Article

Vicariance in a Cryptic Species Pair of European Pseudoscorpions (Arachnida, Pseudoscorpiones, Chthoniidae)

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Abstract

Following the first records of Chthonius (Ephippiochthonius) boldorii Beier, 1934 in central Europe, a species which was previously assumed to occur exclusively in Mediterranean caves, 116 series (595 specimens) of the cryptic taxa C. (E.) boldorii and C. (E.) fuscimanus Simon, 1900 (Syn. C. (E.) austriacus Beier, 1931) were re-examined. Although multivariate analyses suggest specific separation, there is only one unequivocal character for discrimination: the presence or absence of a single isolated tooth on the moveable finger of the chelicerae. The distributions were found to be largely allopatric, therefore it is concluded that the species rank of the two morphospecies is justified. North of the Alps, an almost vicariant pattern emerged: east of 14° E fuscimanus occurs, west of this line boldorii occurs. The results provide a basis for discussing the relevance of minute morphological differences in pseudoscorpion taxonomy.

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... Distribution. Austria, Croatia, Germany, Italy, Slovakia, Slovenia and Switzerland (Muster et al. 2004, Zaragoza 2017, Červená et al. 2020. Remarks. ...
... These localities have habitats and conditions suitable for the occurrence of stable and relatively numerous populations of this species (Christophoryová & Krumpál 2005, Christophoryová 2013. Ephippiochtonius boldorii is an epigeic species occurring in oak-hornbeam forests (Muster et al. 2004, Christophoryová & Krumpál 2007. The highest localities in the Alps exceed 1500 m a.s.l., but on the other hand, this species was also recorded at sea-level in Venice. ...
... The highest localities in the Alps exceed 1500 m a.s.l., but on the other hand, this species was also recorded at sea-level in Venice. The species occurs predominantly in leaf litter in mesophilic habitats (Muster et al. 2004). One female was recorded from compost heaps (Christophoryová et al. 2017b). ...
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... In pseudoscorpion taxonomy, multivariate analyses were used to separate two European Chthoniidae species. Although multivariate analyses suggest specific separation, there was only one unequivocal character for discrimination, the presence or absence of a single isolated tooth on the moveable finger of the chelicerae (Muster et al. 2004). ...
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... Añez et al. 1997;Gebre-Michael and Medhin 1997;Petrarca et al. 1998;Calle et al. 2002;Gamboa and Arrivillaga 2010;Aguirre et al. 2011;Laszlo et al. 2013), including pseudoscorpions (e.g. Harvey 1987;Muster et al. 2004;Christophoryová et al. 2016). ...
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... Cryptic taxa can not only show small different morphological features but also ecological and behavioral ARTICLE IN PRESS differences (Colborn et al. 2001; Muster et al. 2004; Wellborn and Cothran 2004); these could be the result of an adaptation, given that the phenotypic similarity of similar species also implies limited scope to occupy different habitats (Wellborn and Cothran 2004). We observed ecological differences between the 2 similar species, which are sympatric but not syntopic; E. concinnus is more common in natural habitats, such as woods, whereas E. tergestinus was prevalently found in anthropogenic habitats. ...
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... Although there are more detailed morphological or morphometric data in recent descriptions of new taxa, data for populations of common species are mainly limited to species from England (Gabbutt & Vachon 1963, 1965, 1967 Gabbutt 1970) and Switzerland (DeVore-Scribante 1999). Detailed studies have already been published to distinguish Chthonius boldorii and C. fuscimanus (Muster et al. 2004) and to provide measurements for C. heterodactylus (Ducháč et al. 2007), Syarinus strandi (Ducháč 1998), Neobisium slovacum (Ducháč 1996), N. polonicum (Ducháč 1995b) and some other species (Drogla et Lippold 2004; Jost 1982 ). The possibility that cryptic species might exist in this region is supported by preliminary karyological analyses of the common Neobisium carcinoides. ...
