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Angelo Bolzern - One of the best experts on this subject based on the ideXlab platform.
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Revision of the Nearctic Eratigena and Tegenaria species (Araneae: Agelenidae)
The Journal of Arachnology, 2016Co-Authors: Angelo Bolzern, Ambros HänggiAbstract:Abstract Based on specimens from several museum collections and recently sampled spiders during a field excursion to Mexico in 2014, the 11 species of Tegenaria s. l. endemic to the United States of America and Mexico are revised. Morphological characters and mitochondrial DNA sequences (CO1, NADH1, 16S) serve as the basis for proposed new combinations and new species. Tegenaria chiricahuae Roth, 1968 remains the only endemic Tegenaria species in the Western Hemisphere. All other specific names (T. blanda Gertsch, 1971, T. caverna Gertsch, 1971, T. decora Gertsch, 1971, T. flexuosa F.O. Pickard-Cambridge, 1902, T. florea Brignoli, 1974, T. gertschi Roth, 1968, T. mexicana Roth, 1968, T. rothi Gertsch, 1971, T. selva Roth, 1968, and T. tlaxcala Roth, 1968) are transferred to the genus Eratigena Bolzern, Burckhardt & Hanggi, 2013. Six new species are described: E. edmundoi, E. fernandoi, E. guanato, E. queretaro, E. xilitla, and E. yarini. In addition, females of E. flexuosa, and E. gertschi, and the male o...
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phylogeny and taxonomy of european funnel web spiders of the Tegenaria malthonica complex araneae agelenidae based upon morphological and molecular data
Zoological Journal of the Linnean Society, 2013Co-Authors: Angelo Bolzern, Daniel Burckhardt, Ambros HänggiAbstract:The taxonomy and systematics of European house spiders, currently constituting the ill-defined Tegenaria−Malthonica complex (including Aterigena) in the family Agelenidae, are revised. In Europe four monophyletic genera and 81 species are defined. One genus, Eratigena gen. nov., and seven species are described as new; at species level 17 new synonyms and 20 new combinations are proposed, and the original combination of 14 species is reinstated. Five species could not be placed (incertae sedis) because of insufficient material and one taxon is regarded as ‘nomen dubium’. On the basis of a detailed morphological assessment, 88 characters were chosen for a cladistic analysis. Phylogenetically informative characters include mostly spination patterns as well as spinneret and genital structures. In addition to morphology, three gene sections [cytochrome c oxidase subunit 1 (CO1), nicotinamide adenine dinucleotide dehydrogenase subunit 1 (NADH1) 28S] were analysed. Morphological and molecular analyses were performed individually and in combination applying maximum parsimony and Bayesian tree search methods. In all resulting trees Malthonica and Tegenaria in their present composition are either polyphyletic or paraphyletic. Consequently, we redefined the two genera and erected a new genus, Eratigena gen. nov. Identification keys are provided for the European agelenid genera as well as for the European species of Tegenaria and Eratigena gen. nov. The genera and most of the constituent species are described and illustrated. The new classification has also been applied to some extra European members of the Tegenaria-Malthonica complex resulting in additional three new synonyms, seven reversals to the original combination, and four new combinations. © 2013 The Linnean Society of London
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A New Funnel-Web Spider Species (Araneae: Agelenidae, Tegenaria) from Mercantour National Park, France
Arachnology, 2010Co-Authors: Angelo Bolzern, Christophe HervéAbstract:Summary During a survey of the spider fauna of the Mercantour National Park in the French Alps, several interesting and new taxa were found. One of these new taxa, Tegenaria mercanturensis n. sp. (Agelenidae) is described here. The new species shows distinct morphological characters which allow easy separation from all other Tegenaria or Malthonica species. The most important characters in males are the short, truncated embolus (a character shared with T. domestica, the type species of Tegenaria, and T. mirifica), the broad, curved hammerhead-shaped conductor and the RTA. Distinct characters in females are the atrium with wide spiral copulatory openings anterior to an anchorshaped sclerotised ground plate, the strongly sclerotised and irregularly formed “shield” around the tubular-shaped spermathecae, and the conspicuous anterolateral lentiform areas of the spermathecae, covered only by membrane.
