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Vicariance in a Cryptic Species Pair of European Pseudoscorpions (Arachnida, Pseudoscorpiones, Chthoniidae)
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.
... We confirmed an interesting record of Ephippiochtho nius boldorii in Slovakia from Udava in the East Carpathians PLA. This species is mostly present in the leaf litter of mesophilic habitats in oak-hornbeam forests (Muster et al. 2004. Mahnert (1980) distinguished E. boldorii and E. fuscimanus based on the absence (boldorii) or presence (fuscimanus) of an isolated subdistal tooth on the movable finger of the chelicerae. ...
... Mahnert (1980) distinguished E. boldorii and E. fuscimanus based on the absence (boldorii) or presence (fuscimanus) of an isolated subdistal tooth on the movable finger of the chelicerae. Muster et al. (2004) revised extensive European material based on this character. They concluded that E. boldorii was a western species, mostly confined to localities west of 14°E. ...
... Thus the records from Slovakia represent the easternmost outposts of its known range. Further studies are required to determine whether E. boldorii is more widespread in Central Europe than assumed by Muster et al. (2004), or whether another cryptic species could be involved (cf. Muster et al. 2021). ...
The paper presents new data on the diversity and distribution of pseudoscorpions from seven localities in the Stebnícka
Magura massif in the Low Beskids, two localities in the East Carpathians Protected Landscape Area (PLA) and one locality in the Vihorlat
PLA, Slovakia, collected by sieving methods. Altogether, 372 pseudoscorpion specimens belonging to 15 species and four families were
sampled over three years (2007, 2010, 2015). The most abundant families were Neobisiidae (335 specimens, nine taxa) and Chthoniidae
(33 specimens, four taxa); while the families Cheiridiidae and Chernetidae were represented by only one species each. Neobisium sylvaticum
(C. L. Koch, 1835), Neobisium erythrodactylum (L. Koch, 1873), Ephippiochthonius boldorii (Beier, 1934), Ephippiochthonius tetrachelatus
(Preyssler, 1790) and Roncus sp. were recorded only in the East Carpathians PLA within our study. The most abundant species was
Neobisium crassifemoratum (Beier, 1928). The species Ephippiochthonius boldorii and Ephippiochthonius fuscimanus (Simon, 1900) were
found in this area for the first time.
... 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). ...
A total of 667 specimens of pseudoscorpions, belonging to 15 species and four families were collected during 2014. Samples
were taken from leaf litter at five localities in the Western Carpathians in the Slovak Karst National Park, Slovakia. The most abundant families
were Neobisiidae (381 specimens) and Chthoniidae (275 specimens). Furthermore ten chernetid specimens and a single cheliferid
deutonymph were recorded. Four of the 15 pseudoscorpion species were recorded in the Slovak Karst for the first time.
... Although their deep-level phylogenetic relationships are now better understood [37], our knowledge of pseudoscorpion intra-familial relationships and phylogeography is still fairly limited [38][39][40][41][42][43][44][45]. Pseudoscorpions often show a relatively uniform external morphology among closely related species [46,47], but they can exhibit a great plasticity in terms of their karyotypes [48][49][50][51][52][53]. Their interspecific karyotype variability is high; a diploid chromosome number varies from 2n = 7 to 143 [50,51]. ...
... We employed morphometric analyses to complement our morphology-based species validation step [125]. Morphometric approaches were successfully used in pseudoscorpion taxonomy to distinguish Ephippiochthonius Beier, 1930 (Chthoniidae) [46] and Apolpium Chamberlin, 1930 (Olpiidae) [126] species. In Chernetidae, multivariate morphometric techniques successfully detected morphological differentiation of three Lasiochernes Beier, 1932 species and highlighted the most reliable characters for their identification [47]. ...