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... Both cases have been described for spiders (crypticity: Ramirez & Chi 2004;Johannesen et al. 2005;Huber et al. 2005;polymorphism: Pérez-Miles 1989;Huber & Gonzá lez 2001;Jocqué 2002), suggesting that neither may be an exception. Cryptic species complexes can be found in several organisms (Bond & Sierwald 2002;Muster et al. 2004) and they are often revealed by integration of molecular and/or behavioral tools, and not by the definition of morphological apomorphies (Adams & Funk 1997). On the other hand, genitalic polymorphism can be discovered by several methods, the best evidence being the hatching of different morphs from a single egg-sac (Jocqué 2002). ...
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The linyphiid spiders Oedothorax gibbosus (Blackwall 1841) and Oedothorax tuberosus (Blackwall 1841) were formerly described as separate species due to marked differences in prosomal structures of the males. During the last decade it was demonstrated that they are two forms of a single species. However, it remained to be shown whether the former species represent two distinct morphs or extremes of a continuum of variation. A morphometric examination of 246 alcohol-preserved specimens revealed that individual spiders can clearly be assigned to one of two forms. No intermediates were found, demonstrating that there are two distinct morphs.
Article
Complementary abundance gradients of substituting congeneric arachnids along the northern margin of the Alps are described, and the possible historical and contemporary factors that have produced these distributions are explored. The intensities of activity of epigeous arachnids were determined in pitfall traps along seven altitudinal transects approximately 35 km apart in the Bavarian Alps (Germany) and in the county of Salzburg (Austria). No supraregional gradient of physical factors exists within the region investigated. Six pairs of congeners show vicariances or alternating abundance gradients. For each species pair, the abundance gradient is represented in diagrams and compared with a graphic model of substitution types. Three patterns are evident: strict vicariances, abrupt replacement in a narrow belt, and gradual substitution over a wide zone of sympatric occurrence. The different extent of range overlap suggests distinctive classes and intensities of interspecific interactions. With regard to biogeographic histories, one prevailing pattern emerged: endemic species of the eastern Alps are replaced by Eurosiberian counterparts towards the west. It appears that geographical abundance variation reflects colonization history.
Article
All species and higher taxa of Recent Erinaceidae are diagnosed, the character-states most useful in classification are tabulated and the most probable phylogenies presented. Four African species are included in the genus Atelerix. The desert-adapted Paraechinus is believed to be more closely related to Atelerix than to the convergent Hemiechinus. The ranges of all species are mapped, with emphasis upon areas of parapatry and sympatry. Marginal localities are listed if they are not derived from previously published maps. All nominal fossil species that have been allocated to Recent genera are listed. Ecological data relevant to the interpretations of species boundaries, e.g. relating to diet and habitat, are summarized, and the factors likely to determine the species boundaries are reviewed.
Article
Population structure is the result of both present processes and past history. Molecular markers are proving of great value in describing the former, and it is important to similarly determine the latter in order to understand their respective contributions. The study of palaeo-climates has also advanced significantly, and in particular that of the Pleistocene ice ages, which modified species ranges considerably. The last ice age and rapid post-glacial colonization of Europe is summarized. Possible population genetic consequences of expansion northward from southern refugia, and those of remaining in these mountainous regions are discussed. A series of recent case studies are detailed where DNA sequence information has been used to describe species genetic variation and subdivision across Europe. These include a grasshopper, the hedgehog, oak trees, the common beech, the black alder, the brown bear, newts, shrews, water vole, silver fir and house mice. These molecular data confirm southern peninsulas of Europe as major ice age refugia, and in most cases demonstrate that genetically distinct taxa emerged from them. They can thus define genomic differences and so gready augment previous fossil data. The refugial genomes contributed differently in various species to die re-colonization of Europe, with three broad patterns described as paradigms–‘grasshopper’, ‘hedgehog’ and ‘bear’. These different expansion patterns produced clusters of hybrid zones where they made contact, and it is argued that many species genomes may be further cryptically subdivided. A reduction in diversity from southern to northern Europe in the extent of allelic variation and species subdivision is seen; this is attributed to rapid expansion northward and the varied topography of southern refugia allowing populations to diverge through several ice ages. The differences in DNA sequence indicate that some species have been diverging in refugial regions for a few ice ages at most, whilst distinct lineages in other species suggest much more ancient separation.
