Native and naturalized range size in Pinus: relative importance of biogeography,introduction effort and species traits

Aim Pine trees (genus Pinus) represent an ancient lineage, naturally occurring almost exclusively in the Northern Hemisphere, but introduced and widely naturalized in both hemispheres. As large trees of interest to forestry, they attract much attention and their distribution is well documented in both indigenous and naturalized ranges. This creates an opportunity to analyse the relationship between indigenous and naturalized range sizes in the context of different levels of human usage, biological traits and the characteristics of the environments of origin. Location Global. Methods We combined and expanded pre‐existing data sets for pine species distributions and pine species traits, and used a variety of regression techniques (including generalized additive models and zero‐inflated Poisson models) to assess which variables explained naturalized and indigenous range sizes. Results Indigenous and naturalized range sizes are positively correlated but there are many notable exceptions. Some species have large indigenous ranges but small or no naturalized ranges, whereas others have small indigenous ranges, but have naturalized in many regions. Indigenous range is correlated to factors such as seed size (?), age at first reproduction (?), and latitude (+, supporting Rapoport's rule), but also to the extent of coverage of species in the forestry literature (+). Naturalized range size is strongly influenced by the extent of coverage of species in the forestry literature (+), a proxy for propagule pressure. Naturalization was also influenced by average elevation in the indigenous range (?) and age at first reproduction (?). Main conclusions The macroecological and evolutionary pressures facing plant groups are not directly transferable between indigenous and naturalized ranges. In particular, there are strong biases in species naturalization and expansion in invasive ranges that are unrelated to factors determining indigenous range size. At least for Pinus, a new set of macroecological patterns are emerging which are profoundly influenced by humans.     

“Barcoding” and ISSR data illuminate a problematic infraspecific taxonomy for Chrysanthemoides monilifera (Calenduleae; Asteraceae): Lessons for biocontrol of a noxious weed

Chrysanthemoides monilifera Tourn. ex Medik is a noxious weed in Australia and New Zealand. It is a widespread species in southern Africa, where it shows considerable morphological variation that has resulted in a confusing infraspecific taxonomy. We use DNA sequence data from the nuclear Internal Transcribed Spacer (ITS) region from 78 samples of this species from its indigenous distribution range and 10 samples from Australia and New Zealand to determine the regions of origin of the invasive plants. These data are augmented by a smaller study using ISSR markers. Bayesian Inference analysis was somewhat resolved, with many weakly supported nodes. Clades with support tended to correspond to infraspecific taxonomic entities, and were geographically coherent. In contrast, a neighbour-net analysis was not as well resolved and indicated considerable reticulation. All analyses of ITS data retrieved two major clades corresponding to Western and Eastern distributions, with some overlap. Samples from New Zealand and Australia correspond to the taxon C. monilifera subsp. monilifera, and are resolved as most closely related to samples from the greater Cape Town area. Biological control agent populations for C. monilifera subsp. monilifera should be sourced from this region in order to avoid host plant incompatibility problems.     

Avian egg collections: museum collection bias driven by shape and size

Avian eggs exhibit substantial intra- and interspecific variation in shape, size and colour. Considerable efforts have been made to better understand the evolutionary drivers behind such variation, often using museum egg collections. Usually it is assumed that museum collections accurately represent the variation seen in natural populations, but this may not be the case if there is collection bias. Collection bias may lead to the over-representation of certain egg traits in collections, due to the aesthetic (or other) preferences of collectors. The aim of this study is to begin to look for the occurrence of potential collection bias in museum egg collections by comparing three shape indices (pointedness/asymmetry, elongation and polar asymmetry) and egg volume between subsets of eggs in museum collections with those of recently sampled eggs in the field for three different bird species: common guillemot Uria aalge, razorbill Alca torda and northern fulmar Fulmarus glacialis. We found no evidence of collection bias in our sampled razorbill and northern fulmar museum collection eggs, but some evidence for a bias in sampled museum collection eggs of common guillemots. Since the guillemot's egg differs from most bird eggs in being pyriform, we suggest that collection bias by historic egg collectors may be more prevalent in species with extreme egg traits. Researchers using museum egg collections to examine questions relating to egg shape should be aware of collection bias risks and consider how to minimise the effect of these possible biases on accumulated datasets.     