Pseudoscorpions represent an ancient, but homogeneous group of arachnids. The genus Lamprochernes comprises several morphologically similar species with wide and overlapping distributions. We implemented an integrative approach combining molecular barcoding (cox1), with cytogenetic and morphological analyses in order to assess species boundaries in European Lamprochernes populations. The results suggest ancient origins of Lamprochernes species accompanied by morphological stasis within the genus. Our integrative approach delimited three nominal Lamprochernes species and one cryptic lineage Lamprochernes abditus sp. nov. Despite its Oligocene origin, L. abditus sp. nov. can be distinguished from its closest relative only by molecular and cytogenetic differences, or alternatively, by a complex multivariate morphometric analysis involving other Lamprochernes species. The population structure and common haplotype sharing across geographically distant populations in most Lamprochernes species suggest that a phoretic manner of dispersal is efficient in this group.
... The applicability of these methods for differentiation of pseudoscorpion species has been studied on the family of Chthoniidae. Muster et al. (2004) used multivariate analyses to separate two European species of the genus Chthonius. ...
... 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). ...
Morphological variation in three rarely collected European species of the genus Lasiochernes Beier, 1932 is thoroughly examined in the present study. Detailed descriptions of previously ignored morphological characters of L. cretonatus Henderickx, 1998, L. jonicus (Beier, 1929) and L. pilosus (Ellingsen, 1910) are presented. The female of L. cretonatus and the nymphs of L. pilosus are described for the first time. Multi-variate morphometric techniques (principal coordinate analysis and discriminant analyses) were employed to confirm morphological differentiation of the three Lasiochernes species and to identify the most reliable characters for their separation. The usefulness of particular body parts for species identification was evaluated. An identification key for the females of the Lasiochernes species studied is provided. Geographic distribution and habitat preferences of the three species are summarized.
... 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). ...
Pseudoscorpiones is a mesodiverse order of Arachnida, with more than 3600 described species in the world. Species delimitation is sometimes difficult due to the high variability of some characters, difficulty in finding informative and useful characters for species differentiation and the subjectivity of individual specialists. The aim of this work is to explore and to evaluate the role of morphometric characters in species delimitation within the genus Apolpium (Olpiidae). Measurements of 17 structures of 57 individuals were taken and their length/width ratios calculated. Cluster analyses, principal components analyses, discriminant function analyses and non-metric multidimensional scaling were carried out. The results showed low resolution for the number of species currently recognised in the genus. These results highlight the importance of looking for other morphological characters, and also for a more rigorous assessment of previously used characters for defining species.
... Ganz besonders trifft dies auf die sehr kleinen Vertreter der Gattung Chthonius zu, die in der Regel durch ein geringes Sprungvermögen ausgezeichnet sind. Seit dem Erscheinen des Weltkatalogs der Pseudoskorpione (HARVEY 1991) wurden bereits vier Arten neu für Deutschland nachgewiesen: Chthonius (Ephippiochthonius) parmensis Beier, 1963(DROGLA 1990, Chthonius (Chthonius) diophthalmus Daday, 1888(DROGLA & LIPPOLD 1994, Chthonius (Chthonius) alpicola Beier, 1951(MUSTER & LIPPOLD 2003 boldorii Beier, 1934(MUSTER et al. 2004 ...
Chthonius (Ephippiochthonius) nidicola – new to Germany. The first record of this species outside the Alps is presented from the Rhineland-Palatinate, Germany. The locality, Bad Neuenahr, represents a considerable extension of the known range of a species which has up till now been regarded an endemic of southern Switzerland.
... 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. ...
