A new cryptic species of <Emphasis Type="Italic">Aponurus</Emphasis> Looss, 1907 (Digenea: Lecithasteridae) from Mediterranean goatfish (Teleostei: Mullidae)

The status of the trematode Aponurus laguncula Looss, 1907 in the western Mediterranean is re-assessed by means of a comparative morphological study and rDNA sequences based on newly collected material. A. laguncula (sensu stricto) is redescribed from Trachinus draco L. and a new cryptic species of the ‘A. laguncula complex’, Aponurus mulli n. sp., is described on the basis of abundant material from Mullus barbatus L. (type-host) and M. surmuletus L. off the Spanish Mediterranean coasts. The new species is differentiated from A. laguncula (sensu stricto) by its: significantly larger, elongate body, with maximum width at the level of the ventral sucker; shorter forebody; distinctly larger sinus-sac, seminal receptacle and seminal vesicle, with the latter also being more elongate; vesicular pars prostatica; more anteriorly located vitellarium, which consists of eight globular follicles; and distinctly smaller eggs, which are also smaller in relation to body size and have both their opercular and anopercular poles rounded. The variability and the allometric growth of the morphological characters in the new species were studied in detail, resulting in additional distinguishing features. Sequences of the large subunit rRNA (28S) gene (domains D1–D3) and ITS2 rRNA gene region for the new species have been submitted to GenBank in order to enhance future studies on species differentiation within the ‘A. laguncula complex’.     

New and old problems in the taxonomy of the Gelidiales (Rhodophyta)

The order Gelidiales includes over 140 agar-producing taxa. Many species are taxonomically confused; the boundaries of one family and all but four genera recently have been contested, and the controversy over ordinal status has lasted for over 25 years. This study reviews recent developments in the taxonomy of the group and suggests future areas for studies. The order cannot be defined by a few exclusive characters, as intended in the past, but it shows a unique combination of characters. Additional studies on Gelidiella and Acanthopeltis seem advisable to clarify family limits. The segregation of Onikusa and Pterocladiastrum is doubtful. Only three of six characters discriminate Gelidium from Pterocladia. None allows complete generic segregation and all need variability studies. Analysis of nomenclatural types in the light of morphological variation would permit an understanding of species limits in Gelidium and Pterocladia. Similar studies are needed in Gelidiella and Ptilophora.     

Phylogeny,biogeography, and morphological ancestral character reconstruction in the Mediterranean genus Fumana (Cistaceae)

Fumana is a diverse genus of the Cistaceae family, consisting of 21 currently accepted species. In this study, nuclear (ITS) and plastid (matK, trnT‐L) molecular markers were used to reconstruct the phylogeny and to estimate divergence times, including 19 species of Fumana. Phylogenetic analyses (Bayesian Inference, Maximum Parsimony and Maximum Likelihood) confirmed the monophyly of Fumana and did not support the infrageneric divisions previously established. The results support four main clades that group species that differ in vegetative and reproductive characters. Given the impossibility to define morphological characters common to all species within the clades, our proposal is to reject infrageneric divisions. Molecular dating and ancestral area analyses provide evidence for a Miocene diversification of the genus in the north‐western Mediterranean. Ancestral state reconstructions revealed ancestral character states for some traits related to xeric and arid habitats, suggesting a preadaptation to the Mediterranean climate.     

Redescription of Fabriciola tonerella Banse, 1959, and a new record of Novafabricia infratorquata (Fitzhugh, 1983) from the Mediterranean Sea, with a key for the Fabriciinae (Annelida: Polychaeta) of the Mediterranean Sea and the north-east Atlantic

The fabriciin sabellid species Fabriciola tonerella Banse, 1959 living on gastropod shells inhabited by hermit crabs collected in the western Mediterranean Sea is redescribed. Another species of Fabriciinae, Novafabricia infratorquata (Fitzhugh, 1983), is recorded for the first time from the Mediterranean Sea and found in the same habitat is described as well. The variability of characters depending on size and geographical distribution is given for both species. Moreover, selected characters for all known species of Fabriciola and Novafabricia as well as a simple key for all Fabriciinae of the Mediterranean Sea and the north-east Atlantic are provided.     

