North Atlantic phylogeography and large-scale population differentiation of the seagrass Zostera marina L

As the most widespread seagrass in temperate waters of the Northern Hemisphere, Zostera marina provides a unique opportunity to investigate the extent to which the historical legacy of the last glacial maximum (LGM18 000-10 000 years bp) is detectable in modern population genetic structure. We used sequences from the nuclear rDNA-internal transcribed spacer (ITS) and chloroplast matK-intron, and nine microsatellite loci to survey 49 populations (> 2000 individuals) from throughout the species' range. Minimal sequence variation between Pacific and Atlantic populations combined with biogeographical groupings derived from the microsatellite data, suggest that the trans-Arctic connection is currently open. The east Pacific and west Atlantic are more connected than either is to the east Atlantic. Allelic richness was almost two-fold higher in the Pacific. Populations from putative Atlantic refugia now represent the southern edges of the distribution and are not genetically diverse. Unexpectedly, the highest allelic diversity was observed in the North Sea-Wadden Sea-southwest Baltic region. Except for the Mediterranean and Black Seas, significant isolation-by-distance was found from ~150 to 5000 km. A transition from weak to strong isolation-by-distance occurred at ~150 km among northern European populations suggesting this scale as the natural limit for dispersal within the metapopulation. Links between historical and contemporary processes are discussed in terms of the projected effects of climate change on coastal marine plants. The identification of a high genetic diversity hotspot in Northern Europe provides a basis for restoration decisions.     

A Comparative Test of Mechanized and Manual Transplanting of Eelgrass, Zostera marina, in Chesapeake Bay

Abstract The laborious process of manual seagrass transplanting has often limited the size of seagrass restoration efforts. This study tested the efficiency of a mechanized planting boat, previously used for transplanting Halodule wrightii, relative to manual transplanting methods for establishing Zostera marina in Chesapeake Bay. Eelgrass planting was conducted at two sites, one each in the Rappahannock and James rivers, in October 2001. The methods were evaluated by three criteria: (1) initial planting success = proportion of attempted planting units (PUs) initially established (number confirmed in sediment by divers/number attempted); (2) survival = proportion of the initially established PUs persisting over 1, 4, and 24 weeks; and (3) efficiency = labor (in person·seconds) invested in each surviving PU. Initial planting success was significantly lower for the planting boat (24 and 56% at the Rappahannock and James sites, respectively) than for manual transplanting (100% at both sites). At the Rappahannock site, survival of initially established PUs declined over time for both methods, but while mean survival was always higher for manually planted rows, differences in survival between methods were not statistically significant. At the James site, survival to 1 and 4 weeks was significantly lower for the machine than for the manual method, but survival to 24 weeks was not significantly different. While the machine was able to attempt PUs faster than the manual method (2.2 s/PU vs. 5.8 s/PU, respectively), this speed was offset by poorer planting success rates, resulting in a much greater total labor investment for each machine‐planted PU that persisted to 24 weeks than for each similarly persisting manually planted PU (40.6 person·seconds/PU and 22.4 person·seconds/PU, respectively, averaged across sites). In summary, those PUs successfully planted by the machine survived similarly to PUs planted by hand, but as a result of poorer initial planting success, the machine required a greater investment of labor and plant donor stock for each PU surviving to 24 weeks. Therefore, in its tested configuration this planting boat is not a significant improvement over the manual method for transplanting eelgrass.     

Microsatellite loci in eelgrass Zostera marina reveal marked polymorphism within and among populations

Using an enriched genomic library, we developed seven (CT)n/(GA)n microsatellite loci for eelgrass Zostera marina L. Enrichment is described and highly recommended for genomes in which microsatellites are rare, such as in many plants. A test for polymorphism was performed on individuals from three geographically separated populations (N = 15/population) and revealed considerable genetic variation. The number of alleles per locus varied between five and 11 and the observed heterozygosities for single loci ranged from 0.16 to 0.81 within populations. Mean allele lengths were markedly different among populations, indicating that the identified loci will be useful in studying population structure in Z. marina. As the frequency of the most abundant multilocus genotype within populations was always < 1%, these loci have sufficient resolving power to address clone size in predominantly vegetatively reproducing populations.     

