The concept of the taxon cycle in biogeography

Taxon cycles are sequential phases of expansion and contraction of the ranges of species, associated generally with shifts in ecological distribution. The important contribution of the taxon cycle to biogeographical analysis is its emphasis on evolutionary and ecological interactions among colonizing and resident species, which influence their extinction dynamics and establish patterns of geographical distribution. Taxon cycles were inferred originally from the distribution of species across island archipelagos, where a correlation was noted between gaps in island occupancy and the degree of phenotypic differentiation. This pattern implied that phases of colonization were followed by range contraction, while endemic Antillean species that were undifferentiated between islands suggested secondary expansion and the beginning of a new cycle. This interpretation was met with scepticism, but reconstruction of phylogenetic relationships from gene sequences has now permitted us to characterize taxon cycles in Lesser Antillean birds. The relative timing of phases of the cycle can be deduced from genetic divergence between island populations. We have found that taxon cycles have periods in the order of 10⁶ years and that cycles in different lineages occur independently of each other and independently of Pleistocene climate cycles. Individual island populations may persist for several millions of years on the larger islands of the Lesser Antilles; occasional expansion phases lead to the replacement of island populations that have disappeared, thus reducing the archipelago‐wide rate of extinction to nil. What drives taxon cycles is unknown, but we speculate that they may be caused by co‐evolution with enemy populations, and a probable mechanism would involve infrequent mutations influencing parasite virulence and avian host disease resistance. Taxon cycles undoubtedly occur on continents, but the geographical configuration of island archipelagos reveals more clearly their presence and invites their study.     

Intrinsic dynamics of the regional community

Patterns of diversity within large regional biotas express the outcomes of processes, operating on both regional and local scales, that influence evolutionary diversification as well as the distribution and abundance of species. Regional analyses of species distributions suggest that neither ecological sorting of species based on their adaptations to the physical environment, nor interactions between competing species, adequately explain patterns of species richness. Potentially competing species appear to utilise broadly overlapping resources with similar proficiency. Phylogenetic and phylogeographic analyses reveal that species abundances and distributions within regions vary independently of evolutionary relationship. This implies the existence of dynamic, species‐specific controls on population growth, as could be applied by specialised pathogens or other antagonists. Here, I argue that the changing balance of coevolved interactions between hosts and their antagonists shapes the distribution and abundance of individual host populations as well as patterns of local species richness. Geographical expansion creates allopatric populations and thereby could promote diversification; contraction ultimately leads to extinction. This taxon‐cycle dynamic links regional diversity and distribution to intrinsic biological interactions independently of extrinsic ecological conditions. These hypotheses emphasise the central importance of investigating the impacts of pathogens on species abundance and distribution, and the potential consequences of coevolutionary changes in pathogen‐host relationships for species formation and extinction.     

Orchids of the West Indies: predictability of diversity and endemism

Aim We examined phytogeographical patterns of West Indian orchids, and related island area and maximum elevation with orchid species richness and endemism. We expected strong species–area relationships, but that these would differ between low and montane island groups. In so far as maximum island elevation is a surrogate for habitat diversity, we anticipated a strong relationship with maximum elevation and both species richness and endemism for montane islands. Location The West Indies. Methods Our data included 49 islands and 728 species. Islands were classified as either montane (≥ 300 m elevation) or low (< 300 m). Linear and multivariate regression analyses were run to detect relationships between either area or maximum island elevation and species richness or the number of island endemic species. Results For all 49 islands, the species–area relationship was strong, producing a z‐value of 0.47 (slope of the regression line) and explaining 46% of the variation. For 18 relatively homogeneous, low islands we found a non‐significant slope of z = −0.01 that explained only 0.1% of the variation. The 31 montane islands had a highly significant species–area relationship, with z = 0.49 and accounting for 65% of the variation. Species numbers were also strongly related to maximum island elevation. For all islands < 750 km², we found a small‐island effect, which reduced the species–area relationship to a non‐significant z = 0.16, with only 5% of the variation explained by the model. Species–area relationships for montane islands of at least 750 km² were strong and significant, but maximum elevation was the best predictor of species richness and accounted for 79% of the variation. The frequency of single‐island endemics was high (42%) but nearly all occurred on just nine montane islands (300 species). The taxonomic distribution of endemics was also skewed, suggesting that seed dispersability, while remarkable in some taxa, is very limited in others. Montane island endemics showed strong species–area and species–elevation relationships. Main conclusions Area and elevation are good predictors of orchid species diversity and endemism in the West Indies, but these associations are driven by the extraordinarily strong relationships of large, montane islands. The species richness of low islands showed no significant relationship with either variable. A small‐island effect exists, but the montane islands had a significant relationship between species diversity and maximum elevation. Thus, patterns of Caribbean orchid diversity are dependent on an interplay between area and topographic diversity.     

