Pleistocene extinctions as drivers of biogeographical patterns on the easternmost Canary Islands

Subtropical islands are often viewed as refuges where Quaternary climatic shifts driving global episodes of extinction were buffered. Island biodiversity, however, may have been impacted by climatic fluctuations at local scales, particularly in spatially heterogeneous island systems. In this study, we generated a conceptual framework for predicting the potential impact of Pleistocene extinctions on the biogeographical pattern of the Canarian spermatophyte flora, with a focus on the easternmost Canarian islands (ECI). Then, we performed an exhaustive bibliographic revision (270 studies) to examine whether taxonomic, phylogenetic and phylogeographical data support our predictions. Although molecular information is limited for many lineages, the available data suggest that the majority of extant ECI plant taxa may be the result of relatively recent (<1 Ma) dispersal from surrounding insular and mainland areas. Different lines of evidence are compatible with the idea of a Pleistocene period of frequent lineage extirpation on ECI. Extinction may thus have provided new ecological opportunities for recent (re)colonization, with some cases of recent establishment mediated by facilitation. Considering background extinction on ECI, we describe five general patterns of colonization for Canarian plant lineages. In addition to factors related to island ontogeny and long‐distance dispersal, we suggest that Pleistocene extinctions may have significantly contributed to extant biogeographical patterns in the Canarian archipelago, such as the biased distribution ranges of island plants and the low endemic richness on ECI. This new scenario provides testable hypotheses for future studies dealing with the phylogeography, taxonomy and conservation of terrestrial biodiversity on the Canarian islands, and possibly, on other near‐shore islands.     

A taxonomic,biogeographical and ecological overview of invasive woody plants

Abstract. Invasive plants are considered to be a major threat to the earth's biodiversity, but have not been sufficiently investigated. To address this problem a relational database on invasive woody plants has been set up. It is based on 2045 bibliographical references and contains records on 653 species representing 110 families. The families with the largest number of invasive species are: Rosaceae, Mimosaceae, Papilionaceae and Pinaceae. Out of 1060 recorded invasive events an equal number are reported from continents and oceanic islands. The highest number of invasive woody species are recorded from Europe, followed by the Pacific islands, North America, New Zealand, Australia, Indian Ocean islands and southern Africa. Included in these regions are areas which have fewer highly invasive species, e.g. islands on continental shelves, such as the British Isles. Although most invasions occur in disturbed habitats, most natural communities are susceptible to woody plant invasions. Data on species attributes are only available for a minority of species but indicate that invasive woody plant species may be either insect or wind-pollinated, have a wide array of fruit types, fruit and seed sizes, number of seeds per fruit and dispersal agents. The relative frequency of several attributes varies with the degree of invasiveness. Thus it is not yet possible to determine which set of attributes favours invasiveness and therefore it is difficult to make predictions.     

Extended phylogeny of Aquilegia: the biogeographical and ecological patterns of two simultaneous but contrasting radiations

Studies of the North American columbines (Aquilegia, Ranunculaceae) have supported the view that adaptive radiations in animal-pollinated plants proceed through pollinator specialisation and floral differentiation. However, although the diversity of pollinators and floral morphology is much lower in Europe and Asia than in North America, the number of columbine species is similar in the three continents. This supports the hypothesis that habitat and pollinator specialisation have contributed differently to the radiation of columbines in different continents. To establish the basic background to test this hypothesis, we expanded the molecular phylogeny of the genus to include a representative set of species from each continent. Our results suggest that the diversity of the genus is the result of two independent events of radiation, one involving Asiatic and North American species and the other involving Asiatic and European species. The ancestors of both lineages probably occupied the mountains of south-central Siberia. North American and European columbines are monophyletic within their respective lineages. The genus originated between 6.18 and 6.57 million years (Myr) ago, with the main pulses of diversification starting around 3 Myr ago both in Europe (1.25–3.96 Myr ago) and North America (1.42–5.01 Myr ago). The type of habitat occupied shifted more often in the Euroasiatic lineage, while pollination vectors shifted more often in the Asiatic-North American lineage. Moreover, while allopatric speciation predominated in the European lineage, sympatric speciation acted in the North American one. In conclusion, the radiation of columbines in Europe and North America involved similar rates of diversification and took place simultaneously and independently. However, the ecological drivers of radiation were different: geographic isolation and shifts in habitat use were more important in Europe while reproductive isolation linked to shifts in pollinator specialisation additionally acted in North America.     