Coccolithophorids from the west coast of Baja California, Mexico

Studies on coccolithophorids in Mexico are rather scarce, probably due to the use of traditional methods for studying marine phytoplankton (e.g. collection, preservation and analysis). This is the first study of the coccolithophorid flora from west coast of Baja California, Mexico. Bottle samples (4 l) were taken at various depths (5, 25 and 50 m), at fixed stations located on perpendicular transects along the west coast of Baja California, considering three different zones: North, Central and South. A total of 32 samples were studied by optical and scanning electron microscopy, yielding 24 taxa identified (17 species and 5 nominal varieties, plus 2 different varieties). The coccolithophorid assemblages are considered subtropical and coastal (with many cosmopolitan taxa). Emiliania huxleyi and Gephyrocapsa oceanica were the most abundant (relative abundance) and widespread species in the study area. We report 15 new records for the Mexican Pacific. The zonal and vertical distribution of some taxa are briefly discussed, as well as the taxonomy of Rhabdosphaera claviger, Helicosphaera carteri, H. hyalina and related taxa.     

Desiccation,Moisture Content and Germination of Zostera marina L. Seed

Zostera marina is the only seagrass species whose seeds have been successfully used in large‐scale restoration. Although progress has been made in refining Z. marina restoration protocols, additional information on Z. marina seed physiology is necessary as the science of seagrass restoration evolves. We tested the germination rates of Z. marina seeds under different relative humidities and temperatures for different periods of time. Z. marina seed moisture content (MC) and germination rates were also tested when seeds were exposed to a temperature of 25°C and relative humidity of 50%. Z. marina seeds suffered higher mortality when exposed to lower relative humidity and higher temperature for longer period of exposure time. A significant negative correlation was detected between seed germination rate and MC. Z. marina seeds are sensitive to desiccation exposure and long periods of exposure to air should be prevented to minimize seed mortality when seeds are used in restoration projects.     

Genome scans detect consistent divergent selection among subtidal vs. intertidal populations of the marine angiosperm Zostera marina

Genome scans are a powerful tool to detect natural selection in natural populations among a larger sample of marker loci. We used replicated habitat comparisons to search for consistent signals of selection among contrasting populations of the seagrass Zostera marina, a marine flowering plant with important ecological functions. We compared two different habitat types in the North Frisian Wadden Sea, either permanently submerged (subtidal) or subjected to aerial exposure (intertidal). In three independent population pairs, each consisting of one tidal creek and one tidal flat population each, we carried out a genome scan with 14 expressed sequence tag (EST)-derived microsatellites situated in 5'- or 3'-untranslated regions of putative genes, in addition to 11 anonymous genomic microsatellites. By using two approaches for outlier identification, one anonymous and two EST-derived microsatellites showed population differentiation patterns not consistent with neutrality. These microsatellites were detected in several parallel population comparisons, suggesting that they are under diverging selection. One of these loci is linked to a putative nodulin gene, which is responsible for water channelling across cellular membranes, suggesting a functional link of the observed genetic divergence with habitat characteristics.     

Characterization of seven polymorphic microsatellite loci in the Baja California endemic black‐tailed brush lizard Urosaurus nigricaudus

Seven microsatellite loci were developed for the Baja California endemic black‐tailed brush lizard Urosaurus nigricaudus, using an enriched genomic DNA library. All loci were polymorphic and overall presented high levels of variation. Number of alleles ranged from five to 16 (average 12.14), and observed heterozygosities from 0.535 to 0.923 (average 0.752). Cross‐species amplification was successful and polymorphism was detected for all the loci using the congeners Urosaurus lahtelai and Urosaurus ornatus. These markers will be useful to study fragmented populations of U. nigricaudus on agricultural landscape of the Baja California Peninsula.     

