Multiple Pleistocene refugia and post‐glacial colonization in the European chub (Squalius cephalus) revealed by combined use of nuclear and mitochondrial markers

Aim To analyse patterns of nuclear and mitochondrial genetic variation in the European chub, Squalius cephalus (Linnaeus, 1758), in order to understand the evolutionary history of this species and to test biogeographical hypotheses for the existence of co‐distributed European freshwater fish species. Location Rivers in Europe (Finland, Poland, Czech Republic, France, Bulgaria, Spain, Italy). Methods We genotyped 12 polymorphic microsatellite markers derived from 310 individuals collected from across the distribution of S. cephalus in Europe (including a total of 15 populations) and sequenced mitochondrial DNA (mtDNA) from a subset of 75 individuals. Sequences of mtDNA cytochrome b were analysed using both phylogenetic (median‐joining networks) and population genetic methods (tests for demographic history, mismatch distributions, Bayesian coalescent analysis). Geographical structure in microsatellite loci was examined using a distance method (F_ST), factorial correspondence analysis (FCA) and a Bayesian clustering method (structure ). Results The mtDNA network showed a clear split into four different haplogroup lineages: Western (separated into Atlantic and Danubian sublineages), Eastern, Aegean (occurring in two distinct sublineages in the Balkans and in Spain) and Adriatic. Our results indicate recent population expansion in the Eastern and Western Atlantic lineages and the admixture of two previously separate sublineages (Atlantic and Danubian) in the Western lineage. Bayesian structure analysis as well as FCA results roughly corresponded to the mtDNA‐based structure, separating the sampled individuals into almost non‐overlapping groups. Main conclusions Our results support hypotheses suggesting origins of extant lineages of freshwater fishes in multiple refugia and the subsequent post‐glacial colonization of Europe via different routes. We confirmed the previously proposed two‐step expansion scenario from the Danube refuge, the existence of a secondary (Atlantic) refuge during the last glaciation (probably in the Rhone River) and population expansion of this lineage. Conspicuous divergences among Mediterranean populations reflect their different origin, as well as their low contribution to the recent genetic pool of chub in central Europe.     

Multigene molecular systematics confirm species status of morphologically convergent Pagurus hermit crabs

Introduction

In spite of contemporary morphological taxonomy appraisals, apparent high morphological similarity raises uncertainty about the species status of certain Pagurus hermit crabs. This is exemplified between two European species, Pagurus excavatus (Herbst, 1791) and Pagurus alatus (Fabricius 1775), whose species status is still difficult to resolve using morphological criteria alone.

Methodology/Principal Findings

To address such ambiguities, we used combinations of Maximum Likelihood (ML) and Bayesian Inference (BI) methods to delineate species boundaries of P. alatus and P. excavatus and formulate an intermediate Pagurus phylogenetic hypothesis, based upon single and concatenated mitochondrial (cytochrome oxidase I [COI]) and nuclear (16S and 28s ribosomal RNA) gene partitions. The molecular data supported the species status of P. excavatus and P. alatus and also clearly resolved two divergent clades within hermit crabs from the Northeast Atlantic Ocean and the Mediterranean Sea.

Conclusions/Significance

Despite the abundance and prominent ecological role of hermit crabs, Pagurus, in North East Atlantic Ocean and Mediterranean Sea ecosystems, many important aspects of their taxonomy, biology, systematics and evolution remain poorly explored. The topologies presented here should be regarded as hypotheses that can be incorporated into the robust and integrated understanding of the systematic relationships within and between species of the genus Pagurus inhabiting the Northeast Atlantic Ocean and the Mediterranean Sea.     

Phylogenetics and biogeography of the Balkan ‘sand gobies’ (Teleostei: Gobiidae): vulnerable species in need of taxonomic revision

