The giant, the flightless, and the iridescent: seven new endemic genera of Hawaiian leaf-roller moths (Lepidoptera: Tortricidae)

Chromosomal Tracers of Founder Events

The Hawaiian spider genus Orsonwelles , gen. nov. (Araneae : Linyphiidae) is described. All Orsonwelles species are single island endemics: Kauai harbours six species; Oahu has three; Molokai has two; and Maui and Hawaii have one species each. The thirteen species included in Orsonwelles are described and illustrated: O. torosus (Simon), comb. nov., O. malus , sp. nov., O. calx , sp. nov., O. ventus , sp. nov., O. bellum , sp. nov. and O. iudicium , sp. nov. from Kauai; O. polites , sp. nov. (the type species), O. ambersonorum , sp. nov. and O. arcanus , sp. nov. from Oahu; O. othello , sp. nov. and O . macbeth , sp. nov. from Molokai; O. falstaffius , sp. nov. from Maui; and O. graphicus (Simon), comb. nov. from Hawaii. A total of 55 morphological characters (plus one behavioural character) were scored for twelve taxa (four Orsonwelles species plus eight linyphiid outgroups) to test the monophyly of the genus using cladistic methods. The most parsimonious cladograms provide robust character support for the monophyly of Orsonwelles . A single colonisation of the Hawaiian archipelago is hypothesised to explain the presence of these species in the Hawaiian Islands. This genus represents a case of insular gigantism (these are the largest linyphiids described), although the close relatives of Orsonwelles remain unknown. Their web architecture is also described and illustrated.

Two new psyllid species, Ctenarytaina elinae sp. nov. and Anomocephala rosiegeorgeorum sp. nov., are described from French Polynesia. Both are single island endemics known only from a small number of specimens. Ctenarytaina elinae sp. nov. is in the primarily Austro-Pacific genus Ctenarytaina Ferris & Klyver (Aphalaridae), and with the addition of C. elinae there are now three Ctenarytaina species in French Polynesia. Ctenarytaina elinae sp. nov. is endemic to Tahiti in the Society Islands, and although the host is unrecorded there is some evidence to suggest it may be Metrosideros (Myrtaceae). Previously monotypic, Anomocephala Tuthill (Triozidae), with the addition of A. rosiegeorgeorum sp. nov., now includes two species, and both of these are endemic to Rapa island in the southern Austral archipelago. Both the genus, Anomocephala, and the type species, Anomocephala unica Tuthill, 1942, are redescribed. The host plant of Anomocephala rosiegeorgeorum sp. nov. is likely Metrosideros collina. The new species names recognize entomologists working in the Pacific, and this contribution to this special issue honours one of the most prolific psyllid researchers and taxonomists, Daniel Burckhardt.

The subfamily Cheloninae has not been studied in any detail for Australia for more than 80 years, and yet they are an important group of parasitoids of lepidopteran larvae, with many undescribed species, some of which cannot be easily assigned to genera. As a precursor to undertaking taxonomic revisions of specific genera for the Australian fauna, this study assesses the likely species richness of the fauna, provides a key to genera to facilitate their identification, provides a checklist of species and notes on their taxonomy, and discusses their biology. In so doing, we recognise two new genera, Austroascogaster gen. nov., represented by four species, A. gloriousensis sp. nov. (Queensland), A. mellosa sp. nov. (type species) (Queensland, Papua New Guinea, Indonesia), A. nigripropodea sp. nov. (Queensland) and A. varia sp. nov. (Queensland, Indonesia), and Phanaustrotoma gen. nov. represented by P. alba sp. nov. (Queensland) and P. pallina sp. nov. (type species) (Queensland). Following this study, the Australian chelonine fauna comprises 47 species, of which 43 are endemic to the continent; two species, Chelonus phthorimaeae Gahan and Chelonus curvimaculatus Cameron, have been successfully introduced as biological control agents against the potato tuber moth (Phthorimaea operculella (Zeller); Gelechiidae), while three species represent questionable records as there is no reliable evidence that they ever established on the continent, viz, Ascogaster quadridentata Wesmael, Chelonus blackburni Blackburn & Cameron and Phanerotoma hendecasisella Cameron. Based on its description and biology, Phanerotoma coccinellae Girault (Queensland) is undoubtedly not a member of the Cheloninae; however, the holotype of this species is apparently lost.