The functional biogeography of species: biogeographical species roles of birds in Wallacea and the West Indies

Biogeographical systems can be analyzed as networks of species and geographical units. Within such a biogeographical network, individual species may differ fundamentally in their linkage pattern, and therefore hold different topological roles. To advance our understanding of the relationship between species traits and large‐scale species distribution patterns in archipelagos, we use a network approach to classify birds as one of four biogeographical species roles: peripherals, connectors, module hubs, and network hubs. These roles are based upon the position of species within the modular network of islands and species in Wallacea and the West Indies. We test whether species traits – including habitat requirements, altitudinal range‐span, feeding guild, trophic level, and body length – correlate with species roles. In both archipelagos, habitat requirements, altitudinal range‐span and body length show strong relations to species roles. In particular, species that occupy coastal‐ and open habitats, as well as habitat generalists, show higher proportions of connectors and network hubs and thus tend to span several biogeographical modules (i.e. subregions). Likewise, large body size and a wide altitudinal range‐span are related to a wide distribution on many islands and across several biogeographical modules. On the other hand, species restricted to interior forest are mainly characterized as peripherals and, thus, have narrow and localized distributions within biogeographical modules rather than across the archipelago‐wide network. These results suggest that the ecological amplitude of a species is highly related to its geographical distribution within and across bio geographical subregions and furthermore supports the idea that large‐scale species distributions relate to distributions at the local community level. We finally discuss how our biogeographical species roles may correspond to the stages of the taxon cycle and other prominent theories of species assembly.     

The West Indies as a laboratory of biogeography and evolution

Islands have long provided material and inspiration for the study of evolution and ecology. The West Indies are complex historically and geographically, providing a rich backdrop for the analysis of colonization, diversification and extinction of species. They are sufficiently isolated to sustain endemic forms and close enough to sources of colonists to develop a dynamic interaction with surrounding continental regions. The Greater Antilles comprise old fragments of continental crust, some very large; the Lesser Antilles are a more recent volcanic island arc, and the low-lying Bahama Islands are scattered on a shallow oceanic platform. Dating of island lineages using molecular methods indicates over-water dispersal of most inhabitants of the West Indies, although direct connections with what is now southern Mexico in the Early Tertiary, and subsequent land bridges or stepping stone islands linking to Central and South America might also have facilitated colonization. Species-area relationships within the West Indies suggest a strong role for endemic radiations and extinction in shaping patterns of diversity. Diversification is promoted by opportunities for allopatric divergence between islands, or within the large islands of the Greater Antilles, with a classic example provided by the Anolis lizards. The timing of colonization events using molecular clocks permits analysis of colonization-extinction dynamics by means of species accumulation curves. These indicate low rates of colonization and extinction for reptiles and amphibians in the Greater Antilles, with estimated average persistence times of lineages in the West Indies exceeding 30Myr. Even though individual island populations of birds might persist an average of 2Myr on larger islands in the Lesser Antilles, recolonization from within the archipelago appears to maintain avian lineages within the island chain indefinitely. Birds of the Lesser Antilles also provide evidence of a mass extinction event within the past million years, emphasizing the time-heterogeneity of historical processes. Geographical dynamics are matched by ecological changes in the distribution of species within islands over time resulting from adaptive radiation and shifts in habitat, often following repeatable patterns. Although extinction is relatively infrequent under natural conditions, changes in island environments as a result of human activities have exterminated many populations and others--especially old, endemic species--remain vulnerable. Conservation efforts are strengthened by recognition of aesthetic, cultural and scientific values of the unique flora and fauna of the West Indies.     

