Taxonomy and Distribution of the Boreomontane Shore Bugs Salda Sahlbergi and S
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Taxonomy and distribution of the boreomontane shore bugs Salda sahlbergi and S. henschii (Heteroptera, Saldidae) PER LINDSKOG Lindskog, P.: Taxonomy and distribution of the boreomontane shore bugs Salda sahlbergi and S. henschii (Heteroptera, Saldidae). [Taxonomi och utbredning hos de boreomontana strandstinkflyna Salda sahlbergi och S. hensc'hii (Heteroptera, Saldidae).1 Ent. Tidskr. I l2: l-18.'UmeA. Sweden 1991. ISSN 0013-886x. - Additional taxonomic characters are given and illustrated supporting the concept of Salda sahlbergi Reuter and S. henschii (Reuter) as separate species. Their present assignment to the genus Salda (s. str.) is shown to be poorly substantiated. Salient characters of the male genitalia suggest that they may be more closely related b Teloleuco. S. henschii, hitherto only known from the mountains of Central Europe, is recorded for the first time from Sweden and Eastem Fennoscandia (USSR) in northem Europe. It is here confined to freshwater wetlands in lowland coastal areas within the subboreal (boreo-nemoral) zone. Revised data on the geographic distribution of S. henschii and its boreal sister-species, S. sahlbergi, are summaiized. The latter is for the first time reported from the New World (Canada, Newfound- land). Data on habitat and life cycles are given. The biogeography of this species pair is discussed in some detail with respect to vicariance relationships, range disjunctions, and local historic-ecological phenomena reflected in their Fennoscandian distributions. The former issue is centred on ihe nature and interactions of the Angaran and European areas of ende- mism, the latter on the significance of isostatic land uplift. P. Lindskog, Dept. of Entomology, Swedish Museum of Natural History, P.O.Box 50007, S- 104 05 Stockholm, Sweden. Introduction The Salda group of shore bugs includes 2 I species ferous forests (e.g. Schuh 1967, Brooks & Kelton in three genera: Salda L. (Holarctic), 16 spp.; Te- 1967, Wr6blewski 1966, Lindskog 1975, Cobben loleuca Reuter (Holarctic), 4 spp.; and Lampra- 1 98s). canthia Reuter (Nearctic), I sp. (cf. Schuh et al. Among the nine Sa/da species recorded from 1987). They form a strictly Holarctic-boreal and the Palearctic, S. sahlbergi Reuter and S. henschii temperate group, the species appearing as charac- (Reuter) are somewhat isolated with respect to teristic elements of the saldid fauna of the Nort- various taxonomic characters. These two species hem Coniferous (Taiga) zone and corresponding were earlier placed in the genus Saldula until cor- midlatitude montane life-zones in the Old and rectly transferred to the Salda group (Saldini) by New World. Most species are comparatively Cobben (1959, 1960), primarily on the basis of large, ca 5-7 mm, predominantly dark coloured, unmistakably synapomorphic characters of the and exhibiting alary polymorphism, with a preva- male genitalia. Drake & Hoberlandt (195 1) syno- lence for flightless (semibrachypterous) morphs. nymized S. henschii (=Saldula umbrata SchmidQ Salda a;nd Lampracanthia primarily inhabit fresh with S. sahlbergl, a view also followed by Cobben water marshes, bogs, and damp meadows, inclu- (1959, 1960). More recently Hoberlandt (1977) ding alpine sites, in a few cases also saline forma- restored S. henschii as a separate species. tions. The Teloleuca species are primarily associa- According to present knowledge (Hoberlandt ted with the open sandy-stony banks of streams 1977, Schuh et al. 1987, P6ricart 1990) S. henschii and rivers, partly [7r. pellucens (F.)] also occupy- would be restricted to the mountains of Central ing more terrestrial habitats, as moorlands and Europe (mainly the Alps and the Carpathians), as alpine heaths, or sparsely vegetated ground and opposed to S. sahlbergi being a true boreal species decaying mossy logs in montane and boreal coni- ranging from Scandinavia to the Soviet Far East. 2 Per Lindskog However, I have now confirmed the presence of patterns, including centers of melanization and S. henschii in northem Europe (Fennoscandia), spread of dark pigmentation, have definite rela- and revised various materials identified as S. saftl- tionships to and are compartmentalized by wing bergi from northem Europe and Asia. The new topography, primarily the corial veins and the data on the distribution of these species are given cells enclosed by them. As a first step in estab- here, including the first record of S. sahlbergi lishing a nomenclature for identifying these com- from the New World. As a supplement to the ponents, I provide a figure with names for the cells study by Hoberlandt (1977), who separated saftl- in the saldid corium (Fig. 32). I essentially follow bergi and henschii mainly on differences in hem- the practice in naming insect wing cells according elytral colouration and dorsal pilosoty, I present to the identity of the anterior vein enclosing them. additional and more detailed data on discrimina- The identification of veins follows Polhemus tory characters of these species. Special attention (1985) and