Predicting Invasive Plants: Prospects for a General Screening System Based on Current Regional Models

Screening systems for predicting invasive plants have been independently developed for the non-indigenous floras of North America, the South African fynbos, and Australia. To evaluate the performance of these screening systems outside the regions for which they were developed, we tested them for the non-indigenous flora of the Hawaiian Islands. When known invasive plant species in the Hawaiian Islands were evaluated using the North American and Australian systems, 82% and 93% of the species were predicted to be invasive, respectively, and the remainder were classified as requiring further study. The South African fynbos system correctly predicted only 60% of the invasive species in the Hawaiian Islands. All three screening systems correctly classified a majority of the non-invaders as non-invasive. The Australian system has several advantages over the other systems, including the highest level of correct identification of invaders (>90%), ability to evaluate non-woody plants, and ability to evaluate a species even when the answers to some questions are unknown. Nevertheless, with the Australian system, a large fraction of species known not to be invasive were recommended for further study before importing, so there remains room for improvement in identifying non-invasive species. Based on our results for the Hawaiian Islands and a previous evaluation in New Zealand, the Australian system appears to be a promising template for building a globally applicable system for screening out invasive plant introductions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Marine Ostracoda of the Galapagos Islands and Ecuador

Marine algae collected from rock pools on Hood, Fernandina and Mosquera, in the Galapagos Islands, have provided a fauna of 26 ostracod species of which 14 are new. One genus, belonging to the Hemicytheridae, is also new and appears to be endemic to the Islands. From two littoral algal samples collected from Punta Canoa and San Pedro beach, Ecuador, an assemblage of some nine species, four of which are new, is also described. Two ostracods: Touroconcha lapidiscola and Loxoconcha (Lox-ocorniculum) lenticuloides , are the only species so far known to be present both in the Galapagos Islands and off the coast of Central and South America. One species: Cytherelloidea praecipua recorded from off Tobago and Clipperton Islands may be present in the Galapagos but this has not definitely been confirmed. The evolution in the Gulf Coast/Caribbean region of several ostracods and their subsequent dispersal westwards is discussed.     

Multiple origins of Hawaiian drosophilids: Phylogeography of Scaptomyza Hardy (Diptera: Drosophilidae)

Scaptomyza is a highly diversified genus in the family Drosophilidae, having undergone an explosive radiation, along with the Hawaiian‐endemic genus Idiomyia in the Hawaiian Islands: about 60% of 269 Scaptomyza species so far described are endemic to the Hawaiian Islands. Two hypotheses have been proposed for the origin and diversification of Hawaiian drosophilids. One is the “single Hawaiian origin” hypothesis: Scaptomyza and Idiomyia diverged from a single common ancestor that had once colonized the Hawaiian Islands, and then non‐Hawaiian Scaptomyza migrated back to continents. The other is the “multiple origins” hypothesis: Hawaiian Scaptomyza and Idiomyia derived from different ancestors that independently colonized the Hawaiian Islands. A key issue for testing these two hypotheses is to clarify the phylogenetic relationships between Hawaiian and non‐Hawaiian species in Scaptomyza. Toward this goal, we sampled additional non‐Hawaiian Scaptomyza species, particularly in the Old World, and determined the nucleotide sequences of four mitochondrial and seven nuclear genes for these species. Combining these sequence data with published data for 79 species, we reconstructed the phylogeny and estimated ancestral distributions and divergence times. In the resulting phylogenetic trees, non‐Hawaiian Scaptomyza species were interspersed in two Hawaiian clades. From a reconstruction of ancestral biogeography, we inferred that Idiomyia and Scaptomyza diverged outside the Hawaiian Islands and then independently colonized the Hawaiian Islands, twice in Scaptomyza, thus supporting the “multiple origins” hypothesis.     

