Schizymenia jonssonii sp. nov. (Nemastomatales,Rhodophyta): a relict or an introduction into the North Atlantic after the last glacial maximum?

North-Atlantic records of Schizymenia dubyi extend along the eastern shores of the North Atlantic from Morocco to southern Britain and Ireland, and the species is also recorded from Iceland. A study was undertaken to confirm the identity of the specimens from Iceland that were geographically separate from the main distribution of S. dubyi and in contrast to other species of the genus did not have gland cells. We analyzed rbcL and COI molecular sequence data from Icelandic specimens and compared the results with those for Schizymenia specimens available in GenBank. For both markers, Schizymenia was shown to be a monophyletic genus. The Icelandic specimens were clearly genetically distinct from S. dubyi and formed a well-supported clade with Schizymenia species from the Northern Pacific. Based on these results, we have described a new species, Schizymenia jonssonii, which can be distinguished by molecular phylogeny, its lack of gland cells and by being strictly intertidal. Crustose tetrasporophytes with identical COI and rbcL sequences were found at the same locations as foliose plants. Schizymenia apoda is reported for the first time in the UK, its identity confirmed by rbcL sequence data. In light of these findings, it is likely that by further molecular analysis of the genus Schizymenia in the north-eastern Atlantic and the Mediterranean, a higher diversity of Schizymenia spp. will be discovered in this region.     

Phylogeography of the Patagonian otter Lontra provocax: adaptive divergence to marine habitat or signature of southern glacial refugia?

Background  

A number of studies have described the extension of ice cover in western Patagonia during the Last Glacial Maximum, providing evidence of a complete cover of terrestrial habitat from 41°S to 56°S and two main refugia, one in south-eastern Tierra del Fuego and the other north of the Chiloé Island. However, recent evidence of high genetic diversity in Patagonian river species suggests the existence of aquatic refugia in this region. Here, we further test this hypothesis based on phylogeographic inferences from a semi-aquatic species that is a top predator of river and marine fauna, the huillín or Southern river otter (Lontra provocax).     

Introduced mangroves in the Society Islands,French Polynesia (South Pacific): invasive species or novel ecosystem?

Biological Invasions - Biotic mixing, where introduced and native species mix into a newly formed ecosystem, is an increasing phenomenon worldwide. It particularly affects islands, where the number...     

Phylogeography and demographic history of the deep-sea fish Aphanopus carbo (Lowe, 1839) in the NE Atlantic: Vicariance followed by secondary contact or speciation?

Comparative phylogeography for the commercially valuable deep-sea fish Aphanopus carbo from a large area of the NE Atlantic revealed remarkable patterns of concordance using two mtDNA markers. Two strongly supported phylogroups were identified from complete sequences of the control region (731-733 bp) and partial sequences of cytochrome b (414 bp) In one of these groups, all sequences from the Mid-Atlantic Ridge (Faraday seamount), mainland Portugal and Madeira were clustered together. The other group constituted all the sequences from the southern coast of Pico island (Azores, central group). The remaining sampling localities had sequences represented in both phylogroups. Although the two clades were strongly differentiated (Phi(ST) = 0.8281 for the CR and Phi(ST) = 0.9083 for the cytb) no evidence for any geographical pattern in this structure, was found. Historical demography of the mitochondrial control region was analysed to clarify the phylogenetic signals embedded in each phylogroup. Mismatch distributions for both clades suggested that both phylogroups were in agreement with sudden expansion models, and both with similar time estimates of expansion (tau = 4.30 and tau = 3.45 for phylogroup one and two, respectively). A molecular clock based on cytb sequences was enforced and dating of divergence for the two phylogroup was 412.5 KY, a time that coincides with geological events that might have caused a split in the original population of black scabbardfish. Once climatic conditions and sea level were restored, the two separate populations came into contact again, leaving traces of the historical events in the non-recombinant mtDNA genes. An alternative hypothesis suggested is that two species of scabbardfish are present. The outcome from the comparison of the same mtDNA regions of the closely related Aphanopus intermedius from Angola clustered with the ones from phylogroup two (from the southern coast of Pico island, Azores). Therefore, these two species may have overlapping distribution ranges and are found sympatrically in the Azores.     

