Tracking the invasive history of the green alga Codium fragile ssp. tomentosoides

The spread of nonindigenous species into new habitats is having a drastic effect on natural ecosystems and represents an increasing threat to global biodiversity. In the marine environment, where data on the movement of invasive species is scarce, the spread of alien seaweeds represents a particular problem. We have employed a combination of plastid microsatellite markers and DNA sequence data from three regions of the plastid genome to trace the invasive history of the green alga Codium fragile ssp. tomentosoides. Extremely low levels of genetic variation were detected, with only four haplotypes present in the species' native range in Japan and only two of these found in introduced populations. These invasive populations displayed a high level of geographical structuring of haplotypes, with one haplotype localized in the Mediterranean and the other found in Northwest Atlantic, northern European and South Pacific populations. Consequently, we postulate that there have been at least two separate introductions of C. fragile ssp. tomentosoides from its native range in the North Pacific.     

OCCURRENCE OF CODIUM FRAGILE SUBSP. TOMENTOSOIDES (CHLOROPHYTA: BRYOPSIDALES) IN MARINE EMBAYMENTS OF SOUTHEASTERN AUSTRALIA

Codium fragile (Sur.) Hariot subsp. tomentosoides (van Goor) Silva, a widespread weedy species, is reported from Australia for the first time. Specimens were initially found in June 1996 in Corner Inlet, Victoria, where the species fouled fishing nets. This invasive chlorophycean alga was subsequently found in large numbers on 30 January 1997 in Port Phillip Bay and on 9 March 1998 in Western Port, Victoria. Presumably the species is native to Japan. The alga formed dense stands on rocky substrata in intertidal or in subtidal regions. In Port Phillip Bay, the species was also found attached to the common blue mussel Mytilus edulis planulatus Lamarck, and in Western Port it was found attached to the mud oyster Ostrea angasi Sowerby. Results of this study suggest that recruitment of juvenile germlings and growth of C. fragile subsp. tomentosoides differs between populations in Port Phillip Bay and those in Western Port.     

Cryptic speciation in liverworts – a case study in the Aneura pinguis complex

Bryophytes are amongst the most ancestral terrestrial plants and often have large distribution ranges across continents. Recent biochemical and molecular studies have suggested that many worldwide morphological species of bryophyte may represent genetically divergent and reproductively isolated cryptic species. We tested the cryptic species hypothesis in the thalloid liverwort Aneura pinguis complex. We applied analyses of chloroplast DNA (cpDNA) sequence variation and polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) methods to discriminate between specimens of A. pinguis derived from various habitats in five distant geographical regions in Poland. Of the 19 specimens sequence characterized for the cpDNA tRNA^Leu region, seven haplotypes were identified divided into three nonmonophyletic clusters. The application of developed PCR-RFLP markers confirmed the existence of three tRNA^Leu types of A. pinguis (A–C) within the specimens derived from 21 populations. Sympatric populations of different tRNA^Leu types were found in lowland and mountain regions. No clear correlation between stand type and the presence of two tRNA^Leu types (A, B) was observed, as both were growing on soil, humus, and rocks. The tRNA^Leu type C was found only on humus and its distribution was restricted to low-lying northern populations. The above results indicate that the A. pinguis complex is highly differentiated at the molecular level and may represent three cryptic species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 273–282.     

我国植物模式标本的馆藏量

植物标本是物种存在的永久凭证,模式标本在保障命名体系稳定中有不可替代的作用。标本馆藏量和模式标本数量反映一个国家或地区的历史积累。作者通过对比分析,发现我国的标本馆藏量远低于国际平均水平,模式标本馆藏量也相对较少,历史积累不足。在数字化本土馆藏模式标本基础上,标本数字化平台项目应将国外馆藏的中国植物种类的模式标本"引渡"回国。今后我国植物分类学研究者的重要工作之一是加强空白地区和国外标本采集,提高馆藏标本的代表性。     

