Family‐level relationships of the spittlebugs and froghoppers (Hemiptera: Cicadomorpha: Cercopoidea)

The spittlebug superfamily Cercopoidea (Hemiptera: Cicadomorpha) comprises approximately 3000 phytophagous species (including some economically important pests of grass crops) classified among the families Cercopidae, Aphrophoridae, Epipygidae, Clastopteridae and Machaerotidae. However, the monophyly of these taxa has never been tested and the evolutionary relationships among these major lineages are unknown. Presented here are the results of the first ever phylogenetic investigation of the higher‐level relationships within Cercopoidea, based on DNA nucleotide sequence data from six loci (18S rDNA, 28S rDNA, histone 3, wingless, cytochrome oxidase I and cytochrome oxidase II) generated from exemplars of 109 spittlebug species representing all five described families, seven of eight subfamilies and 61 genera (eight additional exemplars, representing a selection of other Auchenorrhyncha taxa, were included as outgroups). The resulting topologies are used to evaluate the monophyly of each cercopoid family, and further to calculate divergence date estimates to examine the chronological origins and historical diversification of Cercopoidea. The results of this investigation suggest that: (i) four of the five described families are monophyletic; Epipygidae was recovered consistently as originating within Aphrophoridae; (ii) the exclusively Old World Machaerotidae is the most anciently diversified family of extant spittlebugs; (iii) New World Cercopidae (i.e. Ischnorhininae) constitute a derived monophyletic lineage; (iv) the genus Microsargane Fowler, classified currently within Aphrophoridae, actually belongs within Cercopidae; and (v) the origins of the major spittlebug lineages probably coincided with the breakup of Pangaea and, subsequently, Gondwana, as well as major floristic diversification such as the rise of angiosperms.     

Historical biogeography of scarabaeine dung beetles

Abstract Aim (1) To review briefly global biogeographical patterns in dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae), a group whose evolutionary history has been dominated by ecological specialization to vertebrate dung in warmer climates. (2) To develop hypotheses accounting for the evolution of these patterns. Location Six principal biogeographical regions: Palaearctic, Oriental, Afrotropical, Australasia, Neotropical, Nearctic and five outlying islands or island groups harbouring endemic genera: Caribbean, Madagascar, Mauritius, New Caledonia, New Zealand. Methods Major patterns of tribal, generic and species distribution are investigated using cluster analysis, ordination, parsimony analysis of endemism and track analysis. Attempts are made to resolve biogeographical patterns with findings in the fields of plate tectonics, fossil and evolutionary history, plus phylogeny of both mammals and dung beetles. Results Because of conflict between published findings, it is uncertain at what point in time density of dinosaur dung, mammal dung or both became sufficiently great to select for specialized habits in dung beetles. However, biogeographical evidence would suggest a Mesozoic origin followed by further taxonomic radiation during the Cenozoic, possibly in response to the increasing size and diversity of mammalian dung types in South America and Afro‐Eurasia. Proportional generic distribution in fourteen tribes and subtribes showed four principal biogeographical patterns: (1) southerly biased Gondwanaland distribution, (2) Americas or (3) Madagascar endemism, and (4) northerly biased, Afro‐Eurasian‐centred distribution with limited numbers of genera also widespread in other regions. Proportional composition of faunas in eleven geographical regions indicated three principal distributional centres, East Gondwanaland fragments, Afro‐Eurasia and the Americas. These patterns probably result from three principal long‐term range expansion and vicariance events (Mesozoic: Gondwanaland interchange and fragmentation, Cenozoic: Afro‐Eurasian/Nearctic interchange and the Great American interchange). It is suggested that old vicariance caused by the Mesozoic fragmentation of Gondwanaland leads to a high degree of regional endemism at generic or tribal level across one or more Gondwanaland tracks. In contrast, it is suggested that the more recent Cenozoic range expansions occurred primarily towards northern regions leading to endemism primarily at species level. These Cenozoic radiations were facilitated by the re‐linking of continents, either because of tectonic plate movements (Africa to Eurasia in Miocene), climatically induced sea‐level change (Afro‐Eurasia to Nearctic in Miocene and Pleistocene), or similar coupled with orogenics (Nearctic to Neotropical in Pliocene). Speciation has followed vicariance either because of climatic change or physical barrier development. These recent range expansions probably occurred principally along an Afro‐Eurasian land track to the Nearctic and Neotropical and an Americas land track northwards from the Neotropics to the Nearctic, with limited dispersal from Eurasia to Australia, probably across a sea barrier. This accounts for the overall, spatially constrained, biogeographical pattern comprising large numbers of species‐poor genera endemic to a single biogeographical region and fewer more species‐rich genera, many of which show wider biogeographical distributions. In most southerly regions (Australasia, Madagascar, Neotropical), faunal composition and generic endemism is primarily dominated by elements with Gondwanaland ancestry, which is consistent with the Gondwanaland origin claimed for Scarabaeinae. In Afro‐Eurasia (Palaearctic, Oriental, Afrotropical), generic endemism of monophyletically derived Afro‐Eurasian and widespread lineages is centred in the Afrotropical region and faunal composition is numerically dominated by Afro‐Eurasian and widespread elements. In the Nearctic region, the fauna is jointly dominated by widespread elements, derived from Afro‐Eurasia, and Gondwanaland and Americas elements derived from the Neotropical region. Main conclusions Global biogeographical patterns in scarabaeine dung beetles primarily result from Mesozoic and Cenozoic range expansion events followed by vicariance, although recent dispersal to Australia may have occurred across sea barriers. Detailed phylogenetics research is required to provide data to support dispersal/vicariance hypotheses.     