Distribution of Ecteinascidia turbinata (Ascidiacea: Perophoridae) in mangroves of the Yucatán Peninsula, Mexico

The ascidian Ecteinascidia turbinata synthesizes some of the most promising substances against solid-type tumors, but the only available source are the natural populations of this tunicate, which is reared or collected in different parts of the world. A total of 33 locations were sampled in the Gulf of Mexico and the Yucatan Peninsula. The tunicate was not found in Veracruz, Tabasco and Campeche, but it was well established on mangrove roots in the Yucatan Peninsula where we estimated densities more or less equal to one colony and an average production of 115 g of biomass per lineal meter of mangrove coastline in one location (Río Lagartos). Sustainable management appears to be possible.     

Phylogeography of Liquidambar styraciflua (Altingiaceae) in Mesoamerica: survivors of a Neogene widespread temperate forest (or cloud forest) in North America?

We investigate the genetic variation between populations of the American sweetgum (Liquidambar styraciflua), a tree species with a disjunct distribution between northeastern Texas and Mexico, by analyzing sequences of two chloroplast DNA plastid regions in Mesoamerica. Our results revealed phylogeographical structure, with private haplotypes distributed in unique environmental space at either side of the Trans‐Mexican Volcanic Belt, and a split in the absence of gene flow dating back ca. 4.2–1.4 million years ago (MYA). Species distribution modeling results fit a model of refugia along the Gulf and Atlantic coasts but the present ranges of US and Mesoamerican populations persisted disjunct during glacial/interglacial cycles. Divergence between the US and Mesoamerican (ca. 8.4–2.8 MYA) populations of L. styraciflua and asymmetrical gene flow patterns support the hypothesis of a long‐distance dispersal during the Pliocene, with fragmentation since the most recent glacial advance (120,000 years BP) according to coalescent simulations and high effective migration rates from Mesoamerica to the USA and close to zero in the opposite direction. Our findings implicate the Trans‐Mexican Volcanic Belt as a porous barrier driving genetic divergence of L. styraciflua, corresponding with environmental niche differences, during the Pliocene to Quaternary volcanic arc episode 3.6 MYA, and a Mesoamerican origin of populations in the USA.     

Endemic or introduced? Phylogeography of Asparagopsis (Florideophyceae) in Australia reveals multiple introductions and a new mitochondrial lineage

The red seaweed Asparagopsis taxiformis embodies five cryptic mitochondrial lineages (lineage 1–5) introduced worldwide as a consequence of human mediated transport and climate change. We compared globally collected mitochondrial cox2‐3 intergenic spacer sequences with sequences produced from multiple Australian locations and South Korea to identify Asparagopsis lineages and to reveal cryptic introductions. We report A. taxiformis lineage 4 from Cocos (Keeling) Islands, Australia, and the highly invasive Indo‐Pacific Mediterranean lineage 2 from South Korea and Lord Howe Island, Australia. Phylogeographic analysis showed a clear haplotype and geographic separation between western Australian and Great Barrier Reef (GBR) isolates belonging to the recently described lineage 5. The same lineage, however, was characterized by a substantial genetic and geographic break between the majority of Australian specimens and Asparagopsis collections from South Solitary Island, Southern GBR, Lord Howe Island, Kermadec Islands, Norfolk Island, New Caledonia and French Polynesia. The disjunct geographic distribution and sequence divergence between these two groups supports the recognition of a sixth cryptic A. taxiformis mitochondrial lineage. As climatic changes accelerate the relocation of biota and offer novel niches for colonization, periodic surveys for early detection of cryptic invasive seaweeds will be critical in determining whether eradication or effective containment of the aliens are feasible.     

Is the incidence of survival in interior Pleistocene refugia (nunataks) underestimated? Phylogeography of the high mountain plant Androsace alpina (Primulaceae) in the European Alps revisited