Characterization of microsatellite loci in the dwarf eelgrass Zostera noltii (Zosteraceae) and cross‐reactivity with Z. japonica

Zostera noltii is an important species of eelgrass occurring along European, north African, Mediterranean, Black Sea and Azov Sea coasts. Nine microsatellite loci were developed and no linkage disequilibrium was observed. Cross‐amplification was observed for all loci (polymorphic) in Z. japonica; only four loci amplified (monomorphic) in Z. marina.     

A microsatellite-based estimation of clonal diversity and population subdivision in Zostera marina, a marine flowering plant

We examined the genetic population structure in eelgrass (Zostera marina L.), the dominant seagrass species of the northern hemisphere, over spatial scales from 12 km to 10 000 km using the polymorphism of DNA microsatellites. Twelve populations were genotyped for six loci representing a total of 67 alleles. Populations sampled included the North Sea (four), the Baltic Sea (three), the western Atlantic (two), the eastern Atlantic (one), the Mediterranean Sea (one) and the eastern Pacific (one). Microsatellites revealed substantial genetic variation in a plant group with low allozyme diversity. Average expected heterozygosities per population (monoclonal populations excluded) ranged from 0.32 to 0.61 (mean = 0. 48) and allele numbers varied between 3.3 and 6.7 (mean = 4.7). Using the expected frequency of multilocus genotypes within populations, we distinguished ramets from genetic individuals (i.e. equivalent to clones). Differences in clonal diversity among populations varied widely and ranged from maximal diversity (i.e. all ramets with different genotype) to near or total monoclonality (two populations). All multiple sampled ramets were excluded from further analysis of genetic differentiation within and between populations. All but one population were in Hardy-Weinberg equilibrium, indicating that Zostera marina is predominantly outcrossing. From a regression of the pairwise population differentiation with distance, we obtained an effective population size Ne of 2440-5000. The overall genetic differentiation among eelgrass populations, assessed as rho (a standardized estimate of Slatkin's RST) was 0.384 (95% CI 0.34-0.44, P < 0.001). Genetic differentiation was weak among three North Sea populations situated 12-42 km distant from one another, suggesting that tidal currents result in an efficient exchange of propagules. In the Baltic and in Nova Scotia, a small but statistically significant fraction of the genetic variance was distributed between populations (rho = 0.029-0. 053) at scales of 15-35 km. Pairwise genetic differentiation between European populations were correlated with distance between populations up to a distance of 4500 km (linear differentiation-by-distance model, R2 = 0.67). In contrast, both Nova Scotian populations were genetically much closer to North Sea and Baltic populations than expected from their geographical distance (pairwise rho = 0.03-0.08, P < 0.01). A biogeographical cluster of Canadian with Baltic/North Sea populations was also supported using a neighbour-joining tree based on Cavalli-Sforza's chord distance. Relatedness between populations may be very different from predictions based on geographical vicinity.     

Pleistocene climatic fluctuations drive isolation and secondary contact in the red diamond rattlesnake (Crotalus ruber) in Baja California

Aim

Many studies have investigated the phylogeographic history of species on the Baja California Peninsula, and they often show one or more genetic breaks that are spatially concordant among many taxa. These phylogeographic breaks are commonly attributed to vicariance as a result of geological or climatic changes, followed by secondary contact when barriers are no longer present. We use restriction‐site associated DNA sequence data and a phylogeographic model selection approach to explicitly test the secondary contact hypothesis in the red diamond rattlesnake, Crotalus ruber.

Location

Baja California and Southern California.

Methods

We used phylogenetic and population clustering approaches to identify population structure. We then used coalescent methods to simultaneously estimate population parameters and test the fit of phylogeographic models to the data. We used ecological niche models to infer suitable habitat for C. ruber at the Last Glacial Maximum (LGM).

Results

Crotalus ruber is composed of distinct northern and southern populations with a boundary near the town of Loreto in Baja California Sur. A model of isolation followed by secondary contact provides the best fit to the data, with both divergence and contact occurring in the Pleistocene. We also identify a genomic signature of northern range expansion in the northern population, consistent with LGM niche models showing that the northern‐most portion of the range of C. ruber was not suitable habitat during the LGM.

