Jaw morphology and ontogeny in five species of Ophryotrocha

Detailed scanning electron microscopy of jaws within the genus Ophryotrocha (Dorvilleidae, Annelida) was performed on 871 jaw parts. The investigations resulted in new understandings of the ontogeny and jaw morphology and have systematic implications for the family. Five species in the genus (Ophryotrocha alborana, O. diadema, O. gracilis, O. hartmanni, and O. labronica pacifica) were kept in culture, and the development of the jaws was studied by sampling throughout their life history. Ophryotrocha species have mandibular plates that remain the same throughout ontogeny, whereas the posterior shafts elongate. Both mandibular plate morphology and shaft ontogeny have species‐specific distinctions. In Ophryotrocha, the maxillae can be assigned to three to four distinct types, which are replaced by moulting. The maxillary morphology and developmental stages at which moults occur are species specific, although with broad intervals. A redefinition is given for some of the basic jaw elements, and new homologies are proposed for structures that are also present across other dorvilleid taxa. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Systematics and biodiversity of Ophryotrocha (Annelida,Dorvilleidae) with descriptions of six new species from deep-sea whale-fall and wood-fall habitats in the north-east Pacific

Six new Ophryotrocha species are described from five whale-falls and two wood-falls off the southern Californian coast. Phylogenetic analyses based on the nuclear gene H3 and the mitochondrial genes COI and 16S using MrBayes and maximum likelihood analyses were performed on 40 dorvilleid taxa and one outgroup. Ophryotrocha batillus sp. nov. is morphologically identical to Ophryotrocha scutellus described from a shallow water whale-fall in the North Atlantic, although the two cryptic species differ genetically. Ophryotrocha langstrumpae sp. nov. is closely related in the molecular phylogenetic analyses to these two sibling species. Ophryotrocha flabella sp. nov. is similar to Ophryotrocha globopalpata, and although there are a few morphological differences, the genetic divergence is low between the two species. Ophryotrocha nauarchus sp. nov. is sexually dimorphic, with males having appendages on the first chaetiger. Ophryotrocha magnadentata sp. nov. and Ophryotrocha longicollaris sp. nov. are sister species in our molecular analyses, and together with O. nauarchus sp. nov. and O. flabella sp. nov. they fall within a clade that includes O. globopalpata and Exallopus jumarsi described from hydrothermal vents in the Pacific Ocean and Ophryotrocha longidentata from the shallow North Atlantic. Our results highlight the remarkable unknown diversity of deep-water habitats and the role of chemosynthetic ecosystems in the evolution of deep-sea life.     

Eine neueOphryotrocha-Art (Polychaeta,Eunicidae) aus Japan

Zusammenfassung 1. Ein japanischer Vertreter der kosmopolitischen PolychaetengattungOphryotrocha, O. notoglandulata n. sp., wird beschrieben.2.O. notoglandulata n. sp. ist eng mitO. labronica Bacci &La Greca verwandt.3. Besonderes Artmerkmal sind sternförmige Drüsenfelder auf den Dorsalflächen der hinteren 6–12 Segmente.4. Die neue Art ist getrenntgeschlechtlich.5. Das Geschlechtsverhältnis schwankt stark von Gelege zu Gelege. Im Mittel beträgt es 1,5:1.6.O. notoglandulata n. sp. ist mitO. labronica nicht kreuzbar.

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A newOphryotrocha species (polychaeta, eunicidae) from Japan

A new Japanese species ofOphryotrocha is described. It is characterized by oblong cells forming star-shaped glands of unknown function on the 6th to 12th of the youngest segments.Ophryotrocha notoglandulata n. sp. is closely related toOphryotrocha labronica Bacci &La Greca; both have the same chromosome number (2 n=6). The new species is gonochoristic with a sex ratio of approximately 1.5:1. In mass culture as well as in couples, no mutual influence on sex determination was found. It is impossible to obtain hybrids betweenO. labronica and the new species.