Diversity despite dispersal: colonization history and phylogeography of Hawaiian crab spiders inferred from multilocus genetic data

The Hawaiian archipelago is often cited as the premier setting to study biological diversification, yet the evolution and phylogeography of much of its biota remain poorly understood. We investigated crab spiders (Thomisidae, Mecaphesa ) that demonstrate contradictory tendencies: (i) dramatic ecological diversity within the Hawaiian Islands, and (ii) accompanying widespread distribution of many species across the archipelago. We used mitochondrial and nuclear genetic data sampled across six islands to generate phylogenetic hypotheses for Mecaphesa species and populations, and included penalized likelihood molecular clock analyses to estimate arrival times on the different islands. We found that 17 of 18 Hawaiian Mecaphesa species were monophyletic and most closely related to thomisids from the Marquesas and Society Islands. Our results indicate that the Hawaiian species evolved from either one or two colonization events to the archipelago. Estimated divergence dates suggested that thomisids may have colonized the Hawaiian Islands as early as ~10 million years ago, but biogeographic analyses implied that the initial diversification of this group was restricted to the younger island of Oahu, followed by back-colonizations to older islands. Within the Hawaiian radiation, our data revealed several well-supported genetically distinct terminal clades corresponding to species previously delimited by morphological taxonomy. Many of these species are codistributed across multiple Hawaiian Islands and some exhibit genetic structure consistent with stepwise colonization of islands following their formation. These results indicate that dispersal has been sufficiently limited to allow extensive ecological diversification, yet frequent enough that interisland migration is more common than speciation.     

Population structure of island‐associated dolphins: Evidence from mitochondrial and microsatellite markers for common bottlenose dolphins (Tursiops truncatus) around the main Hawaiian Islands

We used mitochondrial and nuclear genetic markers to investigate population structure of common bottlenose dolphins, Tursiops truncatus, around the main Hawaiian Islands. Though broadly distributed throughout the world's oceans, bottlenose dolphins are known to form small populations in coastal waters. Recent photo‐identification data suggest the same is true in Hawaiian waters. We found genetic differentiation among (mtDNA Φ_ST= 0.014–0.141, microsatellite F’_ST= 0.019–0.050) and low dispersal rates between (0.17–5.77 dispersers per generation) the main Hawaiian Island groups. Our results are consistent with movement rates estimated from photo‐identification data and suggest that each island group supports a demographically independent population. Inclusion in our analyses of samples collected near Palmyra Atoll provided evidence that the Hawaiian Islands are also occasionally visited by members of a genetically distinct, pelagic population. Two of our samples exhibited evidence of partial ancestry from Indo‐Pacific bottlenose dolphins (T. aduncus), a species not known to inhabit the Hawaiian Archipelago. Our findings have important implications for the management of Hawaiian bottlenose dolphins and raise concerns about the vulnerability to human impacts of pelagic species in island ecosystems.     

Population structure of melon‐headed whales (Peponocephala electra) in the Hawaiian Archipelago: Evidence of multiple populations based on photo identification

Despite the presence of melon‐headed whales in tropical and subtropical waters worldwide, little is known about this species. To assess population structure in Hawai‘i, dedicated field efforts were undertaken from 2000 to 2009. Using only good quality photographs, there were 1,433 unique photo‐identified individuals, of which 1,046 were distinctive. Of these, 31.5% were seen more than once. Resighting data combined with social network analyses showed evidence of two populations—a smaller, resident population, seen exclusively off the northwest region of the island of Hawai‘i, and a larger population, seen throughout all the main Hawaiian Islands (hereafter the “main Hawaiian Islands” population). A Bayesian analysis examining the probability of movements of individuals between populations provided a posterior median dispersal rate of 0.0009/yr (95% CI = 0–0.0041), indicating the populations are likely demographically independent. Depth of encounters with the Hawai‘i Island resident population was significantly shallower (median = 381 m) than those with the main Hawaiian Islands population (median = 1,662 m). Resightings of individuals have occurred up to 22 yr apart for the Hawai‘i Island resident population and up to 13 yr apart for the main Hawaiian Islands population, suggesting long‐term residency to the islands for both populations.     

