Phylogeography and ecology of an endemic radiation of Hawaiian aquatic case-bearing moths (Hyposmocoma: Cosmopterigidae)

The endemic moth genus Hyposmocoma (Lepidoptera: Cosmopterigidae) may be one of the most speciose and ecologically diverse genera in Hawaii. Among this diversity is the Hyposmocoma saccophora clade with previously unrecorded aquatic larvae. I present a molecular phylogeny based on 773 base pairs (bp) of the mitochondrial gene cytochrome c oxidase subunit I and 762 bp of the nuclear gene elongation factor 1-alpha. Topologies were constructed from data using maximum-parsimony, maximum-likelihood and Bayesian search criteria. Results strongly support the monophyly of the H. saccophora clade and the monophyly of the genus Hyposmocoma. The H. saccophora clade has single-island endemic species on Oahu, Molokai and West Maui. By contrast, there are three species endemic to Kauai, two being sympatric. The H. saccophora clade appears to follow the progression rule, with more basal species on older islands, including the most basal species on 11 Myr-old Necker Island, one of the Northwestern Hawaiian Islands. Aquatic behaviour either evolved recently in the species on the main Hawaiian Islands or was secondarily lost on the arid northwestern Necker Island. The phylogeny suggests that Hyposmocoma is older than any of the current main islands, which may, in part, explain Hyposmocoma's remarkable diversity.     

The Hawaiian amphibious caterpillar guild: new species of Hyposmocoma (Lepidoptera: Cosmopterigidae) confirm distinct aquatic invasions and complex speciation patterns

Although aquatic caterpillars are a globally rare lifestyle, we have found them in multiple, independent lineages of the endemic moth genus Hyposmocoma across all of the Hawaiian Islands with flowing water. We formally describe 11 new species of Hyposmocoma that belong to four different larval case types: cone, bugle, medium burrito, and large burrito: Hyposmocoma kahamanoa sp. nov. from Oahu Island, Hyposmocoma kamakou sp. nov. from Molokai Island, Hyposmocoma kahaiao sp. nov. , Hyposmocoma waihohonu sp. nov. , and Hyposmocoma moopalikea sp. nov. from Maui Island, and Hyposmocoma aumakuawai sp. nov. , Hyposmocoma eepawai sp. nov. , Hyposmocoma ipowainui sp. nov. , Hyposmocoma kawaikoi sp. nov. , Hyposmocoma uhauiole sp. nov. , and Hyposmocoma wailua sp. nov. from Kauai Island. We also illustrate and describe in detail the aquatic case‐bearing larva of Hyposmocoma kahamanoa. Despite having similar ecologies as algae and lichen grazers at and below the water line of streams, prior research indicates that species with each case type constitute an independent lineage, with terrestrial sister taxa, and therefore the different groups of species bearing unique case types each represent an independent aquatic invasion. The case‐bearing larvae often occur sympatrically, and on Kauai even species with similar case‐types occur together, suggesting complex patterns of speciation and either past periods of isolation or sympatric speciation. Phylogenetic analysis of 2243 base pairs from two nuclear and one mitochondrial gene for 18 species confirm that each species is endemic to a single volcano, and that morphological divergence within case‐types has not been dramatic. Diversification has been complex, and superficially similar case type lineages are not all monophyletic. Kauai, the oldest but smallest of the major high islands, supports more species in the aquatic guild than any other island, thus island age, rather than size, may be important in generating diversity in this group. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 15–42.     

Phylogeographic patterns of Hawaiian Megalagrion damselflies (Odonata: Coenagrionidae) correlate with Pleistocene island boundaries

