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Modern agricultural pest management systems rely on naturally occurring generalist predators to promote pest suppression. Still, little research has been done to assess their overall effectiveness, especially over the winter period when their potential is high. In this study, we focused on three genera of winter-active spiders Clubiona, Philodromus, and Anyphaena, common predators on pear trees in Central Europe during winter and early spring. We investigated their predation activity, natural diet, and prey preference using molecular gut content and abundance data obtained from cardboard bands, which served as natural shelters. We compared these characteristics between two distinct biocontrol-promoting managements—integrated pest management (IPM) and organic management (ORG). We found the proportion of prey-positive spider individuals during the winter period to be lower compared to the spring period with Anyphaena having by far the highest proportion. The prey composition during winter was more diverse in ORG orchards, but in both managements, it was inclined toward pests, mostly pear psyllids. Conversely, in early spring, despite psyllids still being a part of the diet, spiders in IPM orchards preyed more frequently on neutral prey (mostly dipterans), while the spiders from organic orchards preyed mostly on pests (lepidopterans). Although more data are needed to assess trophic interactions and the overall efficiency of these winter-active predators in complex arthropod food webs present in pear orchards, the results obtained from this research provide the first evidence of higher pest predation during a period of agricultural quiescence. [ABSTRACT FROM AUTHOR]

Philodromus poecilus (Thorell, 1872) is a Transeurasian species reported from most parts of Europe except the British Isles and the Iberian Peninsula. Although widely distributed, this species is extremely rarely recorded in Germany and neighbouring countries. In Bavaria, it has not been found since the time of Ludwig Koch (1825-1908). Recently, several specimens were collected from urban plane trees in Deggendorf (Lower Bavaria), proving that the species is still present in Bavaria. Additionally, images of the copulatory organs and of living and preserved specimens are presented, as is a compilation of records in Germany and neighbouring countries. [ABSTRACT FROM AUTHOR]

Understanding the full diet of natural enemies is necessary for evaluating their role as biocontrol agents, because many enemy species do not only feed on pests but also on other natural enemies. Such intraguild predation can compromise pest control if the consumed enemies are actually better for pest control than their predators. In this study, we used gut metabarcoding to quantify diets of all common arachnid species in Swedish and Spanish apple orchards. For this purpose, we designed new primers that reduce amplification of arachnid predators while retaining high amplification of all prey groups. Results suggest that most arachnids consume a large range of putative pest species on apple but also a high proportion of other natural enemies, where the latter constitute almost a third of all prey sequences. Intraguild predation also varied between regions, with a larger content of heteropteran bugs in arachnid guts from Spanish orchards, but not between orchard types. There was also a tendency for cursorial spiders to have more intraguild prey in the gut than web spiders. Two groups that may be overlooked as important biocontrol agents in apple orchards seem to be theridiid web spiders and opilionids, where the latter had several small-bodied pest species in the gut. These results thus provide important guidance for what arachnid groups should be targets of management actions, even though additional information is needed to quantify all direct and indirect interactions occurring in the complex arthropod food webs in fruit orchards. [ABSTRACT FROM AUTHOR]

Here, the philodromid species Philodromus dispar Walckenaer, 1826 is reported for the first time from Turkey. In addition, detailed descriptions of both male and female samples are provided. [ABSTRACT FROM AUTHOR]

Female mating history can have a strong effect on male fertilization success. Although males often prefer to mate with virgin females, they often also engage with mated females. As the intensity of sperm competition can differ among mated females, males are expected to evolve means to identify their status. In spiders, males often use female silk to gather information about female quality. Males of many spider species deposit mating plugs into female genitalia to hinder further copulations. We tested whether males of the foliage‐dwelling, plug‐producing spider Philodromus cespitum, which is an important natural enemy of pests, discriminate between females of different mating status and whether they can determine the extent of genital plugging in mated females solely on the basis of cues gained from deposited female silk. We presented males with draglines of females that varied in either mating status (virgin vs. mated), the extent of plugging (small vs. big plug), or the age of the plug (fresh vs. old plug) and examined their mate preferences. Additionally, we tested whether males were attracted to volatile cues produced by female bodies. Our experiments revealed that males preferred draglines of virgin females to those of mated females, and mated females with small plugs to those with large plugs. They were also attracted to female volatile cues. This study suggests that males are able to extract fine‐scale information on mating status from female draglines. [ABSTRACT FROM AUTHOR]