A review of Polynesian Carposina Herrich‐Schäffer (Lepidoptera: Carposinidae), with descriptions of four new species

Hawaiian Carposina represent over 17% of the known world fauna of Carposinidae. In contrast, only two species are known for all of French Polynesia in the South Pacific. Here we describe four new species: two from the Hawaiian Islands, C arposina urbanae sp. nov. and C . gagneorum sp. nov. , and two from the Society Islands, C . longignathosa sp. nov. and C . brevinotata sp. nov. We further recognize another new Hawaiian species too worn to describe. Additionally, we present the first phylogeny for Polynesian Carposina, including 19 taxa, using one mitochondrial and two nuclear gene regions. The Hawaiian Carposina sampled thus far form a monophyletic clade. Lastly, we provide a framework to better understand the diversification and phylogeography of this group, and provide a summary of currently known host plant associations. Diversification appears to have resulted from interplay between host switching and geographic isolation across the Hawaiian Archipelago.     

The Hawaiian Archipelago is a stepping stone for dispersal in the Pacific: an example from the plant genus Melicope (Rutaceae)

Aim Pacific biogeographical patterns in the widespread plant genus Melicope J.R. Forst. & G. Forst. (Rutaceae) were examined by generating phylogenetic hypotheses based on chloroplast and nuclear ribosomal sequence data. The aims of the study were to identify the number of colonization events of Melicope to the Hawaiian Islands and to reveal the relationship of Hawaiian Melicope to the Hawaiian endemic genus Platydesma H. Mann. The ultimate goal was to determine if the Hawaiian Islands served as a source area for the colonization of Polynesia. Location Nineteen accessions were sampled in this study, namely eight Melicope species from the Hawaiian Islands, four from the Marquesas Islands, one species each from Tahiti, Australia and Lord Howe Island, two Australian outgroups and two species of the Hawaiian endemic genus Platydesma. To place our results in a broader context, 19 sequences obtained from GenBank were included in an additional analysis, including samples from Australia, Papua New Guinea, New Zealand, Southeast Polynesia and Asia. Methods DNA sequences were generated across 19 accessions for one nuclear ribosomal and three chloroplast gene regions. Maximum parsimony analyses were conducted on separate and combined data sets, and a maximum likelihood analysis was conducted on the combined nuclear ribosomal and chloroplast data set. A broader nuclear ribosomal maximum parsimony analysis using sequences obtained from GenBank was also performed. Geographic areas were mapped onto the combined chloroplast and nuclear ribosomal tree, as well as onto the broader tree, using the parsimony criterion to determine the dispersal patterns. Results Phylogenetic analyses revealed that Platydesma is nested within Melicope and is sister to the Hawaiian members of Melicope. The Hawaiian Melicope + Platydesma lineage was a result of a single colonization event, probably from the Austral region. Finally, Marquesan Melicope descended from at least one, and possibly two, colonization events from the Hawaiian Islands. Main conclusions These data demonstrate a shifting paradigm of Pacific oceanic island biogeography, in which the patterns of long‐distance dispersal and colonization in the Pacific are more dynamic than previously thought, and suggest that the Hawaiian Islands may act as a stepping stone for dispersal throughout the Pacific.     

Redescription of a rare callionymid fish,Paradiplogrammus corallinus,from Hawaii and Japan

Paradiplogrammus corallinus (Gilbert) has been represented by only two female specimens collected from the Hawaiian Islands. The species is redescribed on the basis of male and female specimens from the Hawaiian Islands and Hachijo-jima, Izu Islands, Japan. This species is unique amongParadiplogrammus in having a pair of supraorbital cirri.     