A new endemic psyllid genus from the Hawaiian Islands, Stevekenia gen. nov., is described. Two new species, both single island endemics, feed on host plants in the endemic genus Nothocestrum (Solanaceae). Stevekenia nothocestri sp. nov. is found on Nothocestrum longifolium on the island of Oahu and Stevekenia aiea sp. nov. is found on N. peltatum on the island of Kauai. The host plants are montane rainforest species and both are on the IUCN Red List of Threatened Species. In particular, N. peltatum, which is endemic to the island of Kauai, is considered critically endangered, and therefore a similarly endangered status for these host-specific insects is likely. Only a few specimens of each Stevekenia species have been found, suggesting occurrence at low abundance. The host plants survive either as isolated individuals or small fragmented populations, which may already constitute less than the required critical host plant density for the long term survival of this psyllid genus. Unique morphological characters distinguish Stevekenia from other Hawaiian genera, and the placement of this genus within Triozidae is discussed. Mitochondrial DNA barcodes for the new species are provided.

Seven new species of endemic leaf-roller moths are described from the Hawaiian Islands: Eccoptocera hinanohomauna sp. nov. (O‘ahu), Eccoptocera ohiaha, sp. nov. (O‘ahu), Pararrhaptica kaiona, sp. nov. (O‘ahu), Pararrhaptica pilikuanani, sp. nov. (Kaua‘i), Spheterista huakunana, sp. nov. (O‘ahu), Spheterista hiwakakahi, sp. nov. (Kaua‘i), and Spheterista hakeaiki, sp. nov. (O‘ahu). With the possible exception of S. hakeaiki, all species appear to be single-island endemics. In addition, Pararrhaptica pycnomias (Meyrick), syn. nov. (O‘ahu), is treated as a junior synonym of Pararrhaptica notocosma (Meyrick) (O‘ahu), Macraesthetica Meyrick is transferred from Eucosmini to Olethreutini, and Macraesthetica semicinereana (Swezey), comb. nov. (Hawai‘i), is transferred from Nuritamburia Koçak & Kemal. A conservation status is proposed for each of the newly described species.

