Natural history, biogeography, and endangerment of Hawaiian dry forest trees

We describe the floristic composition of Hawaiian dry forest trees and identify natural history characteristics and biogeographic variables that are associated with risk of endangerment. Hawaiian dry forests are comprised of 109 tree species in 29 families, with 90% of all species endemic, 10% indigenous, and 37% single-island endemics. Forty-five percent of Hawaiian dry forest taxa are at risk of endangerment. Dry forest taxa at risk have a significantly larger range size compared to taxa from other Hawaiian forest types. Dispersal mechanism was a significant predictor of a species occurrence in dry forest compared to other forest types based on logistic regressions clustered by lineage. Among dry forest taxa, hermaphroditic breeding systems, autochorous dispersal mechanisms, conspicuous flowers, and dry fruit were all more likely to be at risk of endangerment. When analyses were clustered by lineage using logistic regressions, only dispersal mechanism and flower size were significant predictors of risk and taxa with autochorous dispersal and conspicuous flowers were more likely to be at risk. The Big Island, Maui, Oahu, and Kauai all have remarkably similar numbers of dry forest taxa (63–65 species) and dry forest taxa at risk of endangerment. However, Big Island and Kauai have the highest number and percentage of single-island endemics. These results demonstrate patterns of endangerment specific to Hawaiian dry forests, the high levels of endangerment in this forest type, and the importance of prioritizing conservation in dry forest regions.     

Molecular biogeography and diversification of the endemic terrestrial fauna of the Hawaiian Islands

Oceanic islands have played a central role in biogeography and evolutionary biology. Here, we review molecular studies of the endemic terrestrial fauna of the Hawaiian archipelago. For some groups, monophyly and presumed single origin of the Hawaiian radiations have been confirmed (achatinelline tree snails, drepanidine honeycreepers, drosophilid flies, Havaika spiders, Hylaeus bees, Laupala crickets). Other radiations are derived from multiple colonizations (Tetragnatha and Theridion spiders, succineid snails, possibly Dicranomyia crane flies, Porzana rails). The geographic origins of many invertebrate groups remain obscure, largely because of inadequate sampling of possible source regions. Those of vertebrates are better known, probably because few lineages have radiated, diversity is far lower and morphological taxonomy permits identification of probable source regions. Most birds, and the bat, have New World origins. Within the archipelago, most radiations follow, to some degree, a progression rule pattern, speciating as they colonize newer from older islands sequentially, although speciation often also occurs within islands. Most invertebrates are single-island endemics. However, among multi-island species studied, complex patterns of diversification are exhibited, reflecting heightened dispersal potential (succineids, Dicranomyia). Instances of Hawaiian taxa colonizing other regions are being discovered (Scaptomyza flies, succineids). Taxonomy has also been elucidated by molecular studies (Achatinella snails, drosophilids). While molecular studies on Hawaiian fauna have burgeoned since the mid-1990s, much remains unknown. Yet the Hawaiian fauna is in peril: more than 70 per cent of the birds and possibly 90 per cent of the snails are extinct. Conservation is imperative if this unique fauna is to continue shedding light on profound evolutionary and biogeographic questions.     

Speciation and phylogeography of Hawaiian terrestrial arthropods

The Hawaiian archipelago is arguably the world's finest natural laboratory for the study of evolution and patterns of speciation. Arthropods comprise over 75% of the endemic biota of the Hawaiian Islands and a large proportion belongs to species radiations. We classify patterns of speciation within Hawaiian arthropod lineages into three categories: (i) single representatives of a lineage throughout the islands; (ii) species radiations with either (a) single endemic species on different volcanoes or islands, or (b) multiple species on each volcano or island; and (iii) single widespread species within a radiation of species that exhibits local endemism. A common pattern of phylogeography is that of repeated colonization of new island groups, such that lineages progress down the island chain, with the most ancestral groups (populations or species) on the oldest islands. While great dispersal ability and its subsequent loss are features of many of these taxa, there are a number of mechanisms that underlie diversification. These mechanisms may be genetic, including repeated founder events, hybridization, and sexual selection, or ecological, including shifts in habitat and/or host affiliation. The majority of studies reviewed suggest that natural selection is a primary force of change during the initial diversification of taxa.     