This work stems from the results of a recent phylogenetic investigation on the Euscorpius carpathicus species complex from the Italian peninsula (Salomone et al. 2004. Phylogenetic relationships between the sibling species Euscorpius tergestinus and E. sicanus (Scorpiones, Euscorpiidae) as inferred from mitochondrial and nuclear sequence data. In: Proceedings of the16th Congress of Arachnology, August 2-7, 2004, Ghent University, Belgium, 268pp.; Salomone et al. in prep.). Molecular investigation produced interesting and unexpected findings on the scorpion Euscorpius tergestinus (C.L. Koch, 1837). Both nuclear and mitochondrial sequence data provided evidence of substantial genetic differentiation in specimens identified as Euscorpius tergestinus according to recent taxonomical changes (Fet andSoleglad2002. Morphology analysis supports presence of more than one species in the '' Euscorpius carpathicus'' complex (Scorpiones: Euscorpiidae). Euscorpius 3, 51pp.). These specimens clearly belong to two well- differentiated evolutionary lineages. Molecular results highlighted the need for a new morphological investigation. The present study undertook the morphological analysis of specimens belonging to both genotypes with the aim of identifying morphological characteristics able to discriminate between the two taxa. The analysis of trichobothria patterns, morphometric ratios, granulation patterns andthe observation of the pectinal sensilla confirm the d ifficulty in distinguishing these two genotypes and the high polymorphism of the subgenus Euscorpius Thorell, 1876. The length of pedipalp segments and dorsal patellar spurs (DPS), as well as femur leg granulation, are the main diagnostic characters; other ratios together with body color also help to distinguish the different genotypes. This study confirms the presence in Italy of two different cryptic species belonging to the ''Euscorpius tergestinus'' complex. Euscorpius tergestinus is a reddish, slender euscorpiid with a large dorsal patellar spine (DPS). A darker and generally squat phenotype with a short DPS, which corresponds to Euscorpius carpathicus concinnus sensu Caporiacco (1950), is elevatedto the species level: Euscorpius concinnus (C.L. Koch, 1837). These two species are sympatric in several Italian regions, and their distribution pattern is possibly determined by intraguild predaction interaction. r 2005 Elsevier GmbH. All rights reserved.
... 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. ...
An illustrated dichotomous key is provided for the identification of adults of the pseudoscorpion species found in the Czech Republic and Slovakia, based on morphological and morphometric characters. The pseudoscorpion fauna of Slovakia includes 51 species in 7 families, and that of the Czech Republic includes 38 species in 7 families. New country records are provided: Mundochthonius carpaticus Rafalski and Microbisium brevifemoratum (Ellingsen) in the Czech Republic and Microbisium suecicum Lohmander in Slovakia.
... 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). ...
For spiders, morphological differentiation within genitalic traits is the main diagnostic criterion of a species. Beside some well-described exceptions of genitalic polymorphism and crypticity, spider genitalic variation is seldom quantitatively analyzed. Using geometric morphometrics landmark analysis, we report clear evidence of quantitative interspecific divergence and intraspecific variation in the genital shape of three species of the genus Paratrechalea (P. azul, P. ornata and P. galianoae). The genitalic species recognition was very consistent with our quantitative data for both sexes. Interspecific variation suggested a character displacement pattern between two syntopic populations of P. azul and P ornata, and also a possible case of species crypticity in P. ornata that will involve splitting the Uruguayan populations from the Brazilian ones.
Nested within the Mediterranean biodiversity hotspot, the Dinaric Karst of the western Balkans is one of the world's most heterogeneous subterranean ecosystems and renowned for its highly diverse and mostly endemic fauna. The evolutionary processes leading to both endemism and diversity remain insufficiently understood, and large-scale analyses on taxa that are abundant in both subterranean and surface habitats remain infrequent. Here, we provide the first comprehensive molecular study on Croatian pseudoscorpions, a lineage of arachnids that is common and diverse in both habitats. Phylogenetic reconstructions using 499 COI sequences derived from 128 morphospecies collected across the Dinaric Karst show that: (i) occurrence in karstic microhabitats boosters speciation and endemism in the most diverse genera Chthonius C.L. Koch, 1843 (37 morphospecies) and Neobisium Chamberlin, 1930 (34 morphospecies), (ii) evidence for ongoing diversification is found in many species and species complexes through low optimal thresholds (OTs) and species delineation analyses, and (iii) landscape features, such as mountain ranges, correlate with patterns of genetic diversity in the diverse genus Neobisium. We present two synonymies: Protoneobisium Ćurčić, 1988 = Neobisium, syn. nov., and Archaeoroncus Ćurčić and Rađa, 2012 = Roncus L. Koch, 1873, syn. nov. Overall, our study suggests that karstic microhabitats promote diversification in soil-and cave-dwelling arthropods at all taxonomic levels, but also provide important refugia for invertebrates in past and present periods of environmental change.