Molecular identification of Chironomus spp. (Diptera) for biomonitoring of aquatic ecosystems

Abstract   Chironomidae larvae often represent a major component of the benthic fauna in inland water bodies and are used frequently as bioindicators of ecosystem health. The genus Chironomus is a recognised indicator of organic enrichment and has been used extensively in the northern hemisphere and New Zealand in ecotoxicological studies. However, similar use of Chironomus in Australia is limited due to the presence of cryptic species that restrict the collection of information on species-specific responses to environmental stress. To address the problems associated with species identification, we have used PCR-RFLP of the mitochondrial COI gene to develop DNA profiles for nine common Australian Chironomus species. Species-specific haplotypes were identified using reference taxa previously identified by cytological analysis, and verified with field specimens collected from seven wetlands around Melbourne. This research provides an effective tool for species identification of this ubiquitous and often abundant genus that will provide the basis of obtaining species-specific information to inform on the health of aquatic ecosystems.     

Multiple introductions and population structure during the rapid expansion of the invasive Sahara mustard (Brassica tournefortii)

The specific mechanisms that result in the success of any species invasion case are difficult to document. Reproductive strategies are often cited as a primary driver of invasive success, with human activities further facilitating invasions by, for example, acting as seed vectors for dispersal via road, train, air, and marine traffic, and by producing efficient corridors for movement including canals, drainages, and roadways. Sahara mustard (Brassica tournefortii) is a facultative autogamous annual native to Eurasia that has rapidly invaded the southwestern United States within the past century, displacing natives, and altering water‐limited landscapes in the southwest. We used a genotyping‐by‐sequencing approach to study the population structure and spatial geography of Sahara mustard from 744 individuals from 52 sites across the range of the species’ invasion. We also used herbaria records to model range expansion since its initial introduction in the 1920s. We found that Sahara mustard occurs as three populations in the United States unstructured by geography, identified three introduction sites, and combined herbaria records with genomic analyses to map the spread of the species. Low genetic diversity and linkage disequilibrium are consistent with self‐fertilization, which likely promoted rapid invasive spread. Overall, we found that Sahara mustard experienced atypical expansion patterns, with a relatively constant rate of expansion and without the lag phase that is typical of many invasive species.     

Invasion of Lepidium draba (Brassicaceae) in the western United States: distributions and origins of chloroplast DNA haplotypes

Advances in phylogeography are of great value for understanding the population structure and origins of invasive genotypes. Such insights provide constructive information for current or future biological control research efforts. In this study, we investigated a highly variable chloroplast DNA (cpDNA) marker for populations of the weed Lepidium draba (Brassicaceae) in its native Eurasian and invasive US ranges. We sequenced DNA from 684 individuals from Eurasia and the US and found 41 different haplotypes. Our comparative study between the native and invasive ranges showed a 33% reduction in allelic richness (A) and a 7% reduction in haplotype diversity (h) since introduction into the US. Most genetic variation in the native range was observed within geographical regions and populations, not between regions, and this result was similar for the invasive range. Assignment tests indicated the most likely origins of many invasive haplotypes. Some of these occurred in western Europe, supporting an expanded native range that had been proposed for the species. Exact locations were identified for a diverse set of invasive haplotypes which can be used in ongoing host-specificity tests of potential biological control agents.     

BIODIVERSITY RESEARCH: Genetic diversity in two introduced biofouling amphipods (Ampithoe valida & Jassa marmorata) along the Pacific North American coast: investigation into molecular identification and cryptic diversity