Notes on the Poaceae of the Robinson Crusoe (Juan Fernández) Islands, Chile

Notes on the Poaceae of the Robinson Crusoe (Juan Fernández) Islands, Chile, Brittonia 54: 154–163. 2001.—Poaceae in the Robinson Crusoe (=Juan Fernández) Islands number 53 species in 32 genera, of which 9% of the species are endemic, 9% indigenous, and 81% adventitious. The endemic taxa (and their conservation status) are:Agrostis masafuerana (rare),Chusquea fernandeziana (not endangered),Megalachne berteroana (not endangered),M. masafuerana (not endangered), andPodophorus bromoides (extinct).Megalachne andPodophorus are endemic genera. Comparisons with Poaceae in the Bonin and Volcano Islands, Canary Islands, Galápagos Islands, and Hawaiian Islands show different levels of endemism: number of endemic taxa, respectively, 5, 10, 12, 40; percent specific endemism, 8, 6, 21, 19. No endemic genera occur.Anthoxanthum odoratum, Avena barbata andHordeum murinum are noxious weeds in the Robinson Crusoe Islands. Many adventives are shared among floras of the archipelagos, with the highest ties of Robinson Crusoe being to the Canaries (53% of total Poaceae known in Juan Fernández) and the Hawaiian Islands (47%). Low levels of adventives occur within the Bonin (5%) and Galápagos (7%) Islands. In contrast, there are many endemic genera of Asteraceae in these same archipelagos: Bonin and Volcano Islands (1), Canary Islands (8), Galápagos Islands (5), and Hawaiian Islands (6); percent of specific endemism is also higher (20, 53, 54 and 56, respectively). Hypotheses for greater levels of endemics among oceanic island Asteraceae include more efficient dispersal mechanisms by wind and birds, animal pollination systems that result in greater degrees of geographic populational genetic isolation, and a vascular cambium that offers enhanced growth-form evolutionary opportunities.     

Molecular biogeography and origins of the Hawaiian fern flora

Located approximately 4000 km from the nearest continent, the Hawaiian Islands comprise the most isolated archipelago on Earth. This isolation has resulted in a unique flora that includes nearly 200 native ferns and lycophytes, 77% of which are endemic to the islands. Because the Hawaiian Islands are volcanic in origin, all abiotically dispersed organisms must have arrived there via the wind or the water. Fern spores are most likely dispersed through the air, and thus patterns of air movement have undoubtedly played a significant role in determining the geographic origins of the ancestors of the Hawaiian ferns. We have identified four possible climate-based or weather-based spore dispersal hypotheses that could have resulted in the movement of ancestral spores to the Hawaiian Islands: (1) the northern subtropical jetstream, moving spores from Indo-Pacific regions; (2) the trade winds, dispersing spores from Central and North America; (3) storms carrying spores from southern Mexico and/or Central America; and (4) a dispersal mechanism carrying spores from the South Pacific across the equator resulting from the combined influence of a seasonal southern shift of the Intertropical Convergence Zone (ITCZ), Hadley Cell air movement, and the trade winds. Utilizing recently published molecular phylogenetic studies of three fern genera (Dryopteris, Polystichum, andHymenophyllum) and new analyses of three additional genera (Adenophorus, Grammitis, andLellingeria), each of which is represented in the Hawaiian Islands by at least one endemic lineage, we reviewed the biogeographical implications for the Hawaiian taxa in light of the possible common dispersal patterns and pathways. We hypothesize that three of the five endemicDryopteris lineages, both of the endemicPolystichum lineages, at least one endemicHymenophyllum lineage in the Hawaiian Islands, and, perhaps, one endemicGrammitis lineage resulted from ancestral spores of each lineage dispersing to the Hawaiian Islands via the northern subtropical jetstream.Adenophorus is sister to a mostly neotropical clade, therefore, it is likely that the ancestor of the Hawaiian clade dispersed to the Hawaiian Islands via the trade winds or a storm system. The ancestor of the endemicLellingeria lineage may have dispersed to the Hawaiian Islands from the neotropics via the trade winds or a storm system, or from the South Pacific across the equator through the combination of a seasonal southern shift of the ITCZ, Hadley Cells, and the trade winds.     