Adaptive radiation describes the divergence of an ancestral taxon into multiple, phenotypically diverse species, adapted to a range of ecological niches by means of natural selection. The process is recognized as a fundamental reason for the origin of biodiversity. The main driver of adaptive radiation is ecological opportunity, though the specific agents are often poorly understood with the exception of some iconic lineages. Many well-studied adaptive radiations are island endemics, which makes island systems an ideal study system for adaptive radiation. Oceanic islands represent discrete replicates of the evolutionary process, as they are isolated, comparatively small, and often topographically complex. Species communities are formed by colonization and in situ diversification. The Hawaiian Islands are the most isolated archipelago on earth and home to a range of adaptively radiating lineages. The islands form as the Pacific plate passes over a magmatic hotspot with the eight current high islands originating within the last ca. 5-6 million years and the majority of the native biodiversity diverging within that time. The genus Melicope colonized numerous archipelagos throughout the Pacific including the Hawaiian Islands, where the lineage comprises currently 54 endemic species and represents the largest radiation of woody plants on the islands. Most species are single-island endemics and adapted to a variety of habitat types and elevational ranges. The lineage is monophyletic with an estimated crown age predating the rise of the current high islands, the oldest of which originated approximately 5 million years ago. As for many adaptively radiating lineages, phylogenetic inference based on Sanger sequencing has not been sufficient to resolve species or deeper level relationships in Hawaiian Melicope. Recent years have seen development of high throughput sequencing methods and their increasing application to solve recalcitrant relationships.
\nIn this thesis, I examined the evolutionary trajectory of the Hawaiian Melicope adaptive radiation. I investigated the so-called ‘island syndrome’, which describes a set of traits commonly characterizing successful island colonizers, including recent polyploidy and shifts associated with subsequent establishment, in Hawaiian Melicope. I utilized restriction site-associated high throughput sequencing (RAD-seq) to reconstruct species relationships and historical biogeography in the lineage and estimate diversification rates and the impact of habitat adaption on species divergence. 
\nRAD-seq datasets provided unprecedented resolution of species relationships in Hawaiian Melicope. However, the size and complexity of high throughput sequencing datasets require a high computational effort, which currently limits the applicability of algorithms for phylogenetic inference to concatenated analysis or site-specific coalescence-based methods. I employed both methods and found them to result in incongruent relationships for the backbone of the Hawaiian Melicope topology. Concatenation violates the assumptions of the multispecies coalescent model, while site-based methods are statistically inconsistent but less accurate in simulated and empirical datasets. Considering the increased accuracy of the concatenated approaches as evaluated by quartet concordance methods and the synergistic effect of concatenation, I concluded that results of concatenated analysis reflect the relationships of Hawaiian Melicope best. 
\nResults of flow cytometric screening of 32 Hawaiian species, representing 66% of the described diversity, and literature searches indicate that the ancestor of Hawaiian Melicope did not show traits associated with successful colonizers. The genus seemingly retained colonization success while exhibiting a combination of traits that typically characterize well-established island specialists. In particular, the ancestral Melicope colonist was not a recent polyploid. Neopolyploidy increases evolutionary flexibility and thus enhances chances for establishment and adaption. In Hawaiian Melicope flexibility is possibly facilitated by introgressive hybridization events.
\nPhylogenetic reconstruction based on RAD-seq datasets provides evidence for two ancient and several recent introgression events. Extant Hawaiian Melicope are divided into five fully supported main clades, two of which correspond to morphologically circumscribed infrageneric groups, whereas three morphologically defined taxonomic units are not monophyletic. All in all, 24 species were included with multiple samples, four of which were resolved as non-monophyletic. Finally, I confirmed that the Melicope radiation endemic to the Marquesas Islands originated from the Hawaiian radiation. These results highlight the necessity for a taxonomic revision in the lineage.
\nEstimated divergence times revealed that the Hawaiian archipelago was colonized prior to the origin of the current high islands. Inter-Island colonization patterns largely follow the progression rule from older to younger islands, but back colonizations to older islands occurred. Extant diversity results from recent divergence of a small number of taxa prevailing through the bottlenecks represented by the origin and colonization of the high islands. Long internal branches and estimated diversification rates indicate a high extinction rate, possibly related to the consequences of volcanic activity and the impact of glacial cycles. Consequently habitat types that are more vulnerable to climatic changes, i.e. dry ranges and bogs show high speciation and extinction rates. Increased rates of diversification are linked to habitat dissection and frequent ecological trait shifts.

The Hawaiian flora, because of its great isolation, high levels of endemism, known lineages, and high rates of endangerment, offers unique opportunities to explore patterns of endangerment related to phylogeny, ecological and life history traits, and geographic patterns. Nine percent of the native flora of 1159 taxa are already extinct, and 52.5% are at risk (extinct, endangered, vulnerable, or rare). Risk is strongly associated with limited geographic distribution at several scales: endemic taxa (native only to the Hawaiian Islands) are at far greater risk than indigenous taxa (with both Hawaiian and extra-Hawaiian ranges); single-island endemics are more at risk than multi-island endemics; small islands have the highest proportion of endemic taxa at risk; and endemics with more limited habitat distributions (elevation, community type) are more at risk. Historic population density is a strong predictor of risk, and taxa with low historic population densities are at greatest risk with rapid anthropogenic changes. Among the major islands, Maui Nui has the highest percent of taxa that are extinct. Kaua'i has the lowest percent of extinct taxa and the highest proportion of single-island endemic taxa that are rare. Endemic taxa at risk are associated with distributions in shrublands, forests, bogs, and cliff habitats. Endemic taxa with distributions in low elevation dry habitats have the highest proportion of taxa at risk, but the greatest absolute numbers of taxa at risk have distributions in mesic lowland and montane forests, and in wet montane forests. The life history patterns associated with risk are complicated, and inclusion of the effects of evolutionary relationships (lineages) changes some of these patterns. Species level analyses without respect to lineage shows risk associated with monomorphic (hermaphroditic) breeding systems and bird pollination because of the large number of hermaphroditic, bird-pollinated species in the Campanulaceae. Analyses incorporating the effect of lineage greatly reduce the impact of large lineages and result in an association of risk with insect pollination, and no effect of breeding system. There is no association of lineage size and the percent of taxa at risk within the lineage; endemic taxa from lineages with large radiations are at no greater risk than endemic single-taxon lineages. The percentages of taxa at risk at the family level in the Hawaiian Islands and worldwide (excluding Hawaiian taxa) are positively correlated, although flowering plant families in the Hawaiian Islands have a much greater proportion of taxa at risk. Some of the approaches described here may be useful to predict geographical and biological patterns of endangerment in island and island-like ecosystems under increasing pressures of endangerment and extinction.