Temperate mountain ranges such as the European Alps have been strongly affected by the Pleistocene glaciations. Glacial advances forced biota into refugia, which were situated either at the periphery of mountain ranges or in their interior. Whereas in the Alps peripheral refugia have been repeatedly and congruently identified, support for the latter scenario, termed “nunatak hypothesis,” is still limited and no general pattern is recognizable yet. Here, we test the hypothesis of nunatak survival for species growing in the high alpine to subnival zones on siliceous substrate using the cushion plant Androsace alpina (Primulaceae), endemic to the European Alps, as our model species. To this end, we analyzed AFLP and plastid DNA sequence data obtained from a dense and range‐wide sampling. Both AFLPs and plastid sequence data identified the southwestern‐most population as the most divergent one. AFLP data did not allow for discrimination of interior and peripheral populations, but rather identified two to three longitudinally separated major gene pools. In contrast, in the eastern half of the Alps several plastid haplotypes of regional or local distribution in interior ranges—the Alpine periphery mostly harbored a widespread haplotype—were indicative for the presence of interior refugia. Together with evidence from other Alpine plant species, this study shows that in the eastern Alps silicicolous species of open habitats in the alpine and subnival zone survived, also or exclusively so, in interior refugia. As the corresponding genetic structure may be lost in mostly nuclear‐derived, rapidly homogenizing marker systems such as AFLPs or RAD sequencing tags, markers not prone to homogenization, as is the case for plastid sequences (Sanger‐sequenced or extracted from an NGS data set) will continue to be important for detecting older, biogeographically relevant patterns.     

Immunohistochemical analysis of adhesive papillae of Clavelina lepadiformis (Müller, 1776) and Clavelina phlegraea (Salfi, 1929) (Tunicata,Ascidiacea)

Almost all ascidian larvae bear three mucus secreting and sensory organs, the adhesive papillae, at the anterior end of the trunk, which play an important role during the settlement phase. The morphology and the cellular composition of these organs varies greatly in the different species. The larvae of the Clavelina genus bear simple bulbous papillae, which are considered to have only a secretory function. We analysed the adhesive papillae of two species belonging to this genus, C. lepadiformis and C. phlegraea, by histological sections and by immunolocalisation of β-tubulin and serotonin, in order to better clarify the cellular composition of these organs. We demonstrated that they contain at least two types of neurons: central neurons, bearing microvilli, and peripheral ciliated neurons. Peripheral neurons of C. lepadiformis contain serotonin. We suggest that these two neurons play different roles during settlement: the central ones may be chemo- or mechanoreceptors that sense the substratum, and the peripheral ones may be involved in the mechanism that triggers metamorphosis.Key words: Settlement, neurotransmitter, serotonin, β-tubulin, papillary nerves, metamorphosis.Ascidians (phylum Chordata; subphylum Tunicata) are sessile filter-feeding organisms that can be found in all benthic marine environments and develop through a swimming tadpole larva. Larvae of colonial ascidians have a short planktonic life that can vary from minutes to hours (Burighel and Cloney, 1997). Prior to metamorphosis, the larva attaches to the substratum by means of peculiar organs of ectodermic origin, located in the anterior region of the trunk. These organs, known as adhesive papillae, secrete sticky substances and effect primary adhesion of the larva to the substrate (Cloney, 1977). They have an important role in the initiation of settlement and metamorphosis and there is evidence that, at least in some species, they participate in substrate selection (Torrence and Cloney, 1983; Svane and Young, 1989; Groppelli et al., 2003). In many species, they are organised in a triangular field, whereas in others they are aligned along the mid-sagittal plane of the trunk. Adhesive papillae have been classified into two types: eversible papillae, typical of some colonial species, composed by several cell types and rapidly changing shape as they touch the substrate, and non-eversible papillae, typical of solitary species, which do not change shape after settlement (Burighel and Cloney, 1997).With few exceptions, all adhesive papillae are formed by elongated secretory and sensory cells, which are recognised as primary neurons (Cloney, 1977, 1979). It has been proposed that sensory cells may detect the chemical and physical characteristics of the substratum at potential sites for settlement and metamorphosis (Young and Braithwaite, 1980; Groppelli et al., 2003).The presence of primary neurons in the papillae has been reported in the larvae of several species such as Distaplia occidentalis, Diplosoma macdonaldi, Phallusia mammillata, Ciona intestinalis and Ascidia malaca (Cloney, 1977;Torrence and Cloney, 1983; Sotgia et al., 1998; Takamura, 1998; Gianguzza et al., 1999).These neurons have axons that join together to form the papillary nerves that enter the central nervous system at the level of the sensory vesicle (Imai and Meinertzhagen, 2007).Recently, different neurotransmitters have been localised in the sensory neurons of the papillae of different species. The presence of GABAergic neurons has been reported in the papillae of Ciona savygni (Brown et al., 2005) and of Ciona intestinalis (Zega et al., 2008), while serotonergic neurons have been localised in the papillae of Phallusia mammillata (Pennati et al., 2001) and Botrylloides leachi (Pennati et al., 2007). Moreover, it has been demonstrated that serotonin plays a role in the mechanism triggering metamorphosis in ascidians (Zega et al., 2005).After attachment, all papillae retract to draw the larva closer to the substratum. In the colonial ascidian Distaplia occidentalis, the process of retraction is reversibly inhibited by cytochalasin B, suggesting that microfilaments are involved in this process (Cloney, 1979).Clavelina lepadiformis is a colonial species, whose larvae bear non-eversible simple bulbous papillae.They have been described as being formed only by columnar glandular cells, whose secreted material is responsible for the sticky properties of the organ. These papillae do not contain sensory cells and were considered the simplest among those studied (Turon, 1991).In this work, the morphology of the adhesive papillae of C. lepadiformis and C. phlegraea was further investigated by histological analysis and immunolabelling techniques in order to clarify the actual cellular composition and function of these organs.     