Effects of climate change on the distribution of plant species and plant functional strategies on the Canary Islands

Aim

Oceanic islands possess unique floras with high proportions of endemic species. Island floras are expected to be severely affected by changing climatic conditions as species on islands have limited distribution ranges and small population sizes and face the constraints of insularity to track their climatic niches. We aimed to assess how ongoing climate change affects the range sizes of oceanic island plants, identifying species of particular conservation concern.

Location

Canary Islands, Spain.

Methods

We combined species occurrence data from single-island endemic, archipelago endemic and nonendemic native plant species of the Canary Islands with data on current and future climatic conditions. Bayesian Additive Regression Trees were used to assess the effect of climate change on species distributions; 71% (n = 502 species) of the native Canary Island species had models deemed good enough. To further assess how climate change affects plant functional strategies, we collected data on woodiness and succulence.

Results

Single-island endemic species were projected to lose a greater proportion of their climatically suitable area (x ̃ = −0.36) than archipelago endemics (x ̃ = −0.28) or nonendemic native species (x ̃ = −0.26), especially on Lanzarote and Fuerteventura, which are expected to experience less annual precipitation in the future. Moreover, herbaceous single-island endemics were projected to gain less and lose more climatically suitable area than insular woody single-island endemics. By contrast, we found that succulent single-island endemics and nonendemic natives gain more and lose less climatically suitable area.

Main Conclusions

While all native species are of conservation importance, we emphasise single-island endemic species not characterised by functional strategies associated with water use efficiency. Our results are particularly critical for other oceanic island floras that are not constituted by such a vast diversity of insular woody species as the Canary Islands.     

Tracing ancestry with chromosomal sequences

Most of the large Drosophila species of Hawaii are single-island endemics. Chromosomal sequences show that species at the new end of the archipelago have been derived stepwise from ancestral populations on older islands. The oldest high island has an endemic species with sequences that match some in the Nearctic-Palearctic robusta species group. Colonization from a continent by long-distance dispersal seems a likely origin for the Hawaiian drosophilids. Telmatogeton, a worldwide genus of marine midges, has five Hawaiian species inhabiting freshwater streams. Chromosomal sequences of a marine species in Hawaiian waters match the freshwater forms, indicating colonization from the ocean.     

Origins of the Hawaiian flora: Phylogenies and biogeography reveal patterns of long‐distance dispersal

Botanists have long considered the origins of the Hawaiian flora in terms of long‐distance dispersal from particular source areas. We extensively reviewed phylogenetic studies of the Hawaiian angiosperm flora to determine the most likely region of origin for each lineage from a defined set of source areas. We also evaluated dispersal modes of each lineage to assess whether certain dispersal modes are associated with a given source area. The largest source category was Widespread (involving related taxa that extend across more than one region), although many of these comprised native non‐endemic species, and accounted for little of the total species diversity (after accounting for in situ speciation). The next largest source regions were Indo‐Malayan and Neotropical. Comparatively few lineages originated from the East Asian region, although these include the single largest lineage. Lineages originating in the Indo‐Malayan region predominantly arrived via Pacific Islands, whereas dispersal from all other regions appears to have been mostly direct. Compared with previous analyses, we found a higher proportion of lineages originating in the Neotropics and temperate North America. Widespread origins were positively associated with dispersal via flotation on water, whereas other origins were associated with dispersal by birds, either through internal transport or external adhesion. We identified thirty‐one potential cases of dispersal out of Hawaii to other islands. Our assessment is complicated by lineages with ancient origins, with further complications likely stemming from hybridization events. Overall, numerous lineages including some distinctive endemic genera have not had sufficient phylogenetic study to determine an origin.     