Article
The unit of adaptation is usually thought to be a gene or set of interacting genes, rather than the whole genome, and this may be true of species differentiation. Defining species on the basis of reproductive isolation (RI), on the other hand, is a concept best applied to the entire genome. The biological species concept (BSC; Mayr, 1963) stresses the isolation aspect of speciation on the basis of two fundamental genetic assumptions – the number of loci underlying species differentiation is large and the whole genome behaves as a cohesive, or coadapted genetic unit. Under these tenets, the exchange of any part of the genomes between diverging groups is thought to destroy their integrity. Hence, the maintenance of each species’ genome cohesiveness by isolating mechanisms has become the central concept of species. In contrast, the Darwinian view of speciation is about differential adaptation to different natural or sexual environments. RI is viewed as an important by product of differential adaptation and complete RI across the whole genome need not be considered as the most central criterion of speciation. The emphasis on natural and sexual selection thus makes the Darwinian view compatible with the modern genic concept of evolution. Genetic and molecular analyses of speciation in the last decade have yielded surprisingly strong support for the neo-Darwinian view of extensive genetic differentiation and epistasis during speciation. However, the extent falls short of what BSC requires in order to achieve whole-genome ‘cohesiveness’. Empirical observations suggest that the gene is the unit of species differentiation. Significantly, the genetic architecture underlying RI, the patterns of species hybridization and the molecular signature of speciation genes all appear to support the view that RI is one of the manifestations of differential adaptation, as Darwin (1859, Chap. 8) suggested. The nature of this adaptation may be as much the result of sexual selection as natural selection. In the light of studies since its early days, BSC may now need a major revision by shifting the emphasis from isolation at the level of whole genome to differential adaptation at the genic level. With this revision, BSC would in fact be close to Darwin’s original concept of speciation.
Article
Das Verhalten, die Samenbertragungsweisen und die Spermatophoren verschiedener Arten von Pseudoscorpionen wurden beobachtet und beschrieben.Alle Pseudoscorpione setzen Spermatophoren ab, denen die Weibchen die Spermien entnehmen. Das Verhalten bei der Samenbertragung ist bei den verschiedenen Arten unterschiedlich; es lt sich eine Evolution von einfachen zu komplizierteren Vorgngen feststellen.Das Verhalten vonChthonius tetrachelatus, Chthonius ischnocheles undNeobisium muscorum ist ursprnglich. Die Mnnchen setzen immer Spermatophoren ab, auch wenn keine Weibchen anwesend rind. Das Mnnchen vonChthonius strzt alte Spermatophoren um und ersetzt sie durch neue. Die Weibchen werden von den Spermatophoren chemotaktisch angelockt. Nach kurzer Prfung berschreiten sie sie hochbeinig und lsen mit einem Tropfen Flssigkeit einen Quellungsvorgang aus, der die Spermien aus den Spermatophoren in das weibliche Genitalatrium treibt.Die Mnnchen der CheiridiidenCheiridium museorum undApocheiridium ferum setzen ihre Spermatophoren dagegen wahrscheinlich nur in Anwesenheit von Weibchen ab. Das Mnnchen vonCheiridium zeigt zuweilen vorher, wenn es einem Weibchen begegnet, eine vibrierende Bewegung einer Palpenhand. Die Weibchen suchen die Spermatophoren chemotaktisch auf. Wie die Mnnchen strzen sie alte Spermatophoren m.Die Chernetiden und Cheliferiden bilden Paare. Die Mnnchen der Chernetiden berfallen jeden Artgenossen und versuchen, ihm einen Paarungstanz aufzuzwingen. Mit anderen Mnnchen gibt es dann einen Commentkampf, mit Weibchen einen Paarungstanz. Das Mnnchen fat seine Partnerin an einer (Chernes cimicoides zu Anfang) oder beiden (Lasiochernes pilosus, Chernes hanhi, Chernes cimicoides) Hnden und geht mit ihr mehrfach vor und zurck. Dann setzt es eine Spermatophore ab und zicht das Weibchen