Correlates of extinction risk of birds from two Indonesian islands

Size of distributional range, position in the range, body size and diet are some of the ecological traits that may correlate with local abundance. Evolutionary phenomena such as taxon cycles, acting over much greater time periods, may also influence abundance and promote species extinction. This paper assesses which of a wide range of ecological and historic traits best predict the variation in abundance of tropical forest birds on Sumba and Buru islands in Wallacea (Indonesia). In addition we seek to determine which traits predict species' ability to adapt to secondary or logged forest. The most important correlates of both abundance and ability to transfer were those related to the evolutionary history of the species within the Wallacean Archipelago and not the traits that were more directly related to species ecology. These relationships are maintained when allowance is made for phylogenetic relationships. Our interpretation of the results is that recent colonists to an island are initially rare in the indigenous forest habitat but concomitant with an adaptation to local conditions they gradually become more abundant and taxonomically distinct from other populations of the same species. These results apparently contradict the taxon cycle hypothesis but this may be a result of our focus on indigenous forest habitats rather than on a wider range dominated by anthropogenic ones.     

History and taxonomy: their roles in the core-satellite hypothesis

Metapopulation models are important in explaining the distribution and abundance of species through time and space. These models combine population dynamics with stochastic variation in extinction and immigration parameters associated with local populations. One of the predictions of metapopulation models is a bimodal distribution of species frequency of occurrence, a pattern that led to the development of the core-satellite species hypothesis. The spatial scale and taxonomic classification of past core-satellite studies has often been undefined. In our study, we have integrated metapopulation dynamics with the roles that differential dispersal ability and history play in the shaping of communities. The differences in distribution patterns between landbridge islands and oceanic islands, and among various taxa (birds, mammals, herptiles, arthropods, fish, and plants) are analyzed. The majority of landbridge islands comprised locally and regionally abundant species (core species), whereas the majority of oceanic islands had a uniform distribution (or no end-peak in their distribution). The patterns of distribution among the taxonomic groups also showed differences. Birds (good dispersers) consistently showed bimodal- and core-distribution patterns. The bimodal prediction of species distribution is best exemplified in the landbridge islands and in birds, and least in oceanic islands and in organisms other than birds. These results illustrate the importance of testing models with various taxonomic groups and at different spatial scales and defining these scales before formally testing the predictions of the models.     

Invasive plant species in the West Indies: geographical,ecological, and floristic insights

The level of invasion (number or proportion of invasive species) in a given area depends on features of the invaded community, propagule pressure, and climate. In this study, we assess the invasive flora of nine islands in the West Indies to identify invasion patterns and evaluate whether invasive species diversity is related to geographical, ecological, and socioeconomic factors. We compiled a database of invasive plant species including information on their taxonomy, origin, pathways of introduction, habitats, and life history. This database was used to evaluate the similarity of invasive floras between islands and to identify invasion patterns at regional (West Indies) and local (island) scales. We found a total of 516 alien plant species that are invasive on at least one of the nine islands studied, with between 24 to 306 invasive species per island. The invasive flora on these islands includes a wide range of taxonomic groups, life forms, and habitats. We detected low similarity in invasive species diversity between islands, with most invasive species (>60%) occurring on a single island and 6% occurring on at least five islands. To assess the importance of different models in predicting patterns of invasive species diversity among islands, we used generalized linear models. Our analyses revealed that invasive species diversity was well predicted by a combination of island area and economic development (gross domestic product per capita and kilometers of paved roadways). Our results provide strong evidence for the roles of geographical, ecological, and socioeconomic factors in determining the distribution and spread of invasive species on these islands. Anthropogenic disturbance and economic development seem to be the major drivers facilitating the spread and predominance of invasive species over native species.     

Distribution anomalies in avian haemosporidian parasites in the southern Lesser Antilles

We compared the haemosporidian parasite faunas (Plasmodium and Haemoproteus) of small land birds on the islands of St Lucia, St Vincent and Grenada in the southern Lesser Antilles. The islands differ in distance from the South American source of colonists, proximity to each other, and similarity of their avifaunas. On each island, we obtained 419–572 blood samples from 22–25 of the 34–41 resident species. We detected parasite infection by PCR and identified parasite lineages by sequencing a portion of the mitochondrial cytochrome b gene. Parasite prevalence varied from 31% on St Lucia to 22% on St Vincent and 18% on Grenada. Abundant parasite lineages differed between the three islands in spite of the similarity in host species. As in other studies, the geographic distributions of the individual parasite lineages varied widely between local endemism and broad distribution within the West Indies, including cases of long‐distance disjunction. St Vincent was unusual in the near absence of Plasmodium parasites, which accorded with low numbers of suitable mosquito vectors reported from the island. Parasites on St Vincent also tended to be host specialists compared to those on St Lucia and Grenada. Similarity in parasite assemblages among the three islands varied in parallel with host assemblage similarity (but not similarity of infected hosts) and with geographic proximity. Parasite prevalence increased with host abundance on both St Lucia and St Vincent, but not on Grenada; prevalence did not vary between endemic and more widespread host species. In addition, the endemic host species harbored parasites that were recovered from a variety of non‐endemic species as well. These results support the individualistic nature of haemosporidian parasite assemblages in evolutionarily independent host populations.     