The fifth insect camp of the Entomological Society
of Austria (ESA) was conducted from April 27 to May 2, 2018. Many of the 39 participants
were recognized experts on different arthropod groups. Fifty-seven localities
within and nearby the Donau-Auen National Park were investigated, and a total of 1265
invertebrate species identified: 8 Collembola, 1 Zygentoma, 19 Odonata, 2 Blattodea,
1 Derma ptera, 9 Ortho ptera, 66 Hetero ptera, 52 Hemi ptera, 2 Neuro ptera, 443
Coleo ptera, 1 Meco ptera, 76 Di ptera, 190 Hymeno ptera, 35 Tricho ptera, 259 Lepidoptera,
11 Diplopoda, 10 Chilopoda, 10 Isopoda, 3 Araneae, 10 Pseudoscorpiones,
2 Opiliones, 5 Acari, 6 Oligochaeta, 3 Bivalvia, and 41 Gastropoda. New to Vienna
and Lower Austria were 9 and 7 species, respectively. The rare hoverfly Epistrophe
cryptica Doczkal & Schmid, 1994 (Di ptera: Syrphidae) was found in Austria for the
first time. One ♀ and one ♂ of the yet undescribed leafhopper Doratura sp. near
paludosa (Hemi ptera: Cicadellidae) were collected.
The updated checklist contains 50 taxa. Since 2004, two species have been recorded as new, one has been split, and two species had to be removed from the list. The red data list is prepared according to the present requirements of the German Federal Agency for Nature Conservation (BfN). For each taxon the frequency in Germany is classified into one of six criteria classes, from extremely rare to very abundant; the long-term and the short-term trend of each taxon’s population is estimated. Based on these estimates the level of threat for each taxon is evaluated using a standardised assessment scheme provided by the BfN. One pseudoscorpion species is considered missing/extinct (Neobisium crassifemoratum), one species is considered endangered (Anthrenochernes stellae), four taxa are categorised as vulnerable (Mundochthonius styriacus, Microbisium brevifemoratum, Syarinus strandi, Dendrochernes cyrneus,), five are generally threatened (without possible estimate of level of threat), seven are extremely rare and for nine species the data is deficient. A high percentage of the taxa (72 % = 36 taxa) has its distribution limits extending through Germany. For seven taxa Germany has a particularly high degree of responsibility, since a major part of the taxa ranges correspond with the country’s territory (Chthonius nidicola, Mundochthonius styriacus, Neobisium hermanni, Syarinus strandi, Dactylochelifer latreillii septentrionalis, Chernes vicinus, Lasiochernes pilosus). Taxonomic changes: A correction of the year of description of Dinocheirus panzeri is conducted.
A taxonomic revision and a key to the species of the subgenus Chthonius (Ephippiochthonius) Beier, 1930 from Italy, Corsica and the Swiss Canton of Ticino are provided. The species are arranged in two species-groups (tetrachelatus group and fuscimanus group) on the basis of the shape of pedipalpal hand and of the type of dentition of the fixed and movable chelal fingers. The following new species are described: i) in the tetrachelatus group: Chthonius (E.) altamurae n. sp. (♀, loc. typ.: Apulia, Bari Prov., Altamura, Grotta Lamalunga 1295 Pu/BA), C. (E.) elymus n. sp. (♂, loc. typ.: Sicily, Trapani Prov., Custonaci, Abisso del Purgatorio 8064 Si/TP), Chthonius (E.) messapicus n. sp. (♂, loc. typ.: Apulia, Brindisi Prov., San Pietro Vernotico, Cerano); ii) in the fuscimanus group: C. (E.) aeneae n. sp. (♂♀, loc. typ.: Liguria, Genoa Prov., Sestri Levante, Punta Manara), C. (E.) etruscus n. sp. (♂♀, loc. typ.: Tuscany, Grosseto Prov., Semproniano, Grotta di Montecchio 254 To/GR), C. (E.) gallii