f/s/f) can be easily distin- directly at the coast, too (Tadoussac, Harpswell, guished from the morphologically similar small Reid State Park). Leptothorax C and D have been brown Leptothorax (f/f/s) of Northeast America collected at various elevations (1000-2000 m) in and the Canadian Rocky Mountains.
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