Systematic revision of the trans-Bassian moriomorphine genus Theprisa Moore (Coleoptera, Carabidae).

The Australian genus Theprisa Moore, 1963, is taxonomically revised to comprise five species, two newly described: Theprisadarlingtoni Liebherr & Porch, sp. nov. of Tasmania, and Theprisaotway Liebherr, Porch & Maddison, sp. nov. from the Otway Ranges, Victoria. Two previously described species, T.australis (Castelnau) and T.montana (Castelnau), are distributed in the mountains of Victoria. The third previously described species, T.convexa (Sloane) is found in Tasmania. A lectotype is designated for T.convexa because the various syntypes are ambiguously labelled. Cladistic analysis based on morphological characters establishes monophyly of Theprisa relative to the Australian genera Sitaphe Moore and Spherita Liebherr. This and a second clade of Australian genera (Pterogmus Sloane, Thayerella Baehr, and Neonomius Moore) do not form a natural group, but are cladistically interdigitated among two monophyletic New Zealand lineages (Tarastethus Sharp, and Trichopsida Larochelle and Larivière) suggesting substantial trans-Tasman diversification among these groups. Hypothesized relationships within Theprisa are consistent with two bouts of speciation involving the Bass Strait; an initial event establishing T.convexa as adelphotaxon to the other four species, and a more recent event establishing the sister species T.darlingtoni and T.montana. Geographic restriction of T.otway to the Otway Ranges is paralleled by Otway endemics in several other carabid beetle genera, as well as by endemics in numerous other terrestrial arthropod taxa. Whereas these numerous Otway endemics support the distinctive nature of the Otway Range fauna, their biogeographic relationships are extremely varied, illustrating that the Otways have accrued their distinctive biodiversity via various means.

Hawaiian Paratachys Casey (Coleoptera, Carabidae): small beetles of sodden summits, stony streams, and stygian voids.

Five Hawaiian species of Paratachys Casey are revised, including four newly described: Paratachys terryli from Kauai; P. perkinsi from Moloka'i; P. haleakalae from Maui; and P. aaa from Hawai'i Island. A lectotype is designated for the fifth Hawaiian species currently combined with Paratachys, Tachys arcanicola Blackburn, 1878 of Oahu. Hawaiian Paratachys spp. known from more than one specimen exhibit some degree of ocular polymorphism, that variation being extreme in P. terryli where individuals range in ocular development from macrophthalmic with broadly convex eyes to microphthalmic with small, flat eyes. All Hawaiian Paratachys species comprise individuals with vestigial wings, with the exception of P. terryli, where a single macropterous, macrophthalmic female complements the other 18 brachypterous specimens. Based on a transformation series of characters from the male aedeagus, the biogeographic history of Hawaiian Paratachys is consistent with progressive colonization of the Hawaiian Island chain. Three of the species do not appear to represent species of conservation concern, with P. terryli and P. haleakalae known from terrestrial deep soil, litter, and streamside microhabitats in montane wet rain forest, and the troglobitic P. aaa occupying the dark zone of numerous, recently developed lava tube caves within the Mauna Loa and Kilauea volcanic massifs. The conservation status of the other two species is much more dire, with P. arcanicola of O'ahu not seen in nature since the early 20th Century, and P. perkinsi known only from a single specimen fortuitously collected in 1894 near sea level on Moloka'i.

Sporades jaechi sp. nov. with comments on classification of the New Caledonian genus Sporades Fauvel (Coleoptera, Carabidae, Trechini, Trechodina).

Sporades jaechi sp. nov. from Poum, New Caledonia is newly described and shown to be a member of the monophyletic Sporades subgenus Perileptosporades Deuve, 2010. Sporades millei Giachino and S. schuhi Donabauer are newly recognized as members of the subgenus Perileptosporades, and a key to its species is provided. Although Perileptosporades can be defined monophyletically relative to the rest of the genus, genitalic variation among species assigned to the nominate subgenus Sporades Fauvel, 1882 leave monophyly of that taxon ambiguously supported. Several morphological characters of long- standing use have been proposed to define the mutual monophyly of Sporades s. l. and its putative adelphotaxon, Trechodes Blackburn, 1901. Increasing knowledge concerning the diversity of male genitalic characters among Sporades spp. lends support to a recently proposed molecular phylogenetic hypothesis positing that Sporades evolved from within Trechodes. The consequences of the alternate phylogenetic hypotheses on their attendant nomenclature are discussed. An additional locality record for S. sexpunctatus Fauvel expands the known distribution of this species to include most of Grande Terre, New Caledonia.