Even though pesticides can have various sublethal effects on behaviour of biocontrol agents, no study to date has investigated the effects of pesticides on the prey choice of generalist predators. Prey choice of generalist predators is among key factors determining the predation pressure they exert on pests, because it influences fitness of predators and consequently their densities and per capita capture rate. Here, we investigated the effect of Integro (a.i. methoxyfenozide) and SpinTor 480 Sc (a.i. spinosad) on prey choice and predatory activity of the spider Philodromus cespitum , which is known significantly to reduce hemipteran and dipteran pests in fruit orchards. We compared the prey preferences of philodromids between the psyllid pest Cacopsylla pyri (Hemiptera, Psyllidae) and beneficial Theridion sp. spiders in laboratory experiments. We found that both pesticides altered the prey preferences of philodromids. While the philodromids in a control treatment preferred theridiid spiders over the psyllid pest, philodromids in the pesticide treatments exhibited no significant prey preferences. The changes in prey preferences were caused by increased predation on the psyllids, while the predation on the theridiid remained similar. We suggest that the changes in prey preferences might theoretically be due to 1) impaired sensory systems, 2) altered taste, and/or 3) altered mobility. In combination with other studies finding reduced predation on fruit flies in P. cespitum after exposure to SpinTor, our results indicate that the sublethal effects of pesticides on predatory behaviour of generalist predators can depend on prey type and/or prey community composition. [ABSTRACT FROM AUTHOR]

Philodromus buchari Kubcová, 2004 is reported for the fi rst time from Hungary. A relatively large and abundant population of Ph. buchari was found in Diósd (near Budapest), and aft er examining other materials, it seems that the species is widespread throughout the country. In the past Ph. buchari was misidentified as Ph. aureolus (Clerck, 1757), Ph. cespitum (Walckenaer, 1802) or Ph. longipalpis Simon, 1870, and possibly it is a native species in Hungary. It may have a wider distribution in Europe and the Mediterranean region as previously was thought. Copulatory/genital organs and habitus of Ph. buchari are illustrated. With 11 fi gures. [ABSTRACT FROM AUTHOR]

The sum of fruit and seed predation by multiple species may strongly affect plant reproduction and population dynamics. We evaluated the combined effects of ungulates, seed-eating rodents and insect pre-dispersal seed predators on the reproductive success of the Mediterranean gum cistus shrub ( Cistus ladanifer ), over two consecutive years within a long-term ungulate-exclusion experiment. We compared fruiting success in shrubs exposed and protected from ungulates by examining fruit abortion and fruit production. We also investigated the effect of insect predation on seed production (i.e. proportion of depredated fruit and seed loss) and measured fruit weight, seed number per fruit, and seed weight of unpredated fruits. Ungulate browsing directly removed 42.3% of the plant reproductive structures, early in the reproductive season and insect predation reduced mature seeds by over 40%. Results also emphasize the additive effects of ungulate browsing on pre-dispersal insect predation and fruit abortion which increased by 74.7% and 60.9%, respectively. Rodents, which only occurred in ungulate-excluded plots, had a limited and later effect on seed production with 6% of mature fruit loss. Fruit weight, seed weight and number were higher in shrubs protected from ungulates. Our study indicated that seed predation by mice was irrelevant, but ungulate and invertebrate seed predation interacted to strongly limit the reproductive success of C ladanifer , potentially affecting plant population dynamics in the long-term. [ABSTRACT FROM AUTHOR]