The Pleistocene geological history of the Hawaiian Islands is becoming well understood. Numerous predictions about the influence of this history on the genetic diversity of Hawaiian organisms have been made, including the idea that changing sea levels would lead to the genetic differentiation of populations isolated on individual volcanoes during high sea stands. Here, we analyse DNA sequence data from two closely related, endemic Hawaiian damselfly species in order to test these predictions, and generate novel insights into the effects of Pleistocene glaciation and climate change on island organisms. Megalagrion xanthomelas and Megalagrion pacificum are currently restricted to five islands, including three islands of the Maui Nui super-island complex (Molokai, Lanai, and Maui) that were connected during periods of Pleistocene glaciation, and Hawaii island, which has never been subdivided. Maui Nui and Hawaii are effectively a controlled, natural experiment on the genetic effects of Pleistocene sea level change. We confirm well-defined morphological species boundaries using data from the nuclear EF-1alpha gene and show that the species are reciprocally monophyletic. We perform phylogeographic analyses of 663 base pairs (bp) of cytochrome oxidase subunit II (COII) gene sequence data from 157 individuals representing 25 populations. Our results point to the importance of Pleistocene land bridges and historical island habitat availability in maintaining inter-island gene flow. We also propose that repeated bottlenecks on Maui Nui caused by sea level change and restricted habitat availability are likely responsible for low genetic diversity there. An island analogue to northern genetic purity and southern diversity is proposed, whereby islands with little suitable habitat exhibit genetic purity while islands with more exhibit genetic diversity.     

Evaluation of relationships within the endemic Hawaiian Platynini (Coleoptera: Carabidae) based on molecular and morphological evidence

Relationships among 69 species of Hawaiian Platynini, a monophyletic beetle radiation, was investigated based on evidence from five data partitions, comprising mitochondrial and nuclear DNA sequences (cytochrome oxidase II, 624 bp; cytochrome b, 783 bp; 28S rDNA, 668 bp; wingless; 441 bp) and morphology (206 features of external and internal anatomy). Results from individual and combined data analyses generally support the monophyly of three putative divisions within Platynini in Hawaii: Division 0 (Colpocaccus species group), Division 1 (Blackburnia species group), and Division 2 (Metromenus species group). However, relationships within and among these three divisions differ from previous morphological hypotheses. An extensive series of sensitivity analyses was performed to assess robustness of recovered clades under a variety of weighted parsimony conditions. Sensitivity analyses support the monophyly of Divisions 0 and 1, but were equivocal for the monophyly of Division 2. A phylogeny based on combined data suggests at least four independent losses/reductions of platynine flight wings. The combined analysis provides corroboration for biogeographic hypotheses, including (1) colonization of Kauai by Hawaiian Platynini with subsequent dispersal and colonization along the island chain from Oahu to Maui Nui to Hawaii Island and (2) incongruent area relationships among Eastern Molokai, West Maui, and Haleakala for two species triplets.     

Molecular phylogeny of Banza (Orthoptera: Tettigoniidae), the endemic katydids of the Hawaiian Archipelago

The extant endemic katydids (Orthoptera: Tettigoniidae) of the Hawaiian Archipelago include one to three species per high island and a single species on Nihoa, all currently placed in the genus Banza. These acoustic insects provide an excellent opportunity for investigating the evolution of reproductive isolation and speciation, but such studies require an understanding of phylogenetic relationships within the group. We use maximum parsimony, likelihood-based Bayesian inference, and maximum likelihood to infer phylogenetic relationships among these taxa, based on approximately 2kb of mitochondrial cytochrome oxidase I and cytochrome b. Our results strongly support two distinct high island clades: one clade ("Clade I") composed of species from Kauai, Oahu, Molokai, and Lanai and another clade ("Clade II") composed of species from Maui and Hawaii (Banza unica, from Oahu, may be basal to both these clades, but its placement is not well resolved). Within these clades, some inferred relationships are strongly supported, such as the sister status of B. kauaiensis (Kauai) and B. parvula (Oahu) within Clade I, but other relationships remain more ambiguous, such as the relative position of B. brunnea (Maui) within Clade II. Although a detailed reconstruction of the historical biogeography of the Hawaiian katydids is difficult, we use our genetic data combined with the known geological history of the Hawaiian Islands to set limits on plausible historical scenarios for diversification of this group. Beyond these historical biogeographic inferences, our results indicate possible cryptic speciation on both Oahu and Hawaii, as well as what may be unusually high average rates of nucleotide substitution. The present work sets the stage for future genetic and experimental investigations of this group.     

