The effect of organic pollution on the abundance and distribution of aquatic oligochaetes in an urban water basin, Taiwan

Aquatic oligochaetes are abundant in polluted areas and are, therefore, commonly used as bioindicators to study organic pollution in rivers and streams. In order to develop a species-level oligochaete biotic index to reflect the River Pollution Index (RPI) in the Taichung Water Basin in Taiwan, we conducted a systematic sampling scheme to collect aquatic oligochaetes from the sediment samples of watercourses in the Taichung Water Basin, Taiwan. We evaluated the relationships between aquatic oligochaetes and the sewage pollution using statistical methods. The distribution of aquatic oligochaetes in relation to environmental variables, such as water quality and sediment characteristics of the regional urban contaminated streams was expressed by Canonical Correspondence Analysis (CCA). We identified 17 species of aquatic oligochaetes (Annelida, Clitellata) including 3 species of Tubificidae, 13 species of Naididae, and 1 species of Enchytraeidae from the watercourses of an urban region in the Taichung Water Basin in Taiwan, during the summer and winter of 2005 and 2006. A positive correlation was found between the total abundance of aquatic oligochaetes and the RPI (r = 0.58, P < 0.05). However, only population density of the most abundant tubificid, Limnodrilus hoffmeisteri, increased with increasing RPI values and a significantly negative correlation was found between the population density of the naidid, Nais communis, and RPI values. The results of CCA indicated that certain naidids, such as Aulophorus furcatus and Allonais gwaliorensis also tolerated extremely polluted environments in upper stream or stony habitats, implying that tubificids should not be the sole representation of simple biotic indices but should also include pollution-tolerant naidids. We found that the community structure of aquatic oligochaetes was influenced by short-term variations in microhabitat rather than according to seasonal factors in our study region. The results proved that aquatic oligochaetes were sensitive enough to provide a supplement for the regional urban pollution assessment applications for biotic indicators at the species-level.     

The distribution of oligochaetes in the Haringvliet-Hollands Diep (Delta region of the Netherlands) before and after the closure in 1970

Summary Samples, collected from 1959 to 1970, were used to describe the distribution and ecology of some oligochaetes in the Delta region. In the brackish and marine parts of the estuary the following species were found;Tubifex costatus, T. pseudogaster, Peloscolex benedeni andMonopylephorus rubroniveus. The other identified species were more or less euryhaline-freshwater organisms. The distribution pattern of these species was correlated with salinity and sediment. The species had a wider chlorinity tolerance than reported in the literature. The preference ofPeloscolex benedeni for clay sediments was due to low oxygen or hydrogensulfide conditions in the sediment.Limnodrilus hoffmeisteri preferred well sorted muddy sand bottoms andL. udekemianus less well sorted clay bottoms.Tubifex costatus occurred abundantly in intertidal regions. The occurrence of most of the collected oligochaetes was positively correlated with the quantity of organic matter in the sediment.An investigation carried out from 1971 to 1977 to study the development of the macrofaunal community in the fresh Haringvliet-Hollands Diep basin after closure of this basin showed a mass development of Tubificidae. This was related to the high organic matter content in the bottom and the high colonization and reproductive capacity of these worms.It appeared that several species of tubificids coexisted for several years in the same habitat at a roughly constant relative abundance. Several investigators showed a selective feeding of tubificids on certain groups of bacteria. A niche delimitation probably exists through the nature of the food used by these tubificids.A detailed account of the results presented here will be given elsewhere.     

Biodiversity of groundwater oligochaetes from a karst unit in northern Iberian Peninsula: ranking subterranean sites for conservation management

The present study suggests that the Santa Eufemia-Ereñozar karst unit in northern Iberian Peninsula is a biodiversity hotspot for groundwater oligochaetes, due to (1) the presence of a high number of stygobiotic species (corresponding to 18% of the total stygobionts known in southern Europe); (2) the comparatively high number of oligochaete species collected (corresponding to 35% of the total epigean and hypogean oligochaete species in the region); and (3) the presence of 5 species endemic to the region. A list of the oligochaete taxa found in the karst unit is presented and a conservation ranking of the cavities in the karst is proposed based on the application of four biodiversity indices (Species richness, Rarity, Vulnerability and Complementarity) to the oligochaete taxa. Vulnerability was evaluated for the first time for groundwater oligochaete taxa and it provided a useful tool to assess the protection status of oligochaetes in karstic systems. Groundwater conservation management strategies could incorporate biodiversity data from the present study.     

