Driven by the West Wind Drift? A synthesis of southern temperate marine biogeography,with new directions for dispersalism

Aim Twentieth century biogeographers developed intriguing hypotheses involving West Wind Drift dispersal of Southern Hemisphere biota, but such models were largely abandoned in favour of vicariance following the development of plate tectonic theory. Here I present a synthesis of southern temperate marine biogeography, and suggest some new directions for phylogeographic research. Location The southern continents, formerly contiguous components of Gondwana, are now linked only by ocean currents driven by the West Wind Drift. Methods While vicariance theory certainly facilitates the development of testable hypotheses, it does not necessarily follow that vicariance explains much of contemporary southern marine biogeography. To overcome the limitations of narratives that simply assume vicariance or dispersal, it is essential for analyses to test biogeographic hypotheses by incorporating genetic, ecological and geological data. Results Recent molecular studies have provided strong evidence for dispersal, but relatively little evidence for the biogeographic role of plate tectonics in distributing southern marine taxa. Despite confident panbiogeographic claims to the contrary, molecular and ecological studies of buoyant macroalgae, such as Macrocystis, indicate that dispersal predominates. Ironically, some of the better supported evidence for marine vicariance in southern waters has little or nothing to do with plate tectonics. Rather, it involves far more localized and recent vicariant models, such as the isolating effect of the Bassian Isthmus during Pleistocene low sea‐level stands (Nerita). Main conclusions Recent phylogeographic studies of southern marine taxa (e.g. Diloma and Parvulastra) imply that passive rafting cannot be ignored as an important mechanism of long‐distance dispersal. I outline a new direction for southern hemisphere phylogeography, involving genetic analyses of bull‐kelp (Durvillaea) and its associated holdfast invertebrate communities.     

Out of the Andes: patterns of diversification in clearwing butterflies

Global biodiversity peaks in the tropical forests of the Andes, a striking geological feature that has likely been instrumental in generating biodiversity by providing opportunities for both vicariant and ecological speciation. However, the role of these mountains in the diversification of insects, which dominate biodiversity, has been poorly explored using phylogenetic methods. Here we study the role of the Andes in the evolution of a diverse Neotropical insect group, the clearwing butterflies. We used dated species-level phylogenies to investigate the time course of speciation and to infer ancestral elevation ranges for two diverse genera. We show that both genera likely originated at middle elevations in the Andes in the Middle Miocene, contrasting with most published results in vertebrates that point to a lowland origin. Although we detected a signature of vicariance caused by the uplift of the Andes at the Miocene–Pliocene boundary, most sister species were parapatric without any obvious vicariant barrier. Combined with an overall decelerating speciation rate, these results suggest an important role for ecological speciation and adaptive radiation, rather than simple vicariance.     

New insight into the phylogenetic and biogeographic history of genus Ficus: Vicariance played a relatively minor role compared with ecological opportunity and dispersal

Studies on the evolution of tropical taxa emphasize the role ofvicariance and the break-up of Gondwana in explaining modern distributions.Earlier studies on figs (Ficus spp.) support this view.In the current study,we used an expanded sample (208 spp.) and improved molecular dating techniques to reconstruct the phylogenetic and biogeographic history of Ficus.Consistent with previous studies,our biogeographic analysis indicated that the ancestor of Ficus was present in Gondwana.However,a relaxed clock analysis relying on uncorrelated rates in BEAST suggested that the Neotropical section Pharmacosycea split-off in South America 86.67 Mya,and that other Ficus lineage ancestors originated in India.Most of the basal lineages appeared to have diverged following KT extinction,then rapidly diversified after India collided with continental Asia.The Afrotropical species most likely evolved initially in the Indian subcontinent then dispersed to Africa,either in the late Cretaceous of Madagascar or even later,following the Eocene collision of India with Asia.The Neotropical section Americana,either islandhopped to South America or took a northern route to the Americas through Europe prior to the terminal Eocene global cooling event.Ficus may have arrived in eastern Malesia following the collision of India with Asia,then widely dispersed thereafter.Given the wide ranges in our date estimates,several other scenarios are possible.However,contrary to earlier reports,our analyses suggest that vicariance played a relatively minor role compared with ecological opportunity and dispersal in the diversification of genus Ficus.     

The Azores diversity enigma: why are there so few Azorean endemic flowering plants and why are they so widespread?