Phylogeographic patterns and demographic history of Schiedea globosa (Caryophyllaceae) on the Hawaiian Islands

Geomorphological changes have been demonstrated to have had profound impacts on biodiversity, often leading to demographic expansions and contractions and allopatric divergence of taxa. We examined DNA sequence variation at two nuclear and one maternally inherited plastid locus among 10 populations of Schiedea globosa on the Hawaiian Islands to assess the primary factors shaping genetic structure, phylogeographic patterns, and the importance of geographic isolation to population divergence. Schiedea globosa has characteristics that may promote gene flow, including wind pollination and rafting of plants in ocean currents. However, we detected significant differentiation among populations on all islands except Hawaii, with the maternally inherited plastid locus having the greatest genetic structure (F(ST) = 0.81). Migration rates across all loci are less than one migrant per generation. We found evidence of growth in several populations and on the islands of Molokai and Maui, which supports population expansion associated with the formation of Maui Nui during the last glacial maximum. Similar to data for many other Hawaiian taxa, these data suggest S. globosa originated on Oahu and subsequently colonized Molokai, Maui, and Hawaii in progression. Given the high level of genetic structure, allopatric divergence will likely contribute to further divergence of populations.     

Bottlenecks and multiple introductions: population genetics of the vector of avian malaria in Hawaii

Avian malaria has had a profound impact on the demographics and behaviour of Hawaiian forest birds since its vector, Culex quinquefasciatus the southern house mosquito, was first introduced to Hawaii around 1830. In order to understand the dynamics of the disease in Hawaii and gain insights into the evolution of vector-mediated parasite-host interactions in general we studied the population genetics of Cx. quinquefasciatus in the Hawaiian Islands. We used both microsatellite and mitochondrial loci. Not surprisingly we found that mosquitoes in Midway, a small island in the Western group, are quite distinct from the populations in the main Hawaiian Islands. However, we also found that in general mosquito populations are relatively isolated even among the main islands, in particular between Hawaii (the Big Island) and the remaining Hawaiian Islands. We found evidence of bottlenecks among populations within the Big Island and an excess of alleles in Maui, the site of the original introduction. The mitochondrial diversity was typically low but higher than expected. The current distribution of mitochondrial haplotypes combined with the microsatellite information lead us to conclude that there have been several introductions and to speculate on some processes that may be responsible for the current population genetics of vectors of avian malaria in Hawaii.     

Ostracoden von Hawaii,insbesondere aus dem marinen Interstitial

The present paper deals with ostracods collected by H. Kunz during October and November 1979 on the Hawaiian Islands (Big Island, Maui and Kauai). 26 species were found, nine species are described, two were already known to science, 15 remain in open nomenclature. Although some of the new species are represented only by a single specimen, the morphologic characters permit to distinguish them from other species of the genus. The ostracods of the Hawaiian Islands are very imperfectly known.      

Cladistic analysis, phylogeny and biogeography of the Hawaiian Platynini (Coleoptera: Carabidae)

The 128 known native Hawaiian species of the tribe Platynini are analysed cladistically. Cladistic analysis is based on 206 unit-coded morphological characters, and also includes forty-one outgroup taxa from around the Pacific Rim. Strict consensus of the multiple equally parsimonious cladograms supports the monophyly of the entire species swarm. The closest outgroup appears to be the south-east Asian-Pacific genus Lorostema Motschulsky, whose species are distributed from India and Sri Lanka to Tahiti, supporting derivation of the Hawaiian platynines from a source in the western or south-western Pacific. The biogeographic relationships of the Hawaiian taxa are analysed using tree mapping, wherein items of error are minimized. The area cladogram found to be most congruent with the phylogenetic relationships, and most defensible based on underlying character data is {Kauai[Oahu(Hawaii{Lanai[East Maui(West Maui + Molokai)]})]}. This progressive vicariant pattern incorporates progressive colonization from Kauai, and vicariance of the former Maui Nui into the present islands of Molokai, Lanai, West Maui and East Maui. The evolution of flightlessness, tarsal structure, pronotal setation and bursal asymmetry are evaluated in the context of the cladogram. Brachyptery is a derived condition for which reversal is not mandated by the cladogram, although repeated evolution of reduced flight wings is required. Tarsal structure supports Sharp's (1903) recognition of Division 1 as a monophyletic assemblage, but exposes his Division 2 as a paraphyletic group requiring removal of the genus Colpocaccus Sharp. Pronotal setation is exceedingly homoplastic, and is not useful for delimiting natural groups. Left-right asymmetry of the bursa copulatrix reversed twice independently, resulting in mirror-image bursal configurations in B. rupicola and Prodisenochus terebratus of East Maui. The amount of character divergence is greater among species comprising Division 1 than among species of its sister group, the redefined Division 2. Based on superior fit of Division 1 relationships to the general biogeographic pattern, a greater speciation rate coupled with more extensive extinction is rejected as the cause for this greater divergence. Intrinsic differentiation in the processes underlying cuticular evolution appears to be more consistent with the observed biogeographic and morphological patterns.     

