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Platynini (Coleoptera: Carabidae) of Vanuatu: Miocene diversification on the Melanesian Arc
Abstract
Vanuatu supports 11 resident species of the carabid beetle tribe Platynini: five indigenous species shared with other Pacific islands and Australia and six newly described precinctive species. Notagonum delaruei, sp. nov. represents a single descendant species of one colonisation event. Helluocolpodes, gen. nov. (type species Colpodes helluo Darlington of New Guinea) is proposed to accommodate a monophylum comprising the type species plus Helluocolpodes discicollis, sp. nov., H. mucronis, sp. nov., H. multipunctatus, sp. nov., H. sinister, sp. nov. and H. vanemdeni, sp. nov., all from Vanuatu. Generic assignments are informed by cladistic analysis of anatomical characters for a variety of Pacific platynine taxa. Metacolpodes Jeannel is redefined cladistically to include seven Pacific and Asian species. Biogeographic relationships among island areas housing platynine taxa on the Australian and Pacific Plates are investigated using a chrono-area cladogram, i.e. a taxon-area cladogram for which terminals are dated based on geological evidence and internal nodes based on non-reversible temporal optimisation. Conclusions reached by constraining the ages of areas within the context of phylogenetic relationships of their resident taxa include: (1) Vanuatu has supported resident platynine taxa since the Middle to Late Miocene; (2) the Hawaiian Blackburnia first colonised that archipelago in the Miocene, long before the present oldest high island, Kauai, came into existence; (3) the New Zealand Ctenognathus most likely arose from Miocene colonisation of New Zealand via Fiji; and (4) the low diversity of the Tahitian platynine fauna is due to relatively recent, Pliocene or later, colonisation of the Society Island chain by this group, also from a Fijian source.
... Vanuatubasis Ober and Staniczek, 2009 is composed of ten species (Ober and Staniczek, 2009;Saxton et al. 2022) and displays high levels of endemism . Endemism is a pattern that appears across both Vanuatu and Fiji (Liebherr, 2005;Beatty et al. 2007). Members of this genus appear to inhabit a very narrow ecological niche and is almost exclusively found on alkaline streams (Saxton et al. 2021). ...
... The few studies that have previously provided faunal ages for clades from Vanuatu are largely in disagreement. Liebherr (2005) suggested a Miocene age for ground beetles found in Vanuatu using the geologic estimates for the older islands (22-25 Ma). In contrast, using a standard substitution rate for gastropods, the age of snails in Vanuatu indicated a Pliocene origin (~5 Ma) (Zielske and Haase, 2014). ...
South Pacific islands provide an ideal study system to explore patterns of speciation, specifically examining the role of dispersal versus vicariance. Dispersal is often the suggested mechanism of diversification in the South Pacific, specifically among remote island chains. Here, we provide a phylogeny of several related genera of Coenagrionidae (Odonata: Zygoptera) from the South Pacific, based on five molecular loci, in order to examine patterns of speciation in the region. We used the endemic damselfly genera Nesobasis, Nikoulabasis, and Vanuatubasis found across both Fiji and Vanuatu. Knowledge of the geologic history of the region was used to inform our understanding of the evolution of these genera. Both archipelagos used to be part of the Vitiaz arc which spanned from the Solomon Islands to Tonga and began to break apart 10-12 Ma. Results of our divergence-time estimations and biogeographic reconstructions support that the breakup of this arc acted as a significant vicariance event in the evolution of these taxa. Specifically, it led to the extant generic diversity seen in these damselflies. We find that within the archipelago of Vanuatu, that Espiritu Santo served as an important source for dispersal to other islands with Malekula acting as a stepping stone to Efate.
... Morphological phylogenetic analysis indicates that the Blackburnia radiation is monophyletic (Liebherr and Zimmerman 1998), with the Australian genus Notagonum (Liebherr 2005(Liebherr , 2017 the best supported outgroup based on Notagonum and Blackburnia being mutual members of a clade characterized by monorchy; i.e., presence of only a single male testis . Via morphological and multilocus phylogenetic analysis, Cryan et al. (2001) corroborated monophyly of major lineages within Blackburnia and resolved many species triplets occupying Molokaʻi, West Maui, and Haleakalā, the former windward faces of Maui Nui (Liebherr 1997). ...
... Via morphological and multilocus phylogenetic analysis, Cryan et al. (2001) corroborated monophyly of major lineages within Blackburnia and resolved many species triplets occupying Molokaʻi, West Maui, and Haleakalā, the former windward faces of Maui Nui (Liebherr 1997). Based on either outgroup, Blackburnia colonized Hawaiʻi from Australia via aerial dispersal, with arrival time hypothesized to be 28-30 Ma based on phylogenetic relationships among Pacific platynine taxa constrained by ages of their occupied islands (Liebherr and Zimmerman 1998;Liebherr 2005Liebherr , 2017. ...
... The former comprises species of the Hawaiian endemic platynine genus Blackburnia Sharp (Liebherr and Zimmerman 2000;Liebherr 2001Liebherr , 2003Liebherr , 2007Liebherr and Short 2007) versus the Tahitian platynine fauna comprising only three species (Perrault 1977). Hawaiian colonisation by Blackburnia is hypothesised to have occurred 28-30 Mya, long before the origin of Kauai at 5.1 Mya, based on the lineage's phylogenetic relationships to other Australian and Pacific taxa (Liebherr 2005). The most generalised Hawaiian Blackburnia are characterised by fully-developed flight wings (Liebherr and Zimmerman 2000). ...
... The most generalised Hawaiian Blackburnia are characterised by fully-developed flight wings (Liebherr and Zimmerman 2000). Conversely, the three platynine species in the Society Islands are arranged phylogenetically into two groups: (1) the 'Colpodes' grade sister species pair 'C.' anachoreta Fairmaire and 'C.' eremita Fairmaire, which both exhibit variously foreshortened though fully venated flight wings; and (2) the fully winged Metacolpodes monticola (Fairmaire) (Liebherr 2005). Tahitian colonisation by these two platynine stocks was hypothesised to have occurred no more than 4.5 Mya, with both sets of species derived from a geographically widespread array of fully winged species. ...
Bembidion (Sloanephila) tahitiense, sp. nov. is described from Mont Mauru, an isolated massif of Tahiti Nui volcano. Based on evidence from seven genes (four nuclear protein-coding, one mitochondrial protein-coding, two nuclear ribosomal), its sister group is the Australian B. jacksoniense Guerin-Meneville, with which it shares a synapomorphic spur on the ostium of the male genitalia. In contrast to B. jacksoniense, B. tahitiense is brachypterous, with rounded humeri, constricted posterior pronotal margins and convex body form. Examination of the seven genes in two species of the Hawaiian subgenus Nesocidium Sharp reveals that the sister group of Nesocidium is subgenus Zecillenus Lindroth from New Zealand. These two subgenera belong to the Ananotaphus complex, a clade inhabiting Australia, New Zealand and Hawaii. The relationships of the second Hawaiian subgenus, Gnatholymnaeum Sharp, are less clear, although Gnatholymnaeum belongs to the Bembidion series (along with Sloanephila and the Ananotaphus complex). Bembidion beetles colonised the Society and Hawaiian islands independently from source areas in the south-west Pacific. Based on parsimonious reconstructions of flight-wing configuration, the Tahitian and Hawaiian colonisations involved winged individuals. Colonisation of the Society and Hawaiian islands by carabid beetles of two other tribes - Platynini and Moriomorphini - follow the dispersal patterns hypothesised for Bembidion.
... Subsequently, two more Blackburnia species were described (Liebherr 2001(Liebherr , 2003 with inclusion of those species in the cladistic analysis enhancing resolution of the phylogenetic hypothesis while corroborating the overall conclusion that the earliest diverging extant Blackburnia species reside on Kauai, with newer islands progressively colonized during evolutionary diversification of the group. Complementary cladistic analysis including taxa from the Australian and Asian biogeographic regions (Liebherr 2005) affirms that the Blackburnia radiation is old amongst those in the Pacific-older than the radiations in Fiji, Vanuatu, Samoa, and Tahiti-suggesting that Blackburnia colonized the Hawaiian chain during the Miocene, relatively soon after the period of continuous Hawaiian island presence began with the emergence of Kure Island, 29-30 Ma (Carson and Clague 1995). ...
... Viewed phylogenetically, the Kauai riparian fauna is derived from three separate stocks. Blackburnia mandibularis is the adelphotaxon to all other Blackburnia (Liebherr and Zimmerman 2000), and so its ancestors were probably from an older Hawaiian island now submerged (Liebherr 2005). This species is characterized by large compound eyes, and is capable of winged flight. ...
Blackburnia riparia, new species is described from the summit of Mt. Waialeale, Kauai, Hawaii, incorporated into a phylogenetic analysis of Blackburnia Sharp, and thereby placed as the adelphotaxon to two other allopatric Kauai species, B. lata Liebherr and B. atra Liebherr. The new species occurs in riparian habitats, including vertical rock-faced seeps and algal mats, and is documented to feed on aquatic larvae of Micropsectra Kieffer (Diptera: Chironomidae). Blackburnia elegans (Sharp) adults also occur in streams on Waialeale summit where they were found walking under water on the undersides of large, flat rocks. Based on documented prey and observed behavior, both species appear to forage under the waterline. The abdominal and elytral anatomy of adult B. riparia and B. elegans is suitable for retention of a subelytral air bubble, suggesting these species respire underwater. Adults and associated larvae of Blackburnia mandibularis Liebherr inhabit moss-mats along falls' margins at Waipoo Falls, Kokee State Park, Kauai, indicating that this species' active life stages also reside in the riparian zone. Attributes of the larval head capsule and mandibles for B. riparia and B. mandibularis are associated with homologous characters expressed in adult anatomy, ontogenetically linking larval and adult head and mouthpart specializations. The various evolutionary origins of riparian habits are examined across Blackburnia. Via one route, occupation of the montane riparian zone has evolved from terrestrial moss-mat habitation, with concomitant evolutionary reduction of adult compound eyes. A second evolutionary route to the riparian zone is based on occupation of open, disturbed or ephemeral habitats, in some cases assisted by adult winged dispersal.
... Subsequently, two more Blackburnia species were described (Liebherr 2001(Liebherr , 2003 with inclusion of those species in the cladistic analysis enhancing resolution of the phylogenetic hypothesis while corroborating the overall conclusion that the earliest diverging extant Blackburnia species reside on Kauai, with newer islands progressively colonized during evolutionary diversification of the group. Complementary cladistic analysis including taxa from the Australian and Asian biogeographic regions (Liebherr 2005) affirms that the Blackburnia radiation is old amongst those in the Pacific-older than the radiations in Fiji, Vanuatu, Samoa, and Tahiti-suggesting that Blackburnia colonized the Hawaiian chain during the Miocene, relatively soon after the period of continuous Hawaiian island presence began with the emergence of Kure Island, 29-30 Ma (Carson and Clague 1995). ...