Phylogenetic placement of two species known only from resting spores: Zoophthora independentia sp. nov. and Z. porteri comb nov. (Entomophthorales: Entomophthoraceae).

Molecular methods were used to determine the generic placement of two species of Entomophthorales known only from resting spores. Historically, these species would belong in the form-genus Tarichium, but this classification provides no information about phylogenetic relationships. Using DNA from resting spores, Zoophthora independentia, infecting Tipula (Lunatipula) submaculata in New York State, is now described as a new species and Tarichium porteri, described in 1942, which infects Tipula (Triplicitipula) colei in Tennessee, is transferred to the genus Zoophthora. We have shown that use of molecular methods can assist with determination of the phylogenetic relations of specimens within the form-genus Tarichium for an already described species and a new species for which only resting spores are available.

The Mecyclothorax beetles (Coleoptera, Carabidae, Moriomorphini) of Haleakala-, Maui: Keystone of a hyperdiverse Hawaiian radiation.

The Mecyclothorax carabid beetle fauna of Haleakala volcano, Maui Island, Hawai'i is taxonomically revised, with 116 species precinctive to Haleakala recognized, 74 newly described. Species are classified into 14 species groups, with the newly described species arrayed as follows: 1, Mecyclothorax constrictus group with Mecyclothorax perseveratus sp. n.; 2, Mecyclothorax obscuricornis group with Mecyclothorax notobscuricornis sp. n., Mecyclothorax mordax sp. n., Mecyclothorax mordicus sp. n., Mecyclothorax manducus sp. n., Mecyclothorax ambulatus sp. n., Mecyclothorax montanus sp. n., Mecyclothorax waikamoi sp. n., Mecyclothorax poouli sp. n., and Mecyclothorax ahulili sp. n.; 3, Mecyclothorax robustus group with Mecyclothorax affinis sp. n., Mecyclothorax anchisteus sp. n., Mecyclothorax consanguineus sp. n., Mecyclothorax antaeus sp. n., Mecyclothorax cymindulus sp. n., and Mecyclothorax haydeni sp. n.; 4, Mecyclothorax interruptus group with Mecyclothorax bradycelloides sp. n., Mecyclothorax anthracinus sp. n., Mecyclothorax arthuri sp. n., Mecyclothorax medeirosi sp. n., Mecyclothorax inconscriptus sp. n., and Mecyclothorax foveolatus sp. n.; 5, Mecyclothorax sobrinus group with Mecyclothorax foveopunctatus sp. n.; 6, Mecyclothorax ovipennis group with Mecyclothorax subtilis Britton & Liebherr, sp. n., Mecyclothorax patulus sp. n., Mecyclothorax patagiatus sp. n., Mecyclothorax strigosus sp. n., Mecyclothorax takumiae sp. n., Mecyclothorax parapicalis sp. n., Mecyclothorax mauiae sp. n., Mecyclothorax subternus sp. n., Mecyclothorax flaviventris sp. n., Mecyclothorax cordaticollaris sp. n., and Mecyclothorax krushelnyckyi sp. n.; 7, Mecyclothorax argutor group with Mecyclothorax ommatoplax sp. n., Mecyclothorax semistriatus sp. n., Mecyclothorax refulgens sp. n., Mecyclothorax