The nature of biodiversity in hypogean waters and how it is endangered

The specialised aquatic hypogean, i.e. stygobiotic, fauna has been recognised in some regions moderately rich. Slovenia, the broader Dinaric region, and Europe are particularly rich with about 7–8% of all Metazoa and about 40% of Crustacea species being stygobionts. The hypogean biotic diversity is in general predominantly a crustacean diversity. The high number of stygobiont Crustacea–Malacostraca species can be explained by the near absence of Insecta as well as by their high endemicity and sometimes additional specialisation, i.e. the spatial and ecological partition of the environment by the species. Although one cave system may exceptionally shelter up to 40 stygobiont species, they are distributed there into separate associations. Among more than 2000 described stygobiontic Malacostraca species, which include close to 950 Amphipoda, the species numbers within some genera are very high (e.g. Niphargus with 275 spp.). With 10 orders represented the higher taxonomic diversity of stygobiont Malacostraca matches that of fresh waters or the sea. Comparison of some faunas shows that the limiting factors for biodiversity might be the lower ecological diversity of habitats and restricted food resources underground, both brought about to a high degree by the darkness and absence of plants. Being K-strategists, stygobionts are endangered by any sudden changes in their environment. In the case of an increased food input by modest organic pollution, they can be outcompeted by energetically demanding but competitively stronger recent immigrants from surface.     

Distribution and ecology of Middle Atlantic Bight Oligochaeta

Distribution and ecology of continental shelf oligochaetes are poorly understood. From 1975 through 1977 baseline ecological studies were conducted in the Middle Atlantic Bight prior to oil exploration activities. A total of 38 species in three oligochaete families were collected. Of this total one family and 29 of the species were undescribed at the time of collection. The most abundant and widely distributed species were Limnodriloides monothecus, L. medioporus, Bathydrilus longus, Phallodrilus biprostatus, and Tubificoides diazi. No single species dominated the oligochaete fauna. All the oligochaete species responded to changes in physical parameters over the shelf similarly to the total macrofauna. Seasonal trends within the oligochaetes were not found. Species that preferred coarser sand sediments were all small interstitial forms. The finer sand and eurytopic species were all burrowers. Zoogeographically the Middle Atlantic Bight oligochaetes were predominantly temperate (16 species) with some southern species (8). Seven species were broadly distributed from the Caribbean to Massachusetts or around the world. Seven species were only known from the area studied.Contribution No. 1397 of the Virginia Institute of Marine Science.     

Species composition,distribution, and abundance of oligochaetes in the Kanawha River,West Virginia

Species composition, distribution, and abundance of oligochaetes in the Kanawha River, West Virginia, were investigated. Eighteen taxa of oligochaetes including 7 tubificids, 8 naidids, and representatives of the families Enchytraeidae, Lumbriculidae, and Aelosomatidae were collected. While most of the species collected are cosmopolitan, Pristina synclites and Stephensoniana sp. previously reported only from Asia and Africa were collected in the Kanawha. Other peculiarities of the Kanawha oligochaete fauna included the common occurrence of specimens of Limnodrilus cervix and Limnodrilus hoffmeisteri with atypical penis sheaths. The dominant and most widely distributed of the Kanawha River oligochaetes were immature tubificids lacking capilliform chaetae, Branchiura sowerbyi, Dero cf. obtusa, Limnodrilus hoffmeisteri, and Aulodrilus pigueti.Oligochaete densities ranged from 401–2403 individuals/m² in areas presumed to be nonpolluted, to 1–671 individual/m² at sites with suspected toxicity. The highest density of oligochaetes (2835 individuals/m²) occurred at a site of organic enrichment.     