... Viewed phylogenetically, the Kauai riparian fauna is derived from three separate stocks. Blackburnia mandibularis is the adelphotaxon to all other Blackburnia (Liebherr and Zimmerman 2000), and so its ancestors were probably from an older Hawaiian island now submerged (Liebherr 2005). This species is characterized by large compound eyes, and is capable of winged flight. ...
Blackburnia kavanaughi, new species, previously misidentified as Blackburnia asquithi Liebherr, is described from the western portion of the Alakai Plateau, Kauai, Hawaii. The species is placed phylogenetically based on cladistic analysis including all known Hawaiian Blackburnia, supporting membership in the subgenus Metromenus Sharp. The new species is hypothesized to be most closely related to B. debilis (Perkins) of Molokai and B. kuiki Liebherr of East Maui, sharing enhanced pubescence of the antennal pedicel, reduction in number of the dorsal elytral setae, and elongate ventrolateral setae of tarsomere 5. Individuals of all three species reside in deep arboreal moss mats associated with ohia lehua trees (Metrosideros polymorpha: Myrtaceae). Blackburnia debilis, previously known only from the single male holotype collected in 1902, was recollected in 2005 from Uapa summit, Molokai, in arboreal moss on the mesic leeward edge of ohia lehua forest. Discovery of B. kavanaughi on Kauai implies an additional colonization event from Kauai to newer islands during the diversification of Blackburnia, while corroborating the previous general biogeographic pattern of progressive colonization of the Hawaiian Islands by this clade. Phylogenetic placement of B. debilis + B. kuiki as adelphotaxon to B. kavanaughi results in a hypothesized sister-group relationship between sympatric sister species—the epigean B. calathiformis (Sharp) and the troglobitic B. howarthi (Samuelson and Liebherr)—corroborating F. G. Howarth's general hypothesis for the evolution of Hawaiian troglobites.
... The Bembidion fauna is hypothesized to have been derived from two founder events (Liebherr 2008), the Blackburnia fauna from one (Liebherr & Zimmerman 1998), and Mecyclothorax most likely also from one (Britton 1948). All three carabid radiations have their roots in the southwest Pacific (Liebherr 2005b(Liebherr , 2007(Liebherr , 2008, though their respective colonists have arrived at different times. The Blackburnia fauna is extensively represented by numerous precinctive species on Kauai, with the common ancestor likely arriving on an earlier emergent island during the Miocene epoch (Liebherr 2005b). ...
... All three carabid radiations have their roots in the southwest Pacific (Liebherr 2005b(Liebherr , 2007(Liebherr , 2008, though their respective colonists have arrived at different times. The Blackburnia fauna is extensively represented by numerous precinctive species on Kauai, with the common ancestor likely arriving on an earlier emergent island during the Miocene epoch (Liebherr 2005b). Mecyclothorax, conversely, is a relatively later immigrant with the most plesiomorphic species in the radiation found on fragments of Maui Nui (Liebherr 2005a(Liebherr , 2007, and the group absent from the oldest high islands; e.g. ...
The Hawaii Island fauna of Mecyclothorax Sharp, 1903 is taxonomically revised and found to comprise 30 species, 18 newly described: M. gagnei sp. n., M. nitidus sp. n., M. maunakukini sp. n., M. punakukini sp. n., M. kaukukini sp. n., M. perivariipes sp. n., M. aa sp. n., M. giffini sp. n., M. hephaestus sp. n., M. funebris sp. n., M. granulipennis sp. n., M. rufipennis sp. n., M. blackburnianus sp. n., M. swezeyi sp. n., M. sinuosus sp. n., M. williamsi sp. n., M. purpuripennis sp. n., and M. footei sp. n. New synonymies include: Mecyclothorax parvus Britton, 1948 = M. subunctus (Perkins), 1917; Thriscothorax munroi Perkins, 1937 = M. karschi (Blackburn), 1882; Thriscothorax gracilis Sharp, 1903 and Mecyclothorax proximus Britton, 1948 = M. konanus Sharp, 1903; Mecyclothorax terminalis Britton, 1948 = M. discedens (Sharp) 1903. Mecyclothorax vulcanus (Blackburn) was described from a mixed series, with the cryptic sibling species M. hephaestus newly described to correct the partial misidentification. Species delimitation for the highly variable M. konanus is achieved using a hierarchical analysis based on infraspecifically variable attributes. Extensive male genitalic variation is documented within M. konanus and M. deverilli (Blackburn), and also among the cryptic sibling species pair M. variipes (Blackburn) and M. perivariipes. The observed variation is consistent with various hypotheses of sexual selection, but not with the genitalic lock and key hypothesis. Areas of endemism are tentatively proposed based on the most restricted distributions of Hawaii Island Mecyclothorax, with various flanks of Mauna Kea and Mauna Loa volcanoes identified as distinctive areas. (© 2008 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)
... ,[80][81][82][83] . The only leaf insect from the Solomon Islands † Eophyllium messelenseFig. ...
The insect order Phasmatodea is known for large slender insects masquerading as twigs or bark. In contrast to these so-called stick insects, the subordinated clade of leaf insects (Phylliidae) are dorso-ventrally flattened and therefore resemble leaves in a unique way. Here we show that the origin of extant leaf insects lies in the Australasian/Pacific region with subsequent dispersal westwards to mainland Asia and colonisation of most Southeast Asian landmasses. We further hypothesise that the clade originated in the Early Eocene after the emergence of angiosperm-dominated rainforests. The genus Phyllium to which most of thẽ 100 described species pertain is recovered as paraphyletic and its three non-nominate subgenera are recovered as distinct, monophyletic groups and are consequently elevated to genus rank. This first phylogeny covering all major phylliid groups provides the basis for future studies on their taxonomy and a framework to unveil more of their cryptic and underestimated diversity.
... The Vitiaz arc has likely contributed to biotic migration processes across the whole Southwest Pacific (Oliver et al., 2018) and has already been considered to explain the colonisation of Fiji (e.g. Duffels & Turner, 2002;Liebherr, 2005). The geological dynamics of the fractured landmasses might have triggered diversification of the obrimine Wallacean lineage and its distribution across the entire arc (Oliver et al., 2018), but subsequently led to the extinction of most species. ...
Stick and leaf insects (Phasmatodea) are large terrestrial herbivorous arthropods known for masquerading as plant parts such as bark, twigs and leaves. Their evolutionary history is largely shaped by convergent evolution associated with adaptive radiations on geographically isolated landmasses that have repeatedly generated ground-dwelling ecomorphs. The members of one lineage, however, the Oriental Heteropterygidae, are morphologically rather uniform, and have a predominantly ground-dwelling lifestyle. The phylogeny of Heteropterygidae that comprises approximately 130 described species is controversial and remains uncertain. In particular, the systematic position of the giant Jungle Nymph Heteropteryx dilatata, whose males are capable of flight and exhibit the most plesiomorphic wing morphology among extant phasmatodeans, is of major interest to the scientific community. Here, we analysed a set of seven nuclear and mitochondrial genes to infer the phylogeny of Heteropterygidae covering the group's overall diversity. The divergence time estimation and reconstruction of the historical biogeography resulted in an ancestral distribution across Sunda-land with long distance dispersal events to Wallacea, the Philippines and the South Pacific. We were able to resolve the relationships among the three principal subgroups of Heteropterygidae and revealed the Dataminae, which contain entirely wingless small forms, as the sister group of Heteropteryginae + Obriminae. Within Heteropteryginae, Haaniella is recovered as paraphyletic in regard to Heteropteryx. Consequently, Heteropteryx must be considered a subordinate taxon deeply embedded within a flightless clade of stick insects. Within Obriminae, the Bornean Hoploclonia is strongly supported as the earliest diverging lineage. Based on this finding, we recognize only two tribes of equal rank among Obriminae, the Hoplocloniini trib. nov. and Obrimini sensu nov. Within the latter, we demonstrate that previous tribal assignments do not reflect phylogenetic relationships and that a basal splitting event occurred between the wing-bearing clade Miroceramia + Pterobrimus and the remaining wingless Obrimini. The Philippine genus Tisamenus is paraphyletic with regard to Ilocano hebardi, thus, we transfer the latter species to Tisamenus as Tisamenus hebardi comb. nov. and synonymize Ilocano with Tisamenus. We discuss character transformations in the light of the new phylogenetic results and conclude that the current taxonomic diversity appears to be mainly driven by allopatry and not to be the result of niche differentiation. This radiation is thus best described as a nonadaptive radiation.
... The earliest insect colonization of eastern Melanesia (Vanuatu and Fiji) was estimated at about 9-17 Ma (Balke et al., 2007;Liebherr, 2005;Lucky and Sarnat, 2010;Sarnat and Moreau, 2011), consistent with the hypothesis that colonization was facilitated by emergent land as part of the subaerial Vitiaz Arc as island chain (Hypothesis 3a). In the Prenolepis genus-group, dispersal has occurred earlier than the recent re-exposure of land in eastern Melanesia during the Pliocene (Hypothesis 3b), whereas early Miocene land has likely remained above water and harboured older colonizers to the present. ...
... Whether the genus Notagonum can be maintained in future, has be to checked by a general taxonomic and phylogenetic revision of the Oriental-Papuan Platynini, that, however, would be a major task and probably will not be undertaken in the near future. Some provisional thoughts on the phylogeny at least of the Platynini from Melanesia have been provided by Liebherr (2005). ...
The ground beetle (Carabidae) fauna of Timor is reviewed based on recent collecting by the Australian Museum, in Timor Leste. 53 species were collected, of which the following nine species are described as new: Mecyclothorax timorensis sp. nov. Baehr; M. reidi sp. nov. Baehr; Rhytisternus externus sp. nov. Baehr; Arhytinus timorensis sp. nov. Baehr; Notagonum reidi sp. nov. Baehr; N. angusticolle sp. nov. Baehr; Perigona timorensis sp. nov. Baehr; Aristolebia timorensis sp. nov. Baehr; Lebia timorensis sp. nov. Baehr The new species are compa red with their nearest relatives. The remaining material is identified to species, or at least to genus, and discussed according to the current classification of Carabidae. Several species and genera are newly recorded for Timor. In spite of the rather small number of species contained, the sample allows some insight into the relations of the Carabid fauna of Timor to the Oriental and Australopapuan Regions, with about two thirds of the species belonging to the Oriental fauna, and one third to the Australopapuan fauna. A few species also belong to subgroups that are restricted to the southeasternmost Oriental Region "Wallacea"-Sulawesi, and the Maluku, Aru and Kei Islands.