Longevity is a key demographic characteristic of herbaceous plants, but often unknown. While root or rhizome growth ring analysis may allow assessment plant longevity directly and conveniently, so far it has only been used in a few case studies of herbaceous dicotyledonous species. To evaluate whether growth ring analysis is applicable to a large spectrum of herbaceous dicotyledonous plant species, we used plant communities of varying species richness in a 12-year-old grassland biodiversity experiment (Jena Experiment). Cross-sections of the oldest available part of the plants were analysed for all available dicotyledonous perennial herb species (S = 37), which represented three functional groups: legumes, small herbs and tall herbs. We studied 1664 individuals representing the genet in clearly distinguishable plant individuals, and the ramet in clonally growing plant species. Roots of eleven species with permanent primary root were harvested. They showed clearly visible growth rings. Longevity was extended with a mean age of 4.0 years (SE = 0.3). Seven species, which also had a permanent primary root, showed less distinct growth rings. They were shorter-lived (mean age 3.0 years (SE = 0.3)). In six species with obligate clonal growth mostly rhizomes were sampled, but individuals were still identifiable due to their growth habit. For these species growth rings were clearly visible. Longevity of rhizomes was extended (mean age 3.3 years (SE = 0.5)). In 13 species with obligate clonal growth also rhizomes were sampled, but plant individuals were not identifiable. For these species longevity was low (mean age 2.1 years (SE = 0.2)). Community mean age was significantly lower when small herbs were present and higher when tall herbs were present, while legumes had no effect on community mean age. In summary, anatomical analysis of roots and rhizomes is a suitable tool to study the population age structure of a large spectrum of perennial dicotyledonous herbaceous species and therefore opens new perspectives for demographic studies at the community level. [ABSTRACT FROM AUTHOR]

Hedgerows are agro-ecological infrastructures that are assumed to enhance biodiversity in an agro-ecosystem and the control of crop pests. However, local movements of arthropod predators from hedgerows to crops remain poorly understood. In this research, these movements were analysed in eleven commercial apple orchards over two weeks in the spring of 2014. Predators were indirectly marked by spraying ovalbumin on the hedgerows. Canopy and ground predators were captured using beating and pitfall traps, respectively, in both the orchards and the hedgerows, and individuals marked by ovalbumin were detected using ELISAs. Approximately 20% of the 1272 captured predators were identified as marked. Movements between the orchards and the hedgerows of the most abundant predator taxa ( Forficula auricularia , Chrysoperla sp., Philodromus spp., Cheiracanthium mildei , and Nebria brevicollis ) and of four guilds (ground spiders, ground beetles, canopy spiders and canopy insects) captured across the eleven locations were estimated using a Bayesian model. On one hand, canopy insects and ground spiders were less likely to stay in the hedgerows than were the canopy spiders and ground beetles. On the other hand, the canopy spiders and ground beetles were less likely to stay in the orchard than were the canopy insects and ground spiders. However, there were exceptions within these groups: F. auricularia and N. brevicollis exhibited a high probability of staying in the hedgerow and in the orchard, respectively. Overall, these results demonstrate the frequent movements that occurred between the orchard and the adjacent hedgerow in a diverse range of predator taxa. The probabilities of movement were further affected by the characteristics of the orchard for most taxa. Therefore, agro-environmental measures that focus on hedgerow management require consideration of the local arthropod predator communities and their characteristics to enhance pest control in apple orchards. [ABSTRACT FROM AUTHOR]

Artificially increased habitat complexity can improve the biocontrol service provided by generalist predators as it can reduce intraguild predation (IGP). However, several counteracting mechanisms can buffer the effect of reduced IGP. Here, we investigated whether the cardboard bands installed in pear trees in early winter improve the suppression of the serious pest pear psylla ( Cacopsylla pyri ) by winter-active spiders ( Anyphaena accentuata and Philodromus spp.). We also investigated in laboratory experiments whether the increased abundances of some predators ( Philodromus sp.) would result in negative or positive predator-predator interactions and whether the type and/or intensity of interaction is temperature dependent. On average, pear trees with cardboard bands hosted less psylla than control trees. This pattern can be attributed to predation by winter-active spiders, as 45% of individuals were psylla positive using molecular gut-content analysis. However, 58% of Anyphaena spiders were also Philodromus positive. The suppression efficiency for psylla increased asymptotically with the increasing density of winter-active spiders. In the laboratory, the per capita capture rate of philodromids decreased with the density of conspecifics, which translated in an asymptotic increase in predation pressure on the pear psylla. Non-consumptive intra-trophic-level interference was more intense at 12 °C than at 7 °C. The results show that artificially increasing habitat complexity by installing corrugated cardboard bands around the trunks and branches of pear trees increased predation pressure on the pear psylla as it most likely reduced IGP among winter-active spiders. However, increased intra-trophic-level interference due to increased abundances of spiders reduced their per capita capture rate and caused only an asymptotic increase in the overall predation pressure on the pest. The installation of corrugated cardboard bands around trees could represent an environmentally-friendly management practice in fruit orchards that improves the biocontrol service provided by generalist predators. [ABSTRACT FROM AUTHOR]

