Genetic variation within and among populations of Rhodiola alsia (Crassulaceae) native to the Tibetan Plateau as detected by ISSR markers

Genetic variation of 10 Rhodiola alsia ( Crassulaceae) populations from the Qinghai - Tibet Plateau of China was investigated using intersimple sequence repeat (ISSR) markers. R. alsia is an endemic species of the Qinghai - Tibet Plateau. Of the 100 primers screened, 13 were highly polymorphic. Using these primers, 140 discernible DNA fragments were generated with 112 (80%) being polymorphic, indicating pronounced genetic variation at the species level. Also there were high levels of polymorphism at the population level with the percentage of polymorphic bands (PPB) ranging from 63.4 to 88.6%. Analysis of molecular variance (AMOVA) showed that the genetic variation was mainly found among populations (70.3%) and variance within populations was 29.7%. The main factors responsible for the high level of differentiation among populations are probably the isolation from other populations and clonal propagation of this species. Occasional sexual reproduction might occur in order to maintain high levels of variation within populations. Environmental conditions could also influence population genetic structure as they occur in severe habitats. The strong genetic differentiation among populations in our study indicates that the conservation of genetic variability in R. alsia requires maintenance of as many populations as possible.

Sexual selection and canine dimorphism in New World monkeys.

Social and ecological factors are important in shaping sexual dimorphism in Anthropoidea, but there is also a tendency for body-size dimorphism and canine dimorphism to increase with increased body size (Rensch's rule) (Rensch: Evolution Above the Species Level. London: Methuen, 1959.) Most ecologist interpret Rensch's rule to be a consequence of social and ecological selective factors that covary with body size, but recent claims have been advanced that dimorphism is principally a consequence of selection for increased body size alone. Here we assess the effects of body size, body-size dimorphism, and social structure on canine dimorphism among platyrrhine monkeys. Platyrrhine species examined are classified into four behavioral groups reflecting the intensity of intermale competition for access to females or to limiting resources. As canine dimorphism increases, so does the level of intermale competition. Those species with monogamous and polyandrous social structures have the lowest canine dimorphism, while those with dominance rank hierarchies of males have the most canine dimorphism. Species with fission-fusion social structures and transitory intermale breeding-season competition fall between these extremes. Among platyrrhines there is a significant positive correlation between body size and canine dimorphism However, within levels of competition, no significant correlation was found between the two. Also, with increased body size, body-size dimorphism tends to increase, and this correlation holds in some cases within competition levels. In an analysis of covariance, once the level of intermale competition is controlled for, neither molar size nor molar-size dimorphism accounts for a significant part of the variance in canine dimorphism. A similar analysis using body weight as a measure of size and dimorphism yields a less clear-cut picture: body weight contributes significantly to the model when the effects of the other factors are controlled. Finally, in a model using head and body length as a measure of size and dimorphism, all factors and the interactions between them are significant. We conclude that intermale competition among platyrrhine species is the most important factor explaining variations in canine dimorphism. The significant effects of size and size dimorphism in some models may be evidence that natural (as opposed to sexual) selection also plays a role in the evolution of increased canine dimorphism.

Relative growth of the limbs and trunk in sifakas: heterochronic, ecological, and functional considerations.