... All in all, nearly 55% of names included in the present supplement are the direct result of taxonomic changes made since 2001. Between 2001 and 2015, several taxonomic groups have been revised, many synonymies and taxonomic changes have been made, and numerous new taxa have been described (20 genera and subgenera, 111 species and subspecies), thanks to the collective efforts of the following researchers:Giachino (2003;Polyderis), Johns (2003;Holcaspis), Leschen et al. (2003;Dromius, Trigonothops), Pawson et al. (2003;Oregus), Emberson (2004;Harpalus), Baehr (2005;Anomotarus), Giachino (2005;Duvaliomimus), Johns (2005;Mecodema, Megadromus), Larochelle and Larivière (2005; Harpalini),Liebherr (2005;" Anchomenus " ), Lorenz (2005;Rhysodini), Toledano (2005;Bembidion, subgenera Zeactedium and Zecillenus), Johns (2007;Anomalobroscus, Mecodema, Megadromus, Onawea, Plocamostethus, Zeopoecilus), Larochelle and Larivière (2007; synopsis of supraspecifi c taxa),Bell and Bell (2009;Rhysodini),Liebherr and Marris (2009;Mecyclothorax, Meonochilus),Cassola and Moravec (2010;Neocicindela), Johns (2010;Migadopini), Townsend (2010; Trechini), Liebherr (2011a; Moriomorphini),Pons et al. (2011;Neocicindela),Seldon and Leschen (2011;Mecodema), Liebherr (2011b;Amblytelina, Meonochilus, Rossjoycea, Moriomorphina), Liebherr et al. (2011;Orthoglymma), Townsend (2011;Cerabilia, Loxandrini), Seldon et al. (2012; ...
A supplement to the “Carabidae (Insecta: Coleoptera): catalogue” published by Larochelle and Larivière (2001; Fauna of New Zealand 43), is provided. A total of 99 genera and 547 species and subspecies belonging to 21 tribes and 8 subfamilies of Carabidae (including Cicindelini and Rhysodini), is recorded for New Zealand. Endemism is high, with 93% of the species and subspecies as well as 62% of the genera recorded from this country only. Changes to the 2001 catalogue are documented. The synonymy and type locality of taxa described between 2001 and 2015 are also provided.
... Among the Carabidae, a monophyletic radiation classifi ed as the Hawaiian endemic genus Blackburnia Sharp occurs on all the high islands from Kauai to Hawaii Island (Liebherr & Zimmerman 2000 ;Liebherr 2001bLiebherr , 2003Liebherr , 2006aLiebherr & Short 2006 ), with all but 4 of the 133 species known from single islands. Phylogenetic relationships of Blackburnia to taxa from across the Pacifi c as well as to taxa occupying Australia, New Guinea, and New Zealand support residence of ancestral Hawaiian Blackburnia since well before the origin of Kauai, 5.1 million years ago (Liebherr 2005a ). A progressive colonization history (Funk & Wagner 1995 ) from Kauai to Oahu (3.7 million years ago) and Maui Nui (1.9 million years ago), to Hawaii Island (0.43 million years ago) (Carson & Clague 1995 ) is shared with other signature Hawaiian radiations, including those in the Drosophilidae (Hardy 1965 ;DeSalle 1995 ), Lygaeidae (Usinger 1942 ;Ashlock & Gagné 1983 ;Polhemus 1998 ), Cosmopterigidae (Zimmerman 1978 ;Rubinoff 2008 ) and Curculionidae (Samuelson 2003 ), with diversifi cation hypothesized to have benefi ted from colonization of previously unoccupied habitats, leading to repeated bouts of community organization (Gillespie & Roderick 2002 ). ...
Mecyclothorax palikea, sp.n. is described from the vicinity of Palikea, southern Waianae Range, Oahu, HI, USA and is assigned to Britton's Mecyclothorax flavomarginatus species group. Cladistic analysis, based on 20 morphological characters and including several outgroup taxa, places Mecyclothorax impunctatus Liebherr of Molokai as adelphotaxon to the other eight species of the group, with subsequent speciation events successively isolating M. sharpi Britton of West Maui versus a clade of seven Oahu species. Phylogenetic relationships among the Oahu clade species posit three historical speciation events vicariating ancestors on the western Waianae and eastern Koolau Ranges. Mecyclothorax palikea is placed as adelphotaxon to M. carteri (Perkins), a species allopatrically distributed to the north in the Waianae, corroborating existence of southern and northern areas of endemism within the Waianae Range. Relative ages of the respective volcanoes housing M. flavomarginatus group species — Waianae (3.7 million years ago), Koolau (2.6 million years ago), Eastern Molokai (1.8 million years ago), West Maui (1.3 million years ago)—imply that ancestral occupation of Oahu by this group occurred subsequent to completion of the shield building phases of Oahu's two volcanoes, Waianae and Koolau. Diversification within the group on Oahu was associated with vicariance events that occurred within a terrestrial environment. Whereas all four species of the M. flavomarginatus group occupying Waianae Range habitats have been observed recently in nature, collection of M. flavomarginatus in 1906 represents the most recent record for any M. flavomarginatus group species in the Koolau Range, indicating the importance of conserving appropriate Waianae Range habitats in order to preserve representative biodiversity in this species group.
... This biogeographic pattern is not restricted to squamate reptiles. Distinctly similar distributional breaks have been documented in plants (Braithwaite 1975, Chew 1975, Gillison 1975, bats (Medway andMarshall 1975, Flannery 1995), birds (Doughty et al. 1999), freshwater gastropods (Haynes 2000), ground beetles (Liebherr 2005), butterflies (Gross 1975, Ackery et al. 1989, and a number of other invertebrates (Gross 1975). Two general distributional patterns can be seen across a broad range of unrelated taxa: (1) a genus is distributed throughout the Vanuatu Archipelago, with the biota of northern and central islands represented by a different assemblage of species than the southern Vanuatu islands; (2) each biotic region in Vanuatu (northern and central versus southern) is represented by a suite of genera, none of which bridges the biotic break between Efate and Erromango islands. ...
The study of distinct biogeographic demarcations has played a pivotal role in our understanding processes responsible for patterns of species distributions and, importantly, the role of geologic processes in promoting biotic diversification. Biogeographic barriers such as Wallace's line have been shown to be the result of old geologic processes shaping ancient faunal or floral diversification events. Based on distributions of birds, bats, reptiles, plants, and invertebrates we identify a distinct biogeographic disjunction in Vanuatu, a geologically nascent oceanic archipelago. We discuss mechanisms contributing to this concordant pattern across these disparate taxonomic groups in light of geologic history, ocean currents, vegetation, soil, and bioclimatic data, and propose the name Cheesman's line to indicate the faunal and floral discontinuity between the northern and southern islands of Vanuatu.
... This conclusion may have been based on the trichotomy of Phanodesta, Gymnochilini and Trogossitini provided in Kolibáč (2006, but see revised tree in 2008), but a more parsimonious explanation of his tree would be a Gondwanan origin because the trichotomy has an immediate sister relationships to Egoliini which consists of three Gondwanan taxa (Acalanthis Erichson, Argentina and Chile; Necrobiopsis Crowson, Australia; Paracalanthis Crowson, Australia) and a derived Neotropical genus Calanthosoma Reitter (Venezuela, Brazil, Antilles) represented as a basal grade in Trogossitinae in both studies by Kolibáč (2006Kolibáč ( , 2008. Meanwhile, the inclusion of Phanodesta pudica from Lord Howe Island and species in New Caledonia indicates a distribution within the Melanesian Arc (or Rift), an area of Miocene-aged fragments (Liebherr, 2005;Michaux, 2009) that may have contained the ancestors of Phanodesta that split from a more widespread Australasian Leperina + Phanodesta lineage. ...
Abstract. Members of Leperina Erichson (Trogossitidae: Gymnochilini) from New Zealand, New Caledonia and Lord Howe Island are morphologically similar to members of the endemic Juan Fernandez Island genus Phanodesta Reitter, sharing at least one obvious character, elytral carinae that are beaded and contain well-defined punctures. To test the monophyly of Leperina and Phanodesta, we reconstructed phylogenetic relationships of the genera of the tribe Gymnochilini by a cladistic analysis of 22 terminals and 47 adult characters rooted with one genus of trogossitine. Leperina is rendered paraphyletic by the placements of Seidlitzella Jakobson and Phanodesta. Kolibacia n.gen. (type species Leperina tibialis Reitter) is described for east Palaearctic species included formerly in Leperina (two new combinations); New Zealand Leperina and other species from New Caledonia and Lord Howe Island (Ostoma pudicum Olliff) are transferred to Phanodesta (six new combinations); and the remaining species are retained in Leperina. The following species are described as new: Phanodesta carinata n.sp., P. manawatawhi n.sp., P. oculata n.sp. and P. tepaki n.sp. Leperina ambiguum Broun is transferred to Grynoma Sharp resulting in a new combination and three new synonymies for New Zealand trogossitines: Leperina interrupta Brookes n.syn. and Leperina sobrina (White) n.syn. [= Phanodesta farinosa (Sharp)], and Trogosita affinis White n.syn. (= Tenebroides mauritanicus Linnaeus). A key to the New Zealand species and a checklist for the species of the Kolibacia, Leperina and Phanodesta are provided. The derived placement of Juan Fernandez Phanodesta in the phylogeny is evidence for long-distance dispersal from Australasia. A tally of all Juan Fernandez Islands Coleoptera shows derivation mostly from Chile and South America, with few from the southern Pacific region, rarely from Australasia.
... This isolation has permitted relatively few successful colonizing taxa resulting in a disharmonic fauna (Zimmerman 1948). Considering the native French Polynesian carabid fauna derived via autochthonous speciation, the 108 Mecyclothorax beetle species share the archipelago with only two Colpodes species and Metacolpodes monticola (Fairmaire) (tribe Platynini : Perrault 1977;Liebherr 2005a;Nishida 2008), and a single species of Bembidion (tribe Bembidiini: Liebherr and Maddison 2013). Disharmony also extends to possible competitors, as Tahiti supports only one potentially native ant species; the rarely encountered Oligomyrmex tahitiensis Wheeler (Wilson and Taylor 1967;Perrault 1976. ...
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.
... Subsequently, two more Blackburnia species were described (Liebherr 2001, 2003) with inclusion of those species in the cladistic analysis enhancing resolution of the phylogenetic hypothesis while corroborating the overall conclusion that the earliest diverging extant Blackburnia species reside on Kauai, with newer islands progressively colonized during evolutionary diversification of the group. Complementary cladistic analysis including taxa from the Australian and Asian biogeographic regions (Liebherr 2005) affirms that the Blackburnia radiation is old amongst those in the Pacific—older than the radiations in Fiji, Vanuatu, Samoa, and Tahiti—suggesting that Blackburnia colonized the Hawaiian chain during the Miocene, relatively soon after the period of continuous Hawaiian island presence began with the emergence of Kure Island, 29–30 Ma (Carson and Clague 1995). ...