Within the Atlantic–Mediterranean region, the ‘sand gobies’ are abundant and widespread, and play an important role in marine, brackish, and freshwater ecosystems. They include the smallest European freshwater fish, Economidichthys trichonis, which is threatened by habitat loss and pollution, as are several other sand gobies. Key to good conservation management is an accurate account of the number of evolutionary significant units. Nevertheless, many taxonomic and evolutionary questions remain unresolved within the clade, and molecular studies are lacking, especially in the Balkans. Using partial 12S and 16S mitochondrial ribosomal DNA sequences of 96 specimens of at least eight nominal species (both freshwater and marine populations), we assess species relationships and compare molecular and morphological data. The results obtained do not support the monophyly of Economidichthys, suggesting the perianal organ to be a shared adaptation to hole‐brooding rather than a synapomorphy, and urge for a taxonomic revision of Knipowitschia. The recently described Knipowitschia montenegrina seems to belong to a separate South‐East Adriatic lineage. Knipowitschia milleri, an alleged endemic of the Acheron River, and Knipowitschia cf. panizzae, are shown to be very closely related to other western Greek Knipowitschia populations, and appear conspecific. A distinct Macedonian–Thessalian lineage is formed by Knipowitschia thessala, whereas Knipowitschia caucasica appears as an eastern lineage, with populations in Thrace and the Aegean. The present study combines the phylogeny of a goby radiation with insights on the historical biogeography of the eastern Mediterranean, and identifies evolutionary units meriting conservation attention. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 73–91.     

Morphological Differentiation Between Geographically Separated Populations of <Emphasis Type="Italic">Neomysis integer</Emphasis> and <Emphasis Type="Italic">Mesopodopsis slabberi</Emphasis> (Crustacea,Mysida)

Morphological variation was examined in Neomysis integer and Mesopodopsis slabberi, two abundant, low dispersal mysid species (Crustacea, Mysida) along the European coasts. Both species dominate the hyperbenthic communities in the northeast Atlantic, and M. slabberi is also widely distributed in the Mediterranean and Black Sea. Three populations of these species were sampled throughout their distribution range; samples of N. integer were collected in the northeast Atlantic Eems-Dollard, Gironde and Guadalquivir estuaries; in the case of M. slabberi, mysids were sampled in two northeast Atlantic estuaries (Eems-Dollard and Guadalquivir) and one Mediterranean site (Ebro Delta). A total of 12 morphometric and 2 meristic characters were measured from 30–64 mysids per sample. Multivariate analysis showed clear morphometric differences between populations of both species. The morphological differentiation within M. slabberi was highly concordant with the available genetic data from mitochondrial loci, pointing to a large divergence between the Atlantic and Mediterranean populations. However, due to some overlap of individuals between the different populations, the present morphometric analysis does not suffice to assign the different populations to a separate (sub)species status. In the case of N. integer, the morphometric patterns showed a divergence of the Gironde population. Differentiation of populations within this mysid, as in M. slabberi, were mainly related to eye and telson morphology. Potential interactions of the mysid morphology and environmental conditions are discussed.     

Identification and characterization of microsatellite loci in the spiny spider crab <Emphasis Type="Italic">Maja brachydactyla</Emphasis>

The spiny spider crab Maja brachydactyla in an important fishery resource throughout its distribution range (Northeast Atlantic). Here we describe the isolation of nine microsatellite loci for this species. These new markers were tested in 20 crabs from NW Spain. The number of alleles ranged from 3 to 20 and expected heterozigosity from 0.57 to 0.95. All loci followed Hardy–Weinberg expectations except mMb33 and mMb307, which could be affected by null alleles. Some of these loci seem to be better fitted by a multi-step substitution process.     

Range‐wide population structure of European sea bass Dicentrarchus labrax

The euryhaline European sea bass Dicentrarchus labrax L., inhabiting the coasts of the eastern Atlantic Ocean and Mediterranean Sea, has had many opportunities for differentiation throughout its large natural range. However, evidence for this has been incompletely documented geographically and with an insufficient number of markers. Therefore, its full range was sampled at 22 sites and individuals were genotyped with a suite of mapped markers, including 14 microsatellite loci (N = 536) and 46 neutral or gene‐linked single nucleotide polymorphisms (SNPs; N = 644). We confirm that the Atlantic and Mediterranean basins harbour two distinct lineages. Within the Atlantic Ocean no pattern was obvious based on the microsatellite and SNP genotypes, except for a subtle difference between South‐eastern and North‐eastern Atlantic sea bass attributed to limited introgression of alleles of Mediterranean origin. SNP genotypes of the Mediterranean lineage differentiated into three groups, probably under the influence of geographical isolation. The Western Mediterranean group showed genetic homogeneity without evidence for outlier loci. The Adriatic group appeared as a distinct unit. The Eastern Mediterranean group showed a longitudinal gradient of genotypes and most interestingly an outlier locus linked to the somatolactin gene. Overall, the spatial pattern fits those observed with other taxa of between‐basin segregation and within‐basin connectivity, which concurs well with the swimming capabilities of European sea bass. Evidence from a few outlier loci in this and other studies encourages further exploration of its regional connectivity and adaptive evolution.     