Main conclusions

We provide the first explicitly model‐based test of the secondary contact model in Baja California and show that populations of C. ruber were isolated before coming back into contact near Loreto, a region that shows phylogeographic breaks for other taxa. Given the timing of divergence and contact, we suggest that climatic fluctuations have driven the observed phylogeographic structure observed in C. ruber and that they may have driven similar patterns in other taxa.     

Not just vicariance: phylogeography of a Sonoran Desert euphorb indicates a major role of range expansion along the Baja peninsula

To examine the generality of population-level impacts of ancient vicariance identified for numerous arid-adapted animal taxa along the Baja peninsula, we tested phylogeographical hypotheses in a similarly distributed desert plant, Euphorbia lomelii (Euphorbiaceae). In light of fossil data indicating marked changes in the distributions of Baja floristic assemblages throughout the Holocene and earlier, we also examined evidence for range expansion over more recent temporal scales. Two classes of complementary analytical approaches — hypothesis-testing and hypothesis-generating — were used to exploit phylogeographical signal from chloroplast DNA sequence data and genotypic data from six codominant nuclear intron markers. Sequence data are consistent with a scenario of mid-peninsular vicariance originating c. 1 million years ago (Ma). Alternative vicariance scenarios representing earlier splitting events inferred for some animals (e.g. Isthmus of La Paz inundation, c. 3 Ma; Sea of Cortez formation, c. 5 Ma) were rejected. Nested clade phylo-geographical analysis corroborated coalescent simulation-based inferences. Nuclear markers broadened the temporal spectrum over which phylogeographical scenarios could be addressed, and provided strong evidence for recent range expansions along the north–south axis of the Baja peninsula. In contrast to previous plant studies in this region, however, the expansions do not appear to have been in a strictly northward direction. These findings contribute to a growing appreciation of the complexity of organismal responses to past climatic and geological changes — even when taxa have evolved in the same landscape context.     

中国北部湾白骨壤红树林的虫害与研究对策

危害北部湾白骨壤的主要害虫是广州小斑螟 (Oligochroacantonella)幼虫、双纹白草螟 (Pseudcatharylladuplicella)幼虫和广翅蜡蝉 (Euricaniasp.)成虫。广州小斑螟喜食害白骨壤叶片 ,造成大量叶片的枯干和枝条的死亡 ,最高可导致林木 73 %的叶面积危害。室内培养发现广大腿小蜂 (Brachymeriasp .)、啮小蜂 (Tetrastichussp .)可寄生广州小斑螟的蛹 ,终止其羽化过程。虫害具有突发性、专一性、快速扩展的特点。海水淹没可明显抑制虫害。讨论了虫害的发源、爆发的原因、生态效应和防治途径 ,并指出深入研究的方向     

Policy plans and management measures to restore eelgrass (Zostera marina L.) in the Dutch Wadden Sea

The Dutch Wadden Sea has been changed dramatically over the last centuries by human activities like land reclamation and different forms of fishery. This has, amongst other things, led to changes in the number of biological communities. One of the changes was the near extinction of eelgrass (Zostera marina L.) in the Dutch Wadden Sea. The deterioration of the area led to policy plans in the late 1980s that aimed at restoring the original natural communities of which the eelgrass community was one. This paper presents a restoration strategy which contains a selection procedure for suitable transplantation sites. The selection procedure is based on factors such as sediment composition, exposure time, current velocity and wave action. These were combined in a GIS-based map integrating these factors. One important action in the restoration process is to increase the number of freshwater discharge points to meet the requirements of the brackish water community in general and the growing conditions for eelgrass in particular. Received: 27 October 1999 / Received in revised form: 3 February 2000 / Accepted: 3 February 2000     

Phylogenetic structure among pocket gopher populations,genus Thomomys (Rodentia: Geomyidae), on the Baja California Peninsula