Blackburnia riparia, new species is described from the summit of Mt. Waialeale, Kauai, Hawaii, incorporated into a phylogenetic analysis of Blackburnia Sharp, and thereby placed as the adelphotaxon to two other allopatric Kauai species, B. lata Liebherr and B. atra Liebherr. The new species occurs in riparian habitats, including vertical rock-faced seeps and algal mats, and is documented to feed on aquatic larvae of Micropsectra Kieffer (Diptera: Chironomidae). Blackburnia elegans (Sharp) adults also occur in streams on Waialeale summit where they were found walking under water on the undersides of large, flat rocks. Based on documented prey and observed behavior, both species appear to forage under the waterline. The abdominal and elytral anatomy of adult B. riparia and B. elegans is suitable for retention of a subelytral air bubble, suggesting these species respire underwater. Adults and associated larvae of Blackburnia mandibularis Liebherr inhabit moss-mats along falls' margins at Waipoo Falls, Kokee State Park, Kauai, indicating that this species' active life stages also reside in the riparian zone. Attributes of the larval head capsule and mandibles for B. riparia and B. mandibularis are associated with homologous characters expressed in adult anatomy, ontogenetically linking larval and adult head and mouthpart specializations. The various evolutionary origins of riparian habits are examined across Blackburnia. Via one route, occupation of the montane riparian zone has evolved from terrestrial moss-mat habitation, with concomitant evolutionary reduction of adult compound eyes. A second evolutionary route to the riparian zone is based on occupation of open, disturbed or ephemeral habitats, in some cases assisted by adult winged dispersal.
... The most diverse lineages, including Hyposmocoma Walsingham (Lepidoptera: Cosmopterygidae) (Zimmerman, 1978b; of the high Hawaiian islands, from Kauai to Hawaii Island. Its phylogenetic position within Pacific Platynini dates the radiation's Hawaiian origin to the Early Miocene (Liebherr, 2005b), considerably before the origin of Kauai, the current oldest high island dated to 5.1 Ma (Carson & Clague, 1995). By contrast, the greater species-level diversity of Mecyclothorax beetles may have evolved subsequent to the lineage's colonization of Maui Nui (Britton, 1948;Liebherr, 2008a): the time of origin of the oldest fragment, West Molokai, is dated at 1.9 Ma. ...
The Mecyclothorax Sharp beetle fauna of Oahu, Hawaii is revised, including the description of 13 new species—M. acherontius, M. uncinus, M. lobatus, M. dentatus, M. calceus, M. euryoides, M. invictus, M. ovatulus, M. lemur, M. satyrus, M. ramsdalei, M. pelops and M. niobespp.n.—to complement the six previously described species. Several cryptic species complexes characterized by similar external anatomy and highly divergent male genitalia are present in the fauna, including the M. brevis (Blackburn) species complex—four species sympatrically distributed on Tantalus, a mountain on the northern margin of Honolulu—and the sympatric sister species M. carteri (Perkins) and M. invictus from Mt. Kaala in the western Waianae Range. This repeated pattern of closely related species characterized by divergent male genitalia and relatively static external anatomy is also observed in the Mecyclothorax fauna of Molokai, an island of moderate geologic age within the Hawaiian archipelago, but not in the Mecyclothorax fauna of the most recently emergent Hawaii Island. Species distributions are generally also much more restricted on the older islands. These results are consistent with the occurrence of rampant, unfixed infraspecific genitalic variation early in an island radiation, with subsequent fixation of divergent genitalic characters occurring as speciation proceeds. Three distinct areas of endemism are defined by distributions of Oahu Mecyclothorax: the Waianae Range, the north Koolau Range and the south Koolau Range. Species distributions in two other carabid beetle genera in Hawaii containing native species—Blackburnia Sharp and Bembidion Latreille—congruently define these areas. The historical range of sampling dates is compared for Mecyclothorax species across the three areas, with the greatest diminution of recently collected species-level diversity characterizing the south Koolau Range, where several Mecyclothorax species are characterized both by restricted geographic distributions within the developmental footprint of Honolulu, and by a lack of recent field collections, suggesting that they may qualify for endangered species status.
... The island hopping hypothesis predicts that close relatives of Fijian taxa are likely to be present on the intermediate islands of the former Vitiaz Arc, and that phylogenetic patterns will reflect stepwise diversification towards Fiji (Burrett et al., 1991). Several previous studies have invoked island hopping across the Vitiaz Arc to explain other Fijian endemic invertebrate groups, including Cosmopsaltriina cicadas (Hemiptera: Cicadidae) (Duffels & Turner, 2002), and Platynini ground beetles (Coleoptera: Carabidae) (Liebherr, 2005). ...
Aim This study addresses the origins of terrestrial biodiversity of the Fijian islands using the ant genus Lordomyrma (Hymenoptera: Formicidae: Myrmicinae) as a model system. We derive the evolution of the genus and determine its closest extra‐Fijian relatives from geological data, molecular phylogenetic reconstruction and divergence estimates .
Location Ant taxa were sampled in the Southwest Pacific, Melanesia, Southeast Asia, Australia and mainland China.
Methods Phylogeny and divergence estimates of the ant genus Lordomyrma based on four nuclear genes (28S, ArgK, LW Rh, CAD) plus data on Indo‐Pacific geological history are used to address current hypotheses regarding the origins of the Fijian biota.
Results The genus Lordomyrma probably originated in mainland Asia, with subsequent colonization of Australia and the Pacific. The Fijian Lordomyrma clade is monophyletic, and originated c. 8.8 Ma, when it diverged from a sister group in Papua New Guinea.
Main conclusions The colonization of Fiji by Lordomyrma is probably a result of long‐distance dispersal from New Guinea, possibly aided by island hopping across the Vitiaz Arc. The timeline of diversification in Lordomyrma is broadly congruent with the Miocene fragmentation of the Vitiaz Arc and the Pliocene emergence of Vanua Levu. The biotic shuttle hypothesis, which posits ‘Eua Island as the source of Fijian endemics, is rejected based on the sister relationship of Fiji and New Guinea lineages, as well as on the Miocene submergence of the terrane below sea level. The diversity of Fijian Lordomyrma results from the radiation of a single lineage, which diverged from a New Guinea sister group. The genus appears to have originated in Asia rather than in Australia.
... It is ironic that the theoretical foundations of both the taxon cycle and island biogeography were built largely upon distributional data of Pacific ants, as native ants are conspicuously absent from many Pacific islands (Wilson & Taylor 1967). While insular radiations are documented for many terrestrial arthropods (Gillespie & Roderick 2002;Liebherr 2005;Bickel 2006;Monaghan et al. 2006;Balke et al. 2007b;Joy et al. 2007), few examples can be drawn from the literature of eusocial insects (Wilson 1988;Fisher 1997Fisher , 2010Lucky & Sarnat 2009). In fact, the absence of eusocial Hymenoptera (ants, bees and wasps) from eastern Pacific islands is implicated as a cause for the ecological release of other arthropods (Gillespie & Roderick 2002;Krushelnycky et al. 2005). ...
While insular radiations are documented for many terrestrial arthropods, few examples are known for eusocial insects. This study seeks to ascertain whether the spinescence observed among Fijian Pheidole ants was inherited from an oversea ancestor or is evidence of ecological release from interspecific competitors. We broaden our understanding of morphological convergence, insular radiation and Pacific biogeography by testing three hypotheses proposed previously for the Fijian Pheidole roosevelti group: (i) the group is monophyletic; (ii) spinescence is a plesiomorphic trait inherited from an overseas ancestor; and (iii) the group is closely related to spinescent New Guinean relatives. The analysis included the fragments of two mitochondrial genes (COI, cytb) and two nuclear genes (H3, EF1α-F2) from 66 taxa, including all members of the roosevelti group, representatives from the spinescent subgenus Pheidolacanthinus, Fijian congeners and widespread Pacific congeners. Our results yield new insights into the biogeographic history of Fiji, reveal a fascinating example of convergent evolution and serve as a novel example of ecological release occurring within an insular eusocial insect lineage. These findings recover the history of a presumably unremarkable ant species that colonized a remote oceanic archipelago in the Miocene (17-10 Ma) and radiated across the emerging islands into niche-space occupied elsewhere in the Pacific by distantly related spinescent congeners. We propose the radiation of Fijian Pheidole into spinescent morphotypes was the consequence of ecological opportunities afforded by the absence of competing ant lineages with conspicuous epigaeic foraging strategies.
An updated checklist of 188 species of the Platyninae (with 139 species under Platynini and 49 species under Sphodrini) known in the Indian subcontinent, along with details of the revisions, literature, and distribution patterns is provided. One hundred twenty-five species of these occur only in the Indian landmass, with 89 in the Platynini and 36 in the Sphodrini. Of the 125 Indian species, 94 occur within the Palaearctic region in India, 22 in the Oriental, and five in both Oriental and Palaearctic regions. Among the 125 Indian species, 111 are endemic to three global hotspots of biodiversity in the Indian subcontinent: (i) 96 endemic to the Himalaya, (ii) eight to the Western Ghats and Sri Lanka, and (iii) seven to the Indo-Burma hotspot of biodiversity. Five genera are endemic to the Indian mainland, and 10 genera to the Himalaya hotspot of biodiversity.
The tribe Platynini (Coleoptera: Carabidae: Harpalinae) is revised for New Zealand. Eight genera and forty-three species are recognized.
Four genera and sixteen species are described as new: Ctenognathus davidsoni Larochelle and Larivière new species, Ctenognathus earlyi Larochelle and Larivière new species, Ctenognathus garnerae Larochelle and Larivière new species, Ctenognathus hoarei Larochelle and Larivière new species, Ctenognathus kaikoura Larochelle and Larivière new species, Ctenognathus marieclaudiae Larochelle new species, Ctenog�nathus perumalae Larochelle and Larivière new species, Ctenognathus takahe Larochelle and Larivière new species, Ctenognathus tawanui Larochelle and Larivière new species, Ctenognathus tepaki Larochelle and Larivière new species, Ctenognathus urewera Larochelle and Larivière new species, Kiwiplatynus Larochelle and Larivière new genus, Kiwiplatynus taranaki Larochelle and Larivière new species, Kupeplatynus Laro�chelle and Larivière new genus, Maoriplatynus Larochelle and Larivière new genus, Maoriplatynus marrisi Larochelle and Larivière new species, Prosphodrus mangamuka Larochelle and Larivière new species, Pros�phodrus sirvidi Larochelle and Larivière new species, Prosphodrus waimana Larochelle and Larivière new species, Tuiplatynus Larochelle and Larivière new genus.
Lectotypes are designated for twelve taxa: Anchomenus adamsi Broun, 1886, Anchomenus colensonis White, 1846, Anchomenus feredayi Bates, 1874, Anchomenus helmsi Sharp, 1881, Anchomenus intermedius Broun, 1908, Anchomenus macrocoelis Broun, 1908, Anchomenus munroi Broun, 1893, Anchomenus sophro�nitis Broun, 1908, Colpodes crenatus Chaudoir, 1878, Colpodes neozelandicus Chaudoir, 1878, Ctenognathus littorellus Broun, 1908, and Ctenognathus pictonensis Sharp, 1886.