Distribution and biogeography of the Paradiaptominae (Copepoda: Calanoida: Diaptomidae)

The Paradiaptominae, a subfamily of the large Diaptomidae family, is a small group of freshwater calanoids of which 24 species in four genera have been described. They are endemic to Africa with the exception of four species, Metadiaptomus asiaticus (Asia and Mongolia), M. gauthieri (Africa and Madagascar), Paradiaptomus greeni (India and Sri Lanka) and Neolovenula alluaudi (Canary Islands, North Africa, Baltic and Mediterranean countries). The North African species Metadiaptomus chevreuxi extends its range into Iraq and Iran. The Paradiaptominae are widely distributed in semi-permanent water bodies in the drier areas of Africa although most individual species have a restricted distribution. Only a few species have the ability to colonise permanent waters and these species are usually located at higher altitudes. The Paradiaptominae have not been recorded from the wet equatorial lowland areas of Africa. It seems likely that they are Gondwanian in origin as evidenced by their distribution (Africa, Madagascar, India and Sri Lanka). Neolovenula alluaudi (Mediterranean) shares morphological characters tenuously linked to the other species in the group and may either have a different origin or have branched off at an early stage in evolution of the group.     

An integrative approach to the taxonomy of the pigmented European Pseudomonocelis Meixner, 1943 (Platyhelminthes: Proseriata)

Only a few species belonging to the Proseriata (Platyhelminthes) show a parenchymatic pigmentation, which may aid identification. Among these, Pseudomonocelis agilis has a yellowish body and is provided with a reddish–brown girdle in front of the statocyst. The species is known for limited areas of northern Europe and the Mediterranean. The present study was conducted to assess both the taxonomic status of populations attributed to the species across the unusually wide range for an interstitial flatworm, which lacks an obvious means of dispersal, and the levels of genetic variability within and among populations, by employing an integrative approach that included the analyses, on six populations, of three molecular markers (small subunit ribosomal 18S‐like gene, inter‐simple sequence repeat, allozymes), karyotypes, and 11 morphological characters. Furthermore, crossbreeding experiments were carried out on the Mediterranean populations. The results obtained revealed the existence of four highly divergent genotypic clusters, accompanied by karyological differences, with complete intersterility among the clusters tested. The combination of approaches adopted strongly supports the conclusion that the wide‐ranging European pigmented species P. agilis is actually composed of four species: P. agilis in the Baltic area; Pseudomonocelis cetinae in the Adriatic; and Pseudomonocelis sp. nov. A and Pseudomonocelis sp. nov. B in the western and eastern Mediterranean, respectively. The latter two species are morphologically indistinguishable for the parameters essayed. Reconstruction of the phylogenetic relationships of these taxa, including congeneric and consubfamilial outgroups, showed that pigmentation is a plesiomorphic condition for the genus Pseudomonocelis and that Pseudomonocelis sp. nov. A shares a previously undetected, sister‐group relationship with species of the unpigmented P. ophiocephala complex. The present study thus depicts complex speciation processes in a mesopsammic species, which involves allopatric divergence operating on different scales and ecological shifts, and highlights that the contribution of microturbellarians to marine biodiversity may be seriously underestimated. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 907–922.     

Morphology of the early zoeal stages of Macropipus tuberculatus (Roux, 1830) (Crustacea, Brachyura, Portunidae)

The first three zoeal stages of the portunid crab Macropipus tuberculatus (Roux, 1830) are described and illustrated from laboratory-reared material obtained from one ovigerous crab captured in the western Mediterranean Sea. Zoeae of M.tuberculatus can be distinguished from other known polybiinid species from the northeastern Atlantic and Mediterranean waters by a combination of meristic and morphometric features, especially by the number of setae in the outer lobe of the maxilla endopod. Larval morphology presents additional evidence supporting the split of the genus Macropipus into the genera Liocarcinus and Macropipus sensu stricto based on adult characters.      