     

To change or not to change sex: A comparison between twoOphryotrocha species (Polychaeta)

Summary The polychaeteOphryotrocha puerilis is a male-to-female sex-changer, whereasO. labronica has separate sexes throughout life. In other respects the two species are remarkably similar: they occur in the same habitat, they look the same, they eat the same things, and in someO. labronica populations sex-changers can actually be found. The size-advantage hypothesis predicts that inO. puerilis males should not benefit reproductively from a size increase as much as males inO. labronica: henceO. puerilis but notO. labronica males should change to the female sex at a certain size. I experimentally compared reproductive success at different body sizes between the two species. In isolated male-female pairs reproductive rate increased significantly with female body size but not with male body size, and this pattern was the same in both species. Hence male fecundityper se cannot account for the difference in reproductive type between the two species. In other experiments I investigated if larger males gained access to more females because they were superior competitors for mates or were preferred by females, compared to small males. InO. puerilis the combined effect of these two factors conferred no size advantage to the males, whereas inO. labronica larger males acquired more females than did smaller males. Hence interactions among males and females, in accordance with the size-advantage hypothesis, can explain why sex change is maintained inO. puerilis, and why separate sexes are maintained inO. labronica.     

Integrative analyses on Western Palearctic Lasiommata reveal a mosaic of nascent butterfly species

Satyrinae butterflies occurring in the Mediterranean apparently have reduced gene flow over sea straits, and for several species, recent wide-scale biodiversity surveys indicate the existence of divergent mitochondrial lineages. Here, we apply an integrative approach and examine the phylogeography of the genus Lasiommata in the Western Palearctic. Our research comprised molecular analyses (mitochondrial and nuclear DNA) and geometric morphometrics (wings and genitalia) for two main species groups, and a comparative GMYC analysis, based on COI, of all the tribes within Satyrinae from this region. The GMYC approach revealed a particularly fast coalescence rate in the Parargina subtribe. The Lasiommata group was divided into 12 evolutionary significant units: six clades for the L. maera species group, five for the L. megera species group, and one for L. petropolitana, with divergences of about 1%. The patterns of COI were mirrored by ITS2 in L. maera, but the two markers were generally inconsistent in L. megera. On the contrary, morphological differences were coherent with the results of COI for L. megera, but less clearly so for L. maera. L. paramegaera and L. meadewaldoi were considerably differentiated for all the analyzed markers and likely proceeded faster in the process of speciation because of geographic isolation and reduced effective population size, rendering the rest paraphyletic. Our study illustrates the continuous nature of speciation and the difficulties of delimiting species. In Lasiommata, the recognition of taxa as diverging lineages or distinct, possibly paraphyletic species, mostly depends on the criteria adopted by different species concepts.     

Ultrastructural investigations on the nuchal organ of the protandric polychaete,Ophryotrocha puerilis (Polychaeta,Dorvilleidae)

Summary The nuchal organs of the protandric hermaphrodite Ophryotrocha puerilis were studied by electron microscopy. Ophryotrocha puerilis is the first species hitherto described which possesses four instead of two nuchal organs. These sensory structures are located as ciliary pits at the posterior margin of the prostomium. Histologically, the nuchal organs are composed of supporting cells with long motile cilia and bipolar sensory cells, the perikarya of which form four distinct nuchal ganglia adjoining the brain. These structural components are concentrically arranged around the central sensory area. This area is covered by a modified cuticle, whereas the cuticle above the peripheral region of the sense organ exhibits the appearance typical for polychaetes. Two types of vesicular material are produced in the basal supporting cells, a dense-cored one within the central supporting cells only and a clear irregular-shaped one in all of these cells. The first type is considered to be responsible for the formation of the modified cuticle. The significance of these most probably long-distance chemoreceptory organs and their possible role in reproductive behaviour is discussed.     