Limb, trunk, and body weight measurements were obtained for growth series of Milne-Edwards's diademed sifaka, Propithecus diadema edwardsi, and the golden-crowned sifaka, Propithecus tattersalli. Similar measures were obtained also for primarily adults of two subspecies of the western sifaka: Propithecus verreauxi coquereli, Coquerel's sifaka, and Propithecus verreauxi verreauxi, Verreaux's sifaka. Ontogenetic series for the larger-bodied P. d. edwardsi and the smaller-bodied P. tattersalli were compared to evaluate whether species-level differences in body proportions result from the differential extension of common patterns of relative growth. In bivariate plots, both subspecies of P. verreauxi were included to examine whether these taxa also lie along a growth trajectory common to all sifakas. Analyses of the data indicate that postcranial proportions for sifakas are ontogenetically scaled, much as demonstrated previously with cranial dimensions for all three species (Ravosa, 1992). As such, P. d. edwardsi apparently develops larger overall size primarily by growing at a faster rate, but not for a longer duration of time, than P. tattersalli and P. verreauxi; this is similar to results based on cranial data. A consideration of Malagasy lemur ecology suggests that regional differences in forage quality and resource availability have strongly influenced the evolutionary development of body-size variation in sifakas. On one hand, the rainforest environment of P. d. edwardsi imposes greater selective pressures for larger body size than the dry-forest environment of P. tattersalli and P. v. coquereli, or the semi-arid climate of P. v. verreauxi. On the other hand, as progressively smaller-bodied adult sifakas are located in the east, west, and northwest, this apparently supports suggestions that adult body size is set by dry-season constraints on food quality and distribution (i.e., smaller taxa are located in more seasonal habitats such as the west and northeast). Moreover, the fact that body-size differentiation occurs primarily via differences in growth rate is also due apparently to differences in resource seasonality (and juvenile mortality risk in turn) between the eastern rainforest and the more temperate northeast and west. Most scaling coefficients for both arm and leg growth range from slight negative allometry to slight positive allometry. Given the low intermembral index for sifakas, which is also an adaptation for propulsive hindlimb-dominated jumping, this suggests that differences in adult limb proportions are largely set prenatally rather than being achieved via higher rates of postnatal hindlimb growth.(ABSTRACT TRUNCATED AT 400 WORDS)

Gene loss, adaptive evolution and the co-evolution of plumage coloration genes with opsins in birds.

BACKGROUND: The wide range of complex photic systems observed in birds exemplifies one of their key evolutionary adaptions, a well-developed visual system. However, genomic approaches have yet to be used to disentangle the evolutionary mechanisms that govern evolution of avian visual systems. RESULTS: We performed comparative genomic analyses across 48 avian genomes that span extant bird phylogenetic diversity to assess evolutionary changes in the 17 representatives of the opsin gene family and five plumage coloration genes. Our analyses suggest modern birds have maintained a repertoire of up to 15 opsins. Synteny analyses indicate that PARA and PARIE pineal opsins were lost, probably in conjunction with the degeneration of the parietal organ. Eleven of the 15 avian opsins evolved in a non-neutral pattern, confirming the adaptive importance of vision in birds. Visual conopsins sw1, sw2 and lw evolved under negative selection, while the dim-light RH1 photopigment diversified. The evolutionary patterns of sw1 and of violet/ultraviolet sensitivity in birds suggest that avian ancestors had violet-sensitive vision. Additionally, we demonstrate an adaptive association between the RH2 opsin and the MC1R plumage color gene, suggesting that plumage coloration has been photic mediated. At the intra-avian level we observed some unique adaptive patterns. For example, barn owl showed early signs of pseudogenization in RH2, perhaps in response to nocturnal behavior, and penguins had amino acid deletions in RH2 sites responsible for the red shift and retinal binding. These patterns in the barn owl and penguins were convergent with adaptive strategies in nocturnal and aquatic mammals, respectively. CONCLUSIONS: We conclude that birds have evolved diverse opsin adaptations through gene loss, adaptive selection and coevolution with plumage coloration, and that differentiated selective patterns at the species level suggest novel photic pressures to influence evolutionary patterns of more-recent lineages.

Variation of phytoplankton assemblages along the Mozambique coast as revealed by HPLC and microscopy

This study is an integrated overview of pigment and microscopic analysis of phytoplankton communities throughout the Mozambican coast. Collected samples revealed notable patterns of phytoplankton occurrence and distribution, with community structure changing between regions and sample depth. Pigment data showed Delagoa Bight, Sofala Bank and Angoche as the most productive regions throughout the sampled area. In general, micro-sized phytoplankton, particularly diatoms, were important contributors to biomass both at surface and sub-surface maximum (SSM) samples, although were almost absent in the northern stations. In contrast, nano- and pico-sized phytoplankton revealed opposing patterns. Picophytoplankton were most abundant at surface, as opposed to nanophytoplankton, which were more abundant at the SSM. Microphytoplankton were associated with cooler southern water masses, while picophytoplankton were related to warmer northern water masses. Nanophytoplankton were found to increase their contribution to biomass with increasing SSM. Microscopy information on the genera and species level revealed the diatoms Chaetoceros spp., Proboscia alata, Pseudo-nitzschia spp., Cylindrotheca closterium and Hemiaulus haukii as the most abundant taxa of the micro-sized phytoplankton. Discosphaera tubifera and Emiliania huxleyi were the most abundant coccolithophores, nano-sized phytoplankton.