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Aterigena, a new genus of funnel-web spider, shedding some light on the Tegenaria-Malthonica problem (Araneae: Agelenidae
2010Co-Authors: Angelo BolzernAbstract:(from Malthonica). The latter two species were originally described in Tegenaria. The new genus is diagnosed by the unique combination of several morphological character states (e.g., notched trochanters III and IV, lateral spines on patellae, shape of vulvae). The monophyly of the new genus is also supported by a molecular analysis based on CO1 sequences of several taxa. Keys are provided for the identification of the recognized genera of Tegenariini and the species of Aterigena n. gen. Several species of PseudoTegenaria Caporiacco 1934, originally described in Tegenaria, are morphologically close to Tegenaria tridentina L. Koch 1872, a species that is grouped in the cladistic analysis using CO1 in the monophyletic taxon ‘‘Tegenari
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Taxonomy and phylogeny of the "Tegenaria-Malthonica"-complex (Araneae, Agelenidae), using morphological and molecular data
2010Co-Authors: Angelo BolzernAbstract:Agelenidae (Araneae) constitutes a spider family currently comprising 42 genera and 514 described species (Platnick, 2010). In Europe Tegenaria and Malthonica are the most species rich genera of the family. The latest version of “World Spider Catalogue” (version 10.5) lists 32 Malthonica and 55 Tegenaria species (plus 1 subspecies). Based upon available specimens from museum collections and field excursions, the taxonomy of the Tegenaria/Malthonica-complex is investigated applying morphological and molecular methods. Parsimony and Bayesian analyses, performed individually or with combined data, of three gene sections (CO1, NADH subunit 1, 28S) and 88 morphological characters showed that the genera Malthonica and Tegenaria, in their present compositions, are poly- or paraphyletic. The constituent taxa of the Tegenaria/Malthonica-complex are form four monophyletic clades. The four clades are recovered in all analyses and are well supported by posterior probability, jackknife or bremer support. The relationships between the four clades are not resolved and remain unknown. Based upon these phylogenetic results a merging of all four clades to one monophyletic group is not possible because then other well established groups (e. g. Histopona or genera of Textricini or Agelenini) would have to be included too. The four clades are attributed, therefore, generic: Tegenaria, Malthonica, Aterigena n. gen. and Eratigena n. gen. While the generic concept defintion for Malthonica by Barrientos & Cardoso (2007) is supported by the present analyses, Tegenaria has to be redefined and the new genera Aterigena n. gen. and Eratigena n. gen. are added here. In addition, eleven species (one in Aterigena n. gen., two in Eratigena and eight in Tegenaria) and the males of three species are newly described and 17 new synonyms are proposed here. The new generic concepts necessitate numerous new or revised combinations (rev. stat. or n. comb.). Due to the lack of specimens or diagnostic descriptions, four species remain unplaced and one taxon is regarded as “nomen dubium”. Additional information is given and taxonomic changes are proposed for some extra European Malthonica and Tegenaria species. For most of the included European taxa, supplementary information or redescriptions with figures are provided. In addition, dichotomous keys for the European agelenid genera and for all Aterigena n. gen., most European Eratigena n. gen. and Tegenaria species are provided.
Marie Trabalon - One of the best experts on this subject based on the ideXlab platform.
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fatty acids mediate aggressive behavior in the spider Tegenaria atrica
Chemoecology, 2005Co-Authors: Grégory Pourié, Fernando Ibarra, Wittko Francke, Marie TrabalonAbstract:The aggressiveness of the house spider (Tegenaria atrica) has a chemical basis in terms of intra- and interspecific chemical communication. Vibrations are used for longer range detection of intruders on the web territory whereas cuticular lipids mediate shorter range agonistic behaviors. Two cuticular acids were shown to promote aggressive responses from adult spiders, whereas mixtures of cuticular alkanes acted as inhibition signals.
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plasticity of agonistic behaviour in relation to diet and contact signals in experimentally group living of Tegenaria atrica
Chemoecology, 2001Co-Authors: Grégory Pourié, Marie TrabalonAbstract:The aim of this work was to investigate whether or not the predominant aggressive behaviour in spiders is stereotypic and whether there is a relation with cuticular chemicals. We compared the intraspecific behaviour of adult females of Tegenaria atrica reared in isolation and grouped. Some plasticity emerges from environmental conditions and allows spiders to tolerate each other. This switch of behaviour, in favour of “acceptance behaviour” vs. “attack”, is promoted by an increase in food resources. Learning during ontogenesis and changes in cuticular lipids could prolong it. A pattern of 6 compounds distinguishes grouped from solitary spiders. We hypothesize that a similar evolution could in part explain the emergence of certain communal species of spiders.
