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Colonization Overseas by Long-Range Aerial Drift in a Formicoxenine Ant (Hymenoptera, Formicidae)

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Colonization Overseas by Long-Range Aerial Drift in a Formicoxenine Ant (Hymenoptera, Formicidae) Ent. Tidskr. 136 (2015) Long distance colonisation of an ant Colonization overseas by long-range aerial drift in a Formicoxenine ant (Hymenoptera, Formicidae) BErnhard SEifErT & andErS hagman Seifert, B. & hagman, a.: Colonization overseas by long-range aerial drift in a formico- xenine ant (hymenoptera, formicidae). [Luftspridning över havet av en liten ettermyra (Hymenoptera, Formicidae).] – Entomologisk Tidskrift 136 (1-2): 5-15. Uppsala, Swe- den 2015. iSSn 0013-886x. Temnothorax crassispinus (Karavajev, 1926) has been found new for Sweden on the island “hästnacken” in the Stockholm archipelago. it is morphologically extremely similar to its Swedish sibling T. nylanderi (förster, 1850). Three different exploratory data analysis methods achieved a full species separation with perfectly congruent classifications in a total of 135 nest samples from the entire European range. all Swedish samples of both spe- cies were clearly classified with posterior probabilities of p>0.998 if run as wild-card in a confirmative linear discriminant analysis. The close association of both species to temper- ate Quercus forest allows to reconstruct time and routes of postglacial immigration of both species from an italo-iberian (T. nylanderi) and Balkan (T. crassipinus) refuge. according to this, T. nylanderi entered the Swedish mainland in about 8300 BP (Skåne) and spread north to the Stockholm area until 5000 BP. Simultaneously, the advance of T. crassipinus from SE Europe was stopped by T. nylanderi along a 900-km long front line running from nW Poland through East germany south to Bavaria. Based on arguments from zoogeog- raphy, dispersal behavior, reproduction biology, meteorology and physiology, long-range aerial drift across the Baltic Sea is by far the most probable way for colonizing hästnacken by T. crassipinus. The counter-hypothesis that nests were introduced with firewood needed for a brickyard run on the island between 1720 and 1830 was rejected by historical reports on wood import to the Stockholm area and the low likelihood of nest microhabitats of get- ting fixed to imported wood materials. Bernhard Seifert, Senckenberg Museum of Natural History Görlitz, Am Museum 1, 02806 Görlitz, Germany; post box 300154. E-mail [email protected] Anders Hagman, Nyodlingsvägen 17, 191 43 Sollentuna, Sweden. E-mail: anders.hag- [email protected] The small formicoxenine ants Temnothorax sibling species having spread from an ibero-ital- nylanderi (förster, 1850) and T. crassispinus ian and Balkan Pleistocene refuge respectively. (Karavajev, 1926) are dominant elements of the These refuge areas probably have been used with forest floor fauna of European temperate wood- similar topography and migration routes during land biomes. T. nylanderi is widely distributed all glaciations since then. data of mtdna (Pusch throughout deciduous forests in Western Europe, et al. 2006a) suggest the splitting of both species whereas T. crassispinus inhabits similar habitats in separate glacial refuges to have occurred al- in Eastern Europe (Seifert 2007). The two species ready in Early Pleistocene ± 1.4 myr BP and very have repeatedly stood in the focus of studies after low within-species sequence divergence (0.14% Seifert (1995) presented evidence of their hetero- in T. nylanderi and 0.29% in T. crassispinus) in- specificity and showed that they form parapatric dicates a rapid postglacial spreading. The species 5 Bernhard Seifert & Anders Hagman Ent. Tidskr. 136 (2015) are ecologically most similar, vicariant species nior author who confirmed that these belonged that have met in germany along a 900-kilome- to T. crassispinus. ters-long front line latest during the atlantic This paper aims to give firm evidence for the some 7500 years BP (Seifert 1995). This front occurrence of T. crassispinus in Sweden, to im- line currently runs from nE germany (anklam) prove the knowledge on the northern distribu- south to Bavaria (münchen). The depth of inter- tion of T. nylanderi and to show that long-range specific geographic overlap along the front line aerial drift across the Baltic Sea is a most prob- is not larger than 25 km whereas the east-west able way for extension of geographical range in extension of both species’ range is 3600 km at these small ants. least. despite frequent occurrence of f1-hybrid samples T. crassispinus x nylanderi within this Material and Methods narrow contact zone (Seifert 1995, Pusch et al. 2006a), the absence of backcrosses saves the Material investigated genetic integrity of both species (Seifert 2007). a total of 77 nest samples of T. crassispinus with recently, a third cryptic species from SE Europe 222 worker individuals has been investigated – T. crasecundus Seifert & Csösz, 2015 – has morphometrically. They originated from austria been added to the T. nylanderi species complex (12 samples), Bosnia (2), Bulgaria (2) Czechia (Seifert & Csösz 2015). (3), germany (37), greece (8), italy (2), Poland T. nylanderi is the only species of this com- (5), Slovenia (2), Sweden (2) and the Ukraine plex reported so far from Sweden and was hith- (3). in T. nylanderi, 58 nest samples with 151 erto known to occur here from Skåne north to worker individuals were investigated. These Östergötland and dalsland (douwes 2012). originated from austria (1 sample), france (3), however, our own investigations showed its germany (39), italy (10), Poland (1), Sweden distribution north to Uppland at least (näsud- (3) and Switzerland (1). hybrid samples from den). The population may reach extremely high the contact zone in germany were excluded nest densities of up to ten nests per square meter from analysis. (e.g. Trollskogen in the north of Öland). Colonies of T. nylanderi and T. crassispinus Equipment and measuring procedures consist of a few dozen workers and a single Spatial positioning of specimens was performed queen (monogyny) and inhabit any microspace by a pin-holding stage, permitting full rotations offering adequate microclimate. This is as a rule around X, Y, and Z axes and a Leica m165C on soil surface and rarely basally at tree stems up high-performance stereomicroscope equipped to 30 cm above ground. most frequent nesting with a 2.0 planapochromatic objective (resolu- substrates are dead wood, hollow acorns, nuts or tion 1050 lines/mm) was used at magnifications galls and bark, occasionally also leaf litter, snail of 120-384x. The mean relative measuring er- shells or soil under stones down to 4 cm depth ror over all magnifications was 0.3%. a Schott (Seifert 2007). however, T. crassipinus has been KL 1500 cold-light source equipped with two shown recently to nest locally in tree canopies flexible, focally mounted light-cables, provid- (Seifert et al. 2013). ing 30°-inclined light from variable directions, T. crassispinus has never been reported from allowed sufficient illumination over the full Sweden before and was not expected to occur magnification range and a clear visualization of here for strong zoogeographical arguments. Yet, silhouette lines. a Schott KL 2500 LCd cold- during a visit to an island in Stockholm archi- light source in combination with a Leica coax- pelago, hästnacken, 19 July 2012, the junior ial polarized-light illuminator provided optimal author examined a rotten oak twig lying on the resolution of tiny structures and microsculpture ground. The ants inhabiting the twig seemed, at at highest magnifications. Simultaneous or al- first glance, to be T. nylanderi. however, when ternative use of the cold-light sources depend- compared with other T. nylanderi collected on ing upon the required illumination regime was Öland, they differed by much stronger propode- quickly provided by regulating voltage up and al spines. a few specimens were sent to the se- down. a Leica cross-scaled ocular micrometer 6 Ent. Tidskr. 136 (2015) Long distance colonisation of an ant with 120 graduation marks ranging over 52 % spines diverge continuously from the tip to the of the visual field was used. To avoid the paral- base, a smallest distance at base is not defined. lax error, its measuring line was constantly kept in this case, SPBa is measured at the level of the vertical within the visual field. bottom of the interspinal meniscus. SPST – distance between the centre of propodeal stigma and spine tip. The stigma centre refers to Method of species delimitation the midpoint defined by the outer cuticular ring Species delimitation was performed on the basis but not to the centre of real stigma opening that of 18 primary morphometric characters. in bilat- may be positioned eccentrically. erally developed characters, arithmetic means of SPTI – the distance of spine tips in dorsal view; if both body sides were calculated. all measure- spine tips are rounded or truncated, the centers of ments were made on mounted and fully dried spine tips are taken as reference points. specimens. measurements of body parts always refer to real cuticular surface and not to the dif- Explorative and supervised data analyses and fuse pubescence surface. Unambiguous charac- classification methods ter definitions with figures are given in Seifert & Before the data were introduced into the ex- Csösz (2015). here we only mention the char- plorative and hypothesis-driven data analyses, acters and describe them with a few words (the removal of allometric variance (raV) was per- four characters needed in the simplified discrim- formed with the procedure described by Seifert inant function are defined thoroughly): (2008). The delimitation of the cryptic species was done by an interaction of nest-Centroid CL – maximum cephalic length in median line. Clustering (nC clustering) and a confirmative CS – cephalic size; the arithmetic mean of CL and linear discriminant analysis (Lda). nC Cluster- CW. ing was run both as hierarchical nC-Ward clus- CW – maximum cephalic width across (including) tering and non-hierarchical nC-K-means clus- eyes. tering. These methods were described in more EYE – the arithmetic mean of the large (EL) and small diameter (EW) of the elliptic compound detail by Seifert et al.
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