Naturalization of alien plants in China

Naturalization (the establishment of a self-sustaining population for at least a decade) is a fundamental precondition for plant invasion and so compiling a complete inventory of naturalized alien species is necessary for predicting and hence preventing such invasion. However, nationwide information on naturalized plants in China is still lacking. We compiled a nationwide list of the naturalized plant species of China, based on various literature reports. The list comprised a total of 861 naturalized plant species belonging to 110 families and 465 genera. The three most dominant families were Compositae, Poaceae, and Leguminosae, accounting for 16, 13 and 12% of naturalized plants, respectively. Among genera, Euphorbia and Solanum had the most naturalized species, followed by Ipomoea, Amaranthus, Oenothera, and Trifolium. Over half of all aliens were of American origin (52%), followed by those with European (14%) and Asian (13%) origins. Annuals and perennial herbs were prevalent among naturalized species; comparison to other studies suggests however that the invasive potential is higher among plants with longer life cycles than those of annuals. The taxonomic pattern of plant naturalization in China is similar to patterns worldwide. However, the low proportion of naturalized plants within the Chinese flora overall suggests that the potential for plant invasions in China may be high. Therefore, greater attention should be focused on naturalization of alien plants in China, especially concerning species of dominant families or genera, and those with a perennial life cycle.     

Coral species richness: ecological versus biogeographical influences

Species richness in communities varies with habitat area, productivity, disturbance level, intensity of species interactions, and regional/historical effects. All of these factors influence coral richness but their effects vary with spatial scale, position on the reef, and regional location. Species richness of corals along depth gradients shows a unimodal, hump-shaped curve that peaks at intermediate depths. Moreover, the peak of the curve is higher in regions with larger species pools. This “regional enrichment” of the local community appears in line transect samples as small as 10 m in length. The pattern suggests that ecological factors operating over scales of tens of meters and regional/historical factors operating over thousands of kilometers can both affect local richness. Regional factors probably include differences in speciation relative to extinction rates among regions and proximity of local sites to richness hotspots. Plausible factors operating at the local scale are species interactions, disturbance, and productivity which combine in different ways to produce the unimodal pattern. Shallow areas support few species because extinction rates are high due to frequent disturbance or because of environmental extremes. In addition, high productivity encourages rapid growth and thus the potential for intense interspecific competition. In areas where branching acroporids are abundant, exclusion by these dominant competitors is possible. Deep areas may be depauperate because few species can tolerate the low light levels found there. Areas of intermediate depth have the richest communities because they are open for colonization by many species and because extinction rates are low. Several theories may explain this “openness” and species persistence: 1. Occasional disturbance coupled with low growth rates results in glacially slow exclusion by the dominant competitor. 2. Aggregation of corals creates spatial variation in the intensity of competition and thus refuges from competition within a spatial landscape. Inferior competitors persist because they are superior at dispersal and refuge colonization. 3. Specialist predators focus on high-density juvenile populations near the parent, creating ecological space for colonization by non-prey. 4. Coral competitive abilities are roughly equal and recruitment into the community is a probabilistic event. The community thus exhibits random drift and exclusion is an extremely lengthy process. Based upon empirical evidence, these theories are listed in order of plausibility, but still need to be rigorously tested. Accepted: 9 September 1999     

Naturalization and invasion of alien plants: concepts and definitions

Abstract.  Much confusion exists in the English-language literature on plant invasions concerning the terms 'naturalized' and 'invasive' and their associated concepts. Several authors have used these terms in proposing schemes for conceptualizing the sequence of events from introduction to invasion, but often imprecisely, erroneously or in contradictory ways. This greatly complicates the formulation of robust generalizations in invasion ecology.
Based on an extensive and critical survey of the literature we defined a minimum set of key terms related to a graphic scheme which conceptualizes the naturalization/invasion process. Introduction means that the plant (or its propagule) has been transported by humans across a major geographical barrier. Naturalization starts when abiotic and biotic barriers to survival are surmounted and when various barriers to regular reproduction are overcome. Invasion further requires that introduced plants produce reproductive offspring in areas distant from sites of introduction (approximate scales: > 100 m over < 50 years for taxa spreading by seeds and other propagules; > 6 m/3 years for taxa spreading by roots, rhizomes, stolons or creeping stems). Taxa that can cope with the abiotic environment and biota in the general area may invade disturbed, seminatural communities. Invasion of successionally mature, undisturbed communities usually requires that the alien taxon overcomes a different category of barriers.
We propose that the term 'invasive' should be used without any inference to environmental or economic impact. Terms like 'pests' and 'weeds' are suitable labels for the 50–80% of invaders that have harmful effects. About 10% of invasive plants that change the character, condition, form, or nature of ecosystems over substantial areas may be termed 'transformers'.     