Karyometric and morphological analysis of two sympatric marine species of the Gyratrix hermaphroditus complex (Platyhelminthes: Kalyptorhynchia) occurring at Roscoff (Brittany,France)

Two marine sibling species of the Gyratrix hermaphroditus complex were found to occur sympatrically at Roscoff (Brittany, France). Both share the same chromosome number 2n = 6, but differ for the values of centromeric indices, length ratio of chromosomes and absolute length of the haploid genome. Differences were also found in body size and male cuticular apparatus morphology. They are furthermore distinct for the above characters from the other two marine species of the complex already known (from the Mediterranean and the North Sea). Chromosomal evolution and phylogenetical relationships within the marine species of the G. hermaphroditus complex are discussed in detail.     

Taxonomic characters from the polyp tubes of coronate medusae (Scyphozoa,Coronatae)

The tubes of the polyp stages of all known species of the family Nausithoidae have been examined to find characters useful for specific identification. Such characters comprise both internal and external structures of the chitinous exoskeleton, measurements of the periderm, and the shape of the tubes defined by the Formquotient. A key for the polyp generation is given based on these characters. The long-term rearing of several species and elucidation of their life cycles has also enabled a key to the medusa generation of the Nausithoidae to be constructed. Stephanoscyphistoma gen. nov. is proposed for the polyps of all species of coronates of uncertain medusa genus.     

Comparative morphology,biology and phylogeny of terminal‐instar larvae of the European species of Toryminae (Hym., Chalcidoidea,Torymidae) parasitoids of gall wasps (Hym. Cynipidae)

We present a phylogenetic and taxonomic study of the morphology and biology of the terminal‐instar larval stage of 19 species representing all the genera of Torymidae parasitoids of gall wasps in Europe, with the single exception of Megastigmus. The genera studied include Adontomerus Nikol'skaya, Idiomacromerus Crawford, Chalcimerus Steffan & Andriescu, Glyphomerus Förster, Pseudotorymus Masi and Torymus Dalman. We primarily used chaetotaxy and some head structures. The terminal‐instar larvae of all studied species are thoroughly described for the first time and illustrated with SEM images. We provide diagnostic characters for the family and the genera studied, and keys to genera and species for the identification of torymid larvae associated with cynipid galls. The majority of the torymid larvae studied are solitary monophagous parasitoids. Finally, to assess the potential use of larval characters in systematic studies of the family, a phylogenetic analysis of the studied taxa based on 42 larval morphological characters is proposed and compared with the current taxonomy of Torymidae. Our results suggest that body chaetotaxy, and characters of the head and mouthparts could be used for genera and species discrimination. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 676–721.     

Variability in the taxonomic characters of Black Sea gyrodactylids (Monogenea)

Variability of the haptoral hard parts of four Black Sea Gyrodactylus species (G. alviga Dmitrieva & Gerasev, 2000; G. crenilabri Zaika, 1966; G. flesi Malmberg, 1957; G. sphinx Dmitrieva & Gerasev, 2000) was studied in relation to season, host and geographical variability. A discriminant analysis was carried out on the measurements of 14 characters of the haptor of all four species. The haptoral hard parts of the investigated gyrodactylids decrease in size with an increase in water temperature. It is known that life-span of gyrodactylids is negatively correlated with water temperature and they exhibit peak intensities at low temperature. At the same time, life-span is considered to be a general indicator of a favourable environment. Therefore, high water temperature is not apparently advantageous for gyrodactylids. In the case of low marine salinity, in adapted species the sizes of the haptoral hard parts are larger at lower levels of salinity and vice versa for typical marine species. Lastly, in specimens parasitising the primary host, these features are larger than in those parasitising a secondary host. It is possible to say that, in general, the more favourable the environment for gyrodactylids the larger their haptoral hard parts. Unfavourable environmental conditions reduce the time of embryogenesis, which has a dual effect on gyrodactylids: positive in relation to their reproduction (by increasing the reproduction rate) and negative in relation to their development (by decreasing the size of the haptoral hard parts).     