Phylogeography of the Qinghai–Tibet Plateau endemic alpine herb Pomatosace filicula (Primulaceae)

In order to trace the response of alpine plants on the Qinghai–Tibet Plateau (QTP) to the Quaternary climate oscillations, the phylogeographic history of Pomatosace filicula Maxim. was investigated in the present study. Based on sequence variations of two maternally inherited plastid markers, matK and trnH-psbA, and the biparentally inherited nuclear ribosomal internal transcribed spacer (nrITS), we estimated the population genetic structure, lineage divergence timescale, and population dynamics of P. filicula. Seven plastid haplotypes and two nrITS genotypes were identified across the range-wide sampling of 200 individuals from 24 populations. Although AMOVA revealed a high level of differentiation among populations (F_ST = 0.560), no significant phylogeographic structure was detected (N_ST = 0.503, G_ST = 0.518, P > 0.05). Molecular dating suggested that the divergences between major plastid lineages and nrITS genotypes occurred during the early and middle Pleistocene, and the middle Pleistocene, respectively. This species most likely survived at multiple unglaciated sites on the QTP during the Last Glacial Maximum, with most of these sites located above 4000 m a.s.l. The species probably experienced range expansion at its distribution fringe, but demographic tests did not suggest significant population size changes. We proposed that pronounced effective gene flow (N_em = 0.393) and short generation time may have obscured the phylogeographic and demographic patterns of this species. Our findings will shed new light on the Quaternary evolutionary history of the alpine flora of the QTP.     

What are the closest relatives of the hot-spring snakes (Colubridae, Thermophis), the relict species endemic to the Tibetan Plateau?

We conducted phylogenetic analyses to identify the closest related living relatives of the Xizang and Sichuan hot-spring snakes (T. baileyi and T. zhaoermii) endemic to the Tibetan Plateau, using mitochondrial DNA sequences (cyt b, ND4) from eight specimens, together with sequences from 95 additional caenophidian and five henophidian genera that were downloaded from GenBank. Phylogenetic trees were obtained using Bayesian Inference and Maximum likelihood methods. Results suggest that hot-spring snakes, which are adapted to high and cold environments, were clustered in the monophyletic Xenodontinae. Xenodontinae is one of the largest subfamilies of colubrid snakes, with about 90 genera and more than 500 species known, and are primarily tropical snakes previously thought to be restricted to the New World. Our data failed to provide any evidence that the New World xenodontines diverged from Thermophis and dispersed into the New World, also failed to suggest a colonization of Asia by New World xenodontines by dispersal from the New World. An alternative plausible scenario may be that Thermophis and the New World xenodontines evolved independently in Asia and America, respectively, after the divergence of their common ancestor. The divergence of the two species in Thermophis was caused by the barrier of the Hengduan Mountains, and the speciation had almost occurred when Tibetan Plateau attained present elevation.     

Have the Harmful Effects of Introduced Rats on Islands been Exaggerated?