darber. Die Mnnchen der beidenChernes-Arten bewegen beim Paarungstanz auffllig ihre Vorderbeine und berhren damit die Weibchen. Das Mnnchen vonLasiochernes hat in der Behaarung seines Palpenfemurs ein zustzliches Reizmittel. Beim Paarungstanz hlt es sein Weibchen so, da dessen Palpenfinger diese Haare berhren.Die hchstentwickelten Verhltnisse zeigen die Cheliferiden. Ihre Mnnchen balzen die Weibchen an, wobei sie mit vorgestreckten zylindrischen Organen vibrierende Bewegungen des Krpers ausfhren. Das Mnnchen vonChelifer besetzt vorher ein Revier, das es wahrscheinlich mit Duftmarken kennzeichnet. Hierfr dienen mglicherweise die Coxalscke. BeiDactylochelifer tanzen beide Partner miteinander vor und zurck, beiChelifer bewegt sich nur das Mnnchen. Zu einem engen Kontakt zwischen beiden Partnern kommt es bei den Cheliferiden erst nach der Bildung der Spermatophore. Das Mnnchen ergreift dann die Palpenfemora des Weibchens und hilft unter krftigen Schubbewegungen mit seinen Vorderbeinen dem Weibchen bei der Aufnahme des Spermas.Die Spermatophoren sind langgestielt. Bei den Chthoniiden stehen sie senkrecht. An der Spitze tragen she einen nicht umhllten Samentropfen, der durch einen Kragen vor Berhrungen geschtzt ist.Auch die Spermatophoren der Neobisiiden und Cheridiiden stehen senkrecht. Sie tragen an der Spitze eine in ein kugelfrmiges Samenpaket eingeschlossene Samenmasse. Durch einen Quellungsvorgang, den das Weibchen mit Hilfe eines aus seiner Geschlechtsffnung austretenden Tropfens auslst, werden die Spermien aus dem Samenpaket in das weibliche Genitalatrium bertragen. Die Spermatophoren vonApocheiridium sind sekundr vereinfacht; she tragen einen nicht umhllten, winzigen Samentropfen.Die Spermatophoren der Chernetiden und Cheliferiden stehen schrg. Sie tragen an der Spitze ein Samenpaket und darunter einen Tropfen Flssigkeit, der bei der Samenbertragung die Quellung im Samenpaket auslst. Bei den Chernetiden hat das Samenpaket die Gestalt zweier konvergierender Schluche, die in einen gemeinsamen Ausfhrgang mnden. Bei der Samenbertragung wird dieses Paket vom Weibchen abgenommen. Doch nur der Ausfhrgang wird in die weibliche Geschlechtsffnung eingefhrt. Gleichzeitig wird der unter dem Samenpaket hngende Tropfen abgestreift und gelangt zwischen die Schluche des Samenpaketes, wo er durch die Wandung eindringt und den Quellungsvorgang auslst, der die Samen wahrscheinlich direkt ins Receptaculum seminis entleert.Die kompliziertesten Spermatophoren haben die Cheliferiden. Ihr Samenpaket trgt flgelfrmige Anhnge, die wahrscheinlich bei der Samenbertragung die gequollene Samenmasse aus dem Samenpaket herauspressen und ins weibliche Genitalatrium entleeren.Die Verhaltensweisen bei Begegnungen mit Artgenossen sind verschieden. Die Chthoniiden stoen gegeneinander vor, ohne einander zu berhren. hnlich verhalten sich zuweilen dieChernes-Arten. Apocheiridium undLasiochernes zucken auffllig mit den Palpen, wenn sie von Artgenossen bedrngt werden. Diese Verhaltensweise wird von anderen Artgenossen mit dem gleichen Palpenzucken beantwortet und pflanzt sich so oft ber alle Tiere einer Gesellschaft fort.Bei den Chernetiden und Cheliferiden gibt es Commentkmpfe zwischen den Mnnchen.The behavior, the methods of sperm transfer, and the structure and function of spermatophores of the Pseudoscorpions have been studied and described.The males of the Pseudoscorpions deposit spermatophores from which the females take up the sperm. The method of sperm transfer is different among the different species; there is an evolution from simple to complicated procedures.The sperm transfer inChthonius tetrachelatus, Chthonius ischnocheles andNeobisium muscorum is primitive. The males deposit spermatophores even when females are not present. The male ofChthonius destroys old spermatophores and replaces them. The females are attracted chemotactically and examine the spermatophores. Then, they step over them on extended legs and release a drop of fluid which triggers a swelling mechanism that forces the spermatozoa out of the spermatophores and into the female genital opening.The males