Biogeographical patterns of the avifaunas of the Caribbean Basin Islands: a parsimony perspective

We analyzed the avifaunas of the Caribbean islands and nearby continental areas and their relationships using Parsimony Analysis of Endemicity (PAE), in order to assess biogeographical patterns and their concordance with geological and phylogenetic evidence. Using distributional information of birds obtained from published literature, a presence/absence matrix for 695 genera and 2026 species of land and freshwater birds was constructed and analyzed. Three different analyses were performed: for species, for genera, and for species and genera combined. In the combined analysis, the Lesser Antilles appear paraphyletic at the base of the cladogram. Then, two major clades are identified: South America (Andes, Venezuelan lowlands, Dutch West Indies and Trinidad and Tobago) and North America, including the Greater Antilles in a clade that is the sister area to Yucatan and the Central American countries nested from north to south. PAE results support Caribbean vicariant models and cladistic biogeographical hypotheses on area relationships, and show relative congruence with available phylogenetic data. Bird biogeography on the Caribbean islands appears to have been caused by both vicariance and dispersal processes. © The Willi Hennig Society 2007.     

Determinants of bird species richness,endemism, and island network roles in Wallacea and the West Indies: is geography sufficient or does current and historical climate matter?

Island biogeography has greatly contributed to our understanding of the processes determining species' distributions. Previous research has focused on the effects of island geography (i.e., island area, elevation, and isolation) and current climate as drivers of island species richness and endemism. Here, we evaluate the potential additional effects of historical climate on breeding land bird richness and endemism in Wallacea and the West Indies. Furthermore, on the basis of species distributions, we identify island biogeographical network roles and examine their association with geography, current and historical climate, and bird richness/endemism. We found that island geography, especially island area but also isolation and elevation, largely explained the variation in island species richness and endemism. Current and historical climate only added marginally to our understanding of the distribution of species on islands, and this was idiosyncratic to each archipelago. In the West Indies, endemic richness was slightly reduced on islands with historically unstable climates; weak support for the opposite was found in Wallacea. In both archipelagos, large islands with many endemics and situated far from other large islands had high importance for the linkage within modules, indicating that these islands potentially act as speciation pumps and source islands for surrounding smaller islands within the module and, thus, define the biogeographical modules. Large islands situated far from the mainland and/or with a high number of nonendemics acted as links between modules. Additionally, in Wallacea, but not in the West Indies, climatically unstable islands tended to interlink biogeographical modules. The weak and idiosyncratic effect of historical climate on island richness, endemism, and network roles indicates that historical climate had little effects on extinction‐immigration dynamics. This is in contrast to the strong effect of historical climate observed on the mainland, possibly because surrounding oceans buffer against strong climate oscillations and because geography is a strong determinant of island richness, endemism and network roles.     

Avian research in the Caribbean: past contributions and current priorities

ABSTRACT The islands of the Caribbean contain habitat of critical importance to a large number of endemic and resident birds, as well as many overwintering Neotropical migrants, and they rank as a globally outstanding conservation priority ecoregion and biodiversity hotspot. Considerable research from the region has focused on the ecology of permanent resident species, and these studies have had particular significance for threatened species management, especially parrot biology and conservation, but also for tropical community ecology in general. Work by ornithologists in the Caribbean has been instrumental in improving our understanding of the ecology of overwintering Neotropical migrants and in developing long‐term monitoring programs. Although Caribbean‐based studies of birds have resulted in significant contributions in many important areas of ecological research, there is a great need for additional research. Especially needed are studies with application to the management of resident species, and studies of how bird populations may be affected by pathogens, parasites, plants, and other types of biotic interactions. Studies focusing on how bird species and populations are affected by global climate change, and cumulative, landscape‐level changes in land use are also needed. Along with additional research, scientists have an important role to play in building capacity to prepare the next generation of biologists in the region who will need to address mounting challenges related to biodiversity protection. As with many conservation efforts, funding is a critical need for almost all organizations and agencies involved in research, conservation action, and capacity building in the West Indies.     