n. sp. (♂♀, loc. typ.: Liguria, Savona Prov., Bergeggi), C. (E.) intemelius n. sp. (♂♀, loc. typ.: Liguria, Imperia Prov., Apricale, Mt Cianela), C. (E.) latellai n. sp. (♂♀, loc. typ.: Latium, Latina Prov., Bassiano, Grotta di Fiume Coperto 1361 La/LT), C. (E.) ligur n. sp. (♂♀, loc. typ.: Liguria, Imperia Prov., near Baiardo), C. (E.) magrinii n. sp. (♂, loc. typ.: Latium, Frosinone Prov., San Giovanni Incarico, Grotta sulla strada per il Santuario della Madonna della Guardia n. c. La/FR), C. (E.) monguzzii n. sp. (♂♀, loc. typ.: Lombardia, Brescia Prov., Sulzano, Oricina de la Pofa del Giardì 438 Lo/BS), C. (E.) sulphureus n. sp. (♂♀, loc. typ.: Marche, Ancona Prov., Genga, Grotta di Frasassi 1 Ma/AN), C. (E.) tyrrhenicus n. sp. (♂♀, loc. typ.: Liguria, Genoa Prov., Genoa, Quinto al Mare, Mt Moro). The following new synonymies are proposed: Chthonius (E.) bauneensis Callaini, 1983 is a junior subjective synonym of C. (E.) berninii Callaini, 1983 (n. syn.), C. (E.) aegatensis Callaini, 1989 is a junior subjective synonym of C. (E.) berninii Callaini, 1983 (n. syn.), Chthonius (E.) maltensis Mahnert, 1975 is a junior subjective synonym of Chthonius (E.) concii Beier, 1953 (n. syn.), Chthonius (E.) bartolii Gardini, 1976 is a junior subjective synonym of Chthonius (E.) concii Beier, 1953 (n. syn.), Chthonius (E.) elbanus Beier, 1963 is a junior subjective synonym of Chthonius (E.) nanus Beier, 1953 (n. syn.), Chthonius (E.) cavicola Gardini, 1990 is a junior subjective synonym of Chthonius (E.) troglophilus Beier, 1930 (n. syn.). Thirty-nine species of Ephippiochthonius are known at present from the above-mentioned areas, of which one is presumably endemic to Sicily, four to Sardinia and four to Corsica. C. (E.) gibbus Beier, 1953 is excluded from the Ital-ian fauna and C. (E.) vachoni Heurtault-Rossi, 1963 is newly recorded from Italy. Chthonius (E.) poeninus Mahnert, 1979 is transferred to the subgenus Globochthonius Beier, 1931.
In the course of a long term survey on the effects of grazing on the diversity of alpine grasslands, the false scorpion Chthonius (Ephippiochthonius) poeninus Mahnert, 1979 was recorded for the first time outside of Switzerland. The preferred habitat at the locality Alpe Einödsberg (Germany, Bavaria) differs strongly from previous findings. Our data suggest an association with Nardus grasslands (Geo montani-Nardetum), whereas earlier records originated from leaf litter, moss and mouldy trunks. Furthermore, the new records between 1540 m and 1973 m above sea level are the first from the subalpine region of the Alps (hitherto C. poeninus was known from 550 m to 1450 m). In the study region, extensive grazing seems to have a positive effect on the abundance of the species, while it is missing from intensively grazed pastures.
An annotated and revised checklist is presented comprising 58 species (8 families) of the arachnid order Pseudoscorpiones from the Czech Republic (38 species) and Slovakia (51 species), formerly Czechoslovakia. According to the latest detailed analyses and revisions Chthonius carinthiacus Beier, 1951, Neobisium validum (L. Koch, 1873), Roncus alpinus L. Koch, 1873 and Pselaphochernes dubius (O. P.-Cambridge, 1892) are excluded from the checklist from Slovakia. Formerly recorded Chthonius diophthalmus Daday, 1888, C. submontanus Beier, 1963, C. kewi Gabbutt, 1966, Neobisium spelaeum (Schiödte, 1847), N. jugorum (L. Koch, 1873), Roncus alpinus and R. transsilvanicus Beier, 1928 have not been included in the list from the Czech Republic. Some generally rare species are discussed.