Population divergence and gene flow in an endangered and highly mobile seabird

Seabirds are highly vagile and can disperse up to thousands of kilometers, making it difficult to identify the factors that promote isolation between populations. The endemic Hawaiian petrel (Pterodroma sandwichensis) is one such species. Today it is endangered, and known to breed only on the islands of Hawaii, Maui, Lanai and Kauai. Historical records indicate that a large population formerly bred on Molokai as well, but this population has recently been extirpated. Given the great dispersal potential of these petrels, it remains unclear if populations are genetically distinct and which factors may contribute to isolation between them. We sampled petrels from across their range, including individuals from the presumably extirpated Molokai population. We sequenced 524 bp of mitochondrial DNA, 741 bp from three nuclear introns, and genotyped 18 microsatellite loci in order to examine the patterns of divergence in this species and to investigate the potential underlying mechanisms. Both mitochondrial and nuclear data sets indicated significant genetic differentiation among all modern populations, but no differentiation was found between historic samples from Molokai and modern birds from Lanai. Population-specific nonbreeding distribution and strong natal philopatry may reduce gene flow between populations. However, the lack of population structure between extirpated Molokai birds and modern birds on Lanai indicates that there was substantial gene flow between these populations and that petrels may be able to overcome barriers to dispersal prior to complete extirpation. Hawaiian petrel populations could be considered distinct management units, however, the dwindling population on Hawaii may require translocation to prevent extirpation in the near future.     

A contribution to the Dicyrtomidae (Collembola) of Hawaii

Ten new species in three subgenera of Dicyrtoma are described from the Hawaiian Islands. Specimens were received from collections made on Hawaii, Maui, Kauai and Oahu. Species definitions are based on chaetotaxy of head, legs and the circumanal region. In addition, presence or absence of lateral spines (neosminthurid setae) on the parafurcular lobes may assist in grouping species within subgenera. Previous records of dicyrtomids from Hawaii include only one species, Ptenothrix ( Papaairioides ) daubia (Folsom). The following new species are described: Ptenothrix ( Ptenothrix ) hawaiicnaeis sp.n., Ptenothrix ( Papirioides ) kauaiensis sp.n., P. ( Papirioides ) serrata sp.n., Dicyrtoma ( Calvatomina ) sylvestratilis sp.n., D. ( Calvatomina ) brevifibra sp.n., D. ( Calvatomina ) tesselata sp.n., D. ( Calvatomina ) longidigita sp.n., D. ( Calvatomina ) bellingeri sp.n., D. ( Calvatomina ) madestris sp.n., and D. ( Calvatomina ) microdentata sp.n.     