Southern Ocean biogeography and taxonomic resolution: what's in the name?

We examined the taxonomic resolution of zooplankton data required to identify ocean basin scale biogeographic zonation in the Southern Ocean. A 2,154 km transect was completed south of Australia. Sea surface temperature (SST) measured at 1 min intervals showed that seven physical zones were sampled. Zooplankton were collected at a spatial resolution of similar to 9.2 km with a continuous plankton recorder, identified to the highest possible taxonomic resolution and enumerated. Zooplankton assemblage similarity between samples was calculated using the Bray-Curtis index for the taxonomic levels of species, genus, family, order and class after first log(10)(x + 1) (LA) and then presence/absence (PA) transformation of abundance data. Although within and between zone sample similarity increased with decreasing taxonomic resolution, for both data transformations, cluster analysis demonstrated that the biogeographic separation of zones remained at all taxonomic levels when using LA data. ANOSIM confirmed this, detecting significant differences in zooplankton assemblage structure between all seven a priori determined physical zones for all taxonomic levels when using the LA data. In the case of the PA data for the complete data set, and both LA and PA data for a crustacean only data set, no significant differences were detected between zooplankton assemblages in the Polar frontal zone (PFZ) and inter-PFZ at any taxonomic level. Loss of information at resolutions below the species level, particularly in the PA data, prevented the separation of some zones. However, the majority of physical zones were biogeographically distinct from species level to class using both LA and PA transformations. Significant relationships between SST and zooplankton community structure, summarised as NMDS scores, at all taxonomic levels, for both LA and PA transformations, and complete and crustacean only data sets, highlighted the biogeographic relevance of low resolution taxonomic data. The retention of biogeographic information in low taxonomic resolution data shows that data sets collected with different taxonomic resolutions may be meaningfully merged for the post hoc generation of Southern Ocean time series.

Have we been underestimating the effects of ocean acidification in zooplankton?

Understanding how copepods may respond to ocean acidification (OA) is critical for risk assessments of ocean ecology and biogeochemistry. The perception that copepods are insensitive to OA is largely based on experiments with adult females. Their apparent resilience to increased carbon dioxide (pCO2 ) concentrations has supported the view that copepods are 'winners' under OA. Here, we show that this conclusion is not robust, that sensitivity across different life stages is significantly misrepresented by studies solely using adult females. Stage-specific responses to pCO2 (385-6000 μatm) were studied across different life stages of a calanoid copepod, monitoring for lethal and sublethal responses. Mortality rates varied significantly across the different life stages, with nauplii showing the highest lethal effects; nauplii mortality rates increased threefold when pCO2 concentrations reached 1000 μatm (year 2100 scenario) with LC50 at 1084 μatm pCO2 . In comparison, eggs, early copepodite stages, and adult males and females were not affected lethally until pCO2 concentrations ≥3000 μatm. Adverse effects on reproduction were found, with >35% decline in nauplii recruitment at 1000 μatm pCO2 . This suppression of reproductive scope, coupled with the decreased survival of early stage progeny at this pCO2 concentration, has clear potential to damage population growth dynamics in this species. The disparity in responses seen across the different developmental stages emphasizes the need for a holistic life-cycle approach to make species-level projections to climate change. Significant misrepresentation and error propagation can develop from studies which attempt to project outcomes to future OA conditions solely based on single life history stage exposures.

Simple shade plots aid better long-term choices of data pre-treatment in multivariate assemblage studies

Shade plots, simple visual representations of abundance matrices from multivariate species assemblage studies, are shown to be an effective aid in choosing an overall transformation (or other pre-treatment) of quantitative data for long-term use, striking an appropriate balance between dominant and less abundant taxa in ensuing resemblance-based multivariate analyses. Though the exposition is entirely general and applicable to all community studies, detailed illustrations of the comparative power and interpretative possibilities of shade plots are given in the case of two estuarine assemblage studies in south-western Australia: (a) macrobenthos in the upper Swan Estuary over a two-year period covering a highly significant precipitation event for the Perth area; and (b) a wide-scale spatial study of the nearshore fish fauna from five divergent estuaries. The utility of transformations of intermediate severity is again demonstrated and, with greater novelty, the potential importance seen of further mild transformation of all data after differential down-weighting (dispersion weighting) of spatially clumped' or schooled' species. Among the new techniques utilized is a two-way form of the RELATE test, which demonstrates linking of assemblage structure (fish) to continuous environmental variables (water quality), having removed a categorical factor (estuary differences). Re-orderings of sample and species axes in the associated shade plots are seen to provide transparent explanations at the species level for such continuous multivariate patterns.