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RELATIONSHIPS AMONG FOOD AND CONTACT SIGNALS IN EXPERIMENTAL GROUP-LIVING YOUNG OF Tegenaria ATRICA
Archives of insect biochemistry and physiology, 1999Co-Authors: Grégory Pourié, Marie TrabalonAbstract:Relationships between the quantity of diet, and group vs. individual rearing toward ontogenesis on degree of cannibalism and cuticular lipid profile of young Tegenaria atrica Araneae, Agelenidae were studied. Ad libitum diet increases the development rate and decreases cannibalism between conspecifics compared to a reduced diet. Individually reared young on the two diets showed no quantitative differences in cuticular lipid profile, but differences were observed in individual vs. group rearing modes. The ad libitum-fed grouped young spiders had notably increased quantities of palmitic acid, 13,17-,11,17-, and 9,17-dimethylhentriacontane, methyl oleate, and n-heptatriacontane over the reduced diet, cannibalistic group. Arch. Copyright 1999 Wiley-Liss, Inc.
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contact sex signals on web and cuticle of Tegenaria atrica araneae agelenidae
Archives of Insect Biochemistry and Physiology, 1999Co-Authors: Olivier Prouvost, Marie Trabalon, Mirjam Papke, Stefan SchulzAbstract:Qualitative changes in lipid content and composition were examined on the web and cuticle of Tegenaria atrica females in relation to sexual receptivity. In this spider species, 78 different compounds were detected by gas chromatography in the cuticle extract and 50 in the web; 28 identical compounds were present both on the spider silk and the cuticle of the female. The components were long-chain aliphatic hydrocarbons, fatty acids, and esters. On the web, sexual receptivity was correlated with changes in eleven polar compounds. On the cuticle, sexual receptivity was correlated with changes in eight polar compounds and 26 hydrocarbons. Bioassays demonstrated that the methanol eluate of webs and females were involved in stimulating the sexual behavior of males. Qualitative and/or quantitative changes in hexadecanoic acid, octadecadienoic acid, octadecenoic acid, methyl palmitate, methyl octadecanoate, and n-tricosane could play a role in the contact sex signals from web and cuticle of T. atrica. Arch. Insect Biochem. Physiol. 40:194–202, 1999. © 1999 Wiley-Liss, Inc.
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relationships among cannibalism contact signals ovarian development and ecdysteroid levels in Tegenaria atrica araneae agelenidae
Insect Biochemistry and Molecular Biology, 1998Co-Authors: Marie Trabalon, Grégory Pourié, Nadine HartmannAbstract:Abstract The present investigation was designed to study the relationships between the reproductive state of Tegenaria atrica females and their agonistic behavior towards conspecific females. Our results show that the virgin females exhibit the lowest levels of agonistic behavior whereas they provoke the highest levels of cannibalism by the mated females. The virgin females can be clearly distinguished from the other females as regards synthesis/release of cuticular compounds: methylesters and hydrocarbons. Furthermore, the ovaries of these females have not yet started vitellogenesis and the haemolymphatic ecdysteroids levels are high. It appears that the variation of females cannibalism parallels ovarian development and could be related to changes in cuticular compounds. Our results show for the first time that changes in spiders sexual and parental behavior during the reproductive cycle are accompanied by characteristic modifications in cuticular compounds, ovarian development as well as haemolymphatic ecdysteroid levels.
Geoff S Oxford - One of the best experts on this subject based on the ideXlab platform.