Complete mitochondrial genome and assembled DNA barcoding analysis of Lutjanus fulgens (Valenciennes, 1830) and its comparison with other Lutjanus species

Lutjanus fulgens (Valenciennes, 1830) is a teleost species classified under the family Lutjanidae which is a native of the Eastern Atlantic Ocean. Though highly commercialized due to its abundance and good taste, the production output has declined in recent years. This is an indication of the need for effective management and conservation measures. However, accurate species identification will ensure strategic management and conservation measure. DNA‐based species identification has proven its reliability in this regard via precise species identification. Several researchers have confirmed the accuracy of DNAbarcode as a species identification tool as well as species phylogeny analysis based on both the complete mitogenome and COI gene. Currently, nine specimens of L. fulgens were sampled from Ghana and subjected to DNA‐based analysis, namely, complete mitochondrial DNAand COI gene (DNA barcoding) analyses. The mitogenomic result revealed that L. fulgens is made up of a 16,500 base pairs (bp) mtDNA which consists of 22 transfer RNAs, 13 protein‐coding genes, and two ribosomal RNAs (GenBank Accession Number: MN398650). Furthermore, a sequence polymorphism analysis of the COIgene (MN986442–MN986450) detected two haplotypes. These haplotypes were both collected from the same fish landing site which suggests a possible cryptic linage diversity in the L. fulgens population at Vodza. According to the phylogeny examination, a close taxonomic relationship exists between L. fulgens and Lutjanus buccanella caused by a recent evolution termed as sympatric speciation. This study serves as a novel study for this species, building the foundation for future molecular‐based study for this species and as a DNA barcode reference data.     

The effect of the Messinian salinity crisis on the early diversification of the Tettigettalna cicadas

The current distribution patterns of many Mediterranean species are often a consequence of large and impactful past geoclimatic events, such as the Messinian Salinity Crisis (MSC) and the Quaternary glacial cycles. Cicadas are flying insects with poor dispersal ability, which have experienced intense local differentiation in the Mediterranean, where the genus Tettigettalna has surfaced as a biogeographic model. The genus includes 10 species with species-specific calling songs but identical morphology. All Tettigettalna species are restricted to Southern Iberia, with the exception of T. estrellae (northwest Iberia), the widespread T. argentata (mainly Iberia, France and Italy), and T. afroamissa (Morocco). With an expanded genetic dataset involving nuclear (EF1α) and mitochondrial (5′ and 3′ COI and ATP) loci, we reconstructed the phylogeny of the genus and estimated divergence dates for Tettigettalna species under a Bayesian framework. Phylogeny with the new mitochondrial dataset was in agreement with previous studies, whereas the nuclear EF1α supported T. josei and T. afroamissa as monophyletic clades but lacked resolution to resolve the remaining taxa. Some sister taxa share mitochondrial haplotypes, hinting for incomplete lineage sorting. Estimates of divergence time settled T. josei as the earliest diverging lineage, likely as a pre- or early-MSC event. As for the origin of T. afroamissa in Morocco, though time estimates could not entirely rule out post-MSC dispersal, the most likely scenario points to isolation of African Tettigettalna after the reopening of the strait of Gibraltar. The Pleistocene glaciations that followed likely impacted on the diversification of the remaining species of the genus in southern Iberia refugia.     

Spermatogenesis and sperm ultrastructure in the polychaete genusOphryotrocha (Dorvilleidae)

The details of spermatogenesis and spermiogenesis are described forOphryotrocha puerilis. The ultrastructure of mature sperm is shown forO. puerilis, O. hartmanni, O. gracilis, O. diadema, O. labronica, andO. notoglandulata. Clusters of sixteen cells each are proliferated by two stem cells in each setigerous segment ofO. puerilis representing the very early stages of both oogenesis and spermatogenesis. In each spermatocyte-I cluster, the cells are interconnected by cytoplasmic bridges. Early, clusters are enveloped by peritoneal sheath cells. These transient gonad walls break down prior to meiosis. The meiotic processes may start in the clusters with the cells still interconnected, or during breakdown of the original cluster, giving rise to smaller subclusters of both spermatocytes I and spermatocytes II with various numbers of cells. Finally, spermatid tetrads are present. As spermiogenesis progresses, the tetrads disintegrate. Golgi vesicles in both spermatocytes and spermatids contain electron-dense material, presumably preacrosomal. The acrosome is formed by such vesicles. In the six species studied here, the acrosomes appear to be of a similar overall structure but are of different shape. Centrioles are usually located beneath the acrosome. The distal centriole forms the basal body of a flagellum-like cytoplasmic process. The microtubules of these flagellar equivalents do not show a normal ciliar arrangement. The flagellar equivalent appears to be non-motile. InO. hartmanni and inO. notoglandulata, a flagellar equivalent is missing. Microtubules originating from the proximal end of the distal centriole stretch to the nuclear envelope. This feature appears to be especially conspicuous inO. puerilis and inO. labronica. InO. labronica and inO. notoglandulata, bundles of microtubules paralleling the cell perimeter appear to stabilise the sperm. Various numbers of mitochondria are either randomly distributed around the nucleus or accumulate on one side, often directly under the acrosome. Parts of the present paper were presented at the 2^nd International Polychaete Conference, Copenhagen 1986 and at the 3^rd International Polychaete Conference, Long Beach, Ca. 1989.     