Seasonal variability of copepod abundance in the Balearic region (Western Mediterranean) as an indicator of basin scale hydrological changes

Spatial and temporal changes of the copepod community have been investigated and related to the environmental variability of the Balearic Sea (Western Mediterranean). The period studied spans from 1994 to 1999 during which we analyzed the abundance and structure copepod variability over a cross-shore transect. Results showed a close link between hydrological changes and the variations of copepod abundance. The synchronous variability of copepods and hydrography indicated the rapid response of this zooplankton group to the inflow of cold and warm water masses coming through the study area. Cluster analysis revealed four main copepod assemblages that distinguished the coastal from the oceanic species and those species with different water masses preference. The copepod assemblage composed of Calanus helgolandicus, Clausocalanus arcuicornis, C. pergens, C. paululus, Calocalanus tenuis and Pleuromamma gracilis was associated with cool salty waters, whereas the assemblage formed by Temora stylifera, C. pavo, C. styliremis, Centropages bradyi and Acartia danae was related to warmer less saline Mediterranean waters. Moreover, it is suggested that changes in sea water temperature and salinity are linked to large-scale changes likely occurring at a basin scale, which is reflected in the Western Mediterranean mesoscale hydrographic changes. Therefore, it is stressed that changes in the Balearic copepod community can be used as potential tracers of the western Mediterranean water masses. Guest editors: S. Souissi & G. A. Boxshall Copepoda in the Mediterranean: Papers from the 9th International Conference on Copepoda, Hammamet, Tunisia     

Contribution to the taxonomy of the family Campulidae Odhner, 1926 (Digenea) by means of a morphometric multivariate analysis

Digeneans of the family Campulidae occur exclusively in marine mammals, particularly in cetaceans. Their taxonomy is confused, being based on adult morphology only. We used a multivariate discriminant analysis of morphometric data to provide new evidence on the taxonomy of the Campulidae. Measurements of 217 specimens from 21 species of all seven genera of the family were taken. The percentage of specimens correctly assigned into their own species was 96.3%. The first three discriminant functions accounted for most of the variation between the species, which were grouped together in suprageneric groups along the first and the second function. The ordination pattern observed conforms partly with the established taxonomy of the Campulidae. Variation along the first discriminant function can be interpreted as a gradient of the vitelline extension and the maximum width to length ratio. This function separates three groups of species, corresponding to the Hunterotrematinae, Orthosplanchninae and Campulinae-Lecithodesminae. The last two subfamilies are separated along the second axis, the Lecithodesminae having longer bodies and suckers than the Campulinae. This study also provides some evidence at the generic and specific level.     

Biased sex ratio and niche restriction in Baruscapillaria obsignata (Madsen 1945) (Nematoda, Capillariidae) from Columba livia (Aves, Columbidae)

In the present study populations of the avian nematode species Baruscapillaria obsignata are described from Columba livia. Male and female individuals were obtained from 27 birds, fixed in alcohol/formalin/acetic acid (AFA) and preserved in 70% ethanol. Nematodes were identified and then counted under a stereoscopic microscope. Baruscapillaria obsignata were much more frequent in the anterior third of the small intestine, and females were more abundant than males in all infra populations. The prevalence was 55.6%, mean intensity was 11.8 (median 11.0; range 1-31) and abundance 6.56. In the present study, we observed an aggregated distribution of parasite infrapopulations, as demonstrated by the value of the exponent of the negative binomial distribution, K?=?0.2773; by the discrepancy index, D?=?0.656 and by the variance/mean ratio, 12.44. The female/male sex ratios found in all infrapopulations were always greater than 1, showing a bias in favour of female abundance. This tendency was especially marked in infrapopulations containing fewer individuals. The sizes of infrapopulations ranged from 5 to 31 individuals. The mean sex ratio observed was 2.69?±?3.28 (median 1.83; range 0-11). In infrapopulations with 5-15 individuals, the sex ratios observed varied from 2.6 to 11, while in those with 17-31 individuals, the sex ratios were lower, ranging from 1.7 to 2.4. There was a negative correlation between the intensity of infection and the sex ratio of infrapopulations. Results are discussed in terms of possible factors influencing the processes that lead to niche restriction and biased sex ratios in parasite infrapopulations.     