Two morphologically similar genera of Penthimiini, Chanohirata Hayashi Machida, 1996 and Reticuluma Cheng Li, 2005 are reviewed. Three new species, Chanohirata yunnana, C. minima and C. plania spp. nov., are described and illustrated and a new synonymy is proposed: C. theae (Matsumura, 1912) equals Reticuluma testacea (Kuoh, 1991) syn. nov. The following new combinations are also proposed for species previously included in Reticuluma: C. bipennata (Xu, Yu, Dai Yang) comb. nov., C. dactyla (Fu Zhang) comb. nov., C. eurya (Fu Zhang) comb. nov., C. hamata (Wang Zhang) comb. nov., C. lini (Cheng Li) comb. nov., C. lageniformia (Wang Zhang) comb. nov. and C. spinata (Cheng Li) comb. nov. As a result, the definition of Reticuluma is restricted to include only the type species, R. citrina Cheng Li, which differs from Chanohirata in having an apical process on the male pygofer. A species checklist and a key to all known species of Chanohirata are presented along with a distribution map for China.

A molecular phylogeny of the Pacific clade of Cyrtandra (Gesneriaceae) reveals a Fijian origin, recent diversification, and the importance of founder events

Tropizodium, a new genus of the spider family Zodariidae belonging to the “femoral organ clade”, is described. It contains three species all known from both sexes: T. peregrinum n. sp., the type species from northern Australia, and T. molokai n. sp. and T. trispinosum (Suman) n. comb., both from Hawaii. It is characterized by somatic characters such as the small intercheliceral triangle and the very small secondary eyes, and by a palp with a straight short embolus and an epigyne with extremely short entrance ducts. The genus is so far known only from the Hawaiian archipelago and northern Australia, extending the distribution of the clade and generating questions regarding its capacity for dispersal.

The 15 species of Mecyclothorax Sharp precinctive to New Caledonia are revised and shown by cladistic analysis to comprise a monophyletic lineage, here treated as subgenus Phacothorax Jeannel. The New Caledonian species of subgenus Phacothorax include Mecyclothoraxfleutiauxi (Jeannel), M.najtae Deuve, and 13 newly described species: M.jeanneli sp. n., M.laterobustus sp. n., M.laterorectus sp. n., M.laterosinuatus sp. n., M.laterovatulus sp. n., M.manautei sp. n., M.megalovatulus sp. n., M.octavius sp. n., M.paniensis sp. n., M.picdupinsensis sp. n., M.plurisetosus sp. n., and two jointly authored species; M.kanak Moore & Liebherr sp. n., and M.mouensis Moore & Liebherr sp. n.. subgenus Phacothorax is one of five subgenera recognized within genus Mecyclothorax based on cladistic analysis of 65 exemplar taxa utilizing information from 137 morphological characters. The four other monophyletic subgenera include the precinctive Australian Eucyclothorax subgen. n. (type species Mecyclothoraxblackburni [Sloane]), the precinctive Queensland Qecyclothorax subgen. n. (type species Mecyclothoraxstoreyi Moore), the precinctive New Zealand Meonochilus Liebherr & Marris status n., and the geographically widespread and very diverse nominate subgenus, distributed from St. Paul and Amsterdam Islands, eastward across Australia and New Guinea, and in the Sundas, Timor Leste, Lord Howe and Norfolk Islands, New Zealand, and the Society and Hawaiian Islands. The biogeographic history of Mecyclothorax can be derived from the parsimony cladogram time-calibrated by times of origin of particular geographic areas inhabited by resident representative species. Based on sister-group status of subgenus Phacothorax and subgenus Mecyclothorax, and occupation of Lord Howe Island–an island originating no earlier than 6 Ma–by the earliest divergent lineage within subgenus Mecyclothorax, the ancestor of present-day Phacothorax spp. is hypothesized to have colonized New Caledonia 6 Ma, subsequent both to Cretaceous Gondwanan vicariance as well as any Oligocene submergence. Area relationships among the New Caledonian Phacothorax point to earliest diversification incorporating the northern massifs, and most recent diversification on the ultramafic volcanic substrates in the south of Grand Terre. Flight wing loss has played an important role in shaping the various island faunas, both in their morphology as well as their diversity. The retention of flight capability in only a few of the many hundred Mecyclothorax spp. is presented in light of how populations evolve from macropterous colonizing propagules to vestigially winged specialists. Interspecific differences in genitalic structures for the sister-species pair M.fleutiauxi + M.jeanneli are shown to involve functional complementarity of male and female structures. Extensive geographic variation of male genitalia is demonstrated for several New Caledonian Mecyclothorax spp. This variation deviates from the geographically uniform male genitalia exhibited by species in the hyperdiverse Mecyclothorax radiation of Haleakalā volcano, Maui, suggesting that extensive sympatry occurring among species in that diverse species swarm selects for stability within this mate recognition system. Conversely, lower levels of sympatry characterizing the depauperate New Caledonian radiation permit the presence of more extensive male genitalic variation, this variation not selected against due to the lower likelihood of interspecific mating mistakes.