Chrysophyte cyst biogeographical and ecological distributions: a synthesis

The distributions of chrysophyte stomatocysts in the sediments of 221 lakes from four study areas located in British Columbia, Canada (two datasets), New York, U.S.A., and south central Siberia were examined. The combined lake set spanned a wide range of chemical and morphometric variables. Approximately 400 cyst morphotypes were identified, from which we selected sixty-three common types for further analysis. Canonical ordination (detrended canonical correspondence analysis (DCCA) and constrained DCCA) showed that four environmental variables (pH, conductivity, depth and total phosphorus (TP) ) were significant (Monte Carlo permutation test, P =0.01 for all four variables) and independent determinants of the cyst distributions. Weighted averaging (WA) inference models were developed for each variable. The best models, in terms of their r ², were developed for pH and conductivity ( r ²=0.78 and 0.73, respectively). The models for depth and TP were less robust ( r ²=0.47 and 0.46, respectively). We discuss the relative importance of conductivity and salinity, lakewater acidity and metal concentrations, lake morphometry and trophic status as determinants of chrysophyte population distributions, and possible causal mechanisms for each variable. We identify assemblages of cysts that appear to be associated with six different groups of lakes: (1) very acidic (pH <6.5) lakes; (2) acidic (pH <7.0), softwater lakes; (3) circumneutral (pH 7–8), oligo- to mesotrophic lakes; (4) alkaline, hardwater, meso- to eutrophic lakes; (5) shallow, alkaline (pH 8–8.5), meso- to eutrophic lakes; and (6) shallow, very alkaline (pH >8.5) lakes, including sub- and hyposaline lakes. Finally, we suggest further possible avenues for research that will improve the usefulness of chrysophyte stomatocysts as paleolimnological indicators.     

SitePainter: a tool for exploring biogeographical patterns

As microbial ecologists take advantage of high-throughput analytical techniques to describe microbial communities across ever-increasing numbers of samples, the need for new analysis tools that reveal the intrinsic spatial patterns and structures of these populations is crucial. Here we present SitePainter, an interactive graphical tool that allows investigators to create or upload pictures of their study site, load diversity analyses data and display both diversity and taxonomy results in a spatial context. Features of SitePainter include: visualizing α -diversity, using taxonomic summaries; visualizing β -diversity, using results from multidimensional scaling methods; and animating relationships among microbial taxa or pathways overtime. SitePainter thus increases the visual power and ability to explore spatially explicit studies. AVAILABILITY: https://sourceforge.net/projects/sitepainter SUPPLEMENTARY INFORMATION: Supplementary data are available at Bioinformatics online. CONTACT: antoniog@colorado.edu, Rob.Knight@colorado.edu.     

Molecules and migration: biogeographical studies in cruciferous plants

In the past two decades our understanding of plant biogeography has been improved substantially by the introduction of various molecular marker systems. Especially within the angiosperms, maternally inherited chloroplast DNA based data sets have elucidated not only genetic relatedness but also geographic structuring of genetic variation. These findings were based on the observation that DNA molecules might mutate during migration, which consequently found its manifestation in the term phylogeography introduced in the late 80s by John Avise. However, other markers such as codominantly inherited allozymes were used before the advent of DNA techniques and were used in theoretical population genetic studies. In actual phylogeographic studies, highly variable markers, such as AFLPs (amplified fragment length polymorphisms), were needed to unravel recent species histories (e.g. pleistocenic differentiation). The levels of molecular variation at such markers are closer to that of allelic variation measured with allozymes. Hence, an increasing number of studies have relied on highly polypmorphic markers, such as DNA microsatellite loci. Herein, we try to present an overview on the various biogeographic and phylogeographic studies using various molecular (including isozyme) markers and methodological approaches to analyse them, concentrating on studies done with representatives of the Brassicaceae family.     