Oligochaete drift in a large river (French Upper Rhône): the effect of life cycle and discharge

In the framework of a macroinvertebrate drift study, the oligochaete community was analysed over one year (1989) at six sites located in a partly impounded reach of the Upper Rhône River (France). Drift composition between sites was somewhat affected by the environmental conditions. These differences were caused by significant deviations in drift rates (P < 0.05) of endobenthic oligochaetes and non-swimming Naididae. There was no significant difference (P > 0.05) among the swimming Naididae. As the latter strongly dominated, global spatial differences in the drift were weak. Temporal changes constituted the main source of variability in drifting oligochaetes. Life cycles and reproductive ecology of four major swimming Naidid species, Nais elinguis, Stylaria lacustris, Nais christinae, and Nais barbata, explained most of the drift structure over the year. Hydrological fluctuations also affected drift structure, both for swimming and non-swimming species. Taxon richness and relative abundance in the drift were very similar to those obtained in the river benthos from a reference reach previously studied. There was a lack of stygobiont and stygophilous species that are frequent in area of groundwater seepage. Biological and environmental implications of oligochaete drift are stressed. The drift phenomenon is appropriated to provide ecological informations on oligochaete communities in fast flowing and deep, large rivers.     

A new species of Typhlatya (Crustacea: Decapoda: Atyidae) from anchialine caves on the French Mediterranean coast

A new species of the thermophylic Tethyan relict prawn Typhlatya is described from two anchialine caves near Perpignan (southern France). The new species is closely related to a congener known only from a freshwater cave at Castellón (eastern Spain), about 400 km to the south-west, differing apparently only in the size and shape of the rostrum and the armature of the dactylus of the fifth pereiopod. Based on palaeogeographical evidence and assuming a sister-group relationship between both species, we suggest that their common ancestor could not be older than early Pliocene in age, and that it was already a stygobiont taxon adapted to live in shallow-water marine crevicular habitats. This ancestor would have vanished from the western Mediterranean after the cooling associated with the onset of northern Hemisphere glaciation, about 3 Mya, as documented for other Mediterranean marine taxa. Indeed, the genus is completely stygobiont and does not occur in fluvial environments. The Pyrenees represent a watershed boundary that eliminates the possibility of the derivation of one species from the other by active dispersal after establishment in continental waters.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 387–414.     

A highly modified stygobiont diving beetle of the genus Copelatus (Coleoptera,Dytiscidae): taxonomy and cladistic analysis based on mitochondrial DNA sequences

Abstract. We report the discovery of a new species of Copelatus that is morphologically highly modified for life in groundwater. Copelatus abditus sp.n. was collected from a 4 m deep bore in central Australia. It was placed in Copelatinae based on morphological evidence. This is the first known stygobiont diving beetle that does not belong to the subfamily Hydroporinae. Sequences from the mitochondrial DNA cytochrome oxidase 1, 16S rRNA, tRNAL and NADH dehydrogenase subunit 1 genes suggest that the species belongs to the subgenus Papuadytes, the morphological apomorphies of which are reduced in the new groundwater species. Copelatus (Papuadytes) abditus very much resembles other world stygobiont dytiscids, which all belong to the subfamily Hydroporinae. We suggest that this striking convergence is due to similar selective pressures imposed on all groundwater beetles. We suggest a scenario in which species from temporary habitats evade drought by entering the hyporheic zone, and may be driven to a subterranean existence by desertification of their ancestral habitats, as can be observed in Australia.     

The Parvidrilidae – a diversified groundwater family: description of six new species from southern Europe,and clues for its phylogenetic position within Clitellata (Annelida)