argutulus sp. n., Mecyclothorax planipennis sp. n., Mecyclothorax planatus sp. n., and Mecyclothorax argutuloides sp. n.; 8, Mecyclothorax microps group with Mecyclothorax major sp. n., Mecyclothorax xestos sp. n., Mecyclothorax orbiculus sp. n., and Mecyclothorax contractus sp. n.; 9, Mecyclothorax scaritoides group with Mecyclothorax scarites sp. n., Mecyclothorax timberlakei sp. n., Mecyclothorax crassuloides sp. n., Mecyclothorax crassulus sp. n., Mecyclothorax gracilicollis sp. n., and Mecyclothorax dispar sp. n.; 10, Mecyclothorax haleakalae group with Mecyclothorax reiteratus sp. n., Mecyclothorax splendidus sp. n., Mecyclothorax bacrionis sp. n., and Mecyclothorax simpulum sp. n.; 11, Mecyclothorax vitreus group with Mecyclothorax kipwilli sp. n., Mecyclothorax kipahulu sp. n., Mecyclothorax kaumakani sp. n., and Mecyclothorax kuiki sp. n.; 12, Mecyclothorax montivagus group with Mecyclothorax rex sp. n.; 13, Mecyclothorax ducalis group with Mecyclothorax aquilus sp. n., Mecyclothorax invisitatus sp. n., Mecyclothorax longidux sp. n., and Mecyclothorax brevidux sp. n.; and 14, Mecyclothorax palustris group with Mecyclothorax hephaestoides sp. n., Mecyclothorax oculellus sp. n., Mecyclothorax bicoloris sp. n., Mecyclothorax bicoloratus sp. n., Mecyclothorax bilobatus sp. n., Mecyclothorax palustroides sp. n., Mecyclothorax filipoides sp. n., Mecyclothorax nanunctus sp. n., Mecyclothorax tauberorum sp. n., and Mecyclothorax pau sp. n. Mecyclothorax integer Sharp, stat. n. is recognized as a species distinct from Mecyclothorax interruptus Sharp. Because type series for species described by Blackburn, Karsch, and Sharp are most often divided among geographically remote collections, lectotypes are designated to stabilize the nomenclature. The radiation includes numerous cryptic sibling species best diagnosed using male genitalia, and photographs are used to represent the male genitalic variability observed among numerous dissected individuals. The large number of new species is based on substantial new collections made from all quarters of the mountain. The dense geographic sampling allows fine-scale discrimination of species boundaries, elucidating the geographic disjunctions that are associated with speciation within this hyperdiverse radiation. Disjunctions between closely related species precinctive to various areas of the mountain are not congruent across the different lineages of the radiation, indicating differential responses by the various lineages to past geological and geographical events. Of the 62 1' latitude × 1' longitude grid cells on Haleakala that are occupied by Mecyclothorax beetles, 22 house 10 or more species, and 9 house 20 or more species. This substantial level of sympatry, associated with occupation of diverse microhabitats by these beetles, provides ample information useful for monitoring biodiversity of the natural areas of Haleakala.

