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.     

Tropical fruitflies (Diptera: Tephritidae) of the Krakatau Archipelago in 1990 and comments on faunistic changes since 1982

Three of the Krakatau islands, in Sunda Strait, some 40 km from both Java and Sumatra, are believed to have been totally devastated by the famous cataclysmic eruption of 1883. The fourth, Anak Krakatau, is an emergent volcanic island which rose from Krakatau's submerged 1883 caldera in 1930, suffered a self-devastating eruption in 1952/3 and several severely damaging ones since then. In 1990 the tephritid fauna was monitored on all the islands with Steiner traps. Nine species were found, five of which appeared to have been found by Yukawa in 1982 (Yukawa 1984) using similar methods and lures, when he noted a total of six species. As in 1982, the fauna in 1990 was characterized by the numerical dominance ofBactrocera albistrigata (de Meijere) although this appeared to be less marked than in 1982 andBactrocera papayae Drew & Hancock appeared to have increased in relative numbers. The distribution of three species over the islands of the archipelago had increased and four species not found in 1982 were present. Rakata, Krakatau's remnant and by far the largest and highest island, carried seven species (six in 1982), Sertung three (two in 1982), Panjang four (one in 1982) and Anak Krakatau four (one in 1982). Capture rates were lower than in 1982 but still higher than those obtained on the Javan mainland in 1982. There was a marked difference between the tephritid faunas of Anak Krakatau's two forelands, which were at different stages of biotic succession.     

Diversity of butterflies on British islands: ecological influences underlying the roles of area, isolation and the size of the faunal source

To distinguish between the influences of area and isolation on the butterfly faunas of British islands two approaches are adopted. First, species richness is related to island area, isolation and the size of the faunal source. Neither area nor isolation account for much variance in species richness, though area is more important than isolation. In contrast, species richness corresponds closely to the size of the faunal source on nearby islands and to that at proximate locations on adjacent mainlands. The second approach relates the incidence of species on islands to their ecological attributes. A very close relationship is found between species incidence on islands and those ecological variables that measure potential for migration and colonization and that resist extinction. The implications are that the majority of British islands in this survey are insufficiently isolated to prevent intermittent migrations of butterflies to them or so small as to generate frequent extinctions. Independent data indicate the capacity of many resident species to migrate distances in excess of the isolation of most of the islands. Some evidence also exists for the long-term survival of species on islands; important considerations in this respect are that most islands in the survey are large compared to habitat patches sustaining species on mainland Britain and that substantial portions of islands are retained in early seral stages or comprise long-lived stable habitats (e.g. peat mosses) that are particularly suitable for many British species.     

Isolation and differentiation of Rivulus hartii across Trinidad and neighboring islands

Diversification of freshwater fishes on islands is considered unlikely because the traits that enable successful colonization—specifically, broad salinity tolerances and the potential for oceanic dispersal—may also constrain post‐colonization genetic differentiation. Some secondary freshwater fish, however, exhibit pronounced genetic differentiation and geographic structure on islands, whereas others do not. It is unclear what conditions give rise to contrasting patterns of differentiation because few comparative reconstructions of population history have been carried out for insular freshwater fishes. In this study, we examined the phylogeography of Hart’s killifish (Rivulus hartii) across Trinidad, with reference to neighboring islands and northern South America, to test hypotheses of colonization and differentiation derived from comparable work on co‐occurring guppies (Poecilia reticulata). Geographic patterns of mitochondrial DNA haplotype variation and microsatellite genotype variation provide evidence of genetic differentiation of R. hartii among islands and across Trinidad. Our findings are largely consistent with patterns of geographically structured ancestry and admixture found in Trinidadian guppies, which suggests that both species share a history of colonization and differentiation and that post‐colonization diversification may be more common among members of insular freshwater fish assemblages than has been previously thought.     

Remarkable new evidence for island radiation in birds

If island biogeography has a sweet spot, it's where islands generate their own species diversity rather than merely taking on mainland immigrants. In birds and other highly dispersive taxa, however, this 'zone of radiation', may be vanishingly small. Darwin's finches and Hawaiian Honeycreepers are among only a handful of examples of island radiation in birds (Price 2008), suggesting that winged powers of dispersal make sufficient isolation from mainland colonists unlikely, while also hindering speciation within and among isolated islands. Nevertheless, two studies in this issue of Molecular Ecology join a string of other recent analyses suggesting that island radiation in birds remains under-appreciated (see also Moyle et al. 2009; Kisel & Barraclough 2010; Rosindell & Phillimore 2011). Melo et al. (2011) use a phylogenetic analysis of white-eyes on islands in the Gulf of Guinea to identify two previously overlooked island radiations, and reveal replicated adaptive divergence on islands where species occur in pairs. Sly et al. (2011), meanwhile, consider possible explanations for speciation and geographic differentiation within a large island, and find the same type of oceanic barriers that are critical to bird speciation across archipelagos may also contribute to divergence that appears to have occurred within a single island.     

Scrophularia arguta,a widespread annual plant in the Canary Islands: a single recent colonization event or a more complex phylogeographic pattern?

Many studies have addressed evolution and phylogeography of plant taxa in oceanic islands, but have primarily focused on endemics because of the assumption that in widespread taxa the absence of morphological differentiation between island and mainland populations is due to recent colonization. In this paper, we studied the phylogeography of Scrophularia arguta, a widespread annual species, in an attempt to determine the number and spatiotemporal origins of dispersal events to Canary Islands. Four different regions, ITS and ETS from nDNA and psbA‐trnH and psbJ‐petA from cpDNA, were used to date divergence events within S. arguta lineages and determine the phylogenetic relationships among populations. A haplotype network was obtained to elucidate the phylogenetic relationships among haplotypes. Our results support an ancient origin of S. arguta (Miocene) with expansion and genetic differentiation in the Pliocene coinciding with the aridification of northern Africa and the formation of the Mediterranean climate. Indeed, results indicate for Canary Islands three different events of colonization, including two ancient events that probably happened in the Pliocene and have originated the genetically most divergent populations into this species and, interestingly, a recent third event of colonization of Gran Canaria from mainland instead from the closest islands (Tenerife or Fuerteventura). In spite of the great genetic divergence among populations, it has not implied any morphological variation. Our work highlights the importance of nonendemic species to the genetic richness and conservation of island flora and the significance of the island populations of widespread taxa in the global biodiversity.