The Parvidrilidae Erséus, 1999 constitute the most recently described family of oligochaete microdriles. Prior to this study, Parvidrilus strayeri Erséus, 1999, and Parvidrilus spelaeus Martínez‐Ansemil, Sambugar & Giani, 2002, found in groundwaters of the USA (Alabama) and Europe (Slovenia and Italy), respectively, were the only two species in this family. In this paper, six new species – Parvidrilus camachoi , Parvidrilus gianii , Parvidrilus jugeti , Parvidrilus meyssonnieri , Parvidrilus stochi , and Parvidrilus tomasini – and Parvidrilus gineti (Juget, 1959) comb. nov. are added to the family. With all species being stygobiont, the Parvidrilidae is unique in being the only family of oligochaetes worldwide comprising taxa that are restricted to groundwater habitats. Parvidrilids are exceedingly small worms whose principal morphological characteristics are the presence of hair setae in ventral bundles, the markedly posterior position of setae within the segments, the presence of mid‐dorsal glandular pouches in mesosomial segments, the lateral development of the clitellum, the presence of a single male pore in segment XII, and the presence (or absence) of a single spermatheca. The phylogenetic relationships of the Parvidrilidae within the Clitellata were investigated using the nuclear 18S rRNA gene, and the most representative and taxonomically balanced data set of clitellate families available to date. The data were analysed by parsimony, maximum likelihood, and Bayesian inference. Irrespective of the method used, Parvidrilidae were placed far from Capilloventridae, one family once suggested to be closely related to parvidrilids. Although closer to Enchytraeidae than Phreodrilidae, two other suggested putative sister families, the exact position of Parvidrilidae within Clitellata still remained uncertain in the absence of branch support. The examination of reproductive structures, together with the similarity of other important anatomical traits of the new species herein described, reinforced the idea that phreodrilids were the best candidate to be the sister group to parvidrilids on morphological grounds. A fragment of the mitochondrial cytochrome oxidase I gene, used as a barcode, also genetically characterized a few Parvidrilus species. The observation that two species diverge from each other by high genetic distances, even though their type localities are more or less only 100 km apart, is interpreted in the context of low dispersal abilities of inhabitants of the subterranean aquatic ecosystem, and habitat heterogeneity. The Parvidrilidae appear to be a diversified, Holarctic, and probably widely distributed family in groundwater, but very often overlooked because of the small size and external similarity with the polychaete family Aeolosomatidae of its members. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 530–558.     

A taxonomic and faunistic survey of the marine Tubificidae and Enchytraeidae (Oligochaeta) of Italy. Introduction and preliminary results

On the basis of material collected along the Ligurian, Tyrrhenian, Ionic and Adriatic coasts of Italy, a tentative list of 26 species (18 tubificids, 8 enchytraeids) of marine Oligochaeta is presented. Most of the species are new to science, and it can therefore be concluded that there is a very high diversity of oligochaetes in the Mediterranean Sea.     

DNA barcode libraries provide insight into continental patterns of avian diversification

Background

The causes for the higher biodiversity in the Neotropics as compared to the Nearctic and the factors promoting species diversification in each region have been much debated. The refuge hypothesis posits that high tropical diversity reflects high speciation rates during the Pleistocene, but this conclusion has been challenged. The present study investigates this matter by examining continental patterns of avian diversification through the analysis of large-scale DNA barcode libraries.

Methodology and Principal Findings

Standardized COI datasets from the avifaunas of Argentina, the Nearctic, and the Palearctic were analyzed. Average genetic distances between closest congeners and sister species were higher in Argentina than in North America reflecting a much higher percentage of recently diverged species in the latter region. In the Palearctic genetic distances between closely related species appeared to be more similar to those of the southern Neotropics. Average intraspecific variation was similar in Argentina and North America, while the Palearctic fauna had a higher value due to a higher percentage of variable species. Geographic patterning of intraspecific structure was more complex in the southern Neotropics than in the Nearctic, while the Palearctic showed an intermediate level of complexity.

Conclusions and Significance

DNA barcodes can reveal continental patterns of diversification. Our analysis suggests that avian species are older in Argentina than in the Nearctic, supporting the idea that the greater diversity of the Neotropical avifauna is not caused by higher recent speciation rates. Species in the Palearctic also appear to be older than those in the Nearctic. These results, combined with the patterns of geographic structuring found in each region, suggest a major impact of Pleistocene glaciations in the Nearctic, a lesser effect in the Palearctic and a mild effect in the southern Neotropics.     

Superficial and hyporheic oligochaete communities as indicators of pollution and water exchange in the River Moselle, France

Benthic oligochaetes were sampled on three occasions (June, August and October 1992) in the upper (0–10 cm) and hyporheic (35–45 cm depths) sediments at five sites of the River Moselle, from upstream of the town of Epinal to Velle-sur-Moselle. The first site (upstream from Epinal) is considered as unpolluted and the four remaining sites are polluted by industrial effluents. The most polluted stations were generally dominated by the pollution tolerant taxon Limnodrilus. Numbers of individuals of this taxon decreased at the less polluted last site in recovery zone, and were also scarce in the first unpolluted site. It is noteworthy that these tendencies were observed in both superficial and hyporheic substrates and to the greatest degree in hyporheic ones. At the unpolluted site, the hyporheic habitat is dominated by the groundwater species Propappus volki, Pristina spp., Pristinella spp. At the less polluted site (last site), the deep sediments are dominated by groundwater species and the Tubificidae without hair setae decrease from June to October. As a result of water exchange between superficial and subterranean waters, superficial substrates of the first and the last stations tend to be colonised by a high proportion of hyporheic species that suggests that flow is primarily from subterranean to superficial waters. The contrary is the case at other polluted stations which are characterised by the invasion of hyporheic substrates by the pollution tolerant superficial taxa Limnodrilus. This suggests that water flows from the river to the deeper groundwater. These two stations are located near drinking water plants which utilise groundwater, thus increasing the vulnerability of groundwater to surface contaminants.     