Subterranean ecosystem studies using multivariate ordination and/or agglomerative classification statistical methods were reviewed in the Science Citation Index (SCI) between 1945 and 2006. Nearly 57,000 publications cited subterranean habitats or their associated biota in the SCI abstracts, however, multivariate statistics applied to strictly hypogean taxa occurred in only 65 papers from 1990 onwards. Over 90% of the multivariate applications were devoted to morphometric or genetic studies of single species and to relationships between the environment and species assemblages. In terms of taxa and ecosystem types, stygobite and waterless cave studies featuring multivariate applications predominated, respectively. Only six different methods (Agglo-merative Clustering, Canonical Correspondence Analysis, Correspondence Analysis, Discriminant Analysis, non-metric Multidimensional Scaling, Principal Component Analysis) were used among the .30 multivariate techniques available within the biostatistical toolbox. The retrieved set of publications was sorted in a simple table by keyword according to type of biota, habitat, research topic and multivariate method, while online biostatistical resources are appended. Further comments are made on the use of statistics in the biological sciences in general.
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.
A systematic revision of the family Sternophoridae is presented. Three genera are recognized: Garyops Banks, Idiogaryops Hoff and Afrosternophorus Beier; the latter is given full generic status. These genera are distinguished solely on the female genitalia. The following generic synonymies are proposed (junior synonym first): Sternophorus Chamberlin = Garyops ; Sternophorellus Beier = Afrosternophorus ; lndogaryops Sivaraman = Afrosternophorus . The following species are synonymized with A. ceylonicus (Beier): Sternophorus transiens Murthy and Ananthakrishnan, S. indicus Murthy and Ananthakrishnan, S. montanus Sivaraman, S. femoratus Sivaraman, S. intermedius Sivaraman and Indogaryops amrithiensis Sivaraman. Four new species are proposed: Afrosternophorus anabates (Australia), A. fallax (Vietnam), A. nanus (Australia) and A. xalyx (Australia). Garyops contains depressus Banks, sini (Chamberlin), centralis Beier and, possibly, ferrisi (Chamberlin). Idiogaryops consists of paludis (Chamberlin) and pumilus (Hoff). Afrosternophorus contains aethiopicus (Beier), anabates , new species, araucariae (Beier), cavernae (Beier), ceylonicus (Beier), chamberlini (Redikorzev), cylindrimanus (Beier), dawydoffi (Beier), fallax , new species, grayi (Beier), hirsti , (Chamberlin), nanus , new species, papuanus (Beier) and xalyx , new species. Two species-groups are proposed in each of the two genera Idiogaryops and Afrosternophorus to accommodate species with differing trichobothrial numbers. Male genitalia was found to be a useful adjunct to traditional characters, and a detailed description of the male genitalia of A. hirsti is presented. Postembryonic development, biogeography and possible evolutionary pathways are discussed.
Twenty-eight species of the Australian and New Zealand genus Synsphyronus are recognised: S. absitus, sp. nov.; S. amplissimus, sp, nov.; S. apimelus, sp, nov.; S. attiguus, sp. nov.; S. bounites, sp. nov.; S. callus Hoff; S. dewae Beier; S. dorothyae, sp. nov.; S. ejuncidus, sp. nov.; S. elegans Beier; S. gigas Beier; S, gracilis, sp. nov.; S. greensladeae, sp. nov.; S. hadronennus, sp. nov.; S. hansenii (With); S. heptatrichus, sp. nov.; S. lathrius, sp. nov.; S. leo, sp. nov.; S. lineatus Beier; S. magnus Hoff; S. meganennus, sp. nov.; S. melanochelatus (Chamberlin); S. mimetus Chamberlin; S. mimulus Chamberlin; S. niger Hoff; S. nullarborensis Beier; S. paradoxus Chamberlin; S. silveirai, sp. nov. The following synonymies are proposed (junior synonym first): S. grayi Beier =S. mimulus Chamberlin; S. gisleni Beier = S. hansenii (With); S. fallaciosus Beier = S. hansenii (With). Maorigarypus viridis Tubb is designated as a nomen dubium. The subgenera Synsphyronus, s. str., and Maorigarypus Chamberlin, which have been distinguished on the presence or absence, respectively, of fusion of the pedal tarsi, are abandoned because the degree of tarsal fusion was found to be intraspecifically variable. Post-embryonic development, biogeography and possible evolutionary pathways are discussed.