We determined the phylogenetic relationships, population history, and hierarchical structure of genetic variation in pocket gophers distributed on the Baja California Peninsula (BCP), based on extensive geographic sampling. Using a fragment of the mitochondrial gene cytochrome b (cyt b), we found three latitudinal structured geographic clades (northern, central, and southern). The northern clade occurs in the border area of the USA and the north of BCP, the central clade occurs from the peninsular highlands through the Central Desert of Baja California, and the southern clade is distributed south of the San Ignacio Lagoon. AMOVA showed that genetic variation is higher among clades (64%) than within populations (18.1%). The deepest divergence among clades is very shallow (～300 000 years), which suggests that climatic changes during the Pleistocene or some inhospitable habitats have affected the structure of this group, rather than influences from older marine transgressions. Phylogenetic groups disclosed by our results do not coincide with the current infraspecific classification; therefore, we propose a change of epithet for BCP gophers (Thomomys nigricans) and a new subspecific taxonomic arrangement with four subspecies: Thomomys nigricans anitae, Thomomys nigricans martirensis, Thomomys nigricans nigricans, and Thomomys nigricans russeolus. © 2013 The Linnean Society of London     

高等植物大叶藻研究进展及其对海洋沉水生活的适应

综述和讨论了目前对海洋沉水植物大叶藻的研究进展 ,主要包括 :(1 )形态解剖结构特点 ,(2 )基本生理研究 ,(3 )耐盐机理 ,(4)生存限制因子 ,(5 )问题与展望。其中着重讨论了大叶藻与海洋沉水生活相适应的一些特点 ,特别是其对海水盐度的适应机理。     

Species richness in macroalgae and macrofauna assemblages on Fucus serratus L. (Phaeophyceae) and Zostera marina L. (Angiospermae) in Skagerrak, Norway

The macrophytes Fucus serratus and Zostera marina form similar substrates for associated flora and fauna in shallow waters in Norway. While F. serratus forms a more or less continuous belt on rocky substrate along the coast, Z. marina forms disjunct populations on sandy or muddy bottoms. This study focused on the organisms associated with these two macrophytes in two localities in the Skagerrak region. In total, 130 taxa of epiphytic organisms were identified: 22 green algae, 41 red algae, 32 brown algae, eight diatoms and 27 sessile animals. One hundred and twenty-seven taxa of mobile macrofauna were registered. The dominant group was crustaceans, with amphipods as the order containing most species. Many species of both plants and animals preferred one or the other habitat. It is concluded that coastal macrophyte systems have high species diversity.     

Genetic structure of Japanese Chattonella marina (Raphidophyceae) populations revealed using microsatellite markers

Chattonella marina var. antiqua and C. marina var. marina (Raphidophyceae) are red tide‐forming, harmful phytoplankton species. We investigated the genetic diversity and genetic relationship among the populations using microsatellite markers to identify putative sources of C. marina var. antiqua and C. marina var. marina in Japanese coastal populations. A positive correlation between genetic divergence and geographical distance (isolation by distance) was recognized for C. marina var. antiqua. The C. marina var. antiqua populations were established throughout a geological time scale, and genetic divergence had progressed in each population with gene flow depending on geographic distances. In contrast, isolation by distance was not observed for C. marina var. marina populations, and the genetic divergence among populations was extremely high. The Tokyo Bay population of C. marina var. marina, which was first recognized in 2008, had many private alleles but was related to the Kagoshima Bay population. The Tokyo Bay population may have been established by several invasions from the Kagoshima Bay population and other regions.     