Upper-montane plant invasions in the Hawaiian Islands: Patterns and opportunities

In the Hawaiian Islands, massive volcanoes have created extreme elevation gradients, resulting in environments ranging from nearly tropical to alpine, spread across a distance of only a few dozen kilometers. Although the Hawaiian Islands are widely recognized for opportunities to study lowland tropical forest invasions, less attention has been paid to invasions of Hawaii's upper-montane forest, sub-alpine and alpine environments. This study synthesizes current knowledge of plant naturalization in upper-montane environments of the Hawaiian Islands in order to (1) determine whether patterns of tropical versus temperate species invasion change with elevation, and (2) evaluate whether upper-montane invaders are having significant impacts on native plant communities. A total of 151 naturalized plant species have been recorded at 2000 m or higher. Most species (93%) are herbaceous, and over half (52%) are native to Europe/Eurasia. Twenty-one species (14%) are reported to be disruptive in native plant communities, mainly by forming dense stands that appear to inhibit recruitment of natives, but also by altering vegetation structure or causing changes in ecosystem processes. Fourteen species (9%) were first recorded within the past 30 years, indicating that new invasions of upper-montane habitats are ongoing. At 1200 m elevation, only 38% of naturalized species are temperate in origin, but the proportion of temperate species increases linearly with elevation up to 3000 m (alpine habitat), where all naturalized species are temperate in origin and over 80% are native to Europe/Eurasia. Declining temperature along the elevation gradient probably drives this pattern. The extreme elevation gradients in the Hawaiian Islands provide specific opportunities for comparative studies on the ecology and evolution of temperate invaders while also creating a unique field environment for understanding interactions between temperate and tropical species.     

Opisthobranchiate Mollusca from Tanzania (Suborder:Doridacea)

Twenty-two species of Doridacea are described from the vicinity of Dar es Salaam, Tanzania. None is new to science, but one species of the genus Gymnodoris was too immature to be fully identified. Ten of the 22 species are known from the Hawaiian Islands, indicating that many species of dorid occur throughout the Indian Ocean and the Pacific as far as Hawaii. One species (Jorunna tomentosa) appears to be cosmopolitan, and two may possibly be confined to the coast of Africa (Chromodoris annulata and C. vicind).
Sixty-nine species of dorid have already been reported from Tanzania by Eliot. Nine species in the present collection were not found by him, so 78 are now known from the area.     

Molecular phylogenetics and historical biogeography of Hawaiian Dryopteris (Dryopteridaceae)

The fern genus Dryopteris (Dryopteridaceae) is represented in the Hawaiian Islands by 18 endemic taxa and one non-endemic, native species. The goals of this study were to determine whether Dryopteris in Hawai'i is monophyletic and to infer the biogeographical origins of Hawaiian Dryopteris by determining the geographical distributions of their closest living relatives. We sequenced two chloroplast DNA fragments, rbcL and the trnL-F intergenic spacer (IGS), for 18 Hawaiian taxa, 45 non-Hawaiian taxa, and two outgroup species. For individual fragments, we estimated phylogenetic relationships using Bayesian inference and maximum parsimony. We performed a combined analysis of both cpDNA fragments employing Bayesian inference, maximum parsimony, and maximum likelihood. These analyses indicate that Hawaiian Dryopteris is not monophyletic, and that there were at least five separate colonizations of the Hawaiian Islands by different species of dryopteroid ferns, with most of the five groups having closest relatives in SE Asia. The results suggest that one colonizing ancestor, perhaps from SE Asia, gave rise to eight endemic taxa (the glabra group). Another colonizing ancestor, also possibly from SE Asia, gave rise to a group of five endemic taxa (the exindusiate group). Dryopteris fusco-atra and its two varieties, which are endemic to Hawai'i, most likely diversified from a SE Asian ancestor. The Hawaiian endemic Nothoperanema rubiginosum has its closest relatives in SE Asia, and while the remaining two species, D. wallichiana and D. subbipinnata, are sister species, their biogeographical origins could not be determined from these analyses due to the widespread distributions of D. wallichiana and its closest non-Hawaiian relative.     