Molecular evidence for adaptive radiation of Micromeria Benth. (Lamiaceae) on the Canary Islands as inferred from chloroplast and nuclear DNA sequences and ISSR fingerprint data

The Canary Islands have been a focus for phylogeographic studies on the colonization and diversification of endemic angiosperm taxa. Based on phylogeographic patterns, both inter island colonization and adaptive radiation seem to be the driving forces for speciation in most taxa. Here, we investigated the diversification of Micromeria on the Canary Islands and Madeira at the inter- and infraspecific level using inter simple sequence repeat PCR (ISSR), the trnK-Intron and the trnT-trnL-spacer of the cpDNA and a low copy nuclear gene. The genus Micromeria (Lamiaceae, Mentheae) includes 16 species and 13 subspecies in Macaronesia. Most taxa are restricted endemics, or grow in similar ecological conditions on two islands. An exception is M. varia, a widespread species inhabits the lowland scrub on each island of the archipelago and could represent an ancestral taxon from which radiation started on the different islands. Our analyses support a split between the "eastern" islands Fuerteventura, Lanzarote and Gran Canaria and the "western" islands Tenerife, La Palma and El Hierro. The colonization of Madeira started from the western Islands, probably from Tenerife as indicated by the sequence data. We identified two lineages of Micromeria on Gomera but all other islands appear to be colonized by a single lineage, supporting adaptive radiation as the major evolutionary force for the diversification of Micromeria. We also discuss the possible role of gene flow between lineages of different Micromeria species on one island after multiple colonizations.     

Floristic biogeography of the Hawaiian Islands: influences of area, environment and paleogeography

Aim A detailed database of distributions and phylogenetic relationships of native Hawaiian flowering plant species is used to weigh the relative influences of environmental and historical factors on species numbers and endemism. Location The Hawaiian Islands are isolated in the North Pacific Ocean nearly 4000 km from the nearest continent and nearly as distant from the closest high islands, the Marquesas. The range of island sizes, environments, and geological histories within an extremely isolated archipelago make the Hawaiian Islands an ideal system in which to study spatial variation in species distributions and diversity. Because the biota is derived from colonization followed by extensive speciation, the role of evolution in shaping the regional species assemblage can be readily examined. Methods For whole islands and regions of each major habitat, species–area relationships were assessed. Residuals of species–area relationships were subjected to correlation analysis with measures of endemism, isolation, elevation and island age. Putative groups of descendents of each colonist from outside the Hawaiian Islands were considered phylogenetic lineages whose distributions were included in analyses. Results The species–area relationship is a prominent pattern among islands and among regions of each given habitat. Species number in each case correlates positively with number of endemics, number of lineages and number of species per lineage. For mesic and wet habitat regions, island age is more influential than area on species numbers, with older islands having more species, more single‐island endemics, and higher species : lineage ratios than their areas alone would predict. Main conclusions Because species numbers and endemism are closely tied to speciation in the Hawaiian flora, particularly in the most species‐rich phylogenetic lineages, individual islands’ histories are central in shaping their biota. The Maui Nui complex of islands (Maui, Moloka‘i, Lāna‘i and Kaho‘olawe), which formed a single large landmass during most of its history, is best viewed in terms of either the age or area of the complex as a whole, rather than the individual islands existing today.     