Phylogeography of the large Myotis bats (Chiroptera: Vespertilionidae) in Europe,Asia Minor,and Transcaucasia

The large Myotis complex in continental Europe, Asia Minor, and Transcaucasia comprises two sibling bat species, the greater mouse‐eared bat, Myotis myotis, and the lesser mouse‐eared bat, Myotis blythii, also referred to as Myotis oxygnathus. Here, we investigate the phylogeography of these bats using two mitochondrial markers: the second hypervariable domain of the control region (HVII) and a fragment of the cytochrome b gene (cyt b). The HVII haplotypes formed six distinct haplogroups associated with different geographical regions. Most of the European HVII haplotypes were exclusive to M. myotis, whereas the majority of HVII haplotypes found in Asia Minor were exclusive to M. blythii/M. oxygnathus. The phylogenetic reconstruction based on the concatenated cyt b and HVII fragments recovered two major lineages. The first lineage comprised samples from Europe (western lineage), and the second lineage included samples from Asia Minor, Transcaucasia, Crimea, Western Ukraine, Thrace, the Balkans, and Eastern Europe (eastern lineage). The mitochondrial lineage of M. blythii, reported from Kyrgyzstan, was not present in Asia Minor and Transcaucasia. Therefore, we consider the possibility that the M. blythii/M. oxygnathus found in Europe, Asia Minor, and Transcaucasia are not recent descendants of the Central Asian M. blythii. Instead, we suggest that M. blythii/M. oxygnathus and M. myotis diverged through allopatric speciation in Asia Minor and Europe, and that they are represented by the eastern and western mitochondrial lineages. We also examine an alternative hypothesis: that the large Myotis complex consists of more than two species that diverged independently in Asia Minor and Europe through ecological speciation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

The curious case of Hermodice carunculata (Annelida: Amphinomidae): evidence for genetic homogeneity throughout the Atlantic Ocean and adjacent basins

Over the last few decades, advances in molecular techniques have led to the detection of strong geographic population structure and cryptic speciation in many benthic marine taxa, even those with long‐lived pelagic larval stages. Polychaete annelids, in particular, generally show a high degree of population divergence, especially in mitochondrial genes. Rarely have molecular studies confirmed the presence of ‘cosmopolitan’ species. The amphinomid polychaete Hermodice carunculata was long considered the sole species within its genus, with a reported distribution throughout the Atlantic and adjacent basins. However, recent studies have indicated morphological differences, primarily in the number of branchial filaments, between the East and West Atlantic populations; these differences were invoked to re‐instate Hermodice nigrolineata, formerly considered a junior synonym of H. carunculata. We utilized sequence data from two mitochondrial (cytochrome c oxidase subunit I, 16S rDNA) markers and one nuclear (internal transcribed spacer) marker to examine the genetic diversity of Hermodice throughout its distribution range in the Atlantic Ocean, including the Mediterranean Sea, the Caribbean Sea, the Gulf of Mexico and the Gulf of Guinea. Our analyses revealed generally low genetic divergences among collecting localities and between the East and West Atlantic, although phylogenetic trees based on mitochondrial data indicate the presence of a private lineage in the Mediterranean Sea. A re‐evaluation of the number of branchial filaments confirmed differences between East and West Atlantic populations; however, the differences were not diagnostic and did not reflect the observed genetic population structure. Rather, we suspect that the number of branchial filaments is a function of oxygen saturation in the environment. Our results do not support the distinction between H. carunculata in the West Atlantic and H. nigrolineata in the East Atlantic. Instead, they re‐affirm the older notion that H. carunculata is a cohesive species with a broad distribution across the Atlantic Ocean.     