The diversity and biogeography of koa-finches (Drepanidini: Rhodacanthis), with descriptions of two new species

All known populations of koa-finches, genus Rhodacanthis , became extinct in the Holocene epoch. Two new species are described here from Quaternary fossil sites in the Hawaiian Islands. One new species, from Kauai and Maui, is roughly the size of the historically known greater koa-finch ( R. palmeri ) but differs in having a more robust skull and in bill morphology. The second new species, from Oahu and Maui, is similar in size to the lesser koa-finch ( R. flaviceps ) but closer to R. palmeri in qualitative osteological traits. The two species of koa-finches known historically from the island of Hawaii are distinct in osteology from the fossil koa-finches on the older Hawaiian islands, indicating that at least two of the four known speciation events in the genus took place within approximately the past 500 kyr. However, the similarity of maxillae from Pleistocene and Holocene sites on Oahu suggests that the Oahu population maintained morphological stasis through the climate changes of the late Quaternary. The evidence that speciation occurred on the youngest island in the archipelago suggests that the process of community assembly on newly emergent Hawaiian landscapes was a stimulus to evolutionary diversification in Rhodacanthis .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 527–541.     

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.     

Disentangling the dynamics of invasive fireweed (<Emphasis Type="Italic">Senecio madagascariensis</Emphasis> Poir. species complex) in the Hawaiian Islands

Studies investigating the genetic variation of invasive species render opportunities to better understand the dynamics of biological invasions from an ecological and evolutionary perspective. In this study, we investigate fine-scale population genetic structure of invasive Senecio madagascariensis (fireweed) using microsatellite markers to determine levels of genetic diversity and how it pertains to introduction history of this species within and among the Hawaiian Islands. Dispersal patterns were interpreted and, together with a habitat suitability analysis, we aim to describe the potential range expansion of S. madgascariensis within the islands. Bayesian and frequency-based analyses revealed genetic structure with two major genetic demes corresponding to the two fireweed-infested islands of Maui and Hawaii. Both these demes showed further genetic sub-structure, each consisting of three genetically distinct subgroups. Overall, fireweed showed significant levels of inbreeding. Major genetic demes (Maui and Hawaii) differed in observed heterozygosities, inbreeding and genetic structure, each harbouring a large proportion of private alleles. In contrast to the current understanding of fireweed’s introduction history between the Hawaiian Islands, fine-scale population genetic parameters suggest that this species has been introduced at least twice, possibly even more, to the archipelago. Spatial analyses also revealed high correlation between genetic similarity and geographical proximity (>2 km apart) followed by a sharp decline. In addition, a single population was identified that likely resulted from a rare human- or animal-mediated extreme long-distance dispersal event from Maui to Hawaii. Bayesian and likelihood estimates of ‘first generation migrants’ also concurred that contemporary dispersal occurs more frequently over smaller spatial scales than larger scales. These findings indicate that spread in this species occurs primarily via a stratified strategy. Predictions from habitat suitability models indicate all Hawaiian Islands as highly suitable for fireweed invasion and the movement of propagules to currently uninfested islands and outlying suitable habitats should be avoided to circumvent further expansions of the invasion.     

Systematics and evolution of Iolania (Hemiptera: Fulgoromorpha: Cixiidae) from Hawai'i

Abstract.  Within the planthopper taxon Cixiidae, which is distributed worldwide, only two lineages have colonized the Hawaiian Islands: Iolania Kirkaldy, 1902, and Oliarus Stål, 1862 , and subsequently given rise to endemic species. Neither radiation has been studied in depth hitherto. Here the degree of speciation within Iolania is assessed and a taxonomic revision including a key to the species based on the male genitalic characters is provided. Six endemic species are recognized: I. perkinsi Kirkaldy, I. koolauensis Giffard, I. oahuensis Giffard, I. lanaiensis Giffard, I. mauiensis Giffard and I. kraussohana sp.n. A lectotype is designated for the type-species I. perkinsi Kirkaldy, and I. perkinsi notata Kirkaldy is interpreted as an invalid name. Morphological arguments for the monophyly of Iolania and phylogenetic relationships among the species are discussed. A plausible scenario for the sequence of speciation events and history of colonization within the Hawaiian Islands is attempted. Combined information from taxon- and area-cladograms suggests progressive inter-island dispersal from older to younger islands in the Hawaiian chain as the major pattern of colonization and speciation.     