The Mecyclothorax beetles (Coleoptera, Carabidae, Moriomorphini) of Tahiti, Society Islands.

The 101 species of Mecyclothorax Sharp known to inhabit Tahiti Island, French Polynesia are taxonomically revised, including 28 species that are newly described: Mecyclothorax claridgeiae sp. n., Mecyclothorax jeanyvesi sp. n., Mecyclothorax poria sp. n., Mecyclothorax aano sp. n., Mecyclothorax papau sp. n., Mecyclothorax manina sp. n., Mecyclothorax everardi sp. n., Mecyclothorax ramagei sp. n., Mecyclothorax pitohitiensis sp. n., Mecyclothorax curtisi sp. n., Mecyclothorax hoeahiti sp. n., Mecyclothorax ninamu sp. n., Mecyclothorax kokone sp. n., Mecyclothorax paahonu sp. n., Mecyclothorax kayballae sp. n., Mecyclothorax ehu sp. n., Mecyclothorax papuhiti sp. n., Mecyclothorax tuea sp. n., Mecyclothorax taatitore sp. n., Mecyclothorax konemata sp. n., Mecyclothorax arboricola sp. n., Mecyclothorax rahimata sp. n., M. oaoa sp. n., Mecyclothorax maninapopoti sp. n., Mecyclothorax hunapopoti sp. n., Mecyclothorax fefemata sp. n., Mecyclothorax maninamata sp. n., and Mecyclothorax niho sp. n. Mecyclothorax muriauxioides Perrault, 1984 is newly synonymized with Mecyclothorax muriauxi Perrault, 1978. Lectotypes are designated for: Thriscothorax altiusculus Britton, 1938; Thriscothorax bryobius Britton, 1938; Mecyclothorax globosus Britton, 1948: and Mecyclothorax sabulicola Britton, 1948. Dichotomous identification keys augmented by dorsal habitus and male aedeagal photographs are provided to the various species-groups and all included species. The spermatophore of Mecyclothorax papau sp. n. is described, with the ampulla and collar found to correspond dimensionally to the length of the internal sac flagellar plate. Variation among characters of the female reproductive tract is presented for all newly described plus other representative species comprising the radiation. Taxa are assigned to species groups, modified from the classification of G.G. Perrault, based on derived character states polarized using the Australian outgroup taxon Mecyclothorax punctipennis (MacLeay). Much of the species-level diversity on this small Pacific island is partitioned allopatrically over very small distributional ranges. No species is shared between Tahiti Nui and Tahiti Iti, and nearly all species in Tahiti Nui are geographically restricted to one ridgelike massif of that volcano. Cladistically similar species are often distributed on different massifs suggesting that vicariance associated with erosional valley formation has facilitated speciation, however several instances in which sister species occupy sympatric distributions on the same ridge system demonstrate that speciation may also occur across extremely localized landscapes. Such localized differentiation is facilitated by the low vagility of these small-bodied, flightless predators whose fragmented populations can persist and diverge within spatially limited habitat patches. The intense philopatry of Tahitian Mecyclothorax spp. coupled with the highly dissected landscape has produced the geographically densest adaptive radiation on Earth. This radiation has occurred very rapidly, with species durations averaging 300,000 yr; a speciation rate similar to that observed in Hawaiian Oliarus planthoppers and Laupala crickets, and East African Rift lake cichlid fishes.

New Mecyclothorax spp. (Coleoptera, Carabidae, Moriomorphini) define Mont Mauru, eastern Tahiti Nui, as a distinct area of endemism.

Seven species of Mecyclothorax Sharp precinctive to Mont Mauru, Tahiti, Society Islands are newly described: Mecyclothorax tuteisp. n., Mecyclothorax tihotiisp. n., Mecyclothorax putaputasp. n., Mecyclothorax toretoresp. n., Mecyclothorax anaanasp. n., Mecyclothorax pirihaosp. n., and Mecyclothorax porosp. n. These seven constitute the first representative Mecyclothorax species recorded from Mauru, and their geographic restriction to this isolated massif defines it as a distinct area of endemism along the highly dissected eastern versant of the Tahiti Nui volcano. Each of the new species has a closest relative on another massif of Tahiti Nui, supporting speciation associated with vicariance caused by extensive erosional valley formation, especially the development of Papenoo Valley. Comparison of the known elevational distributions of the new discoveries on Mont Mauru to the elevational diversity profile of the comparatively well-sampled Mont Marau, northwest Tahiti Nui, suggests that numerous Mecyclothorax species remain to be discovered in higher-elevation habitats of Mont Mauru.

The first precinctive Carabidae from Moorea, Society Islands: new Mecyclothorax spp. (Coleoptera) from the summit of Mont Tohiea.

Seven species of Mecyclothorax Sharp from Moorea, Society Islands are newly described; Mecyclothorax perraultisp. n., Mecyclothorax paheresp. n., Mecyclothorax menemenesp. n., Mecyclothorax mahatahisp. n., Mecyclothorax popotioaoasp. n., Mecyclothorax maposp. n., and Mecyclothorax fatatasp. n. These constitute the first Mecyclothorax species described from Moorea, and the first carabid beetle species shown to be geographically restricted to that island. Each of the newly described species is most similar to a different species on the island of Tahiti, suggesting that none of the seven Moorean taxa are evolutionary end-products of autochthonous speciation within Moorea. The occurrence of precinctive Mecyclothorax species on both Moorea and Tahiti demonstrates that radiation of Mecyclothorax in the Society Islands has been facilitated by speciation events implicating both islands. Whether this speciation has been preceded by vicariance or dispersal is discussed, with the generality of a dispersal hypothesis tested using information from Society Island Nabidae (Hemiptera). Salient morphological characters for taxa in the Society and Hawaiian Islands are compared to those representing a broad survey of southwest Pacific Mecyclothorax spp. This comparison supports the independent founding of each radiation in the Societies and Hawaii from an Australian ancestral propagule, likely drawn from the ecologically general, geographically widespread Mecyclothorax punctipennis (Macleay).