Aim Endemism in the flora of the Azores is high (33%) but in other respects, notably the paucity of evolutionary radiations and the widespread distribution of most endemics, the flora differs markedly from the floras of the other Macaronesian archipelagos. We evaluate hypotheses to explain the distinctive patterns observed in the Azorean endemic flora, focusing particularly on comparisons with the Canary Islands. Location Azores archipelago. Methods Data on the distribution and ecology of Azorean endemic flowering plants are reviewed to ascertain the incidence of inter‐island allopatric speciation and adaptive, ecological speciation. These are contrasted with patterns for the Canary Islands. Patterns of endemism in the Azores and Canaries are further investigated in a phylogenetic context in relation to island age. beast was used to analyse a published molecular dataset for Pericallis (Asteraceae) and to investigate the relative ages of Azorean and Canarian lineages. Results There are few examples of inter‐island allopatric speciation in the Azorean flora, despite the considerable distances between islands and sub‐archipelagos. In contrast, inter‐island allopatric speciation has been an important process in the evolution of the Canary Islands flora. Phylogenetic data suggest that Azorean endemic lineages are not necessarily recent in origin. Furthermore, in Pericallis the divergence of the Azorean endemic lineage from its closest relative pre‐dates the radiation of a Canarian herbaceous clade by inter‐island allopatric speciation. Main conclusions The data presented do not support suggestions that hypotheses pertaining to island age, age of endemic lineages and ecological diversity considered individually explain the lack of radiations and the widespread distribution of Azorean endemics. We suggest that palaeoclimatic variation, a factor rarely considered in macroecological studies of island diversity patterns, may be an important factor. Palaeoclimatic data suggest frequent and abrupt transitions between humid and arid conditions in the Canaries during the late Quaternary, and such an unstable climate may have driven the recent diversification of the flora by inter‐island allopatric speciation, a process largely absent from the climatically more stable Azores. Further phylogenetic/phylogeographic analyses are necessary to determine the relative importance of palaeoclimate and other factors in generating the patterns observed.     

Out of sight, out of mind: high cryptic diversity obscures the identities and histories of geminate species in the marine bivalve subgenus Acar

Aim The rise of the Isthmus of Panama and the formation of ‘geminate’ species pairs serves as an important model of allopatric speciation. However, to function as a model system, hypothesized geminates must first be shown to be each other’s closest living relatives. If the presence of cryptic taxa obscures true relationships, the biogeographical histories of transisthmian taxa are likely to be misinterpreted. We have therefore completed a phylogeographic survey of the transisthmian bivalve subgenus Acar in the genus Barbatia to characterize patterns of tropical American diversity and to place transisthmian taxa in a regional phylogeographic context. Location Tropical America. Methods Mitochondrial cytochrome c oxidase I (COI) and nuclear internal transcribed spacer (ITS) sequences were obtained from 233 specimens of Acar. Sequences were analysed using cladistic and Bayesian methods. Divergence times between species were inferred from net nucleotide divergences and a coalescence‐based method. Results The survey revealed 22 COI clades that were also monophyletic at ITS, indicating that the taxonomy of Acar is potentially greater than a fivefold diversity underestimate. The lone previously recognized geminate [Barbatia (Acar) gradata and Barbatia (Acar) domingensis] is composed of 15 clades. Among the four transisthmian lineages identified, two diverged more than 14 Ma; the two other geminates split just prior to the time of final seaway closure. In addition to a fourfold increase in the number of known geminates, our data show that within‐basin diversification has been more impressive, with one geminate splitting into five monophyletic clades in the Western Atlantic alone since seaway closure. Electron microscopy of the larval shells of specimens indicates that the transisthmian lineage with the greatest rate of post‐Isthmian diversification possesses non‐planktonic larvae, a life‐history feature linked to high speciation rates. Main conclusions Our analyses revealed that the identities of geminate pairs split by the Isthmus of Panama were obscured by extremely high tropical American cryptic diversity. Although we have identified four geminates, only two appear to have been split by the Isthmus. Our uncovering of extensive post‐Isthmian diversification is consistent with the palaeontological perspective that the final closure of the Central American Seaway was followed by high rates of subgeneric diversification, particularly in the tropical Western Atlantic.     