Modelling the introduction and spread of non‐native species: international trade and climate change drive ragweed invasion

Biological invasions are a major driver of global change, for which models can attribute causes, assess impacts and guide management. However, invasion models typically focus on spread from known introduction points or non‐native distributions and ignore the transport processes by which species arrive. Here, we developed a simulation model to understand and describe plant invasion at a continental scale, integrating repeated transport through trade pathways, unintentional release events and the population dynamics and local anthropogenic dispersal that drive subsequent spread. We used the model to simulate the invasion of Europe by common ragweed (Ambrosia artemisiifolia), a globally invasive plant that causes serious harm as an aeroallergen and crop weed. Simulations starting in 1950 accurately reproduced ragweed's current distribution, including the presence of records in climatically unsuitable areas as a result of repeated introduction. Furthermore, the model outputs were strongly correlated with spatial and temporal patterns of ragweed pollen concentrations, which are fully independent of the calibration data. The model suggests that recent trends for warmer summers and increased volumes of international trade have accelerated the ragweed invasion. For the latter, long distance dispersal because of trade within the invaded continent is highlighted as a key invasion process, in addition to import from the native range. Biosecurity simulations, whereby transport through trade pathways is halted, showed that effective control is only achieved by early action targeting all relevant pathways. We conclude that invasion models would benefit from integrating introduction processes (transport and release) with spread dynamics, to better represent propagule pressure from native sources as well as mechanisms for long‐distance dispersal within invaded continents. Ultimately, such integration may facilitate better prediction of spatial and temporal variation in invasion risk and provide useful guidance for management strategies to reduce the impacts of invasion.     

Controls on pathogen species richness in plants' introduced and native ranges: roles of residence time, range size and host traits

Introduced species escape many pathogens and other enemies, raising three questions. How quickly do introduced hosts accumulate pathogen species? What factors control pathogen species richness? Are these factors the same in the hosts' native and introduced ranges? We analysed fungal and viral pathogen species richness on 124 plant species in both their native European range and introduced North American range. Hosts introduced 400 years ago supported six times more pathogens than those introduced 40 years ago. In hosts' native range, pathogen richness was greater on hosts occurring in more habitat types, with a history of agricultural use and adapted to greater resource supplies. In hosts' introduced range, pathogen richness was correlated with host geographic range size, agricultural use and time since introduction, but not any measured biological traits. Introduced species have accumulated pathogens at rates that are slow relative to most ecological processes, and contingent on geographic and historic circumstance.     

The worldwide transfer of ants: geographical distribution and ecological invasions

Aim This is the first comprehensive account of the biogeography of ants transferred and at least temporarily established outside their native habitat. Location Using museum and literature records, I established the distributions of transferred ant species. Methods I used taxonomic and functional groups to assess how geographical spread as a transferred species is affected by taxonomy and life history. Results 147 ant species in forty-nine genera have been recorded outside of their native habitat. The proportion of transferred ants is similar to the number of genera and species in each subfamily. The species-rich subfamily Myrmicinae contains nearly 50% of all transferred species, while many of the species-poor subfamilies have absolutely no transferred species. A disproportionate high number of transferred ants originate from the Neotropical and Oriental biogeographic regions. The Pacific Islands are the recipients of the most transferred ant species. Most transferred ants belong to the CRYPTIC, OPPORTUNIST, and GENERALIZED MYRMICINE functional groups, while there are no recorded transfers of army ants or leaf-cutting ants. Both invasive and human commensal ‘tramp’ ant species are nonrandom subsets of transferred ants. Main conclusions ‘Tramp’ species and invasive species tend to have widespread geographical distributions, and share life history characteristics including queen number, nest structure, and foraging behaviour. Combining observations of functional groups and biogeography may lead to a better understanding of the factors contributing to the spread of transferred species.     

Interactions between introduced and native predatory ladybirds (Coleoptera, Coccinellidae): factors influencing the success of species introductions

Abstract.  1. Why some exotic species invade successfully while others do not is poorly understood. This study focuses on the inability of the exotic predatory ladybird Adalia bipunctata (L.) (Coleoptera, Coccinellidae) to establish and spread in Japan. In particular, this study considers the interactions between A. bipunctata and three native Japanese ladybird species, Harmonia axyridis Pallas, Coccinella septempunctata Linn., and Propylea japonica Mulsant.
2. The number of aphids occurring on the host plants declined dramatically over the course of larval development in the presence of H. axyridis or C. septempunctata . In contrast, many aphids remained at the end of the experiment when only A. bipunctata or P. japonica was present.
3. The survivorship of A. bipunctata from second instar to adult was significantly reduced in the presence of H. axyridis , but not in the presence of C. septempunctata or P. japonica . Most deaths were the result of intra-guild predation when H. axyridis or C. septempunctata larvae were present. Larvae of H. axyridis and C. septempunctata suffered very high mortality in this experiment, but survived better when co-occurring with A. bipunctata than when occurring only with conspecifics. In contrast, P. japonica survived better when occurring with conspecifics than with A. bipunctata .
4. The finding that larvae of the exotic species A. bipunctata died at high rates from intra-guild predation by the native Japanese ladybirds H. axyridis and C. septempunctata may bear on the question of why A. bipunctata has not established more successfully in Japan whereas H. axyridis and C. septempunctata have been very successful in establishing themselves in North America.     