High genetic diversity and low structure in an endemic long-lived tree,Yucca capensis (Asparagaceae)

• Endemic species distributed in fragmented habitats are highly vulnerable to extinction because they may have low genetic diversity. However, some life-history traits can mitigate the effect of genetic drift on populations. We analysed the level and distribution of genetic variation and ancestral population size of Yucca capensis, a long-lived endemic plant of the Baja California Peninsula, Mexico. Its populations are scattered across a habitat that is suffering accelerated transformation.
• We used six nuclear microsatellites to genotype 224 individuals from 17 locations across the entire species' geographic range. We estimated polymorphisms, heterozygosity and genetic structure. We also evaluated the ancestral and recent effective size and time since the population started to change.
• We found high heterozygosity, high polymorphism and low differentiation among locations, suggesting a panmictic population across the range. We also detected a large ancestral effective population size, which suffered a strong reduction in the Mid-Holocene.
• Despite changes in environmental conditions caused by habitat modification, the high diversity and low differentiation in Y. capensis may result from its large ancestral effective size and life-history traits, such as plant longevity, clonal growth and mating system, which reduce the rate of loss of genetic variation. However, the dependence on a specialist pollinator that displays short flight range can reduce gene flow among the plant populations and could, shortly, lead them into an extinction vortex.

     

Eelgrass meadows in the California Channel Islands and adjacent coast reveal a mosaic of two species, evidence for introgression and variable clonality

BACKGROUND AND AIMS: Seagrasses are important facilitator species in shallow, soft-bottom marine environments worldwide and, in many places, are threatened by coastal development and eutrophication. One narrow-leaved species (Zostera marina) and one wide-leaved species, variously designated as Z. marina, Z. pacifica or Z. asiatica, are found off the California Channel Islands and adjacent California-Mexico coast. The aim of the present study was to confirm species identification genetically and to link patterns of genetic diversity, connectivity and hybridization among and within the populations with historical sea levels (Ice Age) or the contemporary environment. METHODS: Samples (n = 11-100) were collected from 28 sites off five California Channel Islands and six sites off the adjacent coast of southern California and Baja California, Mexico. DNA polymorphisms of the rDNA-ITS (internal transcribed spacer) cistron (nuclear), the matK intron (chloroplast) and nine microsatellite loci (nuclear) were examined in a population genetic and phylogeographic context. KEY RESULTS: All wide-leaved individuals were Z. pacifica, whereas narrow-leaved forms were Z. marina. Microsatellite genotypes were consistent with hybridization between the two species in three populations. The present distribution of Z. pacifica follows a glacial age land mass rather than present oceanographic regimes, but no link was observed between the present distribution of Z. marina and past or present environments. Island populations of Z. marina often were clonal and characterized by low genotypic diversity compared with populations along the Baja California coast. The high level of clonal connectivity around Santa Catalina Island indicated the importance of dispersal and subsequent re-establishment of vegetative fragments. CONCLUSIONS: The pristine environmental conditions of offshore islands do not guarantee maximum genetic diversity. Future restoration and transplantation efforts of seagrasses must recognize cryptic species and consider the degree of both genetic and genotypic variation in candidate donor populations.     

Population genetic structure of parthenogenetic flatworm populations with occasional sex

1. Sexual populations are expected to perform better in fluctuating environments than asexuals because recombination provides the potential to adapt to changing environments due to increased genetic variation. Nevertheless, some asexual species show comparably high levels of genotypic diversity. Such diversity might be achieved through gene flow between coexisting sexual and asexual populations or through sexual events within asexual populations. 2. Evidence for occasional sex in the flatworm Schmidtea polychroa was previously found at one specific site that is inhabited by parthenogenetic forms. There, varying rates of sex between subpopulations, reaching up to 12%, were observed. Past recurrent sexual processes left a significant genetic signature in the population genetic structure of this population. In the present study, we examined the population genetic structure of six independent metapopulations (lakes) of the freshwater planarian flatworm S. polychroa, to confirm the presence of occasional sex and that its population genetic consequences can be generalised. 3. Using microsatellites, we found varying rates of occasional sex among subpopulations. Metapopulations showed medium to high levels of genotypic diversity that correlated with the rate of sex. 4. We conclude that occasional sex has considerable consequences for population genetic structure of parthenogenetic species and promotes diversity that might allow response to the particular type of selection that is usually predicted to favour sexual reproduction. This reproductive strategy provides genetic characteristics required for selection to act on, and might, therefore, explain the success of this parthenogenetic species.