Behavioral syndromes, seen as correlations among two or more functionally different behaviors, are well documented in many different animal taxa. They may not be present automatically, however, and their consistency within populations and individuals also varies among studies. Here, we studied a behavioral syndrome comprising foraging aggressiveness and boldness/shyness and its time consistency in the cursorial spider Philodromus albidus. We measured foraging aggressiveness as the number of prey killed per 2-hour period. Boldness/shyness was assessed as a latency of initiating exploration in a novel environment. We found the presence of the behavioral syndrome in P. albidus, as bold individuals were also more aggressive than were shy individuals. The syndrome was consistent through time within the population but not consistent for individuals. We further discuss the possible causes of the pattern. [ABSTRACT FROM AUTHOR]

Although temperature significantly influences pesticides' impact on ectotherms, the joint effect of temperature and pesticides on natural enemies is understudied. In laboratory conditions, we investigated the influence of two commonly used insecticides, spinosad and λ-cyhalothrin, on mortality and functional response of a community of Philodromus spiders ( P. cespitum 80 %, P. albidus 20 %) at different temperatures (10, 17, 25, and 31 °C). Almost all spiders died at 31 °C in the λ-cyhalothrin treatment. Mortality was very low at other temperatures. Mortality in the spinosad treatment did not differ from the control. Both pesticides reduced the prey capture rate and/or changed the type of functional response at all temperatures. The sublethal effects of λ-cyhalothrin were most pronounced at 10 °C, while those of spinosad were most marked at 10 and 25 °C. At 10 °C, the pesticides' effects on the spiders were long lasting. In addition, λ-cyhalothrin influenced the philodromids much more negatively than spinosad suggesting its incompatibility with IPM across all temperatures. The results support the importance of studying the temperature-specific effects of pesticides in order to increase the efficiency of integrated pest management. Philodromids are effective in pest suppression at low temperatures, but application of pesticides at the beginning of the season can significantly disrupt their biocontrol potential. [ABSTRACT FROM AUTHOR]

A number of different generalist (polyphagous) predators occur in agroecosystems. Yet their biocontrol potential has been little investigated in detail. Philodromus species (Philodromidae) belong to the dominant spider species occurring in commercial orchards. We studied in detail the trophic functional traits of Philodromus albidus , Philodromus aureolus , and Philodromus cespitum (Philodromidae) by means of (1) the analysis of natural prey; and (2) experiments on acceptance of a variety of prey taxa. We found that the three philodromids are euryphagous. We classified prey species into three categories according to their function in the orchard: beneficial species, indifferent species, and pests. Philodromid spiders captured mostly other spiders in the field because spiders were most available. As concerns pests, the philodromids preyed mostly on Brachycera and Sternorrhyncha. They selected Acari and Brachycera. Indifferent species, such as Collembola and Nematocera, were also highly selected. In the laboratory, philodromids accepted mostly pests, such as lepidopterans, brachycerans, and aphids, while other spiders were accepted the least. The three philodromids have differentiated trophic niches with respect to prey size not only in the adult stage but throughout their ontogenetic development: P. albidus utilized smaller prey than the other two species. We conclude that the philodromids have a potential as biocontrol agents because they prey mostly on pests but their predation pressure is reduced due to higher selectivity for the indifferent fauna. [ABSTRACT FROM AUTHOR]

1. Several mechanisms can mediate the coexistence of species, such as the neutral dynamic, niche filtering and niche partitioning. The present study investigated which of these mechanisms mediate the coexistence of closely related spider species at the scale of one locality. 2. The coexistence of three spider species of the genus Philodromus (Philodromidae) ( Philodromus albidus, Philodromus aureolus and Philodromus cespitum) was studied. The study area comprised three habitat types, including a deciduous forest, a scrub and a plum tree stand. The spatial niche properties of the Philodromus species were explored by comparing their micro/macrohabitat preferences. The natural diet of these species was analysed. Laboratory experiments involving prey acceptance were conducted to investigate the trophic niche properties. The species' phenologies were studied to compare their temporal niches. 3. The philodromids had differentiated their trophic and habitat niches. The coexistence mechanisms were therefore assessed by studying the relationships between niche overlaps and the pairwise product of relative abundances. A negative relationship was observed between functional niche overlaps and the product of relative abundances, whereas a positive relationship was observed between spatial niche overlaps and the product of relative abundances. 4. The present results suggest that different mechanisms influence different niche dimensions. Niche partitioning influenced functional niches, whereas niche filtering influenced spatial niches. The results also suggest that the filtering process in one dimension was facilitated by niche differentiation in another dimension. [ABSTRACT FROM AUTHOR]