A biogeographical perspective on ecological systems: some personal reflections

During my graduate studies, I characterized patterns of geographical distribution and taxonomic differentiation in birds of the West Indies, which suggested that species undergo phases of expansion and contraction similar to the taxon cycles that E. O. Wilson had described for Melanesian ants. Fieldwork in the early 1970s with George Cox confirmed that these phases were associated with variation in habitat distribution and abundance on individual islands, tying together local ecology and biogeography. Because taxon‐cycle stage was independent of taxonomic or ecological relationships among birds of the West Indies, George and I postulated that whether a species was in a phase of expansion or contraction reflected the outcome of coevolved relationships with antagonists, including pathogens. The taxon cycle concept had a cool reception initially, but subsequent phylogeographical analyses, beginning in the early 1990s with Eldredge Bermingham, provided a time scale that confirmed the relationship between taxon cycle stage and the relative age of the most recent population expansion. The discrete nature of islands allows one to visualize taxon cycles in island systems, but the principle should apply in a continental biota as well. The absence of strong phylogenetic effects in distribution and abundance is consistent with evolutionary lability caused by coevolutionary outcomes with specialized antagonists. Related species appear to compete for resources on a more‐or‐less equal footing across a broad range of environments, and their distribution at any particular time is likely to be determined primarily by their relationships with pathogens, among other antagonists. This model of distribution and abundance within a regional community is consistent with much of what we know about the interactions between pathogens and their host populations, but testing the model will require the development of a new research programme focused on endemic pathogen effects in natural communities.     

Permian ammonoids: Biostratigraphic,biogeographical, and ecological analysis

This monograph analyzes all available data on the biostratigraphic and biogeographical distribution of Permian ammonoids and the ecological interpretation of morphological changes in this group. Ammonoid assemblages of various regions at successive stages of the Permian are examined. Four major stages are substantiated in the evolution of Permian ammonoids. The most contentious stages in the Permian evolution of the group are extensively discussed.     

Liverworts of southwest Ethiopian montane forests: ecological and biogeographical notes

Abstract

Ethiopia has diverse topographic features and climatic conditions with a diverse flora. The liverwort flora of southwest Ethiopia is practically unknown, despite a favourable climate and the occurrence of suitable ecosystems such as montane rainforests. During an ecological study of diversity patterns of bryophytes and vascular plants in relation to land use, we recorded many bryophyte species. In this paper we report the finding of 89 species of liverworts, and give short ecological notes and describe the distribution (locally and in Africa) for each species. Of these, 51 species are newly reported from Ethiopia. It is thus obvious that Ethiopia is considerably richer in liverworts than might be expected from previous checklists.     

Nestedness of Southern Ocean island biotas: ecological perspectives on a biogeographical conundrum

Aim To use patterns of nestedness in the indigenous and non‐indigenous biotas of the Southern Ocean islands to determine the influence of dispersal ability on biogeographical patterns, and the importance of accounting for variation in dispersal ability in their subsequent interpretation, especially in the context of the Insulantarctic and multi‐regional hypotheses proposed to explain the biogeography of these islands. Location Southern Ocean islands. Methods Nestedness was determined using a new metric, d1 (a modification of discrepancy), for the indigenous and introduced seabirds, land birds, insects and vascular plants of 26 Southern Ocean islands. To assess the possible confounding effects of spatial autocorrelation on the results, islands were assigned to 11 major island groups and each group was treated as a single island in a following analysis. In addition, nestedness of the six Southern Ocean islands comprising the South Pacific Province (New Zealand islands) was analysed. All analyses were conducted for species and genera, for each of the taxa on its own, and for the complete data sets. Results Statistically significant nestedness was found in all of the taxa examined, with nestedness declining in the order seabirds > land birds > vascular plants > insects for the indigenous species. Vagility had a marked influence on nestedness and the biogeographical patterns shown by the indigenous species. This influence was borne out by additional analyses of marine taxa and small‐sized terrestrial species, both of which were more nested than the most nested group examined here, the seabirds. Assemblages of non‐indigenous species also showed nestedness, and nestedness was generally more pronounced than in the indigenous species. Surprisingly, vagility had a significant effect on nestedness in these assemblages too. Main conclusions Nestedness analyses provide a quantitative means of comparing biogeographical patterns for groups differing in vagility. These comparisons revealed that vagility has a considerable influence on biogeographical patterns and should be taken into account in analyses. Here, investigations of more vagile taxa support hypotheses for a single origin of the Southern Ocean island biota (the Insulantarctica scenario), whilst those of less mobile taxa support the more commonly held, multi‐regional hypothesis. All biogeographical analyses across the Southern Ocean (and elsewhere) will be influenced by the effects of dispersal ability, with composite analyses dominated by sedentary groups likely to favour multi‐regional scenarios, and those dominated by mobile groups favouring single origins. Mechanisms underlying nestedness in the region range from nested physiological tolerances in more mobile groups to colonization ability and patterns of speciation in less vagile taxa. Considerable nestedness in the non‐indigenous assemblages is largely a consequence of the fact that many of these species are European weedy species.     