Six new combinations are established: Ctenognathus bidens (Chaudoir, 1878) = Kiwiplatynus bidens (Chaudoir, 1878); Ctenognathus crenatus (Chaudoir, 1878) = Kupeplatynus crenatus (Chaudoir, 1878); Ctenog�nathus lucifugus (Broun, 1886) = Kupeplatynus lucifugus (Broun, 1886); Ctenognathus sulcitarsis (Broun, 1880)
= Kupeplatynus sulcitarsis (Broun, 1880); Ctenognathus libitus (Broun, 1914) = Tuiplatynus libitus (Broun, 1914); Ctenognathus sophronitis (Broun, 1908) = Tuiplatynus sophronitis (Broun, 1908).
Fifteen new synonymies are established: Ctenognathus littorellus Broun, 1908 = Ctenognathus adamsi (Broun, 1886); Anchomenus parabilis Broun, 1880 = Ctenognathus cardiophorus (Chaudoir, 1878); Ancho�menus integratus Broun, 1908 = Ctenognathus colensonis (White, 1846); Anchomenus macrocoelis Broun, 1908 = Ctenognathus edwardsii (Bates, 1874); Ctenognathus actochares Broun, 1894 = Ctenognathus elevatus
(White, 1846); Anchomenus punctulatus Broun, 1877, Anchomenus montivagus Broun, 1880, Anchomenus perrugithorax Broun, 1880, Anchomenus politulus Broun, 1880, Anchomenus suborbithorax Broun, 1880, and Colpodes neozelandicus Chaudoir, 1878 = Ctenognathus novaezelandiae (Fairmaire, 1843); Ctenognathus simmondsi Broun, 1912 = Ctenognathus pictonensis Sharp, 1886; Anchomenus (Platynus) cheesemani Broun,
1880 and Calathus deformipes Broun, 1880 = Kupeplatynus crenatus (Chaudoir, 1878); Anchomenus munroi Broun, 1893 = Kupeplatynus lucifugus (Broun, 1886). Ctenognathus elevatus (White, 1846), previously syn�onymized with Ctenognathus novaezelandiae (Fairmaire, 1843), is reinstated as full species.
A revision of all taxa is provided. Descriptions, identification keys, illustrations of male genitalia, hab�itus photos, distributional data and maps are given. Extensive information on ecology, biology, dispersal power, and collecting techniques is included for each species.
The Australian fauna of Mecyclothorax Sharp (Coleoptera: Carabidae: Moriomorphini) is reviewed, with special focus on species assigned to the monophyletic subgenus Eucyclothorax Liebherr: M.isolatus , sp. n. from Western Australia, M.moorei Baehr, M.punctatus (Sloane), M.curtus (Sloane), M.blackburni (Sloane); M.eyrensis (Blackburn); M.peryphoides (Blackburn); M.darlingtoni , sp. n. from Queensland; M.jameswalkeri , sp. n. from Western Australia; M.lophoides (Chaudoir); and M.cordicollis (Sloane). The last six species listed above–the M.lophoides species complex–have been the source of long-term confusion for taxonomists, with male genitalic characters providing trouble-free species circumscription. One new subspecies, M.lewisensisestriatus , subsp. n. from Queensland is added to the seven previously described taxa of the monophyletic subgenus Qecyclothorax Liebherr. The balance of the fauna consists of four species in the subgenus Mecyclothorax: 1 and 2 , the sister-species pair M.lateralis (Castelnau) and M.minutus (Castelnau); 3 , M.ambiguus (Erichson); and 4 , M.punctipennis (MacLeay). Mecyclothoraxfortis (Blackburn), syn. n. , is newly synonymized with M.minutus . Mecyclothoraxovalis Sloane is recombined as Neonomiusovalis (Sloane), comb. n. , and a neotype is designated to replace the destroyed holotype. Phylogenetic relationships for the Australian Mecyclothorax are proposed based on information from 68 terminal taxa and 139 morphological characters. The biogeographic history of Australian Mecyclothorax is deduced based the sister-group relationship between Mecyclothorax and the Amblytelus -related genera, with both groups hypothesized to have originated during the late Eocene. Diversification within Mecyclothorax has occurred since then in montane rainforests of tropical Queensland, temperate forest biomes of the southwest and southeast, and in grasslands and riparian habitats adjacent and inland from those forests. Several species presently occupy interior desert regions, though no sister species mutually occupy such climatically harsh habitats. The M.lophoides species complex exhibits profound male genitalic diversification within the context of conserved external anatomy. This disparity is investigated with regard to the functional interaction of the male internal sac flagellum and female spermathecal duct. Though limited association of flagellar and spermathecal duct configurations can be documented, several factors complicate proposing a general evolutionary mechanism for the observed data. These include: 1 , the occurrence of derived, elongate spermathecal ducts in three species, two of which exhibit very long male flagella, whereas males of the third exhibit a very short flagellum; and 2 , a highly derived and exaggerated male flagellar configuration shared across a sister-species pair even though the two species can be robustly diagnosed using external anatomical characters, other significant genitalic differences involving male parameral setation, and biogeographic allopatry associated with differential occupation of desert versus forest biomes.
Much Australasian biodiversity, lacking the decimating effects of the predominantly northern-hemisphere Pleistocene glaciations, remains intact from a much deeper geological history. This chapter presents some highlights of Australasian insect biodiversity. Selections from Australian insect biodiversity are harvested by Aboriginal Australians. The Arrente people use yarumpa (honey ants) and udnirringita (witchetty grubs), which have now entered the non-indigenous food chain in restaurants catering to wealthy tourists and political entertainment. For the total Australian insect diversity, Taylor estimated about 110,000 species. One of the strongest contrasts between insect biodiversity in New Zealand and Australia is the modest diversity and ecological insignificance of New Zealand ants - the country has only seven genera and 11 species of native ants. The processes threatening insect biodiversity vary somewhat across and within Australasia, but there are several unifying causes. These are habitat loss, introduced animals (including invasive ants), potential effects of climate change, and a minor possibility of overexploitation.
The Mecyclothorax carabid beetle fauna of Haleakalā volcano, Maui Island, Hawai‘i is taxonomically revised, with 116 species precinctive to Haleakalā recognized, 74 newly described. Species are classified into 14 species groups, with the newly described species arrayed as follows: 1, M. constrictus group with M. perseveratus sp. n.; 2, M. obscuricornis group with M. notobscuricornis sp. n., M. mordax sp. n., M. mordicus sp. n., M. manducus sp. n., M. ambulatus sp. n., M. montanus sp. n., M. waikamoi sp. n., M. poouli sp. n., and M. ahulili sp. n.; 3, M. robustus group with M. affinis sp. n., M. anchisteus sp. n., M. consanguineus sp. n., M. antaeus sp. n., M. cymindulus sp. n., and M. haydeni sp. n.; 4, M. interruptus group with M. bradycelloides sp. n., M. anthracinus sp. n., M. arthuri sp. n., M. medeirosi sp. n., M. inconscriptus sp. n., and M. foveolatus sp. n.; 5, M. sobrinus group with M. foveopunctatus sp. n.; 6, M. ovipennis group with M. subtilis Britton & Liebherr, sp. n., M. patulus sp. n., M. patagiatus sp. n., M. strigosus sp. n., M. takumiae sp. n., M. parapicalis sp. n., M. mauiae sp. n., M. subternus sp. n., M. flaviventris sp. n., M. cordaticollaris sp. n., and M. krushelnyckyi sp. n.; 7, M. argutor group with M. ommatoplax sp. n., M. semistriatus sp. n., M. refulgens sp. n., M. argutulus sp. n., M. planipennis sp. n., M. planatus sp. n., and M. argutuloides sp. n.; 8, M. microps group with M. major sp. n., M. xestos sp. n., M. orbiculus sp. n., and M. contractus sp. n.; 9, M. scaritoides group with M. scarites sp. n., M. timberlakei sp. n., M. crassuloides sp. n., M. crassulus sp. n., M. gracilicollis sp. n., and M. dispar sp. n.; 10, M. haleakalae group with M. reiteratus sp. n., M. splendidus sp. n., M. bacrionis sp. n., and M. simpulum sp. n.; 11, M. vitreus group with M. kipwilli sp. n., M. kipahulu sp. n., M. kaumakani sp. n., and M. kuiki sp. n.; 12, M. montivagus group with M. rex sp. n.; 13, M. ducalis group with M. aquilus sp. n., M. invisitatus sp. n., M. longidux sp. n., and M. brevidux sp. n.; and 14, M. palustris group with M. hephaestoides sp. n., M. oculellus sp. n., M. bicoloris sp. n., M. bicoloratus sp. n., M. bilobatus sp. n., M. palustroides sp. n., M. filipoides sp. n., M. nanunctus sp. n., M. tauberorum sp. n., and M. pau sp. n. Mecyclothorax integer Sharp, stat. n. is recognized as a species distinct from M. 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 Haleakalā 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 Haleakalā.
Hawaiianische Käfer der Gattung Blackburnia (Coleoptera, Carabidae, Platynini): ökolo- gische Spezialisierungen und Implikationen für den Artenschutz Die monophyletische hawaiianische Gattung Blackburnia SHARP, 1878 umfasst 132 Arten und kann in vier aufeinander folgende Adelphotaxa unterteilt werden: Untergattung Protocaccus LIEBHERR & ZIMMERMAN, 2000 (1 Art, flugfähig); Untergattung Colpocaccus SHARP, 1903 (4 Arten, flugfähig); Untergattung Blackburnia (52 Arten, viele mit reduzierten Flügeln, Flugfähigkeit aber im Grund- muster); Untergattung Metromenus S HARP, 1884 (75 Arten mit reduzierten Flügeln). Die phylogenetischen Beziehungen der Gattung zu anderen Taxa der Platynini beweisen, dass Blackburnia die hawaiianischen Inseln im Miozän kolonisiert hat, wobei Kauai die zuerst besiedelte Insel darstellt. Die ökologischen Spezialisierungen der einzelnen Arten spiegeln sich in den phylogenetischen Beziehungen der Arten wider. Arten der Untergattung Blackburnia besiedeln weniger Habitattypen im Vergleich zu Arten der Untergattungen Colpocaccus und Metromenus. Die Larven der Untergattungen Blackburnia und Metromenus entwickeln sich vergleichsweise langsam und einige Weibchen der Untergattung Blackburnia legen die größten Eier innerhalb der Platynini. Die größte morphologische und genetische Diversität findet sich innerhalb der Untergattung Black- burnia, was auf eine größere Fragmentierung der Populationen der betreffenden Arten hinweist. Die umfassenderen ökologischen Spezialisierungen der Arten der Untergattung Blackburnia können auf dramatische historische Populationsschwankungen innerhalb der Arten zurückgeführt werden. Anthropogene Einflüsse, wie die Degradierung spezifischer Habitattypen (mesische Acacia koa Wälder) und die Einschleppung von Neozoen (z.B. Formicidae und Isopoda), haben aber die Assoziationen zwischen den phylogenetischen Beziehungen und den ökologischen Spezialisierun- gen verwischt. Diese Einflüsse machen deshalb eine Voraussage des Gefährdungspotentials der Arten mit Hilfe der Ergebnisse der phylogenetischen Analyse unmöglich. Einige Arten, die noch im 19. Jahrhundert Bestandteil der Koa-Wälder auf Maui waren, sind seit Anfang des 20. Jahrhunderts nicht mehr nachgewiesen worden. Diese nicht mehr nachgewiesenen Arten, welche Bestandteil spezifischer ökologischer Assoziationen waren, können bei genauerer Betrachtung die Gefährdungs- kriterien der I.U.C.N. erfüllen. Das Fehlen phylogenetisch verwandter Arten macht die Gefährdung der Arten selbst, als auch die Gefährdung der Koa-Wälder als Ökosysteme deutlich.