Introduced rats are now being eradicated from many islands. Increasingly, these eradications are contested by activists claiming moral, legal, cultural, historic or scientific reasons and poorly documented evidence of effects. We reviewed the global literature on the effects of rats on island flora and fauna. We then used New Zealand as a case study because of its four-decade history of rat eradications and many detailed and innovative studies of how rats affect native species. These include use of exclosures, local manipulations of rat populations, video surveillance, and measurements of responses following eradications. The most intensive studies have been on the Pacific rat (Rattus exulans), a small South-East Asian species spread by Polynesians throughout the Pacific. These and the more recently introduced Norway rat (R. norvegicus) and ship (roof) rat (R. rattus) suppress some forest plants, and are associated with extinctions or declines of flightless invertebrates, ground-dwelling reptiles, land birds, and burrowing seabirds. On islands off France, Norway rats are also implicated in declines of shrews. Globally, ship rats were associated with declines or extinctions of the largest number of indigenous vertebrate species (60), including small mammals such as deer mice and bats. Effects of rats on forest trees and seabird populations are sufficiently pervasive to affect ecosystem structure and function. However, the data are patchy. Deficiencies in our knowledge would be reduced by documenting distribution and abundance of indigenous species before and after eradications. Comprehensive measurements of the responses of indigenous species to rat eradications would enable the development of testable models of rat invasion effects.     

Phylogeography of Schizothorax o’connori (Cyprinidae: Schizothoracinae) in the Yarlung Tsangpo River, Tibet

Schizothorax o’connori is endemic to the Yarlung Tsangpo River on the Tibetan Plateau. We assessed the relative impacts of historical and contemporary factors in organizing genetic variation in S. o’connori populations using mitochondrial cytochrome b sequences. We analyzed 191 samples from 11 populations and identified 78 haplotypes. The phylogenetic analyses and analysis of molecular variance all supported the same conclusions of two well-differentiated east–west phylogroups, separated by the Tsangpo Great Gorge. The split between the two clades accounted for 58% of the genetic variance observed among the examined samples. Waterfalls as effective barriers played an important role in shaping the phylogeographical structure of this species. Analyses of migration rates revealed that upstream dispersal was limited crossing waterfalls. Our study revealed substantial spatial and temporal variation in the influence of landscape features on contemporary patterns of genetic structure in S. o’connori. Interglacial range expansions clearly left their mark on contemporary populations above the Tsangpo Great Gorge.     

Physical Restraint or Stimulation? The Function(s) of the Modified Front Legs of Male Archisepsis diversiformis (Diptera, Sepsidae)

Four hypotheses that could explain the elaborate species-specific morphology of the clasping organs on the front legs of male Archisepsis diversiformis flies were tested: direct male–male combat, mechanical fit, male–female conflict of interests, and male stimulation of the female. Experimental modification of the shape of the male clasper and of the female wing base where the male clasped the female both strongly reduced the chances that a mount would result in copulation. This reduction was not predicted by the male–male combat hypothesis but was predicted by the others. Males in the field did nave to fight other males to remain mounted on females, as expected by the male–male combat hypothesis. Reduced male copulatory success was not due to inferior male ability to grasp and hold onto the female's wings, as predicted by the mechanical fit and male–female conflict hypotheses but to a reduction in the likelihood that the female would allow intromission, as predicted by the stimulation hypothesis. By a process of elimination, and in combination with data from a previous morphological study, the data support the hypothesis that the species-specific aspects of grasping organs in these flies function to stimulate females. Further behavioral data will be needed to test alternative possibilities.     

Establishment of a coastal fish in the Azores: recent colonisation or sudden expansion of an ancient relict population?

The processes and timescales associated with ocean-wide changes in the distribution of marine species have intrigued biologists since Darwin''s earliest insights into biogeography. The Azores, a mid-Atlantic volcanic archipelago located >1000 km off the European continental shelf, offers ideal opportunities to investigate phylogeographic colonisation scenarios. The benthopelagic sparid fish known as the common two-banded seabream (Diplodus vulgaris) is now relatively common along the coastline of the Azores archipelago, but was virtually absent before the 1990s. We employed a multiple genetic marker approach to test whether the successful establishment of the Azorean population derives from a recent colonisation from western continental/island populations or from the demographic explosion of an ancient relict population. Results from nuclear and mtDNA sequences show that all Atlantic and Mediterranean populations belong to the same phylogroup, though microsatellite data indicate significant genetic divergence between the Azorean sample and all other locations, as well as among Macaronesian, western Iberian and Mediterranean regions. The results from Approximate Bayesian Computation indicate that D. vulgaris has likely inhabited the Azores for ∼40 (95% confidence interval (CI): 5.5–83.6) to 52 (95% CI: 6.32–89.0) generations, corresponding to roughly 80–150 years, suggesting near-contemporary colonisation, followed by a more recent demographic expansion that could have been facilitated by changing climate conditions. Moreover, the lack of previous records of this species over the past century, together with the absence of lineage separation and the presence of relatively few private alleles, do not exclude the possibility of an even more recent colonisation event.     