Phylogeography of the parthenogenic ant Platythyrea punctata: highly successful colonization of the West Indies by a poor disperser

Aim We investigated the spatio‐temporal patterns of genetic diversity in West Indian and mainland populations of a widespread parthenogenic ant (Platythyrea punctata F. Smith) to infer source populations and subsequent colonizations across its geographic range. Location Central America, Texas and the West Indies (Florida, the Bahamas, Greater and Lesser Antilles). Methods We employed phylogeographic reconstruction based on 1451 bp of mitochondrial DNA (cytochrome c oxidase subunits I and II) sequenced from 91 individuals of P. punctata. We employed standard population genetic analyses, Bayesian phylogenetic analyses, haplotype networks and molecular dating methods as performed by beast . We also employed phylogenetic analysis using two nuclear markers (970 bp) to understand the placement of P. punctata in the globally distributed genus Platythyrea. Results Based on highly reduced haplotypic variation and temporal estimates, rapid expansion and dispersal from Central America best explains the observed distribution of haplotypes. Platythyrea punctata successfully invaded the West Indies very few times. One haplotype occurred on every island surveyed from the Bahamas and Florida in the north to Barbados at the southern edge of its range. Haplotype diversity in the West Indies is quite low, despite a larger sample size relative to the mainland. Most mainland colonies collected each possessed a unique haplotype, whereas only Florida and the larger islands (the Dominican Republic, Puerto Rico and Guadeloupe) contained more than one haplotype. Island haplotypes were most similar to haplotypes collected in northern Mexico and southern Texas, but genetic distances were nevertheless high. The putative sister species of P. punctata appears to be an endemic of Hispaniola (P. strenua Wheeler & Mann), even though older, mainland populations of P. punctata are sympatric with at least two other congenerics. Main conclusions Dispersal seems very limited on the mainland, with well‐defined clades corresponding to geographical regions. Colonization of the islands from the mainland was extremely rare, but once successful there were very few barriers to expansion to nearly every island in the West Indies. We hypothesize that this invasion occurred during the late Pleistocene as the climate became warmer and less arid.     

Molecular phylogeny and biogeography of West Indian frogs of the genus Leptodactylus (Anura, Leptodactylidae)

Three endemic species of the aquatic-breeding frog genus Leptodactylus are recognized from the West Indies: Leptodactylus albilabris (Puerto Rico and the Virgin Islands), Leptodactylus dominicensis (Hispaniola), and Leptodactylus fallax (Lesser Antilles). DNA sequences were obtained from several mitochondrial genes to resolve taxonomic questions involving these species and to provide insights into their origin and distribution in the islands. We found low levels of sequence divergence between L. dominicensis and L. albilabris, supporting morphological evidence that the former species is a junior synonym of the latter species. Phylogenetic analysis supported previous species-group allocations, finding that L. albilabris is a member of the fuscus group and L. fallax is a member of the pentadactylus group. Molecular time estimates for the divergence of L. albilabris from its closest relative in South America (24-58 million years ago, Ma) and for L. fallax from its closest relative in South America (23-34Ma) indicate that they colonized the West Indies independently by over-water dispersal in the mid-Cenozoic. The absence of detectable sequence divergence between the two extant populations of L. fallax (Dominica and Montserrat), a species used for human food and now critically endangered, suggests that one or both arose by human introduction from an island or islands where that species originated. The relatively minor genetic differentiation of populations of L. albilabris can be explained by vicariance and dispersal in the Pleistocene and Holocene, although human introduction of some populations cannot be ruled out.     

Biogeographical modules and island roles: a comparison of Wallacea and the West Indies