All scientific names of Trachypachidae, Rhysodidae, and Carabidae (including cicindelines) recorded from America north of Mexico are catalogued. Available species-group names are listed in their original combinations with the author(s), year of publication, page citation, type locality, location of the name-bearing type, and etymology for many patronymic names. In addition, the reference in which a given species-group name is first synonymized is recorded for invalid taxa. Genus-group names are listed with the author(s), year of publication, page citation, type species with way of fixation, and etymology for most. The reference in which a given genus-group name is first synonymized is recorded for many invalid taxa. Family-group names are listed with the author(s), year of publication, page citation, and type genus. The geographical distribution of all species-group taxa is briefly summarized and their state and province records are indicated.
One new genus-group taxon, Randallius new subgenus (type species: Chlaenius purpuricollis Randall, 1838), one new replacement name, Pterostichus amadeus new name for Pterostichus vexatus Bousquet, 1985, and three changes in precedence, Ellipsoptera rubicunda (Harris, 1911) for Ellipsoptera marutha (Dow, 1911), Badister micans LeConte, 1844 for Badister ocularis Casey, 1920, and Agonum deplanatum Ménétriés, 1843 for Agonum fallianum (Leng, 1919), are proposed. Five new genus-group synonymies and 65 new species-group synonymies, one new species-group status, and 12 new combinations (see Appendix 5) are established.
The work also includes a discussion of the notable private North American carabid collections, a synopsis of all extant world geadephagan tribes and subfamilies, a brief faunistic assessment of the fauna, a list of valid species-group taxa, a list of North American fossil Geadephaga (Appendix 1), a list of North American Geadephaga larvae described or illustrated (Appendix 2), a list of Geadephaga species described from specimens mislabeled as from North America (Appendix 3), a list of unavailable Geadephaga names listed from North America (Appendix 4), a list of nomenclatural acts included in this catalogue (Appendix 5), a complete bibliography with indication of the dates of publication in addition to the year, and indices of personal names, supraspecific names, and species-group names.
Australasia - the LocaleSome Highlights of Australasian Insect BiodiversityDrowning by Numbers? How Many Insect Species are in Australasia?Australasian Insect Biodiversity - Overview and Special ElementsThreatening Processes to Australasian Insect BiodiversityAustralasian Biodiversity ConservationConclusion
References
Seamounts and islands are quite common features of the deep-ocean basins, particularly the Pacific Basin. A conspicuous but relatively small number of these volcanoes are arranged in long linear chains (Figure 1). The vast majority, however, occur as isolated edifices. Recent work on a small number of these isolated volcanoes shows that many were formed at or very close to mid-ocean spreading axes (Clague and Dalrymple, 1975; Batiza, 1980) but that others are significantly younger than the underlying oceanic crust (G. B. Dalrymple, D. A. Clague, H. G. Greene, unpublished data). These isolated volcanoes require much additional study before their origin is understood. The volcanoes arranged in linear chains have been the focus of a great deal of work, and much has been learned about their origin and evolution. Most of the linear volcanic chains in the Pacific Basin are oriented roughly west-northwesterly and apparently formed sequentially above stationary hot spots during the past 42 m. y. as the Pacific plate rotated clockwise about a pole located near 69°N, 68°W (Clague and Jarrard, 1973a). Another group of linear chains exhibit roughly north-trending orientations and apparently were formed by the same mechanism between at least 80 and 42 m. y. ago as the Pacific plate rotated about a pole of rotation located near 17°N, 107°W (Clague and Jarrard, 1973b). Earlier age data available from linear island and seamount chains were summarized by Jackson (1976), Jarrard and Clague (1977), and McDougall and Duncan (1980); we will briefly review the information presented there and update it with more recent data.
The island of Maupiti is the top of an eroded volcanic shield surrounded by a large lagoon and coral reef. The most important part of this shield is made up of alkali basaltic lavas grading upward into hawaiitic flows. The shield-building event is dated at ca. 4.51 +/- 0.04 Ma. A few mugearitic and benmoreitic late dykes were emplaced at 4.21 +/- 0.04 Ma. These chronological data are consistent with a Pacific plate motion of 11 cm/year with respect to a fixed Society plume. The Maupiti lava series results from the effects of fractional crystallisation of a parental alkali basalt magma displaying a prominent EM II isotopic and trace element signature, in many ways similar to that of the youngest Society hot spot lavas. This feature suggests that the geochemical signature of the Society plume did not change through time.
Electrophoretic allelic frequency data, rearing through two generations, and analysis of field collections from polymorphic populations, all suppport synonymizing Agonum californicum (Dejean) and Agonum thoracicum (Dejean) under Agonum decorum (Say). Agonum decorum is composed of four morphotypes, distributed in different regions of North America. The red-green color polymorphism and hirsute-glabrous setational polymorphism are shown to be determined by unlinked, autosomal diallelic loci. Both presumably derived alleles are dominant to the plesiomorphic alleles. The polymorphisms are discussed in relation to mimicry and the evolution of dominance.
Abstract A consensus cladogram is presented for ninety-seven of the ninety-nine (sub)species of the cicada subtribe Cosmopsaltriina on the basis of a cladistic analysis of forty-nine characters. The consensus cladogram confirms the monophyly of the eight genera constituting the subtribe, and the monophyly of the subtribe as a whole. Cosmopsaltriina occur in Sulawesi, the Moluccas, New Guinea, Cape York, the Solomon Islands, Vanuatu, Fiji, Tonga and Samoa. The subtribe demonstrates a high rate of endemism on genus, species group and species level. The taxon-area and consensus-area cladograms resulting from Brooks parsimony analysis are discussed in relation to the palaeogeography of the area.
The monophyletic, native Hawaiian Platynini have diversified on the Hawaiian Island chain through progressive colonization, mixed with vicariance on the various islands. Single-island endemism stands at 97% of the species, with the few widespread species exhibiting distributions largely congruent with the fundamental area cladogram found using cladistic biogeographic methods. The cost of accepting an ad hoc dispersal hypothesis for individual taxa that conflicts with the fundamental area cladogram is weighed against the savings in items of error when taxa are excluded from the biogeographic analysis. Based on this objective assessment, only one back-dispersal from Maui Nui to O'ahu is supported. Vicariance of Maui Nui, leading to the present-day islands of Moloka'i, Lana'i, and Maui, has resulted in seven resolvable species triplets composed of single-island endemics occupying these areas. These seven triplets represent five biogeographic patterns, necessitating explanation by numerous ad hoc hypotheses of extinction to support a single hypothesis of area relationships. In six of the seven triplets, the cladistically basal species exhibits a higher minimum elevational limit of occupied habitat than either of the more apical sister species. This result is consistent with isolation of more persistent, peripheral populations at higher elevations, leading to speciation. Comparison of higher-elevation endemics to lower-elevation widespread species supports this interpretation. Such a finding affirms the importance of understanding geographic distribution on a scale appropriate to the action of vicariant mechanisms.
This study quantifies long-term landscape changes in the Hawaiian archipelago relating to dispersal, speciation and extinction. Accounting for volcano growth, subsidence and erosion, we modelled the elevations of islands at time intervals of 0.5 Myr for the last 32 Myr; we also assessed the variation in the spacing of volcanoes during this period. The size, spacing and total number of volcanic islands have varied greatly over time, with the current landscape of large, closely spaced islands preceded by a period with smaller, more distantly spaced islands. Considering associated changes in rates of dispersal and speciation, much of the present species pool is probably the result of recent colonization from outside the archipelago and divergence within contemporary islands, with limited dispersal from older islands. This view is in accordance with abundant phylogenetic studies of Hawaiian organisms that estimate the timing of colonization and divergence within the archipelago. Twelve out of 15 multi-species lineages have diverged within the lifetime of the current high islands (last 5 Myr). Three of these, and an additional seven (mostly single-species) lineages, have colonized the archipelago within this period. The timing of colonization of other lineages remains uncertain.
The Parsimony Ratchet1 is presented as a new method for analysis of large data sets. The method can be easily implemented with existing phylogenetic software by generating batch command files. Such an approach has been implemented in the programs DADA (Nixon, 1998) and Winclada (Nixon, 1999). The Parsimony Ratchet has also been implemented in the most recent versions of NONA (Goloboff, 1998). These implementations of the ratchet use the following steps: (1) Generate a starting tree (e.g., a "Wagner" tree followed by some level of branch swapping or not). (2) Randomly select a subset of characters, each of which is given additional weight (e.g., add 1 to the weight of each selected character). (3) Perform branch swapping (e.g., "branch-breaking" or TBR) on the current tree using the reweighted matrix, keeping only one (or few) tree. (4) Set all weights for the characters to the "original" weights (typically, equal weights). (5) Perform branch swapping (e.g., branch-breaking or TBR) on the current tree (from step 3) holding one (or few) tree. (6) Return to step 2. Steps 2-6 are considered to be one iteration, and typically, 50-200 or more iterations are performed. The number of characters to be sampled for reweighting in step 2 is determined by the user; I have found that between 5 and 25% of the characters provide good results in most cases. The performance of the ratchet for large data sets is outstanding, and the results of analyses of the 500 taxon seed plant rbcL data set (Chase et al., 1993) are presented here. A separate analysis of a three-gene data set for 567 taxa will be presented elsewhere (Soltis et al., in preparation) demonstrating the same extraordinary power. With the 500-taxon data set, shortest trees are typically found within 22 h (four runs of 200 iterations) on a 200-MHz Pentium Pro. These analyses indicate efficiency increases of 20×-80× over "traditional methods" such as varying taxon order randomly and holding few trees, followed by more complete analyses of the best trees found, and thousands of times faster than nonstrategic searches with PAUP. Because the ratchet samples many tree islands with fewer trees from each island, it provides much more accurate estimates of the "true" consensus than collecting many trees from few islands. With the ratchet, Goloboff's NONA, and existing computer hardware, data sets that were previously intractable or required months or years of analysis with PAUP* can now be adequately analyzed in a few hours or days.
Four species new to science are described from Vanuatu, namely Holophygdon nishidae sp. n., Idiocysta vanuatuensis sp. n., Nesocypselas minicysta sp. n., and Tingis parvoroe sp. n. New records are provided for six species, five of them being new to Vanuatu: Cysteochila idonea Drake, Stephanitis subfasciata Horvath, Teleonemia scrupulosa Stal, Tingis chazeaui Guilbert, and Tingis irregularis (Montrouzier). A key to genera and species of Vanuatu is added.