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Chemical and other selective insecticides are annually used as foliar treatments against defoliating Lepidoptera in the southern Spanish open woodland (known asdehesa), one of the largest forest ecosystems in the Mediterranean basin. In order to analyse the negative impact of these treatments on beneficial arthropods, the toxic effect of the three authorised insecticides onPhilodromus buxiSimon (Araneae: Philodromidae), the most abundant tree canopy spider in southern Spain, was tested under laboratory conditions. Cypermethrin was the most harmful insecticide 96 hours after application, causing around 90% corrected mortality, while flufenoxuron (an insect growth-regulator) prompted around 45% mortality, andBacillus thuringiensiscaused no mortality at all. In addition, flufenoxuron produced a clear alteration in the predatory capacity ofP. buxi, increasing the reaction times of treated specimens, and additional mortality associated with the moulting process.Bacillus thuringiensiscaused no effects in reaction times, growth or moulting. The results obtained clearly indicate thatB. thuringiensisis the most suitable of the authorised insecticides for conserving populations ofP. buxiindehesawoodland in southern Spain. [ABSTRACT FROM AUTHOR]

Hairy attachment devices that are not supplemented with fluid secretion have evolved independently in lizards and spiders. van der Waals forces have previously been shown to be responsible for excellent adhesive properties of these structures, but it has recently been reported that wetting phenomena also play an important role in such ‘dry adhesives’. To investigate the effect of ambient humidity on the attachment of the living spider Philodromus dispar, traction force was measured on a smooth epoxy resin surface at relative humidities (RHs) of 15, 50, 70, 80 and 99 per cent. The results show that attachment ability is significantly higher at an intermediate humidity compared with that in a dry atmosphere and at high humidity. Water condensation on the substrate surface almost completely abolishes adhesion. Experimental results obtained may be explained by an increase in capillarity or changes in mechanical properties of setae and spatulae owing to water absorption by the cuticle at an intermediate RH. The results obtained show dry adhesion limits under different environmental conditions and are important for understanding spider biology. [ABSTRACT FROM AUTHOR]

Abstract: We studied whether there si intraguild predation among overwintering invertebrates in bark traps installed on apple trees. We found that 86% of the overwintering invertebrates in traps were predators, which were dominated by spiders (72%) of different guilds and different body sizes. Spiders had spatially segregated in the traps so that smaller specimens hid in an interstitial space that was inaccessible to larger specimens. Spiders of the bark-dwelling guild, Anyphaena and Philodromus, are winter-active and can prey on other smaller spiders. Analysis of their abundance in traps indicated a negative relationship between Anyphaena and Philodromus or other spiders. Inspection of traps during winter revealed an influx of spider specimens confirming their winter-activity. In a semi-field experiment, where overwintering in cardboard bands was studied under controlled conditions, we observed mortality of both winter-active Philodromus and Anyphaena spiders due to their interspecific predation. The mortality was affected by their size ratio – larger specimens consumed smaller ones. Presence of alternative prey, theridiid spiders, did not affect their mortalities, but the presence of shelters (interstitial spaces) did. In boards with interstitial spaces smaller spiders suffered lower mortality than in boards without the interstitial spaces. Obtained results suggest that intraguild predation between overwintering spiders can decrease their population density during winter. [ABSTRACT FROM AUTHOR]

Abstract: Anyphaena accentuata and Philodromus spp. are cold adapted and winter-active spider species. Their predation activity was investigated at constant temperatures between –4 and 30°C. The lower temperature threshold for Anyphaena was –3.7°C, while that of Philodromus was –1.2°C. At 1°C the latency to capture and prey consumption was significantly shorter in Anyphaena than in Philodromus. The capture rate increased with temperature and was maximal at 15°C in Anyphaena and at 30°C in Philodromus. At 30°C, the latency to the capture was significantly shorter in Philodromus than in Anyphaena whose mortality significantly increased. [Copyright &y& Elsevier]