Evolutionary history of two allopatric Terricola species (Arvicolinae, Rodentia) from molecular, morphological, and palaeontological data

To investigate the phylogenetic and phylogeographical relationships of arvicolines, we use several Western European ground voles. More particularly, our study is focused on Microtus ( Terricola ) savii and M. ( T. ) pyrenaicus . These two allopatric species are usually considered as having originated from the same ancestor, possibly M . ( T. ) mariaclaudiae . We propose molecular and morphological approaches: nucleotidic data from the mitochondrial cytochrome b and 12S rRNA genes and global morphological analyses from the first lower molar. Four other Terricola species ( multiplex , lusitanicus , duodecimcostatus , subterraneus ) were added to the data set for both analyses, and two other vole species ( Clethrionomys glareolus and Chionomys nivalis ) as outgroup to the molecular analysis, and five fossil populations to the morphological one. Palaeontological data are also widely taken into account. Both molecular and morphological analyses indicate that intra- Terricola relationships reflect the present-day geographical distribution of our data set species. Our results show that M. ( T. ) savii and M. ( T. ) pyrenaicus are from separate speciation events leading to two different biogeographical groups, respectively the Alpine–Italian group and the French–Iberian group, the latter being much more homogeneous. These speciation events could be related to Quaternary climatic changes, which induced southward migration, leading first to M. ( T. ) savii and second to M. ( T. ) pyrenaicus . The classical hypothesis of a geographical speciation for these two taxa from M. ( T. ) mariaclaudiae is invalid. However, the morphological data suggest a potential phylogenetic relationship between M. ( T. ) mariaclaudiae (ancestor) and M. ( T. ) pyrenaicus (descendant).  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 309–323.     

Evolution of oviposition strategies and speciation in the globeflower flies Chiastocheta spp. (Anthomyiidae)

Trollius europaeus (Ranunculaceae) is involved in an intimate interaction with several species of Chiastocheta flies (Anthomyiidae) that are both seed predators and pollinators. In this paper, we analyse the oviposition strategy of the six Chiastocheta species found to coexist on T. europaeus in 19 populations from the French Alps. We show that the species are not equivalent in their oviposition behaviour: C. rotundiventris usually deposits no more than one egg per flower in first-day flowers whereas C. dentifera aggregates its eggs on fruits and thus does not contribute to pollination at all; the four remaining species deposit eggs sequentially during the flowering period from the 2nd to the 7th day. Hence, the outcomes of the interaction in terms of net seed production for the plant greatly depend on the Chiastocheta species visiting it, ranging from a mutualistic to a purely parasitic interaction. We assessed mitochondrial divergence between Chiastocheta spp. by sequencing a 1320-bp mitochondrial DNA fragment. The low divergence observed between species (0–4.15%) suggests that genus diversification took place recently. Unlike in other plant–insect systems where diversification is usually thought to be driven by cospeciation or host shifts, we propose that Chiastocheta speciation took place within the host plant. Basal separation of a particularly mutualistic species provided favourable conditions for plant specialization on this seed-parasite as a pollinator early in the evolution of the association. The parasitic species ovipositing on fruits derived from a species ovipositing on flowers. Diversification of the intermediate strategies probably occurred in relation with the Pleistocene climatic events, reproductive isolation between species being reinforced by niche partitioning for oviposition and/or sexual selection.     

Evolution in caves: Darwin’s ‘wrecks of ancient life’ in the molecular era

Cave animals have historically attracted the attention of evolutionary biologists because of their bizarre ‘regressive’ characters and convergent evolution. However, understanding of their biogeographic and evolutionary history, including mechanisms of speciation, has remained elusive. In the last decade, molecular data have been obtained for subterranean taxa and their surface relatives, which have allowed some of the classical debates on the evolution of cave fauna to be revisited. Here, we review some of the major studies, focusing on the contribution of phylogeography in the following areas: biogeographic history and the relative roles of dispersal and vicariance, colonization history, cryptic species diversity and modes of speciation of cave animals. We further consider the limitations of current research and prospects for the future. Phylogeographic studies have confirmed that cave species are often cryptic, with highly restricted distributions, but have also shown that their divergence and potential speciation may occur despite the presence of gene flow from surface populations. Significantly, phylogeographic studies have provided evidence for speciation and adaptive evolution within the confines of cave environments, questioning the assumption that cave species evolved directly from surface ancestors. Recent technical developments involving ‘next generation’ DNA sequencing and theoretical developments in coalescent and population modelling are likely to revolutionize the field further, particularly in the study of speciation and the genetic basis of adaptation and convergent evolution within subterranean habitats. In summary, phylogeographic studies have provided an unprecedented insight into the evolution of these unique fauna, and the future of the field should be inspiring and data rich.