The use of the generic name Colpodes MacLeay, 1825, is reviewed, along with characteristics previously attributed to the genus by various authors of Carabidae. The single known specimen of the type species, Rembus (Colpodes) brunneus MacLeay, is designated as lectotype, redescribed and figured. Based on comprehensive character analysis, including external anatomical, male genitalic, and female genitalic and reproductive tract characters, application of the generic-level name Colpodes is restricted to a clade of Javanese species also including Colpodes brittoni Louwerens and C. latus Louwerens. In order to further nomenclatural stability within Platynini, Colpodes should best be considered a subgeneric component of an expanded genus Platynus Bonelli, 1810.
Vanuatu (formerly the New Hebrides Anglo—French Condominium) forms one sector of a Cretaceous to Recent island arc system extending from New Britain through the Solomon Islands to Vanuatu, Fiji, Tonga and the Kermadec Islands (Fig. 1). The component arcs originally formed a volcanic chain to the south and west of a subduction zone dipping towards Australia and now seen as a discontinuous line of active and inactive trenches. Fragmentation of this earlier arc—trench system during the late Miocene resulted in the formation of new subduction zones dipping northwards and eastwards beneath the Vanuatu, Solomon Islands and New Britain arc sections.
Characters of the female reproductive tract, ovipositor, and abdomen are analyzed using cladistic
parsimony for a comprehensive representation of carabid beetle tribes. The resulting cIadogram
is rooted at the family Trachypachidae. No characters of the female reproductive tract define the
Carabidae as monophyletic. The Carabidae exhibit a fundamental dichotomy, with the isochaete tribes Metriini and Paussini forming the adelphotaxon to the Anisochaeta, which includes Gehringiini and Rhysodini, along with the other groups considered member taxa in Jeannel's classification. Monophyly of Isochaeta is supported by the groundplan presence of a securiform helminthoid sclerite at the spermathecal base, and a rod-like, elongate laterotergite IX leading to the explosion chamber of the pygidial defense glands. Monophyly of the Anisochaeta is supported by the derived division of gonocoxa IX into a basal and apical portion. Within Anisochaeta, the evolution of a secondary spermatheca-2, and loss of the primary spermatheca-I has occurred in one lineage including the Gehringiini, Notiokasiini, Elaphrini, Nebriini, Opisthiini, Notiophilini, and Omophronini. This evolutionary replacement is demonstrated by the possession of both spermatheca-like structures in Gehringia o/ympica Darlington and Omophron variegatum (Olivier). The adelphotaxon to this spermatheca-2 clade comprises a basal rhysodine grade consisting of Clivinini, Promecognathini, Amarotypini, Apotomini, Melaenini, Cymbionotini, and Rhysodini. The Rhysodini and Clivinini both exhibit a highly modified laterotergite IX; long and thin, with or without a clavate lateral region. This may represent a synapomorphous derivation, or convergence based on a tubular abdomen and burrowing habit. The basal grade gives rise to a grade of taxa sharing the presence of a ramus--a sclerotized portion of the vaginal wall situated medially to each gonocoxal base--along with the plesiomorphic spermatheca-I. The previous interpretation of the gonocoxal rami as a groundplan feature of Carabidae representing portions of abdominal segment VIII is rejected based on its derived occurrence relative to the origin of the family. The gonocoxal ramus or cicindine grade includes I) Cicindini, 2) a monophyletic "carabine" clade including Siagonini, Cychrini, Pamborini, Carabini, and Cicindelini, 3) and the Scaritini and Hiletini which are paraphyletically related to a monophyletic group exhibiting the harpalidian abdominal configuration, including Broscini and all remaining tribes of Carabidae. Broscini retain the separate accessory gland and spermathecal arrangment of Scaritini and Hiletini, and comprise the adelphotaxon to the other remaining tribes. Patrobini represent the next divergent group, exhibiting an appended spermathecal gland but retaining the ramus at the base of gonocoxite I. The remaining tribes, including member tribes of Jeannel's Stylifera (in part) and Conchifera, are ambiguously related based on female characters at a highly polytomous II node. Within this polytomous clade, Pseudomorphini share a derived basal spermathecal sclerite with Geobaenini, Lachnophorini, and Odacanthini. Cnemalobini are placed as the adelphotaxon to Morionini. Other tribal relationships remain ambiguous due to basic homogeneity of the female reproductive tract observed throughout these tribes. The problematic Brachinini appear best placed among these tribes, although a less preferred but equally parsimonious placement is as adelphotaxon to Clivinini. Psydrini and Zolini appear polyphyletic based on female characters, with constituent subtribes placed at various positions between the gonocoxal ramus grade and the highest polytomous clade. The basal position of Clivinini and Apotomini supported by female characters is contrasted with Jeannel's placement in Scrobifera and Stylifera, respectively, based on thoracic structure.
The Parsimony Ratchet1 is presented as a new method for analysis of large data sets. The method can be easily implemented with existing phylogenetic software by generating batch command files. Such an approach has been implemented in the programs DADA (Nixon, 1998) and Winclada (Nixon, 1999). The Parsimony Ratchet has also been implemented in the most recent versions of NONA (Goloboff, 1998). These implementations of the ratchet use the following steps: (1) Generate a starting tree (e.g., a “Wagner” tree followed by some level of branch swapping or not). (2) Randomly select a subset of characters, each of which is given additional weight (e.g., add 1 to the weight of each selected character). (3) Perform branch swapping (e.g., “branch-breaking” or TBR) on the current tree using the reweighted matrix, keeping only one (or few) tree. (4) Set all weights for the characters to the “original” weights (typically, equal weights). (5) Perform branch swapping (e.g., branch-breaking or TBR) on the current tree (from step 3) holding one (or few) tree. (6) Return to step 2. Steps 2–6 are considered to be one iteration, and typically, 50–200 or more iterations are performed. The number of characters to be sampled for reweighting in step 2 is determined by the user; I have found that between 5 and 25% of the characters provide good results in most cases. The performance of the ratchet for large data sets is outstanding, and the results of analyses of the 500 taxon seed plant rbcL data set (Chase et al., 1993) are presented here. A separate analysis of a three-gene data set for 567 taxa will be presented elsewhere (Soltis et al., in preparation) demonstrating the same extraordinary power. With the 500-taxon data set, shortest trees are typically found within 22 h (four runs of 200 iterations) on a 200-MHz Pentium Pro. These analyses indicate efficiency increases of 20×–80× over “traditional methods” such as varying taxon order randomly and holding few trees, followed by more complete analyses of the best trees found, and thousands of times faster than nonstrategic searches with PAUP. Because the ratchet samples many tree islands with fewer trees from each island, it provides much more accurate estimates of the “true” consensus than collecting many trees from few islands. With the ratchet, Goloboff's NONA, and existing computer hardware, data sets that were previously intractable or required months or years of analysis with PAUP* can now be adequately analyzed in a few hours or days.
The use of the generic name Colpodes MacLeay, 1825, is reviewed, along with characteristics previously attributed to the genus by various authors of Carabidae. The single known specimen of the type species, Rembus (Colpodes) brunneus MacLeay, is designated as lectotype, redescribed and figured. Based on comprehensive character analysis, including external anatomical, male genitalic, and female genitalic and reproductive tract characters, application of the generic-level name Colpodes is restricted to a clade of Javanese species also including Colpodes brittoni Louwerens and C. latus Louwerens. In order to further nomenclatural stability within Platynini, Colpodes should best be considered a subgeneric component of an expanded genus Platynus Bonelli, 1810.
Dating of Fijian volcanic rocks enables an estimate of 4.9+/-0.4×106 yr to be made for the age of the Miocene-Pliocene boundary. The change in composition of the volcanism in Fiji between about 5 and 6×106 yr ago may result from migration of the site of subduction.
Knowledge of the time span of arc activity, essential for correct tectonic reconstructions, has been lacking for the Kermadec arc system, but is supplied in this paper through study of microfossils contained in dredge samples, and K–Ar ages on dredged basalt clasts. The Kermadec system at south latitudes 30 to 32° in the southwest Pacific comprises from west to east the Colville Ridge (remnant arc), Havre Trough (backarc basin), Kermadec Ridge (active arc) and Kermadec Trench (site of west-dipping subduction of Pacific plate lithosphere beneath the Australian plate). Data are presented from two traverses (dredge, magnetic, single-channel seismic) across the whole system. An important transverse tectonic boundary, the 32°S Boundary, lies between the two traverse lines and separates distinct northern (32–25°S) and southern (32–36°S) sectors. The northern sector is shallower and well sedimented with broad ridges and a diffuse backarc basin. The southern sector is deeper with narrow ridges and steep escarpments facing inwards to a little-sedimented, rifted backarc basin. The Kermadec Ridge slopes smoothly trenchward to a mid-slope terrace (forearc basin) with minor sediment fill at 5–6 km water depth. A steeper (10–24°) and more rugged lower trench slope is mantled with New Zealand-sourced rhyolitic vitric mud diamictons containing locally derived mafic volcanic clasts; one clast is of late Miocene age (K–Ar age 7.84 Ma). The arc (Kermadec Ridge) is capped by active volcanoes; very young K–Ar ages (
Eighty-nine eucnemid genera occur in the region from South-east Asia to the south-west Pacific. The phylogenies of 84 of these were used together with the present-day distributions of the species to analyse the biogeographical history of the area. Fifty-seven genera shared a pattern coinciding with the traditional model of Laurasia–Gondwana break-up. Six genera showed a pattern contradicting the model. The remaining 21 genera neither supported nor refuted the model. Twenty-five genera were observed to include an Indomalesian clade younger than the South America–Australia connection. This biogeographical unit consisted of present-day South-east Asia and the Sunda islands, but did not include the Philippine Islands and Sulawesi. In addition to this Indomalesian clade, three separate clades involving northern Australia or New Guinea were observed, New Guinea–Australia, New Guinea–Philippines–Sulawesi and New Guinea–Fiji. The possible presence of four separate areas in the general region of New Guinea–north Australia as the result of the Cretaceous geological events is suggested. Three of these, in the area of present-day New Guinea, originally sharing sister-groups with the north-eastern Australian isolate, are regarded as the sources of the New Guinea–Indomalesia, New Guinea–Philippines and New Guinea–Fiji faunas after northward drifting of the Australian continent. During the Oligocene–Miocene these source areas were flooded and their original fauna lost. When the present-day New Guinea emerged, it was invaded from the north-eastern Australian region. This invasion created new New Guinea–Australia connections and brought in the sister-groups of the old New Guinea source areas as well. The eucnemids of Vanuatu, Samoa and Tonga are regarded as having originated in connection with dispersal from Fiji. The New Zealand fauna has strong, old connections with that of south-eastern Australia, but other complex connections are indicated. The Eocene Baltic Amber fauna agrees well with the results obtained from extant species. The species belonging to five fossil genera belong to Gondwanan groups that seem to have invaded the Holarctic via Central America. Four other fossil genera showing discordant patterns belong to the group of six genera exhibiting these aberrant patterns even today. The eucnemid fauna of the region is of Gondwanic origin. Only six Laurasian genera have invaded the area, all of them apparently quite recently.