A phylogenetic assessment of the branchiobdellidan family Branchiobdellidae (Annelida,Clitellata) using spermatological and somatic characters

The spermatozoa of six species belonging to the branchiobdellidan family Branchiobdellidae (i.e. Xironogiton victoriensis, Cirrodrilus kawamurai, Ankyrodrilus legaeus, Xironodrilus formosus, Branchiobdella kobayashii, Branchiobdella orientalis) were studied and compared to the other sperma‐tozoa already described in the group. A parsimony analysis was performed on the spermatozoal data of the species examined, as well as on their somatic characters. The results of the two analyses were contrasted and a further parsimony analysis was run on the matrix comprising both sets of characters. The study of sperm ultrastructure confirmed the genera recognized with traditional somatic characters and the monophyly of the branchiobdellidans. Xironodrilus was proved to be the sister species of Ankyrodrilus and its inclusion into the family Branchiobdellidae was supported. Evolutionary hypotheses on intergeneric differences in the family consistent with its biogeography can be suggested by the cladograms: Xironogiton is an early offshoot of branchiobdellidan lineage migrating to North America and probably radiating only in recent times; Branchiobdella kobayashii has a spermatozoon completely different from that of the other species of the genus, thus suggesting a complex story for this widespread taxon with a disjunct distribution.     

Cryptic speciation and genetic structure of widely distributed brittle stars (Ophiuroidea) in Europe

Pérez‐Portela, R., Almada, V. & Turon, X. (2012). Cryptic speciation and genetic structure of widely distributed brittle stars (Ophiuroidea) in Europe. —Zoologica Scripta, 00, 000–000. The development of molecular techniques has led to the detection of numerous cases of cryptic speciation within widely distributed marine invertebrate species and important taxonomic revisions in all the major marine taxa. In this study, we analysed a controversial marine species complex in the genus Ophiothrix, a widespread taxon in European waters traditionally assigned to two nominal species, Ophiothrix fragilis and O. quinquemaculata. These species are important components of the rocky shores and deep marine benthos along the North Atlantic and Mediterranean littoral. Their status (including variants of both species) has remained contentious due to overlapping variability in morphological characters. In this study, we analysed the genetic and morphological differences of Ophiothrix lineages along the Atlantic and Mediterranean coasts. We also assessed population genetic structure in the Atlantic and Mediterranean basins by sequencing two mitochondrial genes, the 16S rRNA gene and COI gene, of 221 specimens from 13 locations. Phylogenetic analyses demonstrated the existence of two genetically distinct lineages, attributable to two different species although unrelated to previous taxonomic distinctions. Morphological differences could also be detected between these lineages. Samples from the Northeast Atlantic and one from the deep Mediterranean grouped within Lineage I, whereas Lineage II pooled together the southern Atlantic and rocky shallow Mediterranean samples. In the northern region of the Iberian Peninsula and at a deep locality in the Mediterranean, both lineages overlap. Speciation processes likely happened during the Mio–Pliocene transition (about 4.8–7.5 million years ago), when marine‐level oscillations led to the blockage of major marine corridors in Europe and promoted genetic isolation by vicariance. Secondary contact between lineages following sea‐level increases and recolonization during the refilling of the Mediterranean after the Miocene salinity crisis could explain the present‐day distribution of genetic variability. No barriers to gene flow along the Atlanto‐Mediterranean area were detected for Lineage II, and the lack of genetic structure could be caused by a mixture of several factors, such as wide dispersal potential, recent demographic expansion and large population size.     