Adaptive radiation of shrubby tarweeds (Deinandra) in the California Islands parallels diversification of the Hawaiian silversword alliance (Compositae-Madiinae)

Phylogenetic analyses of nuclear rDNA transcribed spacers and cytogenetic studies of interspecific hybrids reported here uphold Carlquist's hypothesis (1965, Island Biology) that shrubby tarweeds (Deinandra) of Guadalupe Island, Mexico, are products of in situ radiation in the California Islands, where evidence of plant diversification has been equivocal. Based on the rDNA findings, the Guadalupe Island endemics (D. frutescens, D. greeneana subsp. greeneana, and D. palmeri) constitute a clade that arose since the late Pliocene, well after the origin of Guadalupe Island and diversification of annual, mainland Californian lineages of Deinandra. High interfertility and normal meiosis in F(1) hybrids between the three endemics contrast with reduced interfertility (to complete intersterility) and meiotic irregularities in F(1) hybrids between other, mostly mainland species of Deinandra. Cloned rDNA sequences provided no convincing evidence of introgression among the Guadalupe Island deinandras; morphological, phenological, and/or habitat differences among those taxa indicate ecological barriers to gene flow and a probable role for ecological divergence in diversification. Biosystematic and molecular phylogenetic data for shrubby tarweeds of Guadalupe Island and another secondarily woody, oceanic-island tarweed lineage, the Hawaiian silversword alliance, reveal strikingly similar evolutionary histories. Both groups violate Baker's Rule by stemming from self-incompatible ancestors in western North America, and each has undergone within-island diversification without evolution of strong sterility barriers among lineages. Evolutionary parallels between these Hawaiian and California Island lineages of Madiinae, first suggested by Carlquist, may reflect characteristics of tarweeds that facilitate insular colonization and adaptive radiation.     

The diversity of life-form type, habitat preference and phenology of the endemics in the Goa region of the Western Ghats, India

Aim To carry out (1) a floristic survey of endemic flowering plants of the Western Ghats occurring in Goa, (2) identify their habitat preference and diversity of life‐form type, (3) observe flowering phenology of the endemics and (4) to correlate factors that affect their phenological pattern. Location Goa state is located between 15°48′ N and 14°53′54′′ N and 74°20′13′′ E and 73°40′33′′ E, in the northern part of the Western Ghats, India. Method A list of endemic plants from the study area was prepared using available floristic works and checklists of endemic plants of India. Based on preliminary field observations carried out in the study area, major habitats such as plateaus, moist deciduous forests, semi‐evergreen forests, evergreen forests and mangroves were identified for the subsequent intensive survey of endemics. Voucher specimens for all the endemic species were collected, processed using conventional herbarium techniques and deposited in the Herbarium, Department of Botany, Goa University. Species were identified using local and regional floras and their identity was confirmed at various herbaria along the Western Ghats. Data on their life‐form types, habitat and phenology was recorded in the field. Phenological observations were made every fortnight. A computerized data base was generated incorporating details on their life‐form type, phenology and habitat. Results A floristic survey of endemic plants of the Western Ghats in Goa resulted in the collection of 113 endemic species. Life‐form analysis reveals that herbaceous endemics are the most dominant followed by trees, shrubs and climbers. Plateaus in the study area harbour the largest number of endemic species, especially herbs. Endemic trees are distributed in the semi‐evergreen and evergreen forests. Endemic species in the study area show different peak and lean seasons of flowering depending on their life‐form type, habitat and ecological factors like temperature and rainfall/moisture content in the soil. Main conclusions The plateaus in the northern part of the Western Ghats are unique, being species rich with herbaceous endemics. These ephemerals are closely associated with the rainfall patterns thus; any change of moisture regime over the long‐term will have an impact on the distribution of these endemics.     