of the Cheiridiids (Cheiridium muscorum andApocheiridium ferum) presumably deposit spermatophores only when females are present. The male of Cheiridium often shows a vibrating movement of one pedipalpal hand when it encounters a female. Males and females destroy old spermatophores and the male replaces them with new ones. The female traces the spermatophores chemotactically.In the Chernetids and Cheliferids there is pairing. The male of the Chernetids attacks all members of its own species and tries to force a courtship dance upon them. With another male, a struggle follows the initial dance while with a female the courtship dance continues. The male grasps one (Chernes cimicoides at the beginning) or both (Lasiochernes pilosus, Chernes hahni, Chernes cimicoides) hands of the female and walks forward and backward several times. Then, it deposits a spermatophore and pulls the female over it. During the mating dance; the male of the two species ofChernes moves its first pair of legs, touching the female with it in a characteristic manner. The male ofLasiochernes has an accessory stimulant, the hair on its pedipalpal femora. During the mating dance, the fingers of the female pedipals touch this hair.The Cheliferids show the most advanced features. Their males court the females, showing extended ram's horn organs and vibrating movements of the body. The male ofChelifer occupies a territory before courting. It presumably marks this territory with an odorous secretion, possibly from the coxal sacs. InDactylochelifer the mates dance forward and backward together. InChelifer only the male moves. In Cheliferids, close contact between mates does not take place before the spermatophore has been deposited. Then, the male grasps the pedipalpal femora of the female. With his forelegs it assists the female in taking up the sperm and shows strong pushing movements.The spermatophores are long-stalked. In the Chthoniids, they stand straight up and have an uncovered sperm drop at the top which is protected against touching by a collar.The spermatophores of Neobisiids and Cheiridiids also stand straight up. At the top they bear a sperm mass enclosed in a globular sperm package. By a swelling mechanism, the spermatozoa are transferred from the sperm package into the female genital opening. The swelling mechanism is released by a drop of fluid, coming out of the female genital opening. InApocheiridium, the spermatophores are simplified, but not primitive. They have a small uncovered sperm drop.In the Chernetids and Cheliferids the stalk of the spermatophore is inclined. It bears a sperm package at the top and under this package a drop of fluid which releases the swelling mechanism in the sperm package during sperm transfer. In Chernetids, the sperm package has the form of two curved and converging tubes which open to a common duct. During sperm transfer, this package is taken by the female, but only the duct enters the female genital opening. At the same time the drop of fluid which was attached to the spermatophore under the sperm package, is pushed between the tubes of the sperm package. There, it enters the sperm mass through the wall of the sperm package and releases the swelling mechanism which presumably forces the sperm directly into the receptaculum seminis of the female.The Cheliferids have the most complicated spermatophores. Their sperm package has wing-like appendages, which presumably press the swelled sperm mass out of the sperm package and into the female genital opening during sperm transfer.The Pseudoscorpions act in different manners when meeting a member of their own species. The Chthoniids show quick movements towards each other without touching. The species ofChernes sometimes act in a similar manner. Apocheiridium andLasiochernes show characteristic short and quick movements of the pedipalps when they are attacked by members of their own species. Members of the same species react with the same movements of the pedipalps. This behavior often spreads to members of the crowd.In Chernetids and Cheliferids, the males sometimes fight against each other.