Blackburnia lata sp. n. is described from the western reaches of the Alakai Swamp Trail, Kauaikinana Stream watershed, Kauai. Blackburnia lata is placed phylogenetically among other Kauai species in the subgenus Blackburnia Sharp, 1878. It is sympatric with three closely related species, all distributed largely within the limits of the primeval Kauai caldera, but allopatric with its adelphotaxon, B. atra Liebherr, found on Mt. Kahili, an isolated southern spur of the Mt. Waialeale massif. All five species occur in arboreal mossmats, at the same time exhibiting substantial variation in tarsal configuration. Tarsal evolution across this mossmat-specific lineage suggests that a single tarsal conformation is not required for occupation of this arboreal ecological zone. Historical transformation patterns of tarsal characters are not entirely congruent with the phylogenetic relationships among these species, and inclusion or exclusion of these characters in cladistic analysis is irrelevant to the definition of phylogenetic relationships. This incongruence argues against the necessity of excluding characters from phylogenetic analysis because they are deemed adaptive, while supporting the total evidence approach, wherein all potential synapomorphies are analyzed simultaneously to estimate phylogeny, regardless of their putative adaptive significance.
The 23 species of Bembidiini revised in this paper were earlier referred variously to subgenus (or genus) Cillenus Sam. or to closely allied taxa. It is shown that the five New Zealand species are taxonomically quite isolated and different from Bembidion; for these the new genus Zecillenus was erected. All remaining species are true members of genus Bembidion belonging to five different subgenera, all previously named. Subgenus Desarmatocillenus Netolitzky contains 13 species distributed from Tasmania to Japan; three other subgenera are restricted to China, Formosa and Japan, respectively. The single European species, B. laterale Samouelle, constitutes subgenus Cillenus s. str.—Six species, Zecillenus embersoni and 5 Bembidion, adelaideae, foochowense, hebridarum, kasaharai, seticorne, are described as new.—None of the subgenera shows clear affinities to any other subgenus of Bembidion. This was judged primarily on the structures of the internal sac of penis.
The fauna of Anthribinae of New Caledonia (inclusive of Art I., Îie des Pins, and Loyalty Islands) and Vanuatu (formerly New Hebrides) is revised. Keys and descriptions are given for all taxa, and 164 line drawings illustrate male and female genitalia and a few other features. The number of species is increased from 14 to 56 for New Caledonia, and from 1 to 9 for Vanuatu representing together 62 species in 24 genera; 9 genera and 43 species are described as new. New Caledonia shares with Vanuatu 5 genera and 3 species, with Australia 5 genera and 1 species, and with New Zealand 5 genera but no species. Araecerus insularis Fauvel is synonymised with A. lutatus (Fairmaire), Acanthopygus rubricollis Montrouzier with A. griseus Montrouzier, Alloderes Jordan with Plintheria Pascoe, Plintheria subtilis Jordan with Helmoreus dufouri (Montrouzier), Jordanopus Elgueta with Dinema Fairmaire, and Jordanopus veitchi (Jordan) and J. Caledoniense (Frieser) with Dinema filicorne Fairmaire. The new genera and their type species are as follows: Haplopygus for H. fauveli sp.n., Pistorhinus for P. fidelis sp.n., Mionus forM creper sp.n., Mesocranius for Proscoporhinus apicatus Jordan, Xylanthribus for Anthribisomus xylophagus Jordan, Etnalidius for is. acuticollis sp.n., Sophronus for S. tibialis sp.n., Aranthribus for A. araucariae sp.n., and Solox for S. vanuatensis sp.n. A list of all species and their distribution is provided.
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.
The World fauna of Agonum Bonelli, 1810 was analyzed by cladistic parsimony implementing Nixon's ratchet, based on 135 terminals representing 131 species, and 138 unit-coded characters, including 98 external morphological characters, 21 characters of the female reproductive tract and ovipositor, and 19 male genitalic characters. The cladogram was outgroup-rooted to the African Agonidium Jeannel, 1948. Phylogenetic data were summarized via strict consensus; this consensus was used as the basis for cladistic classification. A single preferred cladogram was chosen from among 4576 unambiguously optimized, equally parsimonious cladograms using Goloboffs fit value as an optimality criterion, and converted to a taxon-area cladogram for biogeographic analysis. Coxal and femoral setation has generally increased during phylogenesis, and served to define natural groups. The configuration of the female bursa copulatrix helped define the Agonum ingroup. Spermathecal duct length has increased dramatically during diversification of Agonum, though reversal to a short, broad duct has also occurred. A balanced cladistic classification based on relationships maintained in the strict consensus cladogram recognized four subgenera: Platynomicrus Casey, 1920 (2 spp.), Europhilus Chaudoir, 1859 (28 spp.), Agonum (29 spp.), and Agonothorax Motschulsky, 1850 (72 spp.). Nomenclatural changes necessitated by the cladistic analysis include: 1, Megalonychus Chaudoir, 1843 = Agonum Bonelli, 1810; 2, Agonops Bousquet, 2002 = Agonum Bonelli, 1810; 3, 12 species previously combined with Megalonychus are NEWLY COMBINED with Agonidium, whereas the remaining 32 species previously combined with Megalonychus are provisionally assigned to Agonidium pending future study. Fossil locality data were used to determine Pliocene and Pleistocene distributional limits, and to establish minimum ages for extant taxa. Estimates of species duration for three species preserved in the Beaufort Formation of Arctic Canada range from 3–4,000,000 yr. Pliocene and Pleistocene fossil localities in eastern North America and Greenland augmented extant species distributions analyzed biogeographically. Dispersal-vicariance biogeographic analysis (DIVA), which optimized area relationships among major areas of endemism in the northern hemisphere using the taxon-area cladogram, indicated that Agonum diversification began in Eocene time when Europe and Eastern North America were connected by North Atlantic Land Bridges. Eocene origin of Agonum is consistent with a Paleocene-constrained biotic connection to the African outgroup Agonidium. Subsequently, Beringian exchange was associated with Agonum diversification through Late Miocene and Pliocene. Dispersal across Pleistocene-aged Beringia is associated strictly with extant Holarctic and circumpolar species. Plesiomorphically, Agonum beetles were marsh dwellers. Subsequent ecological specialization has involved use umbrotrophic bogs, gravel and cobble streambank habitats, shores of saline lakes and seashores, and moist forest floors. Speciation has followed both an allopatric vicariant mode, and peripatric isolation of peripheral populations. Both Mayr's centripetal speciation and Brown's centrifugal speciation models were supported by geographic relationships of various sister species, indicating that environmental factors have greatly influenced speciation mode, especially of peripheral isolates.
Abstract— Rerent criticisms of component analysis are based on misunderstandings of the relationship between component analysis, parsimony and consensus methods. These criticisms are rebutted, and the appropriateness of applying the Wagner parsimony criterion to the study of biogcography and co-speciation is questioned. An alternative parsimony method, previously applied to mapping gene cladograms onto organism cladograms, is developed.
The New Zealand Subantarctic Islands, emergent remnants of the Campbell Plateau, were given World Heritage status in 1998 in recognition of their importance to global biodiversity. We describe the flora and fauna of these islands and discuss the results of recent phylogenetic analyses. Part of the New Zealand Subantarctic biota appears to be relictual and to be derived from west Gondwana. The relictual element is characterized by genera endemic to the Campbell Plateau that show relationships with taxa of the southern South Island, New Zealand, southern South America, and the north Pacific. In contrast, a younger, east Gondwanan element is composed of species that are either taxonomically identical to widespread mainland species, or endemic species with close New Zealand relatives. Area cladograms support the inclusion of the southern South Island, New Zealand and Macquarie Island (although this is separate geologically) as parts of the Campbell Plateau, but suggest the Chatham Rise and Torlesse terranes of the eastern South Island, New Zealand were originally parts of east Gondwana. East and west Antarctica acted as independent plates during the breakup of Gondwana, and were separated by oceanic crust until a compressive phase sutured them along the trace of the trans-Antarctic mountains during the early Tertiary. The Campbell Plateau microcontinent was connected to west Antarctica until its separation at 80 Mya, contemporaneous with the separation of the southern portion of the Melanesian rift from east Gondwana. Presently the Campbell Plateau is joined to the Melanesian Rift along the Alpine Fault. Cenozoic plate tectonic reconstructions place the Campbell Plateau adjacent to the Melanesian Rift throughout the rift–drift phase, relative motion being confined to strike–slip movement over the last 20 Myr. Our synthesis of phylogenetic and plate tectonic evidence suggests that the Alpine Fault is the most recent development of a much older extensional rift/basin boundary originally separating west and east Gondwana. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86, 95–115.
Abstract— Branch support is quantified as the extra length needed to lose a branch in the consensus of near-most-parsimonious trees. This approach is based solely on the original data, as opposed to the data perturbation used in the bootstrap procedure. If trees have been generated by Farris's successive approximations approach to character weighting, branch support should be examined in terms of weighted extra length needed to lose a branch. The sum of all branch support values over the tree divided by the length of the most parsimonious tree[s] provides a new index, the total support index. This index is a measure of tree stability in terms of supported resolutions, which is of prime importance in cladistic analysis.
Opening mode of the Japan Sea is studied from the viewpoint of geologic structure around the western marginal fault zone of the back-arc basin. Latest reflection seismic survey in the Fukue Basin, located at the western end of the rifted sliver of southwest Japan, delineates an NNE–SSW transcurrent fault that is an extension of the Yangsan Fault running along the eastern coast of the Korean Peninsula. As suggested by paleomagnetic studies in Korea, the Yangsan Fault was activated with dextral motions during the early Miocene rifting event of the Japan Sea. Our offshore seismic study indicates that the NNE–SSW fault zone was simultaneously active with dextral sense because it constitutes a pull-apart depression on a right stepping of the fault. The Fukue Basin was under continuous transtensional stress regime during the Japan Sea rifting accompanied with a clockwise rotation of southwest Japan. It is discordant from the previous tectonic model of two-stage (parallel and fan-shaped) back-arc opening, which requires compressive regime with sinistral fault motions to the west of rotation pivot of southwest Japan in the second stage. The southward transportation of the rifted sliver with formation of pull-apart basins was the dominant mode along the western margin of the Japan Sea throughout the opening event. The southern part of the marginal fault zone has been immune from post-opening tectonic events, such as the late Miocene strong inversion on the back-arc margin of southwest Japan or the Quaternary E–W tectonic stress in the Japan Sea reactivating the northern sector of the fault zone, Yangsan Fault.
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.
The complex biogeographic relations of the region reflect its geologic history.
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