Rarity and extinction risk in coral reef angelfishes on isolated islands: interrelationships among abundance, geographic range size and specialisation

Determining the species most vulnerable to increasing degradation of coral reef habitats requires identification of the ecological traits that increase extinction risk. In the terrestrial environment, endemic species often face a high risk of extinction because of an association among three traits that threaten species persistence: small geographic range size, low abundance and ecological specialisation. To test whether these traits are associated in coral reef fishes, this study compared abundance and specialisation in endemic and widespread angelfishes at the remote Christmas and Cocos Islands in the Indian Ocean. The interrelationships among traits conferring high extinction risk in terrestrial communities did not apply to these fishes. Endemic angelfishes were 50–80 times more abundant than widespread species at these islands. Furthermore, there was no relationship between abundance and ecological specialisation. Endemic species were not more specialised than widespread congeners and endemics used similar resources to many widespread species. Three widespread species exhibited low abundance and some degree of specialisation, which may expose them to a greater risk of local extinction. For endemic species, high abundance and lack of specialisation on susceptible habitats may compensate for the global extinction risk posed by having extremely small geographic ranges. However, recent extinctions of small range reef fishes confirm that endemics are not immune to the increasing severity of large-scale disturbances that can affect species throughout their geographic range.     

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.     

Molecular insights into the phylogenetic structure of the spider genus Theridion (Araneae, Theridiidae) and the origin of the Hawaiian Theridion-like fauna

The Hawaiian happy face spider ( Theridion grallator Simon, 1900), named for a remarkable abdominal colour pattern resembling a smiling face, has served as a model organism for understanding the generation of genetic diversity. Theridion grallator is one of 11 endemic Hawaiian species of the genus reported to date. Asserting the origin of island endemics informs on the evolutionary context of diversification, and how diversity has arisen on the islands. Studies on the genus Theridion in Hawaii, as elsewhere, have long been hampered by its large size (> 600 species) and poor definition. Here we report results of phylogenetic analyses based on DNA sequences of five genes conducted on five diverse species of Hawaiian Theridion , along with the most intensive sampling of Theridiinae analysed to date. Results indicate that the Hawaiian Islands were colonised by two independent Theridiinae lineages, one of which originated in the Americas. Both lineages have undergone local diversification in the archipelago and have convergently evolved similar bizarre morphs. Our findings confirm para- or polyphyletic status of the largest Theridiinae genera: Theridion , Achaearanea and Chrysso . Convergent simplification of the palpal organ has occurred in the Hawaiian Islands and in two continental lineages. The results confirm the convergent evolution of social behaviour and web structure, both already documented within the Theridiidae. Greater understanding of phylogenetic relationships within the Theridiinae is key to understanding of behavioural and morphological evolution in this highly diverse group.     

Genetic differentiation and speciation in Stomion (Goleoptera: Tenebrionidae): flightless beetles of the Galápagos Islands, Ecuador

Stomion is a swarm of 13 tenebrionid beetle taxa endemic to the Galápagos Islands. The taxa show marked morphological variation, and all are flighdess. They can be divided into three divergent morphological groups based on external sculpturing of the elytra. Eleven taxa were examined electrophoretically at 20 enzyme loci to measure the extent of genetic differentiation which accompanied morphological differentiation and speciation. Stomion are shown to be ideal candidates for allopatric and founder event speciation, with their limited dispersal ability, narrow distributions, and incidence of single-island endemics. Accordingly, individual heterozygosities were low in most taxa, and gene pool divergence was apparent among populations of single taxa occupying different islands. Both interspecific and intraspecific genetic identities were high, with an average 1 = 0.88 and I = 0.96, respectively. The relatively young age of the genus and the allopatric mode of speciation may account for high genetic relatedness. No clear relationship was found between allozyme phenotypes and the three morphological groups, thus, the evolution of these traits may have been facilitated by similar selection among islands of similar age and geography. The geographical distributions of the morphological groups can be used to suggest a general pattern of speciation for the group.     

Evolutionary biogeography of the terrestrial biota of the Marquesas Islands,one of the world's remotest archipelagos

Aim

Here I review phylogenetic studies concerning the biogeography of the Marquesas Islands, an oceanic hotspot archipelago in the Pacific Ocean formed <5.5 Ma, and compare patterns (particularly pertaining to colonization and diversification) within the archipelago to those reported from the Hawaiian and Society Islands.

Location

Marquesas Islands, French Polynesia (Pacific Ocean).