The marine ascomycete Dryosphaera tropicalis Kohlm. & Volkm.-Kohlm., sp.nov., is described from the Caribbean (Tobago), the Indian Ocean (Sri Lanka, Thailand), and the Pacific Ocean (Hawaiian Islands: Hawaii, Kauai, Maui, and Molokai). The new species occurs on intertidal and supratidal wood on sandy beaches. It is compared with the type species, Dryosphaera navigans from temperate waters, and differs mainly by ascospore dimensions and appendages. Key words: arenicolous fungi, ascomycetes, Dryosphaera, marine fungi, tropics.

Spatangoids are probably the least resolved group within echinoids, with known topological incongruencies between phylogenies derived from molecular (very scarce) and morphological data. The present work, based on the analysis of 270 specimens of Spatangidae (Echinoidea, Spatangoida) trawled in the Sardinian seas (Western Mediterranean), allowed us to verify the constancy of some characters that we consider to be diagnostic at the genus level —such as the path of the subanal fasciole and the relationship between labrum and adjacent ambulacral plates —and to distinguish two distinct forms within the studied material. Based on morphological characters, morphometrics, and molecular analyses (sequencing of two mitochondrial markers: cytochrome c oxidase subunit1 (COI) and 16S), most of the individuals were classified as morphotype A and attributed to the species Spatangus purpureus, the most common spatangoid in the Mediterranean Sea, while a few corresponded to a different morphotype (B), genetically close to the species Spatangus raschi. Preliminary morphological analyses seemed to indicate that morphotype B specimens from Sardinia are slightly different from S. raschi and from Spatangus subinermis individuals, the second species of the family known to occur in the Mediterranean Sea. On the basis of morpho-structural observations and molecular analyses, comparing Mediterranean living forms with species from other areas (Central Eastern Atlantic, North Sea and neighboring basins, South African Sea, Philippines and Indonesian Archipelago, New Zealand, and Hawaiian Islands), the clear distinction of S. purpureus from several other species classified as Spatangus was confirmed. Based on the morphological and genetic differences, we propose to maintain the genus Spatangus including in it only the type species S. purpureus among the living species and to establish the new genus Propespatagus nov. gen. to include several other species previously classified as Spatangus. The clear distinction among different genera was also detected in fossil forms of Spatangus, Propespatagus nov. gen., and Sardospatangus (†) from the European Oligo-Miocene sedimentary rocks of Germany; the Miocene of Ukraine, Italy, and North Africa; the Plio-Pleistocene of Italy; and the Mio-Pliocene of Florida (USA). The new data can help in addressing taxonomic ambiguities within echinoids, as well as in improving species identification, and hence biodiversity assessments in the Mediterranean region.

AbstractThe genus Dudresnaya is reviewed and a key to its 12 species is given. The European type species, D. verticillata (Withering) Le Jolis, and the three species known from Australia (D. australis Setchell, D. hawaiiensis R.K.S. Lee, D. capricornica sp. nov.) are described in detail. Dudresnaya australis and D. verticillata are similar in habit but distinguishable by their internal structure. Dudresnaya hawaiiensis, previously recorded only from the Hawaiian Islands, is now known from several eastern Australian localities and is the most morphologically variable and geographically widespread of all the Dudresnaya species. The newly described species, D. capricornica, is also extremely variable in habit and occurs in both eastern and western Australia.