Length–weight relationships for common juvenile fishes and prey species in Hawaiian estuaries

Length–weight relationships (LWR) using 3876 individuals were determined for 39 species sampled in estuarine systems in the main Hawaiian Islands, including 33 species of fish from 21 families, five species of crustaceans (prey) from two families, and one mollusk species. For 17 species, LWRs were estimated for the first time. For another 14 species their LWRs were newly reported for Hawaiian waters, including seven introduced taxa that are well established in the islands' estuaries. The LWRs for all taxa include juvenile and sub‐adult size ranges commonly encountered in Hawaiian estuaries.     

Phylogenetic position and biogeography of Hillebrandia sandwicensis (Begoniaceae): a rare Hawaiian relict

The Begoniaceae consist of two genera, Begonia, with approximately 1400 species that are widely distributed in the tropics, and Hillebrandia, with one species that is endemic to the Hawaiian Islands and the only member of the family native to those islands. To help explain the history of Hillebrandia on the Hawaiian Archipelago, phylogenetic relationships of the Begoniaceae and the Cucurbitales were inferred using sequence data from 18S, rbcL, and ITS, and the minimal age of both Begonia and the Begoniaceae were indirectly estimated. The analyses strongly support the placement of Hillebrandia as the sister group to the rest of the Begoniaceae and indicate that the Hillebrandia lineage is at least 51-65 million years old, an age that predates the current Hawaiian Islands by about 20 million years. Evidence that Hillebrandia sandwicensis has survived on the Hawaiian Archipelago by island hopping from older, now denuded islands to younger, more mountainous islands is presented. Various scenarios for the origin of ancestor to Hillebrandia are considered. The geographic origin of source populations unfortunately remains obscure; however, we suggest a boreotropic or a Malesian-Pacific origin is most likely. Hillebrandia represents the first example in the well-studied Hawaiian flora of a relict genus.     

Multiple cave invasions by species of the planthopper genus Oliarus in Hawaii (Homoptera: Fulgoroidea: Cixiidae)

Although geologically young, the Hawaiian Islands harbour a rich and remarkably diverse fauna of terrestrial troglobites: more than 70 cave species are known from Kauai, Oahu, Molokai, East Maui, and Hawaii Island. Among the more speciosc groups that have invaded the subterranean biome are the planthoppers (Homoptera Cixiidae) of the genus Oliarus. Five new obligately cavernicolous (troglobitic) Oliarus species which differ in their degree of troglomorphy and male genital structures, are described from lava tubes on the Hawaiian Islands: O. lorettae sp. nov. and O. makaiki sp. nov. from Hawaii Island, O. gagnei sp. nov. and O. ivaikau sp. nov. from Maui Island, and O. kalaupapae sp. nov. from Molokai Island. Short diagnoses of the two troglobitic species already known, 0. polyphemus Fennah, 1973 from Hawaii Island and 0. priola Fennah, 1973 from Maui are provided. Notes on the ecology and distribution of all cavernicolous species are given. Morphological evidence suggests that each of the seven cavernicolous Oliams species from Hawaii represents a separate, independent adaptive shift to underground environments. Potential relationships to the extant epigean species are discussed.     

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.     

Repeated range expansion and niche shift in a volcanic hotspot archipelago: Radiation of C4 Hawaiian Euphorbia subgenus Chamaesyce (Euphorbiaceae)

Woody perennial plants on islands have repeatedly evolved from herbaceous mainland ancestors. Although the majority of species in Euphorbia subgenus Chamaesyce section Anisophyllum (Euphorbiaceae) are small and herbaceous, a clade of 16 woody species diversified on the Hawaiian Islands. They are found in a broad range of habitats, including the only known C₄ plants adapted to wet forest understories. We investigate the history of island colonization and habitat shift in this group. We sampled 153 individuals in 15 of the 16 native species of Hawaiian Euphorbia on six major Hawaiian Islands, plus 11 New World close relatives, to elucidate the biogeographic movement of this lineage within the Hawaiian island chain. We used a concatenated chloroplast DNA data set of more than eight kilobases in aligned length and applied maximum likelihood and Bayesian inference for phylogenetic reconstruction. Age and phylogeographic patterns were co‐estimated using BEAST. In addition, we used nuclear ribosomal ITS and the low‐copy genes LEAFY and G3pdhC to investigate the reticulate relationships within this radiation. Hawaiian Euphorbia first arrived on Kaua`i or Ni`ihau ca. 5 million years ago and subsequently diverged into 16 named species with extensive reticulation. During this process Hawaiian Euphorbia dispersed from older to younger islands through open vegetation that is disturbance‐prone. Species that occur under closed vegetation evolved in situ from open vegetation of the same island and are only found on the two oldest islands of Kaua`i and O`ahu. The biogeographic history of Hawaiian Euphorbia supports a progression rule with within‐island shifts from open to closed vegetation.     