Wouon & Beus (1986). This permits is paid to some biogeographical pattems pertai- a topographic definition of components of saldid ning to this species pair. wing patterns by referring to the "radial spot", New important data on the taxonomy and distri- "median eye spot", "basicostal centre of melaniza- bution of the Eurasian species of the Salda group tion", etc. One should note that the preradial vein were recently provided by the late Dr Ren6 H. identifiable as subcosta is only more exceptionally Cobben (Cobben 1985). He deliberately refrained plainly visible or completely developed in the Sal- from considering the present two species in his didae. Generally, the course of Sc and thus the paper, instead referring to my study for further border between the costal and subcostal cells or details (cf. Cobben 1985:262, foot-note). fields (the two terms being used interchangeably here), is indicated as a more or less clearly defined depression of exocorium delimiting the reflexed costal margin (cf. Fig. 32, stippled line), most Material and methods evident in the proximal part of exocorium. This Material has been studied from the following col- typically corresponds to a demarcation line be- lections (abbreviations in parenthesis): Naturhis- tween a different surface texture (e.g. costal field toriska Riksmuseet, Stockholm, Sweden (NRS); shiny /subcostal field dull) or pigmentation of ex- Entomologiska Museet, Lunds Universitet, Lund, ocorium. The latter pattem is especially apparent Sweden (EML); Zoologisk Museum, Oslo, Nor- in the eunomy of species like Saldula opacula way (ZMO); Universitetets Zoologiska Museum, (Zetterstedt) and others, where dark pigment spre- Helsinki, Finland (Zl|lIH); Zoologicheskiy Institut, ads along this line and then gradually expands AN SSSR, Leningrad, U.S.S.R. (ZIL); Laborato- inwards over the subcostal field, leaving a contin- rium voor Entomologie van de Landbouwho- uous and neatly delimited, light costal stripe (e.g. geschool, Wageningen, the Netherlands (WAG); Cobben 1960:figs 142-149). Department of Biology, Nankai University, Tian- jin, Peoples Republic of China (NUT). In addition to the new material from Northem Europe listed further below, S. henschii was also Taxonomy studied from Austria: Nordtirol, Seefeld, Wild- Relationships of the sahlbergi group moosalm, ca I 300 m,5d7?, partly reared from larvae (L5), l.vii.l973, P. Lindskog (NRS); Kiirn- The present paper is an off-shoot of my still ongo- ten, Weissensee, 30.vii. 1958,4629, H.-H. We- ing revision of supraspecific relationships in the ber (NRS). Salda-group and studies of the relationships be- Eunomies of hemelytral pigmentation are at- tween this group and other saldine taxa. Suffice tributed a decisive role as characters in the differ- it to note here that the current assignment of the entiation of species in saldid taxonomy (e.g. Cob- Salda-group to a separate tribe, Saldini, which ben 1960). There is an obvious need, both at would form the sister group of Saldoidini (=Char- taxonomic and evolutionary studies of the group, toscirtini of Cobben) (Cobben 1959, Polhemus for developing a more strict basis for the compara- 1985), is poorly substantiated by cladistic data at tive analysis of saldid wing patterns. Accordingly, hand. The Saldoidini encompasses all remaining it is clear that the various components of these saldine genera minus Salduncula (Saldunculini) Taxonomy of Salda sahlbergi and S. henschii 3 9 Figs l-l 1. Salda group, male endosomal sclerites. 14. Frontal view. 5-8. Lateral view; arrow directed - - (Reuter). anteriorly. - 9-1 1. Basal view. - 1, 5. Salda sahlbergl Reuter. -2,6,9. S. henschii -3,7,10. Teloleuca pellucens (F.). - 4, 8, 11. T. bifasciata (Thomson). Scale line 0.1 mm. Hanens endosomala skleriter. - 14. Framifr6n. - 5-8. FrAn sidan; pilen pekar framit. - 9-1 1. Frin basen. according to the most recent higher classification 1959, 1960), or upholding their status as separate of the Saldidae (Polhemus 1985). Instead, as will genera (Polhemus 1985, Schuh et al. 1987), is of be demonstrated elsewhere, the Saldini is pro- secondary concem here.] I have earlier noted that bably more closely related to some subgroup of S . henschii (Lindskog 197 5:166, as a form of sahl- Saldoidini, the latter thus standing out as paraphy- bergi) more agrees with Teloleuca thanwrth Salda letic (Lindskog & Chen, in prep.). with respect to the structure of the male endoso- One further point concems the actual delinea- mal (phallic) sclerites. The same applies to S. tion of the three genera of the Salda group. [The sahlbergi. Unlike all other Old or New World question of their rank, i.e. either classifying them Sa/da species the upper sclerotized pieces of the as subgenera of the single genus Salda (Cobben complex median sclerite are not united basally 4 Per Lindskog with the inner furciform sclerite in sahlbergi- +henschii, a condition shared with Teloleuca (see A Figs 1, 2,9) and Lampracanthia. (The presence A /,4\ of an inner furciform sclerite is a synapomorphy / //\ of the Salda group.) Further and most importantly, μ the upper pieces are not connected basally in sahl- ヽ bergi+henschii, i.e.