Article
This paper focuses on the relationship between population genetic structure and speciation mechanisms in a monophyletic species group of Appalachian cave spiders (Nesticus). Using mtDNA sequence data gathered from 256 individuals, I analyzed patterns of genetic variation within and between populations for three pairs of closely related sister species. Each sister-pair comparison involves taxa with differing distributional and ecological attributes; if these ecological attributes are reflected in basic demographic differences, then speciation might proceed differently across these sister taxa comparisons. Both frequency-based and gene tree analyses reveal that the genetic structure of the Nesticus species studied is characterized by similar and essentially complete population subdivision, regardless of differences in general ecology. These findings contrast with results of prior genetic studies of cave-dwelling arthropods that have typically revealed variation in population structure corresponding to differences in general ecology. Species fragmentation through both extrinsic and intrinsic evolutionary forces has resulted in discrete, perhaps independent, populations within morphologically defined species. Large sequence divergence values observed between populations suggest that this independence may extend well into the past. These patterns of mtDNA genealogical structure and divergence imply that species as morphological lineages are currently more inclusive than basal evolutionary or phylogenetic units, a suggestion that has important implications for the study of speciation mechanisms.
Article
The neotropical pseudoscorpion Cordylochernes scorpioides (Chernetidae: Lamprochernetinae) is currently described as a single species ranging from Central America to northern Argentina. However, interpopulation crosses have recently demonstrated that C. scorpioides actually represents a complex of cryptic species. Here we present mitochondrial COI gene sequence data from C. scorpioides individuals from Panama, Trinidad, and French Guiana which demonstrate little or no intrapopulation variability but divergence ranging from 2.6 to 13.8% between geographic populations. Phylogenetic analysis provides evidence of a major split between C. scorpioides lineages from Central and South America. Levels of interpopulation mtDNA divergence correspond well with previously established patterns of postzygotic reproductive incompatibility between geographically distinct units within the C. scorpioides complex. By contrast, multivariate morphometric analysis demonstrates that extensive sequence divergence has occurred in the absence of appreciable morphological differentiation between the populations. To provide a framework for assessing the scale of geographic divergence in C. scorpioides, Cordylochernes sequences were compared with homologous sequence from its presumed sister taxon, Lustrochernes, and from Parachernes and Semeiochernes, representatives of the second chernetid subfamily, the Chernetinae. Our preliminary, generic-level analysis suggests that COI sequence data may prove useful in resolving relationships within this problematic family.
Article
European hedgehog populations belonging to Erinaceus europaeus and E. concolor have been investigated by mitochondrial DNA analysis. A 383 bp fragment of the cytochrome b gene has been sequenced and maximum parsimony and neighbour-joining trees of Tamura-Nei genetic distance values have been constructed. Similar topologies have been produced by both methods, showing a deep divergence between E. europaeus and E. concolor and a further subdivision of each species into a western and an eastern clade. A comparison with previously published allozyme data is made, and concordant and discordant patterns are discussed. The influence of Pleistocene glaciations on the observed pattern of divergence is inferred.