Climate change, species range limits and body size in marine bivalves

We use data on the Pleistocene and modern range limits of Californian marine bivalves to show that species that shifted their geographical ranges in response to Pleistocene climatic fluctuations were preferentially drawn from the large end of the regional body size–frequency distributions. This difference is not due to phylogenetic effects (i.e. dominance of extralimital species by a few large-bodied clades), differences among major ecological categories (burrowing versus surface-dwelling, or suspension feeding versus non-suspension feeding), or differences in modes of reproduction and larval development. In addition, we show that successful invasive species of bivalves in present-day marine habitats also tend to be large-bodied, despite the difference in mechanisms between present-day and Pleistocene range expansions. These results indicate that range limits of large-bodied bivalve species are more unstable than small-bodied ones, and that body size and its correlates need to be considered when attempting to predict the responses of marine communities to climate change, biotic interchanges and human-mediated invasions.     

Well known does not mean well studied: morphological and molecular support for existence of sibling species in the Javanese gliding frog Rhacophorus reinwardtii (Amphibia, Anura)

Morphological, morphometrical and molecular data support the existence of two sibling species in the taxon previously known as Rhacophorus reinwardtii. The two species can be distinguished by the coloration pattern of webbing, the size of adult specimens and the relative size of various morphometric characters. This long and commonly known taxon should be separated into two species. As a consequence, the conservation status of the new species Rhacophorus kio n. sp. as well as of the redefined species Rhacophorus reinwardtii must be re-evaluated and, considering the new distribution data and the particular ecological demands, both species should be considered as 'endangered'.     

Assessing the vulnerability of riparian vegetation to invasion by Mimulus guttatus: relative importance of biotic and abiotic variables in determining species occurrence and abundance

Riparian habitats are particularly susceptible to invasion by non‐native plants. At present, attempts to build consensus as to what the primary drivers of plant invasion in riparian ecosystems might be is hindered by the absence of common standards for data collected on plant species (e.g. occurrence, or relative abundance). Mimulus guttatus L., a non‐native riparian plant species, was used as a model to determine how environmental drivers influence two aspects of invasibility: species occurrence and abundance (assessed in relation to three variables number of patches, patch area and number of stems per patch). Mimulus occurrence and abundance, together with 20 environmental variables, were surveyed in almost 700 contiguous 50‐m‐long riverbank segments within a catchment in north‐east Scotland. More than half of the segments had been colonized by Mimulus. Occurrence and number of patches responded to similar environmental gradients, particularly bare sediment, boulders, high soil moisture, short‐statured ruderal communities, and open canopies, and tended to be highest downstream where the river was widest. In contrast to occurrence and patch number, patch area and stem number per patch were higher in the upper reaches of the catchment and were positively associated with low tree canopy and vegetation dominated by light‐demanding species and smaller‐statured species. Patch area and stem number per patch were also positively related to grazing. This study has highlighted the importance of assessing more than one measure of invasion success (occurrence or patch number and either patch area or stem number per patch), as they are each determined by a different suite of environmental variables. Abiotic factors, such as sediment availability and presence of boulders, appeared to be the major determinants of occurrence and patch number, whereas biotic factors, such as interspecific competition and grazing, were more important ecological determinants underlying area and stem number per patch.     

What it takes to invade grassland ecosystems: traits,introduction history and filtering processes

Whether the success of alien species can be explained by their functional or phylogenetic characteristics remains unresolved because of data limitations, scale issues and weak quantifications of success. Using permanent grasslands across France (50 000 vegetation plots, 2000 species, 130 aliens) and building on the Rabinowitz's classification to quantify spread, we showed that phylogenetic and functional similarities to natives were the most important correlates of invasion success compared to intrinsic functional characteristics and introduction history. Results contrasted between spatial scales and components of invasion success. Widespread and common aliens were similar to co‐occurring natives at coarse scales (indicating environmental filtering), but dissimilar at finer scales (indicating local competition). In contrast, regionally widespread but locally rare aliens showed patterns of competitive exclusion already at coarse scale. Quantifying trait differences between aliens and natives and distinguishing the components of invasion success improved our ability to understand and potentially predict alien spread at multiple scales.     

The illusion of nature: perception and the reality of natural landscapes,as illustrated by vertebrate fauna in the Northern Territory,Australia

Biomass figures are compared for common native vertebrate species and common non‐native vertebrate species (including fish, amphibian, reptile, bird and mammal species) across the Northern Territory, an area of ostensibly, largely unmodified natural system. The total biomass of the considered non‐native species very substantially surpasses that of the common native vertebrate species considered; hence, the perception that this is a largely intact natural system is, to some extent, an illusion. This preponderance of biomass of non‐native species may help explain the increasing evidence of ecosystem stress and decline of native species. An integrity index based on the relative biomass of native and non‐native species may provide insight into the status and trends of ecological systems across different regions.     