Two new species of Pseudoniphargus (Amphipoda), in Bermudian groundwaters*

The genus Pseudoniphargus comprises a number of stygobiont Amphipoda, distributed on both sides of the Atlantic. In Bermuda, groundwaters are inhabited by two species. P. grandimanus and P. carpalis, both new. They occur throughout the entire salinity spectrum, from infrahalne to euhaline, but are most abundant in oligohaline waters. Large adults, but no ovigerous specimens. arc found at greater distance from the sea coast than juveniles, which may indicate that Pseudoniphargus needs anchihaline/marine waters for its reproduction and that juveniles migrate inland to grow up.     

Models of stream habitat characteristics associated with tubificid populations in an intermountain watershed

Habitat assessments were conducted in an intermountain watershed at three spatial extents to explore ways to predict the presence of tubificid oligochaetes likely to support the parasite Myxobolus cerebralis, cause of salmonid whirling disease. Stream reaches with six different reach slope characteristics were selected using GIS. The aquatic habitat in 60 reaches selected at random was measured and classified into distinct habitat units. Within the habitat units, areas of microhabitat with depositional fine sediments were chosen, measured, and core samples were removed to characterize the sediments and benthic oligochaetes. Two tubificids, Tubifex spp. and Limnodrilus hoffmeisteri, were abundant and co-occurred in silt-clay and fine sand sediments in these habitats. Models were posed and tested to predict the presence and relative abundance of tubificids using habitat characteristics from the three spatial extents: reach, habitat unit, and microhabitat. At the reach extent, tubificids were associated with low-reach slope and with slow water habitats. Within habitat units, tubificids were associated with higher percentages of fine sediments and higher stream width:depth ratios. In microhabitat cores, the presence of silt-clay sediments was positively associated with higher average stream width:depth ratios. Since ecological relationships are often scale dependent and stream systems have a natural hierarchy, predictive habitat models such as these that use measures from several scales may help researchers and managers more efficiently identify and quantify aquatic communities at highest risk of infection by the M. cerebralis parasite. Handling editor: J. Trexler     

Distribution,Abundance and Life Cycles of Oligochaeta from the Marine Hygropsammal with Special Reference to the Phallodrilinae (Tubificidae)

Tubificid and enchytraeid oligochaetes are common members of the interstitial fauna of sandy beaches. Their abundance dynamics, life cycles and distribution patterns were investigated at two beaches on the western Baltic Sea and at a North Sea beach (Isle of Sylt). Populations of 115,000 ind./m² were found at protected Baltic beaches. Reproduction of the tubificids, Phallodrilus monospermathecus and Spiridion insigne takes place within a well defined breeding period once a year. The market horizontal and vertical distribution patterns of various tubificid and enchytraeid species are elucidated by preference reactions to several physiographic parameters, such as oxygen and water content of the sediment, and salinity.     

Epizootiology of Eustrongylides ignotus in Florida: distribution,density, and natural infections in intermediate hosts

A total of 63,451 fish, representing 39 species, was collected from 176 foraging sites used by ciconiiform wading birds in peninsular Florida (USA) and examined for larvae of Eustrongylides ignotus. Infected fish were identified from 30 (17%) of the sites, all of which had been altered by human disturbance such as removal of sediment to construct ditches and dikes, improve water flow, or increase storage capacity and had a history of receiving anthropogenic nutrients such as sewage effluent, urban runoff, or agricultural runoff. The mosquitofish (Gambusia holbrooki) and several species of sunfish (Centrarchidae) were the most important intermediate hosts. Infected fish were not collected at any of the unaltered sites. A total of 10,508 oligochaetes (representing 36 species) was identified from 22 sites that had fish infected with E. ignotus and 36 sites where no infected fish were collected. None of the oligochaetes was infected with larvae of E. ignotus. Immature tubificids without hair setae (probably Limnodrilus sp.), Dero digitata, and L. hoffmeisteri were the most abundant oligochaetes at sites where infected fish occurred, making up 78% of the total collected. Compared to unaltered sites, altered sites were characterized by higher mean densities of fish and oligochaetes; surface waters with decreased dissolved oxygen and increased total nitrogen, total phosphorus, and chlorophyll-a; sediments with higher soil oxygen demand and total phosphorus; larger grain sizes; and higher percentage emergent vegetation and grasses.     