Aim We investigated patterns of genetic diversity among invasive populations of Ampithoe valida and Jassa marmorata from the Pacific North American coast to assess the accuracy of morphological identification and determine whether or not cryptic diversity and multiple introductions contribute to the contemporary distribution of these species in the region. Location Native range: Atlantic North American coast; Invaded range: Pacific North American coast. Methods We assessed indices of genetic diversity based on DNA sequence data from the mitochondrial cytochrome c oxidase subunit I (COI) gene, determined the distribution of COI haplotypes among populations in both the invasive and putative native ranges of A. valida and J. marmorata and reconstructed phylogenetic relationships among COI haplotypes using both maximum parsimony and Bayesian approaches. Results Phylogenetic inference indicates that inaccurate species‐level identifications by morphological criteria are common among Jassa specimens. In addition, our data reveal the presence of three well supported but previously unrecognized clades of A. valida among specimens in the north‐eastern Pacific. Different species of Jassa and different genetic lineages of Ampithoe exhibit striking disparity in geographic distribution across the region as well as substantial differences in genetic diversity indices. Main conclusions Molecular genetic methods greatly improve the accuracy and resolution of identifications for invasive benthic marine amphipods at the species level and below. Our data suggest that multiple cryptic introductions of Ampithoe have occurred in the north‐eastern Pacific and highlight uncertainty regarding the origin and invasion histories of both Jassa and Ampithoe species. Additional morphological and genetic analyses are necessary to clarify the taxonomy and native biogeography of both amphipod genera.     

Can herbarium records be used to map alien species invasion and native species expansion over the past 100 years?

Aim To determine if the temporal and spatial pattern of alien plant invasion and native plant expansion can be observed using 100 years of herbarium data from Oklahoma, USA, and to eliminate herbarium collection biases in such analyses. Location Oklahoma, USA. Methods Using herbarium records from the Oklahoma Vascular Plants Database from 1903 to 2004, we reconstructed the spatial and temporal collection history of two alien invasive taxa (Lonicera japonica and Tamarix spp.) and three native expansive species (Ambrosia psilostachya, Amphiachyris dracunculoides and Juniperus virginiana). To compare the overall collecting trend, groups of native non‐expansive taxa were selected as counterparts. We recorded the year of the first collection in each township in Oklahoma for all taxa. The cumulative number of occupied townships was log‐transformed, plotted against time and modelled with linear regression. The slope of the linear regression represented collection trend over time for the non‐expansive counterpart group. However, for the invasive and expansive species, the regression slope represented the collection effort plus the invasion or expansion rate. We calculated the proportion of invasive and expansive species to non‐expansive species by dividing the cumulative number of townships for each invasive or expansive species by the cumulative number of townships occupied by the counterpart group (proportion curve). Results Maps of the collection records of invasive and expansive taxa illustrated no discernible spatial invasion or expansion pattern. The slopes of the linear regression for alien invasive taxa were significantly steeper than those of their associated native non‐expansive counterparts, indicating an increase in abundance. Juniperus virginiana, L. japonica and Tamarix spp. exhibited one or more periods during which they were collected at a disproportionately higher rate than their native non‐expansive counterparts. Main conclusions Patterns of species invasion and expansion in Oklahoma were detected using techniques developed for regions with longer collecting plant histories. The proportion curve analysis eliminated some biases inherent in herbarium data by reducing the effect of collecting effort. Both the regression model and proportion curve analyses illustrate the temporal invasion patterns of alien invasive species. The native species did not show a clear expansion pattern. The information found in recently established herbaria may not be sensitive enough to detect the increase in abundance of native species.     

Invasion genetics of the Ciona intestinalis species complex: from regional endemism to global homogeneity