Next Generation Sequencing Reveals the Hidden Diversity of Zooplankton Assemblages

Zooplankton play an important role in our oceans, in biogeochemical cycling and providing a food source for commercially important fish larvae. However, difficulties in correctly identifying zooplankton hinder our understanding of their roles in marine ecosystem functioning, and can prevent detection of long term changes in their community structure. The advent of massively parallel next generation sequencing technology allows DNA sequence data to be recovered directly from whole community samples. Here we assess the ability of such sequencing to quantify richness and diversity of a mixed zooplankton assemblage from a productive time series site in the Western English Channel. Methodology/Principle Findings Plankton net hauls (200 µm) were taken at the Western Channel Observatory station L4 in September 2010 and January 2011. These samples were analysed by microscopy and metagenetic analysis of the 18S nuclear small subunit ribosomal RNA gene using the 454 pyrosequencing platform. Following quality control a total of 419,041 sequences were obtained for all samples. The sequences clustered into 205 operational taxonomic units using a 97% similarity cut-off. Allocation of taxonomy by comparison with the National Centre for Biotechnology Information database identified 135 OTUs to species level, 11 to genus level and 1 to order, <2.5% of sequences were classified as unknowns. By comparison a skilled microscopic analyst was able to routinely enumerate only 58 taxonomic groups. Conclusions Metagenetics reveals a previously hidden taxonomic richness, especially for Copepoda and hard-to-identify meroplankton such as Bivalvia, Gastropoda and Polychaeta. It also reveals rare species and parasites. We conclude that Next Generation Sequencing of 18S amplicons is a powerful tool for elucidating the true diversity and species richness of zooplankton communities. While this approach allows for broad diversity assessments of plankton it may become increasingly attractive in future if sequence reference libraries of accurately identified individuals are better populated.

Lamellibrachia anaximandrin. sp., a new vestimentiferan tubeworm (Annelida) from the Mediterranean, with notes on frenulate tubeworms from the same habitat

A new species of lamellibrachiid vestimentiferan, Lamellibrachia anaximandri n. sp., has been found in the Eastern Mediterranean, close to cold seeps of fluid carrying dissolved methane and sources of sulfide in superficial sediments. It occurs at about 1100 to 2100 m depth, on some of the mud volcanoes on the Anaximander Mountains, south of Turkey, on the Mediterranean Ridge, south of Crete, and on the Nile deep-sea fan. In addition, it has been obtained from rotting paper inside a sunken ship, torpedoed in 1915 and lying at 2800 m depth, southeast of Crete. Some frenulate pogonophores also occur on the mud volcanoes (including a species of Siboglinum resembling S. carpinei and tubes of other unidentified genera). The new Lamellibrachia is the first vestimentiferan species to be described from the Mediterranean. It differs from L. luymesi taken from the Gulf of Mexico population in the very weak development of collars on its tube and in having a smaller number of pairs of branchial lamellae in the branchial plume. Sequencing of the COI and the mt16S genes confirms a difference at the species level between the new species and L. luymesi, and a difference between these two species and four described species of Lamellibrachia from the Pacific Ocean. The largest individuals of L. anaximandri n. sp. may be many years old, but there are numerous young individuals at some sites, showing that favourable conditions are available for settlement and early growth. The development of the branchial plume in a series of young stages reveals that the sheath lamellae, which are characteristic of the genus Lamellibrachia, begin to form only after the establishment of several pairs of branchial lamellae. Examination of the adult trophosome by transmission electron microscopy shows Gram-negative bacteria without internal stacked membranes, indicating that the symbionts are most probably sulfide oxidizing.