The first quantitative phylogenetic analysis within the spider family Philodromidae (using 65 morphological characters from 40 ingroup taxa) does not corroborate Schick's division into the tribes Thanatini (ApollophanesO. P.-Cambridge, 1898, PelloctanesSchick, 1965, ThanatusC. L. Koch, 1837; Tibellus, Simon, 1875) and Philodromini (EboKeyserling, 1884, PhilodromusWalckenaer, 1826; RhysodromusSchick, 1965). Instead, Ebois sister to all other genera in the family, and a fraction of Philodromus(the histriospecies-group=RhysodromusSchick) are contained with the Thanatini. The forgotten genus ArtanesThorell, 1870 constitutes a well-defined distal taxon in a clade that contains the majority of Philodromusspecies. Here Artanesis considered a subgenus of Philodromus, and includes the margaritatusand the poecilusspecies-groups. The western Palearctic species of the subgenus are revised. Twelve species are (re-)described, keyed and illustrated: Philodromus blanckei(Wunderlich, 1995) (first description of ♀); P. calidusLucas, 1846; P. femurostriatus, sp. nov. from the eastern Mediterranean; P. fuscomarginatus(De Geer, 1778); P. johani, sp. nov. from Crete; P. laricium(Simon, 1875), removed from synonymy with P. corticinus(C. L. Koch); P. maghrebi, sp. nov. from northern Africa (♀ only); P. margaritatus(Clerck, 1757); P. parietalisSimon, 1875; P. pentheri, sp. nov. from the Caspian region and Albania (♀ only); P. pinetorum, sp. nov. from the Mediterranean; and P. poecilus(Thorell, 1872). [ABSTRACT FROM AUTHOR]

Additional data on the biogeography and habitat of Philodromus ruficapillus Simon, 1885 are presented. New drawings of the epigyne and vulva are shown in order to complete the knowledge about the variability of these structures in a species with a wide distribution. [ABSTRACT FROM AUTHOR]

Spider field collections often consist of a high percentage of immature specimens that are not identifiable to species: in many studies these juveniles are discarded and not used in analyses. To evaluate if this practice affects the results of a community study, we sampled foliage-dwelling spiders in two habitats, reared the collected immature spiders until maturity, and identified them to species. We tested if measurements of species richness, evenness, and assemblage composition changed with the exclusion of data from immature specimens by analyzing two datasets: one including mature spiders only. the other including both mature and immature spiders (complete dataset). Nine of the total 49 spider species were collected only as juveniles, but only one of these nine species, Philodromus praelustris Keyserling 1880. was common (ge; 10% of collection). The distribution of individuals among species was more even in the complete dataset than the mature-only dataset, which could either indicate differences in composition or reflect sampling effort. However, species richness estimates were similar regardless of dataset, and there were only small changes in species composition of the samples between datasets, suggesting that there were not important compositional differences between the samples in each dataset. The inclusion of immature spiders in the data in this study yielded the same results that would occur with increased sampling effort. [ABSTRACT FROM AUTHOR]

The Philodromus pulchellus species-group is defined and diagnosed. Eleven species are included, described or redescribed, keyed and illustrated: P. afroglaucinus, sp. nov. from Algeria; P. bistigma Simon, 1870, P. glaucinus Simon, 1870, P. lamellipalpis, sp. nov. from Algeria; P. medius O. P.-Cambridge, 1872; P. pardalis, sp. nov. from northern Africa and the Iberian peninsula; P. pulchellus Lucas, 1846, P. punctigerus O. P.-Cambridge, 1908, P. ruficapillus Simon, 1885, P. simony de Mello-Leito, 1929, and P. wunderlichi, sp. nov. from the western Canary Islands. The validity of P. bistigma and P. medius is re-established (formerly in synonymy with P. pulchellus); neotype and lectotype, respectively are newly designated. The following new synonymies are proposed: P. torquatus O. P.- Cambridge, 1908 = P. pulchellus; P. salinarum Denis, 1939 = P. glaucinus; P. glaucinoides Wunderlich, 1987 = P. punctigerus; P. marionschmidti (Schmidt, 1990) = P. pulchellus. Philodromus albopictus Simon, 1875 and P. rubidus Simon, 1870 are considered nomina dubia. Determination of phylogenetic relationships within the group is difficult owing to continuous character variation, resulting in partially incongruent reconstructions using morphological and molecular data (partial mitochondrial cytochrome c oxidase I gene). Dispersal vicariance analysis provides support for a western Mediterranean origin of the group. [ABSTRACT FROM AUTHOR]