Do different dispersal capacities influence the biogeography of the western Palearctic dragonflies (Odonata)?

The biogeography of the western Palearctic has been intensively studied for more than a century. Recent advances in genetics have allowed the testing of old theories based on distribution patterns, although these analyses are obviously restricted to a reduced number of specific genetic data sets. On the other hand, an increased knowledge on the distributions of species and advances in computer capacities have allowed more detailed biogeographical analyses based on species presence/absence. In the present study, we selected the Odonata as the study group. For all 162 species native to the western Palearctic, we compiled their respective presence or absence in 97 predefined biogeographical regions. Using cluster analyses and principal component analyses, both based on Jaccard similarity coefficients, we analysed the differentiation among these regions and species. In subsequent analyses, the data set was reduced to the Zygoptera, Anisoptera, and the western Palearctic endemics. All analyses consistently showed different faunal regions and faunal elements. In particular, the (1) western and (2) eastern Mediterranean; (3) Central and (4) Northern Europe; and (5) the British Isles were invariably found in all cases. Although the two major Mediterranean regions were characterized by several endemic faunal elements, Northern Europe and the British Isles lacked such elements, but were characterized by faunal compositions strongly deviating from the rest of the western Palearctic region. Moderate differences between Zygoptera and Anisoptera existed, with the latter more clearly redrawing the Mediterranean refuge areas, whereas the former reflected to a greater extent the postglacial expansion patterns from these regions. In general, our findings underline the old biogeographical theories, but refine especially our understanding of the Atlanto‐ and Ponto‐Mediterranean region. Central Europe, comprising the area with the highest species numbers of our whole study region, unravels as a crossroad of postglacial immigrations, but might also represent a region of in situ glacial survival. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 177–195.     

Systematics of the Saharo‐Arabian clade of the Palearctic naked‐toed geckos with the description of a new species of Tropiocolotes endemic to Oman

Geckos are one of the most species‐rich, abundant, and widely distributed of all Squamata lineages and present several characteristics that have made them favorite model organisms for biogeographical, ecological, physiological, and evolutionary studies. One of the key aspects of any comparative study is to have a robust, comprehensive phylogeny, and an updated taxonomy. Recently, the Infraorder Gekkota has been the subject of several phylogenetic analyses and taxonomic revisions at different levels. Despite all these phylogenetic and taxonomic advances, there are still some groups whose systematics and taxonomy remain highly problematic. Maybe one of the most poorly resolved groups in spite of decades of intensive research by many herpetologists are the so‐called Palearctic naked‐toed geckos of the family Gekkonidae. This group of nocturnal geckos distributed from Mauritania across North Africa, Arabia, southwestern and central Asia to northern India, western China and southern Mongolia is characterized by the synapomorphy of lack of adhesive subdigital pads. Within the Palearctic naked‐toed geckos, the Saharo‐Arabian clade comprised by the genera Pseudoceramodactylus, Stenodactylus, and Tropiocolotes is the clade with the largest distribution range. At the same time, it is one of the problematic groups, presenting poorly supported phylogenetic relationships, with the genus Tropiocolotes being recovered non‐monophyletic in all analyses despite its morphological uniformity. To reassess the phylogeny of the Palearctic naked‐toed geckos with a special interest in the systematics of Tropiocolotes, we assembled a dataset comprising 298 gecko specimens from 283 different species (including all Tropiocolotes species but one) belonging to 122 of a total of 124 described gecko genera. This dataset included the nuclear c‐mos, ACM4, RAG1, RAG2, and PDC and the mitochondrial ND2 gene. To further investigate the relationships within Tropiocolotes and to revise the systematics of the south Arabian endemic species Tropiocolotes scorteccii, we used an integrative approach including information from the nuclear MC1R and c‐mos, the mitochondrial 12S, 16S, cytb genes, and morphological data from nine of the 10 described Tropiocolotes species. The phylogenetic analyses of the Gekkota dataset recovered a similar topology for the Palearctic naked‐toed geckos to previous studies, but in this case, Tropiocolotes was recovered monophyletic in all analyses, with high support in two of them. The results of the analyses of three datasets specifically assembled to test the effect of both gene sampling and taxon sampling in the monophyly of Tropiocolotes, and the internal relationships of the Palearctic naked‐toed geckos clearly showed that both the number and kind of characters (nuclear or mitochondrial data) and the number of taxa played a fundamental role in recovering the correct phylogenetic relationships. The phylogenetic analyses within Tropiocolotes suggested the existence of high levels of undescribed diversity in the south Arabian T. scorteccii, including a new genetically and morphologically distinct species endemic to Oman (Tropiocolotes confusus sp. nov. ). Our study using a large dataset, including several loci and a dense taxon sampling within Gekkota and especially within Tropiocolotes, has proved a valuable strategy to address the monophyly of Tropiocolotes and the relationships within the Saharo‐Arabian Palearctic naked‐toed geckos. The integrative systematic approach including several samples of south Arabian T. scorteccii based on many years of fieldwork has, once more, uncovered a new species endemic to this region. This highlights the importance of this area of Arabia as a reservoir of reptile endemicity and biodiversity, which is likely linked to the high degree of habitat heterogeneity and the effect of the monsoons. Obviously, based on this and previously published evidence, south Arabia represents an area with still high levels of undiscovered diversity.     