Molecular approaches to the study of invasive seaweeds

A wide range of vectors is currently introducing a plethora of alien marine species into indigenous marine species assemblages. Over the past two decades, molecular studies of non-native seaweeds, including cryptic invaders, have successfully identified the species involved and their sources; we briefly review these studies. As yet, however, little research has been directed towards examining the genetic consequences of seaweed invasions. Here we provide an overview of seaweed invasions from a genetic perspective, focusing on invader species for which the greatest amount of information is available. We review invasion processes, and rationalize evolutionary and genetic consequences for the indigenous and invader species into two main groups: (1) changes in gene-pool composition, in population structure and allele frequencies; and (2) changes in genome organization at the species level through hybridization, and in individual gene expression profiles at the levels of expressed messenger RNA and the proteome (i.e., all proteins synthesized) and thus the phenotype. We draw on studies of better-known aquatic and terrestrial organisms to point the way forward in revealing the genetic consequences of seaweed invasions. We also highlight potential applications of more recent methodological and statistical approaches, such as microarray technology, assignment tests and mixed stock analysis.

Extinction risk from climate change

Climate change over the past ,30 years has produced numerous shifts in the distributions and abundances of species1,2 and has been implicated in one species-level extinction3. Using projections of species’ distributions for future climate scenarios, we assess extinction risks for sample regions that cover some 20% of the Earth’s terrestrial surface. Exploring three approaches in which the estimated probability of extinction shows a powerlaw relationship with geographical range size, we predict, on the basis of mid-range climate-warming scenarios for 2050, that 15–37% of species in our sample of regions and taxa will be ‘committed to extinction’. When the average of the three methods and two dispersal scenarios is taken, minimal climate-warming scenarios produce lower projections of species committed to extinction (,18%) than mid-range (,24%) and maximum change (,35%) scenarios. These estimates show the importance of rapid implementation of technologies to decrease greenhouse

Estimating trophic link density from quantitative but incomplete diet data

The trophic link density and the stability of food webs are thought to be related, but the nature of this relation is controversial. This article introduces a method for estimating the link density from diet tables which do not cover the complete food web and do not resolve all diet items to species level. A simple formula for the error of this estimate is derived. Link density is determined as a function of a threshold diet fraction below which diet items are ignored (

Ceramium Roth (Ceramiales, Rhodophyta) from Venice lagoon (Adriatic Sea, Italy): Comparative studies of Mediterranean and Atlantic taxa

The cosmopolitan genus Ceramium (Ceramiaceae, Rhodophyta) is a large and systematically complex group. The taxonomy of this genus remains in a chaotic state due to the high degree of morphological variation. Culture studies, suggesting a strong influence of environment on phenotype, and the use of molecular tools have recently questioned the validity of morphological features used in species recognition. Here we compare three Ceramium taxa from Venice lagoon with samples from northwest Europe using the plastid ribulose-1,5-bisphosphate carboxylase/oxygenase gene (rbcL) and the rbcL-rbcS intergenic spacer combined with morphological observations. A strongly banded species, previously identified as member of a poorly understood and misnamed group, the Ceramium diaphanum complex sensu Feldmann-Mazoyer, is probably conspecific with British samples of Ceramium diaphanum sensu Harvey, for which no valid name has been identified up to now. We show that Ceramium polyceras (Kutzing) Zanardini is a valid name for this species. A fully corticated Ceramium species morphologically resembling C. secundatum differs at the species level from Atlantic C. secundatum; a valid name for this entity is Ceramium derbesii Solier ex Kutzing, described from Mediterranean France. A third species characterized by cortical spines, previously listed as Ceramium ciliation var. robustum (J. Agardh) Mazoyer, is shown to be Ceramium nudiusculum (Kutzing) Rabenhorst, originally described from Venice.

Four new exotic red seaweeds on European shores

We report the occurrence of four red macroalgae new to Europe. Two species were unambiguously determined to the species level with a DNA barcoding approach, while the remaining two species could only be assigned to a genus. Gelidium vagum was found in the Oosterschelde estuary (the Netherlands). Gracilariopsis chorda, Chondracanthus sp. and Solieria sp. were found in the Gulf of Morbihan in Brittany (France); Solieria sp. was also subsequently observed in the Thau Lagoon (France). Gelidium vagum and Gracilariopsis chorda are species originating from the north-western Pacific, around the Japanese archipelago. Phylogenetic analyses also show a likely Pacific origin for Chondracanthus sp. and Solieria sp. Three of these species are likely to have been introduced after 2008, indicating some active transport pathways between the Pacific and the north-eastern Atlantic. These findings also underline the importance of consistent and continuous local expertise (versus rapid assessment) in early warning systems.