Phylogenetic analysis of the north‐east Atlantic and Mediterranean species of the genus Stenosoma (Isopoda,Valvifera, Idoteidae)

Xavier, R., Santos, A. M., Harris, D. J., Sezgin, M., Machado, M., Branco, M. (2012). Phylogenetic analysis of the north‐east Atlantic and Mediterranean species of the genus Stenosoma (Isopoda, Valvifera, Idoteidae). —Zoologica Scripta, 41, 386–399. The marine isopod genus Stenosoma is widespread in the northern hemisphere. However, 12 of its 14 known species are found within the Mediterranean basin and adjacent regions of the north‐east Atlantic and the Black Sea. Such a high level of diversity confined to a limited region of a much larger circumglobal distribution suggests that the Mediterranean region may have played a crucial role in the evolutionary history of this genus. In the present work, the phylogeny of the genus Stenosoma was investigated on the basis of DNA sequencing data from one nuclear (28SrRNA) and two mitochondrial (COI, ND4) gene fragments obtained for nine of 12 Atlantic–Mediterranean species. Divergence time estimates point to a Tethyan origin of Stenosoma and suggest that the speciation events from which stem most of the extant species took place well before the Messinian Salinity Crisis. Stenosoma spinosum and Stenosoma appendiculatum are the only exceptions, as they apparently arose within the Mediterranean during the Pleistocene. Phylogenetic reconstruction agrees with current taxonomic status of most species. However, Stenosoma capito clustered in two distinct and well‐supported clades, one composed of eastern Mediterranean and Black Sea specimens and the other by western Mediterranean and Atlantic ones. Such polyphyly suggests the existence of a previously unrecognized species, Stenosoma sp., which so far has been confounded with S. capito.     

Phylogeography of the marine isopod Stenosoma nadejda (Rezig, 1989) in North African Atlantic and western Mediterranean coasts reveals complex differentiation patterns and a new species

The transition zone between the Mediterranean and Atlantic basins has been extensively addressed in phylogeographical studies of marine species. However, biases exist towards the analysis of highly dispersive species, and there is a higher sampling effort in European coasts compared to North Africa. This may be hindering a detailed understanding of the historical and contemporary processes that shaped patterns of population genetic structure in the region. In the present study, we investigated the phylogeographical and phylogenetic patterns of mitochondrial cytochrome c oxidase subunit I sequences from a species with direct development and low dispersal abilities, Stenosoma nadejda (Rezig, 1989). The study area included 13 localities along the Atlantic and Mediterranean North African coasts, as well as the Alboran Sea. A new Stenosoma species, from the coasts of Algeria and Alboran Island, was discovered. For S. nadejda, phylogeographical analyses revealed three distinct clades: one in the Iberian Atlantic plus the Alboran Sea, one in the western Mediterranean, and another in the Atlantic coast of Africa. Haplotypes from the Alboran Island were more related to those from the western Mediterranean coast (east of the Almeria–Oran Front). Given the strong differentiation, it is probable that this species survived in multiple glacial refugia during the Pleistocenic glaciations. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 419–431.     

Natural or Naturalized? Phylogeography Suggests That the Abundant Sea Urchin Arbacia lixula Is a Recent Colonizer of the Mediterranean

We present the global phylogeography of the black sea urchin Arbacia lixula, an amphi-Atlantic echinoid with potential to strongly impact shallow rocky ecosystems. Sequences of the mitochondrial cytochrome c oxidase gene of 604 specimens from 24 localities were obtained, covering most of the distribution area of the species, including the Mediterranean and both shores of the Atlantic. Genetic diversity measures, phylogeographic patterns, demographic parameters and population differentiation were analysed. We found high haplotype diversity but relatively low nucleotide diversity, with 176 haplotypes grouped within three haplogroups: one is shared between Eastern Atlantic (including Mediterranean) and Brazilian populations, the second is found in Eastern Atlantic and the Mediterranean and the third is exclusively from Brazil. Significant genetic differentiation was found between Brazilian, Eastern Atlantic and Mediterranean regions, but no differentiation was found among Mediterranean sub-basins or among Eastern Atlantic sub-regions. The star-shaped topology of the haplotype network and the unimodal mismatch distributions of Mediterranean and Eastern Atlantic samples suggest that these populations have suffered very recent demographic expansions. These expansions could be dated 94–205 kya in the Mediterranean, and 31–67 kya in the Eastern Atlantic. In contrast, Brazilian populations did not show any signature of population expansion. Our results indicate that all populations of A. lixula constitute a single species. The Brazilian populations probably diverged from an Eastern Atlantic stock. The present-day genetic structure of the species in Eastern Atlantic and the Mediterranean is shaped by very recent demographic processes. Our results support the view (backed by the lack of fossil record) that A. lixula is a recent thermophilous colonizer which spread throughout the Mediterranean during a warm period of the Pleistocene, probably during the last interglacial. Implications for the possible future impact of A. lixula on shallow Mediterranean ecosystems in the context of global warming trends must be considered.     