Zoogeography of the southern African ascidian fauna

Aim To describe the biogeography of the ascidian fauna of southern Africa, to compare the results obtained with those reported for other fauna and flora of the same region, and to speculate about the origin of ascidians in the region. Location Southern Africa extending over 4000 km from Mossâmedes (15° S–12° E) to Inhaca Island (26°30′ S–33° E), including Vema Seamount (31°40′ S–8 °20′ E), Amsterdam‐Saint Paul Islands (38° S–77°30′ E) and the Tristan‐Gough Islands (38° S–12°20′ W). Methods We constructed a presence/absence matrix of 168 species for 26 biogeographical divisions, 21 classical biogeographical regions described by Briggs (Marine zoogeography, McGraw‐Hill, New York, 1974) and five provinces within the southern African region. We considered the following limits and divisions into provinces for the southern African region: Namibia, Namaqua, Agulhas and Natal as proposed by Branch et al. (Two oceans. A guide to the marine life of southern Africa, David Philip Publishers, 1994), and the West Wind Drift Islands province (WWD) according to Briggs (Global biogeography, Elsevier Health Sciences, Amsterdam, 1995). To examine the biogeographical structure, species and divisions were classified using cluster analysis (based on UPGMA as the aggregation algorithm) with the Bray–Curtis index of similarity. This classification was combined with MDS ordination. Main conclusions Four main groups were obtained from the analysis of affinities among species: (1) species present in the WWD, separated by a high percentage of endemisms and a low number of species with a southern African distribution. Moreover, in the light of the species distribution and the results of further analysis, which revealed that they are completely separated and not at all related to the southern African region, it appears that there are no close relationships among the different islands and seamounts of the West Wind Drift Island province. This province was therefore removed from the remaining analyses; (2) species with a wide distribution; (3) species of colder waters present in Namaqua and Agulhas provinces, a transitional temperate area in which gradual mixing and replacement of species negate previous hypotheses on the existence of a marked distributional break at Cape of Good Hope; (4) species of warmer waters related to Natal province. The classification into biogeographical components was dominated by the endemic (47%), Indo‐Pacific (25%) and cosmopolitan (13%) components. The analysis of affinities among biogeographical areas separated Namibia from the rest of the southern African provinces and showed that it was related to some extent to the Antarctic region because of the cold‐temperate character of the province and the low sampling effort; Namaqua, Agulhas and Natal were grouped together and found to be closely related to the Indo‐West Pacific region. In general, our results were consistent with those obtained for other southern African marine invertebrates. The frequency distribution of solitary/colonial strategies among provinces confirmed the domination of colonial organisms in tropical regions and solitary organisms in colder regions. Finally, we speculate that the southern African ascidian fauna mainly comprises Indo‐Pacific, Antarctic and eastern Atlantic ascidians.