Cladistic assessment of subtribal affinities within the tribe Moriomorphini with description of Rossjoycea glacialis, gen. n. and sp. n. from the South Island, and revision of Meonochilus Liebherr and Marris from the North Island, New Zealand (Coleoptera, Carabidae).

Phylogenetic relationships within the tribe Moriomorphini Sloane, 1890 are analyzed cladistically based on 75 morphological characters and 21 ingroup terminal taxa rooted at a Trechus obtusus Erichson outgroup. Based on the resultant cladistic relationships, two subtribes-Moriomorphina and Amblytelina Blackburn, 1892-are recognized, with the following new synonymies proposed: Meonides Sloane, 1898 = Amblytelina (NEW SYNONYMY); Tropopterides Sloane, 1898 = Amblytelina (NEW SYNONYMY); Mecyclothoracitae Jeannel, 1940 = Amblytelina (NEW SYNONYMY). Monophyly of Moriomorphina is based on presence of elongate, parallel-sided and glabrous to nearly glabrous male parameres, whereas Amblytelina are defined most broadly by possession of conchoid parameres with narrowed, setose apices, subtending a clade defined by a more derived parameral configuration whereby elongate styloid parameres terminate in a whip-like apical extension. Representatives of all New Zealand moriomorphine genera are included in the analysis, with cladistic results necessitating description of Rossjoycea glacialis, gen. n. and sp. n., known from a single locality near the Franz Josef Glacier, Westland, South Island, New Zealand. Monophyly of Meonochilus Liebherr and Marris, 2009 is demonstrated, and its six species are taxonomically revised: Meonochilus amplipennis (Broun), Meonochilus eplicatus (Broun), Meonochilus placens (Broun), Meonochilus bellorum, sp. n., Meonochilus rectus, sp. n., and Meonochilus spiculatus, sp. n. Geographic restriction of Meonochilus to the North Island of New Zealand, coupled with its sister-group status to an Australian-based Amblytelus Erichson-Mecyclothorax Sharp clade reinforce the interpretation that Meonochilus was isolated in New Zealand by vicariance along the Norfolk Ridge, subsequent to New Zealand's initial Cretaceous isolation from Tasmania and southeastern Australia via opening of the Tasman Sea.

Absence asymmetry: the evolution of monorchid beetles (Insecta: Coleoptera: Carabidae).

Asymmetrical monorchy, or the complete absence of one testis coupled with the presence of its bilateral counterpart, is reported for 174 species of the carabid beetle tribes Abacetini, Harpalini, and Platynini (Insecta: Coleoptera: Carabidae) based on a survey of over 820 species from throughout the family. This condition was not found in examined individuals of any other carabid beetle tribes, or of other adephagan beetle families. One monorchid taxon within Platynini exhibits symmetrical vasa deferentia at the beginning of the pupal stadium, suggesting that developmental arrest of the underdeveloped vas deferens takes place in pupation. The point at which development of the testis is interrupted is unknown. Complete absence of one organ of a bilateral pair--absence asymmetry--is rarely found in any animal clade and among insects is otherwise only known for testes in the minute-sized beetles of the family Ptiliidae, ovaries in Scarabaeinae dung beetles, and ovaries of some aphids. Based on current phylogenetic hypotheses for Carabidae, testis loss has occurred independently at least three times, and up to five origins are possible, given the variation within Abacetini. Clear phylogenetic evidence for multiple independent origins suggests an adaptive or functional cause for this asymmetry. A previously posited taxon-specific hypothesis wherein herbivory in the tribe Harpalini led to testis loss is rejected. Optimal visceral packing of the beetle abdomen is suggested as a general explanation. Specifically, based on the function of various organ systems, we hypothesize that interaction of internal organs and pressure to optimize organ size and space usage in each system led to the multiple origins and maintenance of the monorchid condition. Testes are the only redundant and symmetrically paired structures not thought to be developmentally linked to other symmetrical structures in the abdomen. Among all possible organs, they are the most likely--although the observed frequency is very small--to bypass constraints that maintain bilateral symmetry, resulting in absence asymmetry. However, based solely on our observations of gross morphology of internal organs, no function conclusively explains the ontogenetic loss of one testis in these taxa. Unlike the analogous absence asymmetry of organs in other animal groups, no dramatic body-form constraint--e.g., snakes and lung loss, ptiliid beetles' small body-size and relatively giant sperm--or adaptive scenario of improved locomotory performance--e.g., birds and ovary loss due to flight constraints-applies to these carabid beetles. We tentatively suggest that testis loss is driven wholly by an interaction among the internal organs of these beetles, possibly due to selective pressure to maximize the comparatively large accessory glands found in these taxa. However, as the ordering of these evolutionary events of testis loss and accessory gland size increase is not known, large accessory glands might have secondarily evolved to compensate for a decreased testicular output.