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many a slip dissecting the causes of reproductive isolation in two species of Tegenaria spiders agelenidae
Biological Journal of The Linnean Society, 2014Co-Authors: Geoff S Oxford, Peter J P CroucherAbstract:Key to our understanding of the mechanisms underlying the process of speciation is the determination of the nature of the barriers to gene flow between related taxa. Species that show zero gene flow in sympatry or parapatry are of little use in this respect. In the present study, we used two closely-related species of large house spider, Tegenaria saeva and Tegenaria gigantea, which hybridize to a limited extent along a natural contact zone in southern Britain. The species are apparently indistinguishable with respect to habitat utilization and phenology. Laboratory crosses using individuals from both allopatric and parapatric populations suggest that, although male and female courtship, as well as web and cuticular-borne pheromones, are conserved between the species, mechanical difficulties are experienced during interspecific copulation. Copulation bouts are, on average, significantly shorter during interspecific matings because of these difficulties, and are probably not sufficiently long for effective sperm transfer to take place. In the two cases of successful interspecific crossing, and in subsequent F1 and backcross generations, there are few indications of differential fertility, fecundity or viability, suggesting little post-zygotic incompatibility. The high success rate of crosses between F1 hybrids and both parental species underlines the principally mechanical barrier to gene flow between these taxa. Once this is breached, there appears to be little impediment to continuing introgression, which could, in some geographical areas at least, ultimately lead to the fusion of the two species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 355–367.
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distributions of species in the Tegenaria atrica group araneae agelenidae in north east england chance contingency and priority
Arachnology, 2014Co-Authors: Geoff S Oxford, C D SmithAbstract:Summary Tegenaria atrica is a species common in continental Europe and the Republic of Ireland but on the British mainland it has long been regarded as a relatively rare import that has failed to establish. The recent discovery of this species in some numbers and over several years in Burnopfield, County Durham, represented the first-reported, self-sustaining population. We show here that the species is widespread across a large swathe of country comprising parts of north County Durham, Tyne and Wear, and south-east Northumberland — an area of some 710 km2. Within a core range of approximately 400 km2, the other species in the Tegenaria atrica group, T. saeva and T. gigantea, tend to be uncommon or absent. The distributions of these three closely related species raise questions regarding the origins of the T. atrica population, the effect of chance and priority in initial establishment, and the role of possible competitive interactions between them. Although determining ecological processes from present p...
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historical distributions in britain of two species of large house spiders Tegenaria saeva and t gigantea araneae agelenidae and their evolutionary implications
Arachnology, 2009Co-Authors: Geoff S OxfordAbstract:Summary Current relative distributions and evolutionary interactions of two species of large house spiders, Tegenaria saeva and T. gigantea, vary geographically in Britain. To understand fully their underlying causes requires knowledge of species ranges in the past. Here, information from five sources: (a) historical distribution maps based on county and vice county lists; (b) a re-examination of museum and other collections; (c) catalogues of museum specimens recently re-examined by experts; (d) scrutiny of the literature; (e) a request published in a Wildlife Trust magazine, is used to determine historical distribution patterns. It is shown that the northern limits of both species have generally increased through time and that this phenomenon is not a result of variable recorder effort. In the south of England, the species boundary in Dorset has apparently been stable for at least a century. The ecological and evolutionary implications of these results are discussed.
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patterns of sperm usage in large house spiders Tegenaria spp genetics of esterase markers
Heredity, 1993Co-Authors: Geoff S OxfordAbstract:Patterns of sperm usage in large house spiders ( Tegenaria spp.): genetics of esterase markers
Ambros Hänggi - One of the best experts on this subject based on the ideXlab platform.
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Revision of the Nearctic Eratigena and Tegenaria species (Araneae: Agelenidae)
The Journal of Arachnology, 2016Co-Authors: Angelo Bolzern, Ambros HänggiAbstract:Abstract Based on specimens from several museum collections and recently sampled spiders during a field excursion to Mexico in 2014, the 11 species of Tegenaria s. l. endemic to the United States of America and Mexico are revised. Morphological characters and mitochondrial DNA sequences (CO1, NADH1, 16S) serve as the basis for proposed new combinations and new species. Tegenaria chiricahuae Roth, 1968 remains the only endemic Tegenaria species in the Western Hemisphere. All other specific names (T. blanda Gertsch, 1971, T. caverna Gertsch, 1971, T. decora Gertsch, 1971, T. flexuosa F.O. Pickard-Cambridge, 1902, T. florea Brignoli, 1974, T. gertschi Roth, 1968, T. mexicana Roth, 1968, T. rothi Gertsch, 1971, T. selva Roth, 1968, and T. tlaxcala Roth, 1968) are transferred to the genus Eratigena Bolzern, Burckhardt & Hanggi, 2013. Six new species are described: E. edmundoi, E. fernandoi, E. guanato, E. queretaro, E. xilitla, and E. yarini. In addition, females of E. flexuosa, and E. gertschi, and the male o...