Infrapopulations of Sclerocollum saudii Al-Jahdali, 2010 (Acanthocephala: Cavisomidae) in the rabbitfish Siganus rivulatus (Teleostei, Siganidae) from the Saudi coast of the Red Sea

In infrapopulations of helminth parasites, density-dependent effects, through some form of intra- and interspecific competition, play an important role in shaping and regulating the infrapopulations. The mechanisms responsible for these processes have often been observed in laboratory studies and rarely studied under natural conditions. Here, 24 natural infrapopulations (77-447 individuals) of the acanthocephalan Sclerocollum saudii Al-Jahdali, 2010 from the fish Siganus rivulatus consisted of cystacanths, newly excysted juveniles, immature and mature worms, distributed in a well-defined fundamental niche (anterior 60% of the intestine). Each of these stages exhibited a significantly different longitudinal distribution within this niche. In small infrapopulations, cystacanths and newly excysted juveniles were found in the sixth 10% of the intestine, immature worms in the fifth 10% and mature worms in the anterior 40% of the intestine. However, their proportions followed a clear ascending order in each infrapopulation, and the female-male ratios of both immature and mature worms were distinctly female-biased. In large infrapopulations, mature worms existed partially in the site of immature ones, where a differential mortality among immature females was constantly observed. However, the proportions of immature worms increased significantly and those of mature worms decreased significantly, the mean lengths of immature and mature females decreased dramatically and the female-male ratios were distinctly male-biased. The mean sizes of immature and mature males seemed stable through all infrapopulations. The distribution of mature males and females suggests intense male-male competition for access to females, and reveals that larger females are copulated prior to the smaller ones. The results are statistically significant and suggest that infrapopulation self-regulation is through density-dependent mechanisms, in which immature females may play a key role.     

Systematics ofPrimula veris (Primulaceae)

Primarily, 50 characters taken from herbarium specimens, cultivated plants, and whole populations ofPrimula veris L. were critically evaluated. Only 24 characters were found to be suitable for taxonomic use. The geographical and multivariate analysis of about 1000 herbarium specimens from the whole area lead to a reduction of the number of subspecies. Only a less restricted subsp.veris (incl. subsp.canescens, subsp.inflata and probably including subsp.macrocalyx) and subsp.columnae (incl. subsp.suaveolens) can be distinguished considering the variability of characters within an individuum, under different culture conditions, and during the growth period. The status of the taxon macrocalyx cannot be cleared presently.     

Spatio-temporal distribution of recruits (0 group) of Merluccius merluccius and Phycis blennoides (Pisces, Gadiformes) in the Strait of Sicily (Central Mediterranean)

Aspects of the recruitment of hake (Merluccius merluccius L., 1758) and greater fork beard (Phycis blennoides Brunnich, 1768) in the Strait of Sicily (Central Mediterranean) are presented. Data were collected from 1994 to 1999 during the international bottom trawl survey program MEDITS. In view of the available literature on juvenile growth of these two species in the Mediterranean area, a length-based criterion was adopted to separate fish belonging to the 0 group (recruits). Recruit density indices (R/km²) by haul were calculated based upon the likely variability of recruit growth among years and used to study abundance variability and spatial pattern of recruitment and to identify the main nursery areas. Although there was inter-annual variability, two stable areas for M. merluccius were identified on the eastern side of the Adventure Bank and the Malta Bank at depths ranging between 100 and 200 m. The main nursery areas for P. blennoides were deeper (from 200 to 400 m) and two stable nursery areas were identified on the western and eastern side of the Adventure Bank; other nurseries were found in the easternmost part of the Strait in 1998 and 1999. Recruitments of the two species were significantly correlated, with the strongest recruitment occurring in 1998 and 1999 for each species.