Genetic and morphological variation in an ecosystem engineer,Lithophyllum byssoides (Corallinales,Rhodophyta)

Lithophyllum byssoides is a common coralline alga in the intertidal zone of Mediterranean coasts, where it produces biogenic concretions housing a high algal and invertebrate biodiversity. This species is an ecosystem engineer and is considered a target for conservation efforts, but designing effective conservation strategies currently is impossible due to lack of information about its population structure. The morphological and molecular variation of L. byssoides was investigated using morphoanatomy and DNA sequences (psbA and cox2,3) obtained from populations at 15 localities on the Italian and Croatian coasts. Lithophyllum byssoides exhibited a high number of haplotypes (31 psbA haplotypes and 24 cox2,3 haplotypes) in the central Mediterranean. The psbA and cox2,3 phylogenies were congruent and showed seven lineages. For most of these clades, the distribution was limited to one or a few localities, but one of them (clade 7) was widespread across the central Mediterranean, spanning the main biogeographic boundaries recognized in this area. The central Mediterranean populations formed a lineage separate from Atlantic samples; psbA pair‐wise divergences suggested that recognition of Atlantic and Mediterranean L. byssoides as different species may be appropriate. The central Mediterranean haplotype patterns of L. byssoides were interpreted as resulting from past climatic events in the hydrogeological history of the Mediterranean Sea. The high haplotype diversity and the restricted spatial distribution of the seven lineages suggest that individual populations should be managed as independent units.     

Unmasking alpha diversity,cladogenesis and biogeographical patterning in an ancient panarthropod lineage (Onychophora: Peripatopsidae: Opisthopatus cinctipes) with the description of five novel species

Speciation and biogeographical patterning in the velvet worm Opisthopatus cinctipes was examined under a null hypothesis that numerous discrete lineages are nested within the species. A total of 184 O. cinctipes specimens, together with a single specimen of each of the two congeneric point endemic sister species (O. roseus and O. herbertorum), were collected throughout the forest archipelago in the Eastern Cape, KwaZulu‐Natal and Mpumalanga provinces of South Africa. All specimens were sequenced for two partial mitochondrial DNA loci (COI and 12S rRNA), while a single specimen from each locality was sequenced for the nuclear 18S rRNA locus. Evolutionary relationships were assessed using maximum‐likelihood and Bayesian inferences, while divergence time estimations were conducted using BEAST. A Bayesian species delimitation approach was undertaken to explore the number of possible novel lineages nested within Opisthopatus, while population genetic structure was examined for the COI locus using ARLEQUIN. Phylogenetic results revealed that O. cinctipes is a species complex comprising seven geographically discrete and statistically well‐supported clades. An independent statistical approach to species delimitations circumscribed ca. 67 species. Results from divergence time estimation and rate constancy tests revealed near constant net diversification occurring throughout the Eocene and Oligocene with subdivision of ranges during the Miocene. Gross morphological characters such as leg pair number within O. cinctipes were invariant, while dorsal and ventral integument colour was highly polymorphic. However, scanning electron microscopy revealed considerable differences both between and within clades. The caveats associated with both morphological and algorithmic delineation of species boundaries are discussed. The five novel Opisthopatus species are described.     