Phylogeny and biogeography of Muusoctopus (Cephalopoda: Enteroctopodidae)

Deep‐sea octopuses of the genus Muusoctopus are thought to have originated in the Pacific Northern Hemisphere and then diversified throughout the Pacific and into the rest of the World Ocean. However, this hypothesis was inferred only from molecular divergence times. Here, the ancestral distribution and dispersal routes are estimated by Bayesian analysis based on a new phylogeny including 38 specimens from the south‐eastern Pacific Ocean. Morphological data and molecular sequences of three mitochondrial genes (16S rRNA, COI and COIII) are presented. The morphological data confirm that specimens newly acquired from off the coast of Chile comprise two species: Muusoctopus longibrachus and the poorly described species, Muusoctopus eicomar. The latter is here redescribed and is clearly distinguished from M. longibrachus and other closely related species in the region. A gene tree was built using Bayesian analysis to infer the phylogenetic position of these species within the species group, revealing that a large genetic distance separates the two sympatric Chilean species. M. longibrachus is confirmed as the sister species of Muusooctopus eureka from the Falkland Islands; while M. eicomar is a sister species of Muusoctopus yaquinae from the North Pacific, most closely related to the amphi‐Atlantic species Muusoctopus januarii. Molecular divergence times and ancestral distribution analyses suggest that genus Muusoctopus may have originated in the North Atlantic: one lineage dispersed directly southward to the Magellan region and another dispersed southward along the Eastern Pacific to the Southern Ocean and Antarctica. The Muusoctopus species in the Southern Hemisphere have different phylogenetic origins and represent independent invasions of this region.     

Seed morphology in Moehringia L. and its taxonomic significance in comparative studies within the Caryophyllaceae

Seeds of 30 species of Moehringia and 12 representatives of the Caryophyllaceae were examined with the Scanning Electron Microscope. High diversity was found in seed coat micromorphology, and a number of novel morphological features have been observed. Comparison with other representatives of the Caryophyllaceae has provided new insight into the potential taxonomic value of discrete morphological characters. Though many characters appear informative within Moehringia, they are affected by a high level of homoplasy when considered in a wider phylogenetic context. Smooth seed is the common condition in Moehringia, except for Eastern Balkan and Iberian species that maintain a secondary ornamentation of the testa cells resembling the plesiomorphic Caryophyllaceae-type. Five types of strophioles were recognized, in accordance with previous literature. Among these, the strophiole of the Iberian species appears very distinctive and a proper strophiole could not be verified in M. glochidisperma and M. fontqueri.     

TESTS OF PHYLOGEOGRAPHIC MODELS WITH NUCLEAR AND MITOCHONDRIAL DNA SEQUENCE VARIATION IN THE STONE CRABS,MENIPPE ADINA AND MENIPPE MERCENARIA

Evolutionary relationships among stone crabs (Menippe) from the Gulf of Mexico and western Atlantic were investigated by comparisons of restriction sites within anonymous nuclear DNA sequences and nucleotide sequences of both mitochondrial and a duplicated nuclear form of the mitochondrial large subunit ribosomal RNA (LSrDNA) gene. A survey of over 100 restriction sites by Southern blot analysis with 10 anonymous nuclear DNA sequence probes failed to reveal any differences between Menippe adina and M. mercenaria. Sequence comparisons of both mitochondrial and nuclear forms of the LSrDNA gene also did not distinguish these species. Although both LSrDNA gene sequences were variable, some haplotypes were shared by the two species, implying either incomplete gene lineage sorting or introgressive hybridization. Based on molecular clock calibrations, we estimate that all of the observed mitochondrial LSrDNA sequences share a common ancestor between 1.5 and 2.7 million years before present (M.Y.B.P.). However, because identical sequences are shared by the two species, these data are also compatible with a more recent common ancestry. These findings conflict with a previously proposed biogeographic scenario for North American Menippe, which featured a relict hybrid zone on the Atlantic Coast. We suggest an alternative scenario based on relatively recent events and ongoing, rather than historical, gene flow.     