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phylogeny and taxonomy of european funnel web spiders of the Tegenaria malthonica complex araneae agelenidae based upon morphological and molecular data
Zoological Journal of the Linnean Society, 2013Co-Authors: Angelo Bolzern, Daniel Burckhardt, Ambros HänggiAbstract:The taxonomy and systematics of European house spiders, currently constituting the ill-defined Tegenaria−Malthonica complex (including Aterigena) in the family Agelenidae, are revised. In Europe four monophyletic genera and 81 species are defined. One genus, Eratigena gen. nov., and seven species are described as new; at species level 17 new synonyms and 20 new combinations are proposed, and the original combination of 14 species is reinstated. Five species could not be placed (incertae sedis) because of insufficient material and one taxon is regarded as ‘nomen dubium’. On the basis of a detailed morphological assessment, 88 characters were chosen for a cladistic analysis. Phylogenetically informative characters include mostly spination patterns as well as spinneret and genital structures. In addition to morphology, three gene sections [cytochrome c oxidase subunit 1 (CO1), nicotinamide adenine dinucleotide dehydrogenase subunit 1 (NADH1) 28S] were analysed. Morphological and molecular analyses were performed individually and in combination applying maximum parsimony and Bayesian tree search methods. In all resulting trees Malthonica and Tegenaria in their present composition are either polyphyletic or paraphyletic. Consequently, we redefined the two genera and erected a new genus, Eratigena gen. nov. Identification keys are provided for the European agelenid genera as well as for the European species of Tegenaria and Eratigena gen. nov. The genera and most of the constituent species are described and illustrated. The new classification has also been applied to some extra European members of the Tegenaria-Malthonica complex resulting in additional three new synonyms, seven reversals to the original combination, and four new combinations. © 2013 The Linnean Society of London
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Aterigena, a new genus of funnel-web spider, shedding some light on the Tegenaria-Malthonica problem (Araneae: Agelenidae)
Journal of Arachnology, 2010Co-Authors: Angelo Bolzern, Ambros Hänggi, Daniel BurckhardtAbstract:Abstract Aterigena n. gen. is erected for four Palearctic species of funnel-web spiders previously placed in Tegenaria Latreille 1804 or Malthonica Simon 1898 (Agelenidae: Tegenariini) and A. aspromontensis n. sp., an Italian species described here. The following new combinations are proposed: Aterigena aculeata (Wang 1992), A. ligurica (Simon 1916), n. comb. (from Tegenaria), as well as A. aliquoi (Brignoli 1971) and A. soriculata (Simon 1873), n. comb. (from Malthonica). The latter two species were originally described in Tegenaria. The new genus is diagnosed by the unique combination of several morphological character states (e.g., notched trochanters III and IV, lateral spines on patellae, shape of vulvae). The monophyly of the new genus is also supported by a molecular analysis based on CO1 sequences of several taxa. Keys are provided for the identification of the recognized genera of Tegenariini and the species of Aterigena n. gen. Several species of PseudoTegenaria Caporiacco 1934, originally descr...
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Funnel web spiders from Sardinia: Taxonomical notes on some Tegenaria and Malthonica spp. (Araneae: Agelenidae)
Revue suisse de zoologie., 2008Co-Authors: Angelo Bolzern, Ambros Hänggi, Daniel BurckhardtAbstract:Funnel web spiders from Sardinia: Taxonomical notes on some Tegenaria and Malthonica spp. (Araneae: Agelenidae). - Based on spe - cimens collected by hand during several field trips to Sardinia and on specimens examined in several collections, the female of Tegenaria hen roti Dresco and the male of Malthonica sardoa Brignoli are described for the first time. Malthonica eleonorae (Brignoli) is redescribed. It can be distin- guished from T. henroti, which we regard as its sister species without yet proposing a new combination, by the number of teeth on the upper margin of the cheliceral groove and by the spine formulae of all leg tibiae. Additionally, the rim of the atrium, the form of the spermathecae, as well as the shape of the tegular apophysis and the ridge on the male bulb are important characters for separating these species. The female described under T. henroti by Wunderlich actually belongs to M. dalmatica (Kulczynski). The latter species would be new to Sardinia but has been recorded before on this island under the name T. drescoi Brignoli, which we consider as a new junior synonym of M. dalmatica. In addition, Malthonica sicana Brignoli is recorded for the first time from Sardinia, and the species is redescribed here. Males of the three species, M. sardoa, M. sicana and M. arganoi (Brignoli), can be distinguished by the shape of their retro lateral tibia apophysis, the shape of the terminal end of the conductor and the shape of the median apophysis.