Recent emergence and worldwide spread of the red tomato spider mite, <Emphasis Type="Italic">Tetranychus evansi</Emphasis>: genetic variation and multiple cryptic invasions

Plant biosecurity is increasingly challenged by emerging crop pests. The spider mite Tetranychus evansi has recently emerged as a new threat to solanaceous crops in Africa and the Mediterranean basin, with invasions characterized by a high reproductive output and an ability to withstand a wide range of temperatures. Mitochondrial (868 bp of COI) and nuclear (1,137 bp of ITS) loci were analyzed in T. evansi samples spanning the current geographical distribution to study the earliest stages of the invasive process. The two sets of markers separate the samples into two main clades that are only present together in South America and Southern Europe. The highest COI diversity was found in South America, consistent with the hypothesis of a South American origin of T. evansi. Among the invaded areas, the Mediterranean region displayed a high level of genetic diversity similar to that present in South America, that is likely the result of multiple colonization events. The invasions of Africa and Asia by T. evansi are characterized by a low genetic variation associated with distinct introductions. Genetic data demonstrate two different patterns of invasions: (1) populations in the Mediterranean basin that are a result of multiple cryptic introductions and (2) emerging invasions of Africa and Asia, each likely the result of propagules from one or limited sources. The recent invasions of T. evansi illustrate not only the importance of human activities in the spread of agricultural pests, but also the limits of international quarantine procedures, particularly for cryptic invasions.     

Colonization of the Mediterranean basin by the vector biting midge species Culicoides imicola: an old story

Understanding the demographic history and genetic make‐up of colonizing species is critical for inferring population sources and colonization routes. This is of main interest for designing accurate control measures in areas newly colonized by vector species of economically important pathogens. The biting midge Culicoides imicola is a major vector of orbiviruses to livestock. Historically, the distribution of this species was limited to the Afrotropical region. Entomological surveys first revealed the presence of C. imicola in the south of the Mediterranean basin by the 1970s. Following recurrent reports of massive bluetongue outbreaks since the 1990s, the presence of the species was confirmed in northern areas. In this study, we addressed the chronology and processes of C. imicola colonization in the Mediterranean basin. We characterized the genetic structure of its populations across Mediterranean and African regions using both mitochondrial and nuclear markers, and combined phylogeographical analyses with population genetics and approximate Bayesian computation. We found a west/east genetic differentiation between populations, occurring both within Africa and within the Mediterranean basin. We demonstrated that three of these groups had experienced demographic expansions in the Pleistocene, probably because of climate changes during this period. Finally, we showed that C. imicola could have colonized the Mediterranean basin in the Late Pleistocene or Early Holocene through a single event of introduction; however, we cannot exclude the hypothesis involving two routes of colonization. Thus, the recent bluetongue outbreaks are not linked to C. imicola colonization event, but rather to biological changes in the vector or the virus.     

Salinity appears to be the main factor shaping spatial COI diversity of Corbicula lineages within the Chinese Yangtze River Basin

Aim

To date, few studies have examined the phylogenetics of Corbicula clams in their native range and the environmental parameters influencing their distribution, although this could provide great insights into the biological adaptation and invasion dynamics of Corbicula clams. We sought to identify the genetic lineages of native Corbicula clams and elucidate the environmental factors shaping the distributions of identified lineages.

Location

China, mainly the Yangtze River Basin.

Methods

The alignment comprised 558 COI sequences including samples from China and 222 COI sequences from published studies. This dataset was used to generate phylogenetic trees and compare population diversity. We used dbRDA method to assess the relationship between these COI data and environmental factors measured to identify the important factors affecting Corbicula's distribution.

Results

The COI phylogenetic tree delineated the monophyly of 3 major COI clades and 77 distinct COI haplotypes in the Yangtze River Basin. The invasive lineage C/S (FW17) was not identified in our sampling in China, while invasive lineages A/R (FW5 = Hap6), B (FW1 = Hap17) and Rlc (FW4 = Hap43) were identified, abundant and widely distributed in the middle and lower reaches of Yangtze River. Focusing on populations from Yangtze River Basin, both the COI haplotype and nucleotide diversity in the lakes along the Yangtze River increased with longitude, except for the river mouth population. The AMOVA tests showed significant differentiation between the middle and lower reaches of the Yangtze River and among populations. The dbRDA results suggested that the parameter chloride explained most of the spatial COI haplotype distribution variation in the Yangtze River Basin, with the three invasive lineages tolerating broad fluctuations of salinity (chloride levels ranging from 4 to 60 mg/L).