A CLADISTIC TEST OF THE TAXON CYCLE AND TAXON PULSE HYPOTHESES.

Abstract- A species' habitat preference is intcrpretablc both as a response to present-day conditions and as a result of evolutionary response to historical conditions. The taxon cycle and taxon pulse have been proposed as hypotheses that allow prediction of patterns of habitat specialization within a lineage. Both are based on common assumptions: (1) habitat specialization is largely irreversible in a lineage, (2) ecological specializations arise in a center of origin, and (3) dispersal events leading to current distributions can be ascertained. Eight taxa of Antillean, Mexican, and Central American Carabidae, for which cladistic hypotheses of relationship have been proposed, are used to test the generality of the taxon cycle and pulse. The patterns of habitat utilization predicted by the taxon cycle and taxon pulse hypotheses are tested by comparing cladistic transformations of habitat preference to randomly generated patterns of data generated under a null hypothesis. Evolutionary changes in habitat are interpreted using Camin-Sokal coding, which assumes irreversible habitat shifts and a predetermined ecological ground state (assumptions of the taxon cycle and taxon pulse). An observed pattern is considered to demonstrate the taxon cycle or pulse when it results in an explanation of the habitat shifts that is more parsimonious than 95 % of the randomly generated patterns. Of the eight carabid groups, only one exhibits a statistically significant pattern supporting the taxon cycle and pulse. The failure of the taxon cycle and pulse as generally predictive hypotheses may be due to historical changes in climate that permit episodes of range expansion for species previously restricted to small ranges, and habitat shifts and specialization that do not progress in a linear transformation series. Habitat shifts are also analyzed using Farris optimization, resulting in the most parsimonious transformation series of habitats, subject to a predefined ordering of habitats, while allowing reversals. Significance of an observed pattern of habitat shifts under Farris optimization implies habitat constancy relative to cladogenesis, and step-wise changes in habitat preference. Two of the eight groups exhibit significant patterns of habitat utilization under Farris optimization, indicating that vicariance of areas of like habitat has been the predominant factor generating species diversity in these groups. The other groups do not exhibit habitat constancy, suggesting rapid changes in habitat preference relative to speciation.

GENE FLOW IN GROUND BEETLES (COLEOPTERA: CARABIDAE) OF DIFFERING HABITAT PREFERENCE AND FLIGHT-WING DEVELOPMENT.

Estimates of gene flow vary 100-fold among five carabid species, ranging from the winged lowland subtropical Agonum elongatulum to the flightless montane temperate Platynus angustatus. Results based on Wright's (1943) FST method, and Slatkin's (1981) graphical and (1985a) private-allele methods are concordant. Genetic heterogeneity, measured by Wright's FST , is not correlated with degree of flight-wing development; one fully winged species exhibits heterogeneity of the same order as a vestigially winged species. Genetic heterogeneity is positively correlated with the average elevation of collection sites for these species. Lower levels of gene flow associated with greater genetic subdivision may occur in upland areas because of habitat fragmentation (due to topographic diversity) and habitat persistence (leading to a lower extinction rate for populations). In at least one species, the distribution of stable infraspecific polymorphisms indicates that the high estimate of present-day gene flow is likely to be due to historical gene flow and not to present-day conditions.