Determining the degree of population connectivity and investigating factors driving genetic exchange at various geographical scales are essential to understanding population dynamics and spread potential of invasive species. Here, we explore these issues in the highly invasive vase tunicate, Ciona intestinalis, a species whose invasion history has been obscured by its poorly understood taxonomy and population genetics. Recent phylogenetic and comparative genomic studies suggest that C. intestinalis is a cryptic species complex consisting of at least three species. We reconstructed phylogenies based on both mitochondrial (cytochrome c oxidase subunit 3—NADH dehydrogenase subunit 1 region and NADH dehydrogenase subunit 4 gene) and nuclear (internal transcribed spacer 1) sequences, results of which support four major phylogroups corresponding to the previously reported spA, spB and Ciona spp. (spC) as well as an undescribed cryptic species (spD). While spC and spD remain restricted to their native ranges in the Mediterranean Sea and Black Sea, respectively, the highly invasive species (spA and spB) have disjunct global distributions. Despite extensive interspecific divergences, we identified low phylogeographical structure within these two invasive species. Haplotype network analyses revealed comparatively limited mutation steps among haplotypes within each species. Population genetic analyses based on two mtDNA fragments and eight unlinked microsatellites illustrated relatively low population differentiation and high population connectivity at both regional and continental scales in the two invasive species. Human‐mediated dispersal coupled with a high potential for natural dispersal is probably responsible for the observed genetic homogeneity.     

Higher genetic diversity in introduced than in native populations of the mussel Mytella charruana: evidence of population admixture at introduction sites

Aim  Levels of genetic diversity can be used to determine haplotype frequency, population size and patterns of invasive species distribution. In this study, we sought to investigate the genetic structure of the invasive marine mussel Mytella charruana and compare variation from invasive populations with variation found within three native populations.
Location  Invaded areas in the USA (Florida, Georgia); native areas in Ecuador, Colombia and Brazil.
Methods  We sequenced 722 bp of the mitochondrial COI gene from 83 M. charruana samples from four invasive populations (USA) and 71 samples from two natural populations (Ecuador, Columbia). In addition, we sequenced 31 individuals of a congeneric species, Mytella guyanensis , from Salvador, Brazil. We constructed the phylogenetic relationship among all haplotypes and compared diversity measures among all populations.
Results  We found significantly higher levels of nucleotide diversity in invasive populations than in native populations, although the number of haplotypes was greater in the native populations. Moreover, mismatch distribution analyses resulted in a pattern indicative of population admixture for the invasive populations. Conversely, mismatch distributions of native populations resulted in a pattern indicative of populations in static equilibrium.
Main conclusion  Our data present compelling evidence that the M. charruana invasion resulted from admixture of at least two populations, which combined to form higher levels of genetic diversity in invasive populations. Moreover, our data suggest that one of these populations originated from the Caribbean coast of South America. Overall, this study provides an analysis of genetic diversity within invasive populations and explores how that diversity may be influenced by the genetic structure of native populations and how mass dispersal may lead to invasion success.     

The molecular systematics of Leiopotherapon unicolor (Günther, 1859): testing for cryptic speciation in Australia's most widespread freshwater fish

Leiopotherapon unicolor is the most widespread freshwater fish species in Australia. A comprehensive allozyme and mitochondrial DNA 16S rRNA data set was assembled from 141 specimens of L. unicolor collected Australia-wide in order to test for cryptic speciation in this far-ranging species. Surprisingly, little genetic diversity was observed within L. unicolor and provided no evidence for the existence of cryptic species within this lineage. In contrast, a small sample set of L. aheneus used as the outgroup showed two highly divergent haplotypes strongly suggestive of cryptic speciation. L. unicolor has a number of ecological and life history attributes that may explain the lack of significant genetic divergence over substantial geographical distances. The occurrence of other widespread fish and crustacean species that also display only limited genetic diversity indicate that climate conditions more favourable to dispersal across central and northern Australia than is suggested by the extent of present-day aridity have occurred in the relatively recent geological past.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 537–552.     