Peter J P Croucher - One of the best experts on this subject based on the ideXlab platform.
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many a slip dissecting the causes of reproductive isolation in two species of Tegenaria spiders agelenidae
Biological Journal of The Linnean Society, 2014Co-Authors: Geoff S Oxford, Peter J P CroucherAbstract:Key to our understanding of the mechanisms underlying the process of speciation is the determination of the nature of the barriers to gene flow between related taxa. Species that show zero gene flow in sympatry or parapatry are of little use in this respect. In the present study, we used two closely-related species of large house spider, Tegenaria saeva and Tegenaria gigantea, which hybridize to a limited extent along a natural contact zone in southern Britain. The species are apparently indistinguishable with respect to habitat utilization and phenology. Laboratory crosses using individuals from both allopatric and parapatric populations suggest that, although male and female courtship, as well as web and cuticular-borne pheromones, are conserved between the species, mechanical difficulties are experienced during interspecific copulation. Copulation bouts are, on average, significantly shorter during interspecific matings because of these difficulties, and are probably not sufficiently long for effective sperm transfer to take place. In the two cases of successful interspecific crossing, and in subsequent F1 and backcross generations, there are few indications of differential fertility, fecundity or viability, suggesting little post-zygotic incompatibility. The high success rate of crosses between F1 hybrids and both parental species underlines the principally mechanical barrier to gene flow between these taxa. Once this is breached, there appears to be little impediment to continuing introgression, which could, in some geographical areas at least, ultimately lead to the fusion of the two species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 355–367.
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mitochondrial differentiation introgression and phylogeny of species in the Tegenaria atrica group araneae agelenidae
Biological Journal of The Linnean Society, 2004Co-Authors: Peter J P Croucher, G S Oxford, Jeremy B SearleAbstract:The relationships between the three members of the Tegenaria atrica group (T. atrica, T. saeva and T. gigantea) were examined with DNA sequence data from mitochondrial CO1, 16S rRNA, tRNAleu(CUN) and ND1 genes. Members of this group of large house spiders have overlapping distributions in western Europe and hybridize with each other to a variable degree. The close relatedness of all three species was supported by all analyses. T. saeva and T. gigantea are more closely affiliated than either is to T. atrica. Haplotypes clearly assignable to T. gigantea were also present in many specimens of T. saeva suggesting asymmetrical introgression of mtDNA from T. gigantea into T. saeva. Molecular clock calibrations (CO1) suggest that deeper divisions within the genus Tegenaria may be in excess of 10 million years old, and that the evolutionary history of the T. atrica group has been moulded by Quaternary glacial-interglacial cycles.
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mitochondrial differentiation introgression and phylogeny of species in the Tegenaria atrica group araneae agelenidae
Biological Journal of The Linnean Society, 2004Co-Authors: Peter J P Croucher, G S Oxford, Jeremy B SearleAbstract:The relationships between the three members of the Tegenaria atrica group (T. atrica, T. saeva and T. gigantea) were examined with DNA sequence data from mitochondrial CO1, 16S rRNA, tRNAleu(CUN) and ND1 genes. Members of this group of large house spiders have overlapping distributions in western Europe and hybridize with each other to a variable degree. The close relatedness of all three species was supported by all analyses. T. saeva and T. gigantea are more closely affiliated than either is to T. atrica. Haplotypes clearly assignable to T. gigantea were also present in many specimens of T. saeva, suggesting asymmetrical introgression of mtDNA from T. gigantea into T. saeva. Molecular clock calibrations (CO1) suggest that deeper divisions within the genus Tegenaria may be in excess of 10 million years old, and that the evolutionary history of the T. atrica group has been moulded by Quaternary glacial–interglacial cycles. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81, 79–89.