Article
The origins and development of the study of speciation, hybrid zones and phylogeography are outlined using evolutionary iconography. This traces the ideas in this field from Lamarck and Darwin through to the present as represented in diagrams and figures. A 'tree of trees' summarizes this growth and current vitality. The new facility to use various DNA sequences from nuclear, mitochondrial and chloroplast genomes to determine genetic variation throughout a species range is examined particularly. There is great genomic subdivision across species distributions, which can be interpreted in the light of the recent demonstrations of severe palaeoclimatic oscillations. Refugia and postglacial colonization routes are proposed for several organisms across Europe. The role of geography in speciation through the Pleistocene is considered. These emerging principles and analyses are applied to data available on a variety of organisms in other regions of the world, such as the Arctic, North America and the Tropics, and including the progress of Homo sapiens through the last ice age. Some suggestions are made for future research directions.
Article
European hedgehogs, Erinaceus europaeus and E. concolor, are among the many European plant and animal taxa that have been subjected to cyclical restriction to glacial refugia and interglacial expansion. An analysis of 95 mitotypes, comprising partial cytochrome b and control region sequences, shows deep divergence between the two hedgehog species. Three europaeus and two concolor clades are clearly identified and are consistent with previously identified refugia for Europe: the Iberian peninsula, Italy, and the Balkans. The degree of mitochondrial divergence among these clades suggests pre-Pleistocene separation of the refugial populations. In contrast, analysis of two nuclear introns clearly separates the two concolor clades, as in the mitochondrial data, but cannot discriminate the three europaeus clades. This discrepancy between nuclear and mitochondrial data is attributed to historical differences in the refugial population size of europaeus and concolor. The geographical distribution of mitotypes is analysed using nested clade analysis. This method, by including unobserved ('missing') mitotypes, can identify mitotype groupings that remain undetected in conventional analyses. However, the application of nested clade analysis to the study of refugial populations may be hampered by such factors as the loss of haplotypes from the refugial areas by repeated contractions of the population and the recent time scale of colonization relative to mutation rate.
Rote Liste der Pseudoskor-pione Bayerns (1. Fassung) (Arachnida: Pseudoscor-piones)
  • T Blick
  • C Muster
BLICK, T. & MUSTER, C. (2003): Rote Liste der Pseudoskor-pione Bayerns (1. Fassung) (Arachnida: Pseudoscor-piones). SchrR. Bayer. Landesamt f. Umweltschutz, in press
Molecular phylogenetics at the popula-tion species interface in cave spiders of the southern Appalachians (Araneae: Nesticidae: Nesticus) Male dimorphism in Oedothorax gibbosus (Araneae, Linyphiidae): a morpho-metric analysis
  • M C C Hedin
  • T Muster
  • T Schmardaand
  • S Rheinemann
  • G Uhl
HEDIN, M. C. (1997): Molecular phylogenetics at the popula-tion species interface in cave spiders of the southern Appalachians (Araneae: Nesticidae: Nesticus). Mol. Biol. Evol. 14: 309–324. 308 C. MUSTER, T. SCHMARDAand T. BLICK rHEINEMANN, S. & UHL, G. (2000): Male dimorphism in Oedothorax gibbosus (Araneae, Linyphiidae): a morpho-metric analysis. J. Arachnol. 28: 23–28
Vergleichende Untersuchungen zur Fortpflanzungsbiologie der Pseudoscorpione. Beobachtungen über das Verhalten, die Samenübertragungsweisen und die Spermatophoren einiger einheimischer Arten
  • Weygoldt
WEYGOLDT, P. (1966): Vergleichende Untersuchungen zur Fortpflanzungsbiologie der Pseudoscorpione. Beobach-tungen über das Verhalten, die Samenübertragungsweisen und die Spermatophoren einiger einheimischer Arten. Z. Morphol. Ökol. Tiere 56: 39–92.