Genetic diversity of rodent species sold in South African pet shops

Murid rodents are considered globally important invasive species, yet they are still sold in the pet trade. Little is known about the genetic diversity of traded rodents, and many species are incorrectly identified in the pet trade. We used mitochondrial gene regions to assess the taxonomy and genetic diversity of 149 rodents sold in pet shops across eight South African provinces. We identified a total of 112 specimens as Mus musculus, while 31 were Rattus norvegicus, and six were identified as the southern African endemic, southern multimammate mouse Mastomys coucha. Phylogenetic analyses revealed that the three species were monophyletic. Mus musculus and R. norvegicus showed higher levels of genetic diversity, with 19 unique mtDNA haplotypes recovered for M. musculus and eight haplotypes for R. norvegicus. KwaZulu-Natal, Western Cape and Gauteng Provinces had the most unique haplotypes than other provinces. Our findings showed that non-native species are widely distributed in the South African pet trade industry, while M. coucha was not widely traded, although recorded in three provinces. This suggests that most provinces comply with the trade regulations on native species, but the threat of invasive rodents to South Africa's unique biodiversity is highlighted.     

Genetic networking of the Bemisia tabaci cryptic species complex reveals pattern of biological invasions

Background

A challenge within the context of cryptic species is the delimitation of individual species within the complex. Statistical parsimony network analytics offers the opportunity to explore limits in situations where there are insufficient species-specific morphological characters to separate taxa. The results also enable us to explore the spread in taxa that have invaded globally.

Methodology/Principal Findings

Using a 657 bp portion of mitochondrial cytochrome oxidase 1 from 352 unique haplotypes belonging to the Bemisia tabaci cryptic species complex, the analysis revealed 28 networks plus 7 unconnected individual haplotypes. Of the networks, 24 corresponded to the putative species identified using the rule set devised by Dinsdale et al. (2010). Only two species proposed in Dinsdale et al. (2010) departed substantially from the structure suggested by the analysis. The analysis of the two invasive members of the complex, Mediterranean (MED) and Middle East – Asia Minor 1 (MEAM1), showed that in both cases only a small number of haplotypes represent the majority that have spread beyond the home range; one MEAM1 and three MED haplotypes account for >80% of the GenBank records. Israel is a possible source of the globally invasive MEAM1 whereas MED has two possible sources. The first is the eastern Mediterranean which has invaded only the USA, primarily Florida and to a lesser extent California. The second are western Mediterranean haplotypes that have spread to the USA, Asia and South America. The structure for MED supports two home range distributions, a Sub-Saharan range and a Mediterranean range. The MEAM1 network supports the Middle East - Asia Minor region.

Conclusion/Significance

The network analyses show a high level of congruence with the species identified in a previous phylogenetic analysis. The analysis of the two globally invasive members of the complex support the view that global invasion often involve very small portions of the available genetic diversity.     

Giants invading the tropics: the oriental vessel fern, Angiopteris evecta (Marattiaceae)

The Oriental vessel fern, Angiopteris evecta (G.Forst.) Hoffm. (Marattiaceae), has its native range in the South Pacific. This species has been introduced into other localities since the 18th century and is now listed as an invasive species in several regions (Jamaica, Hawaii and Costa Rica). The purpose of our study is (1) to trace the distributional history of the species, and (2) to model its potential future range based on climatic conditions. The native range and the history of introduction are based on the existing literature and on 158 specimens from 15 herbaria, together with field observations. As there are taxonomic problems surrounding A. evecta, we limited our analysis to samples from the Pacific, most closely resembling the type from Tahiti. We modelled the potential range using GARP species distribution modelling with basic climatic variables, elevation, and location in relation to the coast. Analysis of past records shows that the species is able to colonise new ecosystems with relative ease. The modelling reveals that the species could be cultivated over a much wider range than where it currently is grown. The escape of cultivated plants into nature is probably due to distance from natural areas and is limited by local ecological factors, such as soil conditions or competitors. The predicted distribution in Asia and Madagascar is similar to the native distribution of the entire genus Angiopteris. It can therefore be assumed that most Angiopteris species have similar climatic preferences, and the absence of A. evecta in this predicted region may be due to dispersal limitation. In the Americas there is no native Angiopteris, but our climatic model predicts a vast potential habitat in tropical America; an invasion of A. evecta should be anticipated here in localities where the species is cultivated. Vessel ferns are known to alter the natural environment, which may reduce local biodiversity. Since A. evecta is not yet widely cultivated, it is advisable to restrict the trade and spread of the species and to discourage its cultivation as an ornamental. The global climate data available for modelling is however not detailed enough to predict the spread of A. evecta on a local or regional scale.     