Aim In order to advance our understanding of the assembly of communities on islands and to elucidate the function of different islands in creating regional and subregional distribution patterns, we identify island biogeographical roles on the basis of the distribution of the islands’ biota within the archipelago. We explore which island characteristics determine island biogeographical roles. Furthermore, we identify biogeographical subregions, termed modules. Location Wallacea in Indonesia, and the West Indies in the Caribbean Sea. Methods We use a network approach to detect island biogeographical roles and avian biogeographical modules. To designate the biogeographical role of an island, each island is assigned two coordinates, l and r. The position of an island in l–r space characterizes its role, namely as peripheral, connector, module hub, or network hub. Island characteristics are tested as predictors of l and r. Results Both Wallacea and the West Indies were found to be significantly modular and divided into four biogeographical modules. The four modules identified within Wallacea each contain all existing island roles, whereas no module in the West Indies represents all possible roles. Island area and elevation appeared to be the most important determinants of an island’s l score, while measurements of isolation essentially determined the r score. Main conclusions In both Wallacea and the West Indies, the geographic structuring into biogeographical modules corresponds well with our knowledge of past connections and contemporary factors. In both archipelagos, large, mountainous islands are identified as hubs and are thus responsible for faunal coherence within modules (module hubs) and across the entire archipelago (network hubs). We thus interpret these as source islands for the surrounding islands in their module (module hubs) or for the entire archipelago (network hubs). Islands positioned marginally in their module and distant from the mainland are identified as connectors or network hubs, behaving as sinks and stepping stones for dispersing species. Modularity and predictors of biogeographical roles are similar for Wallacea and the West Indies, whereas the build‐up of biogeographical modules and the assortment of roles depend on the spatial constellation of islands in each archipelago.     

Regional variation in the historical components of global avian species richness

Aim Using a global data base of the distribution of extant bird species, we examine the evidence for spatial variation in the evolutionary origins of contemporary avian diversity. In particular, we assess the possible role of the timing of mountain uplift in promoting diversification in different regions. Location Global. Methods We mapped the distribution of avian richness at four taxonomic levels on an equal‐area 1° grid. We examined the relationships between richness at successive taxonomic levels (e.g. species richness vs. genus richness). We mapped the residuals from linear regressions of these relationships to identify areas that are exceptional in the number of lower taxa relative to the number of higher taxa. We use generalized least squares models to test the influence of elevation range and temperature on lower‐taxon richness relative to higher‐taxon richness. Results Peaks of species richness in the Neotropics were congruent with patterns of generic richness, whilst peaks in Australia and the Himalayas were congruent with patterns of both genus and family richness. Hotspots in the Afrotropics did not reflect higher‐taxon patterns. Regional differences in the relationship between richness at successive taxonomic levels revealed variation in patterns of taxon co‐occurrence. Species and genus co‐occurrence was positively associated with elevational range across much of the world. Taxon occurrence in the Neotropics was associated with a positive interaction between elevational range and temperature. Conclusions These results demonstrate that contemporary patterns of richness show different associations with higher‐taxon richness in different regions, which implies that the timing of historical effects on these contemporary patterns varies across regions. We suggest that this is due to dispersal limitation and phylogenetic constraints on physiological tolerance limits promoting diversification. We speculate that diversification rates respond to long‐term changes in the Earth's topography, and that the role of tropical mountain ranges is implicated as a correlate of contemporary diversity, and a source of diversification across avian evolutionary history.     

Dimorphism and divergence in island and mainland Anoles

Relative to the West Indies, the ecology and evolution of anoles inhabiting islands off Central and South America have received little attention. The paucity of studies on continental islands has limited our ability to generalize and extend results based on the West Indian paradigm, as well as our understanding of the profound differences between the adaptive radiations of continental vs. Greater Antillean anoles. Here we compare the morphological, ecological, behavioural and genetic divergence between Anolis nebulosus populations inhabiting a small island in the Bay of Chamela, Mexico, and a nearby mainland forest. Notably, the two populations exhibit intra‐sexual dimorphism with respect to head and limb sizes, the first such polymorphism documented for an Anolis species. We also compare the shape of island and mainland A. nebulosus with each other, the six West Indian ecomorphs and a hypothetical generalist species. Finally, we address the generalist convergence hypothesis for anoles on single species islands. We conclude that convergence on a generalist morphology is widespread among solitary anoles in the West Indies. We present data on a limited sample of solitary anoles with mainland ancestors that suggest a parallel convergence on a similar generalist morphology, probably due to similar adaptive landscapes shaped by selective forces common to small island environments.     