Molecular differentiation between races or closely related species is often incongruent with the reproductive divergence of the taxa of interest. Shared ancient polymorphism and/or introgression during secondary contact may be responsible for the incongruence. At loci contributing to speciation, these two complications should be minimized (1, 2); hence, their variation may more faithfully reflect the history of the species' reproductive differentiation. In this study, we analyzed DNA polymorphism at the Odysseus (OdsH) locus of hybrid sterility between Drosophila mauritiana and Drosophila simulans and were able to verify such a prediction. Interestingly, DNA variation only a short distance away (1.8 kb) appears not to be influenced by the forces that shape the recent evolution of the OdsH coding region. This locus thus may represent a test case of inferring phylogeny of very closely related species.
Troglochthonius doratodactylus Helv. is recorded from two caves near Triest discrediting the original (erroneous) record from Sardinia. Chthonius (E.) boldorii Beier is recorded from Krk, its status is discussed. A few other cave-dwelling species are mentioned-.
From a study of the geographic occurrences of contemporary hybridization among North American animals, it has become apparent that most of the hybrids are produced in a few relatively localized zones, with little hybridizing in the vast areas between these zones of mixing. The hybrids tend to be at least moderately fertile and therefore to be a source of significant gene exchange between the typically allopatric pairs of species and semispecies. There is a wide variety of consequences from this introgression, with greater or lesser influence on the parental populations, and large portions of the fauna and probably flora are involved. An appropriate term for such a belt of interfaunal and interfloral linkage is suture-zone.1
Differences in secondary sexual characteristics of males often provide the most conspicuous means of distinguishing between closely related species. Does this therefore imply that the absence of differentiation in exaggerated male traits between allopatric populations provides evidence of a single, genetically cohesive species? We addressed this question with a comprehensive investigation of two populations (French Guiana and Panama) of the harlequin beetle-riding pseudoscorpion, Cordylochernes scorpioides. This highly sexually dimorphic pseudoscorpion is currently described as a single species, ranging throughout the Neotropics. Our morphometric analyses detected minimal differentiation between the two populations in all nine external morphological characters measured, including sexually dimorphic traits in males. Only in traits of the spermatophore was there any appreciable level of differentiation. Behavior differentiation and prezygotic reproductive isolation were also limited: 78.3% of males successfully transferred sperm to "foreign" females, and in 63.9% of these cases, females' eggs were successfully fertilized. By contrast, extensive divergence existed in two of nine electrophoretic loci, including an essentially fixed-allele difference at the Ldh locus. Most significantly, postzygotic reproductive isolation was complete, with heteropopulation zygotes invariably aborting early in development. These results strongly suggest that the two populations are, in fact, sibling species, a conclusion supported by our recently published findings on their marked divergence in minisatellite DNA. How can such interpopulation homogeneity in male sexually dimorphic traits exist in the face of strong genetic divergence? We propose that sexual selection, oscillating between favoring small and then large males, maintains such high levels of male variability within each population that it has obscured a speciation event in which genetic divergence and postzygotic incompatibility have clearly outpaced the evolution of prezygotic reproductive isolation.
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 greatly augment previous fossil data. The refugial genomes contributed differently in various species to the 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.
The two male morphs of the dimorphic jumping spider, Maevia inclemens, differ dramatically in morphology and courtship behavior. The purpose of this study was to examine and compare the mating success of the two male types. Mating success was measured by the number and duration of copulation events, the latency of visual orientation by the female to a courting male, signals of female receptivity, risk of predation by the female, and the number of offspring produced by each morph. The morphs did not differ significantly with respect to copulation success, mating events, mating duration, signals of receptivity or the number of offspring produced. However, males did differ with respect to latency of visual orientation as a function of distance from the female. Near to the female, the gray males attracted female attention in significantly less time than tufted males. Conversely, at far distances from the female, the tufted males attracted female attention in less courtship time. This study suggests that males attain equal levels of mating success and that the two male morphs may have evolved alternative reproductive strategies for courtship at different distances from the female.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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.
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);