Record error and range contraction, real and imagined, in the restricted shrub Banksia hookeriana in south-western Australia

Banksia hookeriana Meissn. (Proteaceae) is a fire‐killed shrub endemic to the northern sandplains of south‐western Australia that could be described as endangered based on its small geographical range (< 5000 km²) and area of occupancy (～500 km²). Impacts on the species’ geographical range by land clearance for farming and mining, and by altered fire regime, were investigated using three lines of evidence: records of herbarium collections, a comprehensive field survey of extant populations, and air photo and satellite images revealing the recent history of land clearance and fires. These show that the species’ range has contracted by up to 40% in area and 26% latitudinally through the loss of outlier and range limit populations since 1960. In addition, 22% of remaining native shrubland on the Eneabba sandplain has been lost over this period through clearing for farming and mining, representing further habitat loss for B. hookeriana. Detailed investigation of B. hookeriana herbarium collections (n = 46) revealed important errors that artificially affected the geographical range of the species and emphasized the importance of close examination of all data captured from collection records. Recorded locations occurred hundreds of kilometres outside the current geographical range of the species in areas with different climate and substrate. Incorrect species identification of herbarium specimens further extended the apparent geographical range of the species. On the other hand, credible records indicated the loss of the species from localities at the limits of its range. Overall, a disconcertingly high proportion of records contained errors that may be difficult to identify without close examination of the original collections and detailed ground‐truthing. Were these records to be used to model climate envelopes, identify potential habitat where the species might occur, or might migrate to either as pests or under climate change scenarios, or to analyse evolutionary or ecological theory (for example) — as is now becoming commonplace — large errors may ensue.     

Molecular genetic approaches to elucidate the ecological and evolutionary issues associated with biological invasions

Biological invasions may cause serious damage to the native environments and threaten the native biodiversity. Molecular genetic approaches have been found to be powerful tools for investigating the ecological and evolutionary aspects of biological invasions because the genetic structure and level of genetic variation of an invasive species are changed following its invasion. The present article reviews the use of molecular markers in addressing various aspects of invasive species. The application of these techniques has shown that many invasive species are actually "cryptic" species – species whose uniqueness is only recognizable at the genetic level. An estimation of the actual number of invasive species is essential when evaluating its ecological and economic impacts. Molecular genetic approaches have also enabled the source populations of invasive species to be identified. Reconstructions of invasion histories are crucial to preventing future invasions and conserving the native biodiversity, while comparisons of genetic variations between the native and introduced populations provide valuable opportunities to elucidate the mechanisms of rapid adaptation demonstrated by many invasive species.     

CURRENT STATUS OF THE INVASIVE GREEN ALGA CODIUM FRAGILE IN EASTERN CANADA

Ten years after the initial discovery of Codium fragile in eastern Canada on the Atlantic coast of Nova Scotia, C. fragile has extended its range considerably to northern Nova Scotia, southern New Brunswick and Prince Edward Island (P.E.I.) in the Gulf of St. Lawrence. In all principal areas C. fragile subsp. tomentosoides is present. In two regions of Prince Edward Island, Codium is becoming extremely abundant, and is known by local fishers as the "oyster thief". A potential second taxon of Codium was collected in 1999 at three sites in Malpeque Bay where it is locally abundant. These plants are distinguished from C. fragile subsp. tomentosoides by their smaller frond size, flatter utricle tops, smaller mucrons, and smaller gametangia. Principal component analysis distinguishes plants of the two types. The abundance of C. fragile on both hard bottom and soft bottom substrata suggests that it will become the dominant alga in many subtidal algal communities in eastern Canada.