Molecular Phylogenetic Studies of Caribbean Palms (Arecaceae) and Their Relationships to Biogeography and Conservation

The Caribbean Islands are one of the world’s 34 biodiversity hotspots, remarkable for its biological richness and the high level of threat to its flora and fauna. The palms (family Arecaceae) are well represented in the West Indies, with 21 genera (three endemic) and 135 species (121 endemic). We provide an overview of phylogenetic knowledge of West Indian Palms, including their relationships within a plastid DNA-based phylogeny of the Arecaceae. We present new data used to reconstruct the phylogeny of tribe Cryosophileae, including four genera found in the West Indies, based on partial sequences of the low-copy nuclear genes encoding phosphoribulokinase (PRK) and subunit 2 of RNA polymerase II (RPB2). Recently published phylogenetic studies of tribe Cocoseae, based on PRK sequences, and tribes Cyclospatheae and Geonomateae, based on PRK and RPB2 sequences, also provide information on the phylogenetic relationships of West Indian palms. Results of these analyses show many independent origins of the West Indian Palm flora. These phylogenetic studies reflect the complex envolutionary history of the West Indies and no single biogeographical pattern emerges for these palms. The present day distributions of West Indian palms suggest complicated evolutionary interchange among islands, as well as between the West Indies and surrounding continents. We identified six palm lineages that deserve conservation priority. Species-level phylogenies are needed for Copernicia, Sabal, and Roystonea before we can build a more complete understanding of the origin and diversification of West Indian palms. An erratum to this article can be found at     

A multi-scale test for dispersal filters in an island plant community

Constraints on plant distributions resulting from seed limitation (i.e. dispersal filters) were evaluated on two scales of ecological organization on islands off the coast of British Columbia, Canada. First, island plant communities were separated into groups based on fruit morphology, and patterns in species diversity were compared between fruit‐type groups. Second, abundance patterns in several common fleshy‐fruited, woody angiosperm species were compared to species‐specific patterns in seed dispersal by birds. Results from community‐level analyses showed evidence for dispersal filters. Dry‐fruited species were rare on islands, despite being common on the mainland. Island plant communities were instead dominated by fleshy‐fruited species. Patterns in seed dispersal were consistent with differences in diversity, as birds dispersed thousands of fleshy‐fruited seeds out to islands, while dry fruited species showed no evidence of mainland‐island dispersal. Results from population‐level analyses showed no evidence for dispersal filters. Population sizes of common fleshy‐fruited species were unrelated to island isolation, as were rates of seed dispersal. Therefore, island isolation distances were not large enough to impose constraints on species’ distributions resulting from seed limitation. Rates of seed dispersal were also unrelated to island area. However, several species increased in abundance with island area, indicating post‐dispersal processes also help to shape species distributions. Overall results suggest that seed dispersal processes play an important role in determining the diversity and distribution of plants on islands. At the community‐level, dry‐fruited species were seed limited and island communities were instead dominated by fleshy‐fruited species. At the population‐level, common fleshy‐fruited species were not seed limited and showed few differences in distribution among islands. Therefore, although evidence for dispersal filters was observed, their effects on plant distributions were scale‐dependent.     

Slaying dragons: limited evidence for unusual body size evolution on islands

Aim Island taxa often attain forms outside the range achieved by mainland relatives. Body size evolution of vertebrates on islands has therefore received much attention, with two seemingly conflicting patterns thought to prevail: (1) islands harbour animals of extreme size, and (2) islands promote evolution towards medium body size (‘the island rule’). We test both hypotheses using body size distributions of mammal, lizard and bird species. Location World‐wide. Methods We assembled body size and insularity datasets for the world’s lizards, birds and mammals. We compared the frequencies with which the largest or smallest member of a group is insular with the frequencies expected if insularity is randomly assigned within groups. We tested whether size extremes on islands considered across mammalian phylogeny depart from a null expectation under a Brownian motion model. We tested the island rule by comparing insular and mainland members of (1) a taxonomic level and (2) mammalian sister species, to determine if large insular animals tend to evolve smaller body sizes while small ones evolve larger sizes. Results The smallest species in a taxon (order, family or genus) are insular no more often than would be expected by chance in all groups. The largest species within lizard families and bird genera (but no other taxonomic levels) are insular more often than expected. The incidence of extreme sizes in insular mammals never departs from the null, except among extant genera, where gigantism is marginally less common than expected under a Brownian motion null. Mammals follow the island rule at the genus level and when comparing sister species and clades. This appears to be driven mainly by insular dwarfing in large‐bodied lineages. A similar pattern in birds is apparent for species within orders. However, lizards follow the converse pattern. Main conclusions The popular misconception that islands have more than their fair share of size extremes may stem from a greater tendency to notice gigantism and dwarfism when they occur on islands. There is compelling evidence for insular dwarfing in large mammals, but not in other taxa, and little evidence for the second component of the island rule – gigantism in small‐bodied taxa.