Methods

I reviewed 37 phylogenetic studies incorporating Marquesas‐endemic taxa. I asked the following questions: (a) where are the sister‐groups of Marquesas lineages distributed? (b) are Marquesas‐endemic “radiations” monophyletic or polyphyletic? (c) what major between‐island phylogeographic barriers are seen in the Marquesas? (d) what evidence exists for diversification within islands? (e) how old is the Marquesas biota compared to the archipelago's age? Finally, these patterns are compared with those seen in the Society Islands and Hawaii.

Results

Most Marquesan lineages have their closest known relatives on other Pacific plate archipelagos (particularly the Society, Hawaiian, and Austral islands). Most Marquesas‐endemic radiations are found to be monophyletic, and among‐island diversification appears to be common. There is limited evidence for within‐island diversification. Some radiations may be consistent with a weak progression rule in which younger lineages are on younger islands. Crown ages of no Marquesas radiations appear to be older than the age of the archipelago (with one exception).

Main conclusions

Diversification of the Marquesas biota resembles that of the Hawaiian Islands more than that of the Society Islands. Many radiations are monophyletic and some appear to diversify in parallel with the formation of the archipelago.

     

Allozyme diversity within and divergence among species ofTolpis(Asteraceae-Lactuceae) in the Canary Islands: systematic, evolutionary, and biogeographical implications

Plants endemic to oceanic islands represent some of the most unusual and rare taxa in the world. Enzyme electrophoresis was used to assess genetic diversity within and divergence among all endemic species of a small genus of plants on the Canary Islands. Our results show that the genus Tolpis is similar to many other island groups in having generally low allozyme divergence among species, with the highest divergence found among four groups of endemics. The two rare and highly localized species T. glabrescens and T. crassiuscula are each divergent from all other species in the Canaries. Tolpis coronopifolia is also divergent at allozyme loci; this is the only endemic species that is a self-compatible annual (or weak biennial). A large, morphologically variable species complex consisting of T. laciniata and T. lagopoda together with several named and unnamed morphological variants shows low allozyme divergence among its elements. The evolution of polyploidy from diploid ancestors in situ in oceanic archipelagos is uncommon, but the tetraploid T. glabrescens is an exception. Allozyme data do not implicate any extant diploid Tolpis species as parents of the polyploid. It is possible that T. glabrescens originated early in the evolution of Tolpis in the Canary Islands and that its parents are now extinct. The nonendemic T. barbata shows no greater divergence from the Canary Island endemics than some endemics exhibit among themselves. Both changes in allele frequencies and unique alleles are responsible for genetic divergence among species of Tolpis.     

Correlates of vulnerability among arthropod species threatened by invasive ants

Invasive species are causing population declines and extinctions of native species worldwide. Correlates of species vulnerability, which help identify at-risk taxa, are not well developed for arthropods, particularly with respect to threats from invasive species. At five sites undergoing invasion by ants in the Hawaiian Islands, we assessed body size, population density, trophic role and provenance (introduced or endemic to the Hawaiian Islands) as potential correlates of vulnerability for 300 arthropod species. Among rare species, provenance was the most important factor associated with absence from invaded plots, with endemic species much more commonly absent. Trophic role was also important, but only when interacting with provenance: endemic carnivores were by far the most vulnerable group, followed by endemic detritivores. For non-rare species, Hawaii endemics were significantly more reduced in invaded plots compared to introduced species. In addition, species that occurred at lower population densities were more vulnerable than those occurring at higher densities. Body size did not correlate with vulnerability for either rare or non-rare species. Despite these trends, there was relatively high variability in responses to invasion among species in many taxonomic orders, as well as among populations of particular species at different sites. While the consideration of additional intrinsic traits might increase predictive ability to some degree (e.g., intrinsic traits only explained 21% of the variation in impact among non-rare species), community-specific extrinsic factors appear to play a large role in influencing outcomes for many species, making prediction substantially more difficult.