Sperm competition and the evolution of precopulatory weapons: Testis size and amplexus position,but not arm strength,affect fertilization success in a chorusing frog

Trade‐offs between pre‐ and postcopulatory traits influence their evolution, and male expenditure on such traits is predicted to depend on the number of competitors, the benefits from investing in weapons, and the risk and intensity of sperm competition. Males of the chorusing frog Crinia georgiana use their arms as weapons in contest competition. Previously, we showed that increased numbers of rivals elevated the risk and intensity of sperm competition due to multimale amplexus, and caused a reversal in the direction of precopulatory selection on arm girth. Here, we focused on the factors affecting postcopulatory fertilization success during group spawning, using paternity data from natural choruses. Competitive fertilization success depended on the time spent amplexed and amplexus position. Relative testes size but not arm girth, contributed to fertilization success, but the effect of testes size depended on amplexus position. Our findings offer within species empirical support for recent sperm competition models that incorporate precopulatory male–male competition, and show why an understanding of the evolution of animal weapons requires a consideration of both pre‐ and postcopulatory episodes of sexual selection.     

The relationships between habitat suitability,population size and body condition in a pond-breeding amphibian

The ecological niche of a species determines whether a species can persist and reproduce in a patch or not. The niche of a species is often described using habitat suitability models and indices. Accordingly, one may expect tight links between demography, phenotypes of individuals, population size, and habitat suitability. However, such links are not always found. Here, we study the relationship between a habitat suitability index that is commonly used for conservation assessments and metrics describing the performance at the level of populations and individuals. Using data from a metapopulation of a pond-breeding amphibian, the Great Crested Newt (Triturus cristatus), we show that habitat suitability predicts population size but not body condition. Ponds with higher suitability had a higher population size of newts, whereas population size correlated negatively with body condition of individuals. Our results are in line with previous studies showing no straightforward relationship between habitat suitability and body condition (a measure of individual performance) and the performance of populations. We suggest that a population size-dependent reduction of body condition may be a regulatory mechanism in newt populations.     

The Upper Tremadocian (Ordovician) graptolite Bryograptus: taxonomy, biostratigraphy and biogeography

Abstract:  The taxonomy, biostratigraphical and palaeogeographical distribution of the Lower Ordovician graptolite genus Bryograptus is evaluated. Bryograptus is recognized as a distinct triradiate anisograptid with a multiramous, pendent rhabdosome. The species of the genus Bryograptus can be interpreted as shallow water faunal elements with a strongly limited biogeographical distribution to the Atlantic Faunal Realm. Bryograptus is restricted to a narrow interval in the Upper Tremadocian, the Bryograptus Biozone of Scandinavia and South America (Argentina), making it a taxon with a high potential for precise biostratigraphical correlations. The proximal end development can be used to differentiate the genus Bryograptus from other pendent multiramous graptoloid genera with a homoplastic rhabdosome development. Characteristics of the proximal end development and structure easily differentiate these genera in relief specimens, but not in flattened material.     

The population genetics of a biological control introduction: mitochondrial DNA and microsatellie variation in native and introduced populations of Aphidus ervi, a parisitoid wasp

Introductions of biological control agents may cause bottlenecks in population size despite efforts to avoid them. We examined the population genetics of Aphidius ervi (Hymenoptera: Braconidae), a parasitoid that was introduced to North America from Western Europe in 1959 to control pea aphids. To explore the phylogeographical relationships of A. ervi we sequenced 1249 bp of mitochondrial DNA (mtDNA) from 27 individuals from the native range and 51 individuals from the introduced range. Most individuals from Western Europe, the Middle East and North America shared one of two common haplotypes, consistent with the known history of the introduction. However, some A. ervi from the Pacific Northwest have a haplotype that is most similar to haplotypes found in Japan, raising the possibility of a second accidental introduction. To examine population structure and assess whether a bottleneck occurred upon introduction to North America, we assayed variation at 5 microsatellite loci in 62 individuals from 2 native populations and 230 individuals from 6 introduced populations. Introduced samples had fewer rare alleles than native samples (F1,34 = 13.5, P = 0.0008), but heterozygosity did not differ significantly. These results suggest that a mild bottleneck occurred in spite of the introduction of over 1000 individuals. Using a hierarchical Bayesian approach, the founding population size was estimated to be 245 individuals. amova showed significant genetic differentiation between the European and North American samples, and a Bayesian assignment approach clustered individuals into four groups, with most European individuals in one group and most North American individuals in the other three. These results highlight that genetic changes are associated with founder events in rapidly growing natural populations, even when the founding population size is relatively large.