Origin of the parasites of an invading species,the Australian cane toad (Bufo marinus): are the lungworms Australian or American?

Phylogeographical analyses that identify the geographical origin of parasites in invading species can clarify the parasites' potential for biological control of the invader and the risks posed by the parasite to native species. Our data on nuclear and mitochondrial genetic sequences show that the nematode lungworms (Rhabdias spp.) in invasive Australian populations of cane toads (Bufo marinus) are Rhabdias pseudosphaerocephala, a South American species. We did not find this lungworm species in any Australian frogs sympatric with cane toads, suggesting that the parasite does not attack Australian frogs and hence may offer potential as a biocontrol agent of the toad.     

Occurrence of species of the genus Pityophthorus Eichhoff (Coleoptera,Curculionidae, Scolytinae) in?the?province of Quebec,Canada

Twig beetles in the genus Pityophthorus Eichhoff, 1864 include more than 300 species worldwide, with maximum diversity in tropical and subtropical regions. To date, approximately 50 species of Pityophthorus have been recorded in Canada, and these species are associated mainly with coniferous trees. Since 1981, no comprehensive study on this difficult taxonomic group has been conducted in Quebec, Canada, most likely due to their limited significance as forest pests. Based on data gathered from five years of field sampling in conifer seed orchards and compiled from various entomological collections, the distribution of Pityophthorus species in Quebec is presented. Approximately 291 new localities were recorded for the Pityophthorus species. Five species-group taxa, namely Pityophthorus puberulus (LeConte, 1868), Pityophthorus pulchellus pulchellus Eichhoff, 1869, Pityophthorus pulicarius (Zimmermann, 1868), Pityophthorus nitidus Swaine, 1917,and Pityophthorus cariniceps LeConte&Horn, 1876 were the most widespread. In contrast, Pityophthorus consimilis LeConte, 1878, Pityophthorus intextus Swaine, 1917, Pityophthorus dentifrons Blackman, 1922, Pityophthorus ramiperda Swaine, 1917, and Pityophthorus concavus Blackman, 1928 display a notably limited distribution. In addition, the first distribution records of Pityophthorus intextus and Pityophthorus biovalis Blackman, 1922 are furnished, and the subspecies Pityophthorus murrayanae murrayanae Blackman, 1922is reported from Quebec for the second time. Moreover, distribution maps are provided for all Pityophthorus species recorded in the province of Quebec.     

Phylogeography of an alpine aquatic herb Ranunculus bungei (Ranunculaceae) on the Qinghai–Tibet Plateau

Many phylogeographic studies of terrestrial plant species on the Qinghai–Tibet Plateau (QTP) have been carried out to elucidate the range shifts in response to climatic oscillations in the Quaternary. However, patterns of range shifts for aquatic plants following the climate change on the QTP are poorly understood. Here, we studied the historical range shifts of the aquatic herb Ranunculus bungei Steud. on the QTP using four chloroplast (cpDNA) non-coding spacers. We revealed low within-population genetic diversity (H_S = 0.052) and high interpopulation genetic differentiation (G_ST = 0.914; N_ST = 0.954). But the high population differentiation was not coupled with a distinct phylogeographic structure (N_ST > G_ST, P > 0.05). Phylogenetic analyses revealed two main cpDNA haplotype lineages and the split between these two lineages can be dated back to the late Tertiary (3.84–11.90 Ma). Two independent range expansions within the two intraspecific lineages at approximately 0.15–0.46 and 0.17–0.50 Ma were revealed. Our results suggested that R. bungei survived the Last Glacial Maximum and/or previous glacial periods on the QTP. Colonization or recolonization during the repeated range expansions may have replaced the early haplotypes and the pre-existing genetic structure and could explain the non-significant phylogeographical structure.