Conclusions

The Corbicula COI haplotypes found within the Yangtze River Basin had distinct distribution preferences, with the invasive androgenetic lineages being the most abundant and widely distributed. Genetic diversity was higher in this native region than in invaded areas in Europe and America, while it decreased with increasing distance from the river mouth. Salinity appeared to be the main environmental factor shaping the COI haplotype distribution of Corbicula lineages within their native range.     

Phylogeography of the temperate marine bivalve Cerastoderma edule (Linnaeus, 1758) (Bivalvia: Cardiidae) in the Subarctic: Unique diversity and strong population structuring at different spatial scales

Using mitochondrial COI sequencing, we explored the genetic diversity and population structuring of the common cockle Cerastoderma edule (Linnaeus, 1758) in the Norwegian and Barents Seas. Phylogeographic diversity and hence the evolutionary history of C. edule on the Scandinavian and Russian coastlines were found to be richer than expected for populations of temperate species in postglacially colonized seas. A major phylogeographic break at Lofoten Islands separated a group of subarctic populations dominated by a distinct star‐shaped clade of haplotypes from those to the south, extending to the North Sea and having highest gene diversities (h). At the northeastern edge of the range of C. edule, the Russian Murman coast, populations show a mosaic structure with considerable admixture of haplotypes from the south and high local‐scale variation in haplotype diversity (ranging between 0 and 0.8). To explain this mosaic we refer to the core‐satellite metapopulation model, with Norwegian populations as core, and Murman populations as satellites. Our results contradict the conventional biogeographic paradigm implying lack of metapopulation structuring in marine broadcast spawning invertebrates. Hypotheses considered to explain the origin of the unique variation in cockles from Northern Norway involve an early postglacial colonization and establishment of these populations (10–12 ka ago), a persistent oceanographic break at Lofoten, and a mitochondrial selective sweep associated with the postglacial recolonization of the subarctic seas by the boreal C. edule.     

Axon terminals with unusual vesicles in the brain of the polychaete,Ophryotrocha puerilis

Summary In the brain of Ophryotrocha puerilis swollen nerve endings filled with electron-lucent vesicles and aggregates of vesicles were observed. The vesicles do not resemble elementary neurosecretory granules. Tests for biogenic amines were negative; no dense-core vesicles were found. The vesicle type described here cannot be related to any of the types thus far found in nerve cells.Supported by Deutsche Forschungsgemeinschaft Pf116/3     

Comparative phylogeography of two sympatric beeches in subtropical China: Species‐specific geographic mosaic of lineages

In subtropical China, large‐scale phylogeographic comparisons among multiple sympatric plants with similar ecological preferences are scarce, making generalizations about common response to historical events necessarily tentative. A phylogeographic comparison of two sympatric Chinese beeches (Fagus lucida and F. longipetiolata, 21 and 28 populations, respectively) was conducted to test whether they have responded to historical events in a concerted fashion and to determine whether their phylogeographic structure is exclusively due to Quaternary events or it is also associated with pre‐Quaternary events. Twenty‐three haplotypes were recovered for F. lucida and F. longipetiolata (14 each one and five shared). Both species exhibited a species‐specific mosaic distribution of haplotypes, with many of them being range‐restricted and even private to populations. The two beeches had comparable total haplotype diversity but F. lucida had much higher within‐population diversity than F. longipetiolata. Molecular dating showed that the time to most recent common ancestor of all haplotypes was 6.36 Ma, with most haplotypes differentiating during the Quaternary. [Correction added on 14 October 2013, after first online publication: the time unit has been corrected to ‘6.36’.] Our results support a late Miocene origin and southwards colonization of Chinese beeches when the aridity in Central Asia intensified and the monsoon climate began to dominate the East Asia. During the Quaternary, long‐term isolation in subtropical mountains of China coupled with limited gene flow would have lead to the current species‐specific mosaic distribution of lineages.     

Revisiting the phylogeography of Asellus aquaticus in Europe: insights into cryptic diversity and spatiotemporal diversification

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