First record of troglobitic Hemiptera (Fulgoromorpha: Cixiidae) from La Réunion Island

A new species of troglobitic Cixiidae (Hemiptera: Fulgoromorpha), Brixia briali n. sp. is described from a lava tube on La Réunion Island. Information on its ecology, distribution, and comments on its tentative systematic placement into the genus Brixia is provided. The new discovery from La Réunion is placed into the context of cavernicolous Fulgoromorpha of the world, their zoogeography, ecology and biology.     

Evolutionary relationships of four species of hawaiian Drosophila as measured by DNA reassociation

Four species of the Hawaiian Drosophila planitibia subgroup which are homosequential in their polytene chromosomes are resident on the islands of Molokai, Maui and Hawaii. Comparisons of DNA sequence divergence in these four have been made by hybridization of total single-copy radiolabeled tracer DNA from each of the species with excess nonlabeled DNA from each of the species, and measurement of the reduction of average melting temperature (DeltaTma) was made in 2.4 m tetraethyl ammonium chloride. The mean DeltaTma between either D. heteroneura or D. silvestris and either D. planitibia or D. differens was found to be 1.06 degrees , whereas the difference between D. planitibia and D. differens in 0.65 degrees and between D. heteroneura and D. silvestris is 0.75 degrees . These measurements taken together with the distances calculated from isozyme studies, chromosomal relationships, as well as the island locations indicate that the ancestor of these species diverged from other planitibia subgroup flies on Molokai [age 1.8 million years before present, (My BP)]. We hypothesize that one line became the present-day D. differens and diverged probably at the time of formation of East Maui (0.8-1 My BP) to form the species D. planitibia. Flies from the other line migrated to Hawaii soon after its formation (0.7 My BP) to form the two species D. heteroneura and D. silvestris.     

Rapid evolution of cuticular hydrocarbons in a species radiation of acoustically diverse Hawaiian crickets (Gryllidae: trigonidiinae: Laupala)

Understanding the origin and maintenance of barriers to gene exchange is a central goal of speciation research. Hawaiian swordtail crickets (genus Laupala) represent one of the most rapidly speciating animal groups yet identified. Extensive acoustic diversity, strong premating isolation, and female preference for conspecific acoustic signals in laboratory phonotaxis trials have strongly supported divergence in mate recognition as the driving force behind the explosive speciation seen in this system. However, recent work has shown that female preference for conspecific male calling song does not extend to mate choice at close range among these crickets, leading to the hypothesis that additional sexual signals are involved in mate recognition and premating isolation. Here we examine patterns of variation in cuticular lipids among several species of Laupala from Maui and the Big Island of Hawaii. Results demonstrate (1) a rapid and dramatic evolution of cuticular lipid composition among species in this genus, (2) significant differences among males and females in cuticular lipid composition, and (3) a significant reduction in the complexity of cuticular lipid profiles in species from the Big Island of Hawaii as compared to two outgroup species from Maui. These results suggest that behavioral barriers to gene exchange in Laupala may be composed of multiple mate recognition signals, a pattern common in other cricket species.     

A new genus and two new species of Peltogastridae (Crustacea: Cirripedia: Rhizocephala) parasitizing hermit crabs from Okinawa Island (Ryukyu Archipelago, Japan), and their DNA-barcodes

A new genus and two new species of Peltogastridae, Peltogaster postica sp. nov. and Ommatogaster nana gen. et sp. nov., are described from Okinawa Island, Ryukyu Islands, southwestern Japan. The two new rhizocephalans were found to be parasitic on the estuarine hermit crabs, Pagurus minutus Hess, 1865 and Diogenes leptocerus Forest, 1956, respectively. Peltogaster postica sp. nov. is allied to P. curvata Kossmann, 1874, P. paguri Rathke, 1842 , and P. reticulata Shiino, 1943 , but is distinguished by its relative length and internal and external structures of the mature externa. Ommatogaster gen. nov. is established for the present new species O. nana based on the morphologies of the visceral mass of the externa and the presence of a nauplius eye in the larvae. Partial COI sequences were obtained from the two new species and one known species, Dipterosaccus indicus Van Kampen and Boschma, 1925, to test the possible usefulness of the sequences as tags for species identification.