Botryosphaeriaceae associated with Acacia heterophylla (La Réunion) and Acacia koa (Hawaii)

Acacia koa and A. heterophylla are commonly occurring native trees on the Hawaiian Islands and La Réunion, respectively. A recent phylogenetic study suggested that A. heterophylla renders A. koa paraphyletic, and that the former likely arose from the Hawaiian Islands around 1.4 million years ago. An intriguing question is whether their microbiota is similar, although they occur naturally in two very distant geographical locations. In this study, we compared the fungi in the Botryosphaeriaceae isolated from natural populations of A. koa and A. heterophylla. These fungi were chosen because they commonly occur on woody plants and some are important pathogens. They are also known to have been moved globally on asymptomatic plant materials. Isolates were identified based on comparisons of DNA sequence data for the rDNA-ITS, TEF1-α and β-tubulin loci. Ten Botryosphaeriaceae species were identified, of which four species were specific to A. koa from the Hawaiian Islands and five to A. heterophylla in La Réunion. Only one species, Neofusicoccum parvum, which is known to have a wide global distribution, was common to both hosts. The overall results of this study suggest that although A. koa and A. heterophylla share a recent evolutionary history, they have established independent microbiota, at least in terms of the Botryosphaeriaceae.     

PACT in practice: comparative historical biogeographic patterns and species–area relationships of the Greater Antillean and Hawaiian Island terrestrial biotas

Aim To compare the evolutionary and ecological patterns of two extensively studied island biotas with differing geological histories (the Hawaiian Islands and the Greater Antilles). We evaluated the results from PACT (phylogenetic analysis for comparing trees), an innovative approach that has been proposed to reveal general patterns of biotic expansion (between regions) and in situ (within a region) diversification, as well as species–area relationships (SAR) and the taxon pulse dynamic. Location The Hawaiian Islands and Greater Antilles. Methods We used the PACT algorithm to construct general area cladograms and identified biotic expansion and in situ nodes. We analysed the power‐law SAR and relative contribution of biotic expansion and in situ diversification events using power‐law and linear regression analyses. Results Both biotic expansion and in situ nodes were prevalent throughout the PACT general area cladograms (Greater Antilles, 55.9% biotic expansion, 44.1% in situ; Hawaiian Islands, 40.6% biotic expansion, 59.4% in situ). Of the biotic expansion events, both forward and backward events occurred in both regions (Greater Antilles, 85.1% forward, 14.9% backward; Hawaiian Islands, 65% forward, 35% backward). Additionally, there is a power‐law SAR for the Greater Antilles but not for the Hawaiian Islands. However, exclusion of Hawai'i (the youngest, largest Hawaiian Island) produced a power‐law SAR for the Hawaiian Islands. Main conclusions The prevalence of in situ events as well as forward and backward biotic expansion events reveals that both Hawaiian and Greater Antillean biotas have evolved through alternating episodes of biotic expansion and in situ diversification. These patterns are characteristic of the taxon pulse dynamic, for which few data have previously been recorded on islands. Additionally, our analysis revealed that historical influences on the power‐law SARs are pronounced in both assemblages: old, small islands are relatively species rich and young, large islands are relatively species poor. Thus, our PACT results are consistent with hypotheses of geological influence on the evolution of island biotas and also provide greater insight into the role of the taxon pulse dynamic in the formation of island equilibria.