Drift in the tropics: Phylogenetics and biogeographical patterns in Combretaceae

Aim

The aim of this study was to further advance our understanding of the species-rich, and ecologically important angiosperm family Combretaceae to provide new insights into their evolutionary history. We assessed phylogenetic relationships in the family using target capture data and produced a dated phylogenetic tree to assess fruit dispersal modes and patterns of distribution.

Location

Tropical and subtropical regions.

Time Period

Cretaceous to present.

Major Taxa Studied

Family Combretaceae is a member of the rosid clade and comprises 10 genera and more than 500 species, predominantly assigned to genera Combretum and Terminalia, and occurring on all continents and in a wide range of ecosystems.

Methods

We use a target capture approach and the Angiosperms353 universal probes to reconstruct a robust dated phylogenetic tree for the family. This phylogenetic framework, combined with seed dispersal traits, biome data and biogeographic ranges, allows the reconstruction of the biogeographical history of the group.

Results

Ancestral range reconstructions suggest a Gondwanan origin (Africa/South America), with several intercontinental dispersals within the family and few transitions between biomes. Relative abundance of fruit dispersal types differed by both continent and biome. However, intercontinental colonizations were only significantly enhanced by water dispersal (drift fruit), and there was no evidence that seed dispersal modes influenced biome shifts.

Main Conclusions

Our analysis reveals a paradox as drift fruit greatly enhanced dispersal distances at intercontinental scale but did not affect the strong biome conservatism observed.     

Distributional patterns of Neotropical seasonally dry forest birds: a biogeographical regionalization

Neotropical seasonally dry forests (NSDFs) are widely distributed and possess high levels of species richness and endemism; however, their biogeography remains only partially understood. Using species distribution modelling and parsimony analysis of endemicity, we analysed the distributional patterns of the NSDF avifauna in order to identify their areas of endemism and provide a better understanding of the historical relationships among those areas. The strict consensus trees revealed 17 areas of endemism for NSDFs, which involve four large regions: Baja California, Caribbean–Antilles islands, Mesoamerica and South America. These well-resolved clades are circumscribed by geographical and ecological barriers associated with the Gulf of California, the leading edge of the Caribbean plate, the Tehuantepec Isthmus, the Polochic–Motagua fault, the Nicaragua Depression, the Chocó forest, the Amazon basin and the Andean Cordillera. Relationships among groups of NSDFs found here suggest that evolution of their avifauna involved a mixture of vicariance and dispersal events. Our results support the idea of independent diversification patterns and biogeographical processes in each region, including those previously associated with the Pleistocene Arc Hypothesis for NSDFs of south-eastern South America. This study provides a biogeographical framework to open new lines of research related to the biotic diversification of NSDFs.     

Analysis of introduction history of invasive plants in Brazil reveals patterns of association between biogeographical origin and reason for introduction

Knowledge of historical factors associated with biological invasions in a region can help identify source regions, vectors and pathways more likely to originate potential invaders as well as prioritize resource allocation for selective prevention and early detection strategies. In Brazil, little is known about the introduction history of many invasive plant species, and analyses of historical factors associated with invasions are lacking. To fill this gap, I used a dataset of 117 invasive alien plants across 13 habitats in Brazil to identify potential patterns of continent of origin, reason for introduction, and to test the hypotheses that (i) more Eurasian species are invasive in Brazil than species native from other continents, that (ii) more horticultural species are invasive in Brazil than species introduced for other reasons, and that (iii) continent of origin and reason for introduction are associated. I found that significantly more invasive plant species in Brazil are native to Africa and Asia, were introduced for horticulture and forage, and are part of the families Poaceae, Fabaceae, and Pinaceae. I also found a significant association between continent of origin and reason for introduction, with more invasive species than average being African forage grasses and Asian agroforestry or ornamental plants. In conclusion, the results suggest that the current invasive alien flora of Brazil results from the combination of patterns of recent human migration waves and deliberate species introductions for technological and commercial reasons. These results can help prioritize invasive species and vectors in prevention, early detection, and control strategies.