Molecular phylogeny of the crab genus Brachynotus (Brachyura: Varunidae) based on the 16S rRNA gene

The crab genus Brachynotus de Haan, 1833 is restricted to the intertidal and shallow subtidal of the Mediterranean and northeastern Atlantic. It is presently recognized to consist of four species, of which three (B. foresti, B. gemmellari and B. sexdentatus) are endemic to the Mediterranean. The fourth species, B. atlanticus, is found along the Atlantic coasts of northern Africa and southern Europe, but also extends into the western Mediterranean. This high level of endemism suggests that speciation within Brachynotus is strongly correlated with the geography and geology of the Mediterranean Sea. A molecular phylogeny based on the mitochondrial large subunit (16S) rRNA gene indicates that the four species of Brachynotus form a monophyletic group within Atlantic Varunidae. The DNA sequence data also show that the genus Brachynotus can be subdivided into two species groups, one comprising B. atlanticus and B. foresti, and the other one B. gemmellari and B. sexdentatus. While B. atlanticus and B. foresti are clearly genetically distinct, B. gemmellari and B. sexdentatus are identical in the studied region of the 16S rRNA gene, suggesting a recent separation or continuing gene flow.     

A global invader at home: population structure of the green crab, Carcinus maenas, in Europe

The European green crab, Carcinus maenas, has a native distribution that extends from Norway to Mauritania. It has attracted attention because of its recent invasions of Australia, Tasmania, South Africa, Japan and both coasts of North America. To examine the population structure of this global invader in its native range, we analysed a 502-base-pair fragment of the mitochondrial cytochrome c oxidase I (COI) gene from 217 crabs collected in the North Atlantic and 13 specimens from the Mediterranean. A clear genetic break (11% sequence divergence) occurs between the Mediterranean and Atlantic, supporting the species-level status of these two forms. Populations in the Faeroe Islands and Iceland were genetically distinct from continental populations (F(ST) = 0.264-0.678), with Iceland represented by a single lineage also found in the Faeroes. This break is consistent with a deep-water barrier to dispersal in green crabs. Although there are relatively high levels of gene flow along the Atlantic coast of Europe, slight population structure was found between the central North Sea and populations to the south. Analysis of variance, multidimensional scaling, and the distribution of private haplotypes support this break, located between Bremerhaven, Germany, and Hoek van Holland. Similar biogeographical and genetic associations for other species, such as benthic algae and freshwater eels, suggest that the marine fauna of Europe may be generally subdivided into the areas of Mediterranean, western Europe and northern Europe.     

Phylogenetic reconstruction of the genusBrachypodium P. Beauv. (Poaceae) from combined sequences of chloroplastndhF gene and nuclear ITS

A phylogenetic reconstruction of eight species of the genusBrachypodium P. Beauv. (Poaceae) has been obtained combining sequence data from the chloroplastndhF gene and the nuclear ITS, and using five representatives of tribes Triticeae (Secale), Poeae (Lolium), Meliceae (Melica, Glyceria) and Oryzeae (Oryza) as out-groups. Similar numbers of informative substitutions for the ingroup species were provided by both the 3 region of the chloroplastndhF gene and the nuclear ITS region. The Mediterranean annualBrachypodium distachyon appears to be the basal lineage, followed by the divergence of the New World non-rhizomatousB. mexicanum, which antedates the separation of a core of six European and Eurosiberian rhizomatous perennials (Brachypodium arbuscula, B. retusum, B. rupestre, B. phoenicoides, B. pinnatum, andB. sylvaticum). The evolutionary reconstruction based on sequences of the chloroplast and the nuclear genomes is congruent with topologies obtained from analysis of RAPD data.     