550 m, 1 …, 3 juv. (12.08.1947, leg. Strouhal, NHMW); E Puchenstuben
  • E Puchenstuben
  • ° Langseit
  • ° 56′ N
E Puchenstuben, Langseit, 47° 56′ N, 15° 19′ E, 550 m, 1 …, 3 juv. (12.08.1947, leg. Strouhal, NHMW); E Puchenstuben, Rotte Berg, 47° 56′ N, 15° 19′ E, 600 m, 1 …, 2 , 1 juv. (12.08.1947, leg. Strouhal, NHMW);
Höhlenpseudoskorpione aus Nordital-ien und der dalmatinischen Insel Krk. Atti e Memorie della Commissione Grotte
  • V Mahnert
MAHNERT, V. (1980): Höhlenpseudoskorpione aus Nordital-ien und der dalmatinischen Insel Krk. Atti e Memorie della Commissione Grotte " E. Bogan " 20: 95–100.
Neue cavernicole und subterrane Pseu-doscorpione. Mitt. Höhlen-und Karstforschung
  • References Beier
REFERENCES BEIER, M. (1934): Neue cavernicole und subterrane Pseu-doscorpione. Mitt. Höhlen-und Karstforschung 1934: 53–59.
1 juv. (01.01.1948, leg. Strouhal, NHMW) Note on nomenclature: GARDINI (1980) detected the syn-onymy of Chthonius tetrachelatus fuscimanus Simon with Chthonius (E.) austriacus Beier. Because the name fusci-manus Simon had not been mentioned in literature for more than 80 years
  • Tennengebirge
  • ° Wassergraben Ne Werfen
  • ° 29′ N
Tennengebirge, Wassergraben NE Werfen, 47° 29′ N, 13° 12′ E, 1 juv. (01.01.1948, leg. Strouhal, NHMW). Note on nomenclature: GARDINI (1980) detected the syn-onymy of Chthonius tetrachelatus fuscimanus Simon with Chthonius (E.) austriacus Beier. Because the name fusci-manus Simon had not been mentioned in literature for more than 80 years, he proposed regarding it a nomen oblitum.
1 … (02.07.1950 (?), leg. Strouhal, NHMW); SE Puchberg am
  • E Puch-Berg
  • Schneeberg
  • ° Reitzenberg
  • ° 48′ N
E Puch-berg am Schneeberg, Reitzenberg, 47° 48′ N, 15° 57′ E, 1 … (02.07.1950 (?), leg. Strouhal, NHMW); SE Puchberg am Schneeberg, SE, Kienberg, SW-Hang, 47° 46′ N, 15° 54′ E, 700 m, 1 … (13.08.1951, leg. Strouhal, NHMW);
D-95503 Hummeltal, Germany Received: 27. 03. 2003 Returned for revision: 05. 06
  • Theo Blick Heidloh
Theo BLICK, Heidloh 8, D-95503 Hummeltal, Germany Received: 27. 03. 2003 Returned for revision: 05. 06. 2003
500 m, 3 … (07.07.1986, leg. Schawaller, SMNS 1623) Baden Württemberg: Friedrichshafen
  • ° Nw Kiefersfelden
  • ° 37′ N
NW Kiefersfelden, 47° 37′ N, 12° 11′ E, 500 m, 3 … (07.07.1986, leg. Schawaller, SMNS 1623). Baden Württemberg: Friedrichshafen, 47° 40′ N, 09° 30′ E, 4 …, 4  (01.05.1996, leg. Lippold, Coll. Lippold 1925); SW Berau, 47° 42′ N, 8° 15′ E, 1 … 1  (18.06.1993, leg. Konzel-mann, SMNS 3240);
270 m, 6 …, 3  (04.05.1952, leg. Strouhal, NHMW); Höflein a. d
  • Marbachtal
  • ° Kierling
  • ° 19′ N
Marbachtal nördl. Kierling, 48° 19′ N, 16° 17′ E, 270 m, 6 …, 3  (04.05.1952, leg. Strouhal, NHMW); Höflein a. d. Donau, S Graben, 48° 21′ N, 16° 16′ E, 300 m, 5 …, 5  (27.06.1948, leg. Strouhal, NHMW);