Molecular phylogeny of the Acre clade (Crassulaceae): Dealing with the lack of definitions for Echeveria and Sedum

The phylogenetic relationships within many clades of the Crassulaceae are still uncertain, therefore in this study attention was focused on the “Acre clade”, a group comprised of approximately 526 species in eight genera that include many Asian and Mediterranean species of Sedum and the majority of the American genera (Echeveria, Graptopetalum, Lenophyllum, Pachyphytum, Villadia, and Thompsonella). Parsimony and Bayesian analyses were conducted with 133 species based on nuclear (ETS, ITS) and chloroplast DNA regions (rpS16, matK). Our analyses retrieved four major clades within the Acre clade. Two of these were in a grade and corresponded to Asian species of Sedum, the rest corresponded to a European–Macaronesian group and to an American group. The American group included all taxa that were formerly placed in the Echeverioideae and the majority of the American Sedoideae. Our analyses support the monophyly of three genera – Lenophyllum, Thompsonella, and Pachyphytum; however, the relationships among Echeveria, Sedum and the various segregates of Sedum are largely unresolved. Our analyses represents the first broad phylogenetic framework for Acre clade, but further studies are necessary on the groups poorly represented here, such as the European and Asian species of Sedum and the Central and South American species of Echeveria.     

Multiple Recurrent Evolution of Trophic Types in Northeastern Atlantic and Mediterranean Seabreams (Sparidae, Percoidei)

Seabreams are among the most valuable fish, not only for small-scale and semiindustrial fisheries but also for aquaculture throughout the Mediterranean. Nevertheless, their phylogenetic relationships are not at all clear. The current taxonomy is based solely on trophic morphology and rests on the assumption that each trophic type evolved only once from a less specialized ancestral condition. We analyzed a 486-bp segment of the mitochondrial 16S rDNA of all 24 seabream species described for the northeastern Atlantic and the Mediterranean to elucidate their generic and subfamily-level relationships. Three major mitochondrial lineages, each comprising species of different feeding strategy and dentition, were found that do not agree with the present taxonomic assignments. Most of the investigated genera were resolved paraphyletically, indicating that the structure and arrangement of oral teeth must have repeatedly evolved from a less specialized ancestral condition. Further, the genus Sparus was resolved as distantly related to the genus Pagrus, in that it was assigned to a different major mitochondrial lineage. Oblada melanura was consistently placed within the Diplodus radiation as sister group to Diplodus puntazzo. Our phylogenetic hypothesis thus suggests multiple independent origins of similar trophic specializations within the Sparidae and indicates that the currently recognized three or four subfamilies need to be redefined. Received: 5 October 1999 / Accepted: 9 November 1999     

Evolution of Nemertesia hydroids (Cnidaria: Hydrozoa,Plumulariidae) from the shallow and deep waters of the NE Atlantic and western Mediterranean

Moura, C. J., Cunha, M. R., Porteiro, F. M., Yesson, C. & Rogers, A. D. (2011) Evolution of Nemertesia hydroids (Cnidaria: Hydrozoa, Plumulariidae) from the shallow and deep waters of the NE Atlantic and Western Mediterranean. —Zoologica Scripta, 41, 79–96. Hydroid species from the genus Nemertesia develop some of the largest and most complex hydrozoan colonies. These colonies are abundant and ecologically important in both shallow and deep waters worldwide. Here, we analyse the systematics of most Nemertesia species from the NE Atlantic and Mediterranean using morphology and phylogenetic inferences of 16S rRNA haplotype data. Phylogeographical analysis revealed multiple movements of taxa to and from the Mediterranean after the Messinian salinity crisis through shallow and deep waters. The nominal species Nemertesia belini and Nemertesia antennina revealed multiple genetic lineages representing cryptic species diversity. Molecular phylogenetic evidence was supported by consistent phenotypic differences between lineages, and three and seven putative species were resolved within the N. belini and N. antennina complexes, respectively. Three putative species of the N. antennina complex found at different seamounts of Azores grouped in a clade clustered amongst the other four cryptic species present at neighbouring bathyal localities of the Gulf of Cadiz. These cryptic species, mostly from the deep sea, form a clade distantly related to the typical N. antennina from European coastal waters. Depth or environmental correlates of depth seem to influence the reproductive strategies of Nemertesia colonies and ultimately speciation. In particular, speciation of these hydroids must have been influenced by hydrography, habitat heterogeneity, isolation by distance and larval dispersal capacity. The deep sea is shown as an important environment in the generation and accumulation of lineages that may occasionally invade coastal waters in the NE Atlantic. Glacial cycles of cooling, along with changes in sea level, and eradication of some coastal faunas likely facilitated speciation and evolutionary transitions from deep to shallow waters.