On the Proteaceae—the evolution and classification of a southern family*

A scheme of phylogeny in the Proteaceae is derived from analysis of new and previously available data on morphological, anatomical and chromosomal characters. The probable attributes of a common ancestor are indicated and it is concluded that the family has no close relatives, although it possibly diverged early from the Rosiflorean line. Chromosomal evolution has been complex, with early polyploidy, repeated dysploid reductions and great change in chromosome size, from a probable base of x = 7. Habit, wood anatomy, leaf form and sequence, chemical features, inflorescences, pollination and pericarp anatomy are discussed in relation to adaptation, phylogeny, parallelism and convergence. Distribution is analysed ecogeographically and in relation to the fossil record and chronology of continental movement. Ancestors of major groups may have differentiated before the break-up of Gondwanaland. Early members of almost all tribes were probably trees of mesothermic closed forests; xeromorphic characters found in many modern genera are secondary and of multiple origin in Australia, South Africa, New Caledonia and to a lesser degree in South America. The Proteaceae, with 75 genera, are divided into 5 subfamilies (3 new) comprising 14 tribes, further subdivided into 33 units of subtribal level. New genera described are Toronia, Acidonia, Pycnonia, Triunia, Malagasia, Athertonia, Virotia, Floydia.     

Taxonomy, cladistics and biogeography of Coenosopsia Malloch (Diptera, Anthomyiidae) and its significance to the evolution of anthomyiids in the Neotropics

Abstract.  The anthomyiid fly genus Coenosopsia Malloch contained five species geographically restricted to the forests of the New World, from southern U.S.A. to southeastern Brazil and Paraguay. Two new species are described here from the Brazilian Cerrado: C. ferrari sp.n. and C. michelseni sp.n. Viviparity in C. brasiliensis Michelsen and C. peruviana Michelsen is reported and discussed. A cladistic analysis of the genus was performed using Fannia bahiensis Albuquerque (Fanniidae), Polietina orbitalis (Stein) (Muscidae), Anthomyia pluripunctata (Albuquerque), Delia platura (Meigen) and Phaonantho benevola Couri (Anthomyiidae) as outgroups. The analysis was carried out using three character weighting schemes: equal, successive and implied weighting. In the phylogenetic relationship (( C. ferrari , C. brasiliensis ) ( C. peruviana ( C. prima ( C. michelseni ( C. floridensis , C. mexicana ))))), two major clades were found, one distributed mainly in southeastern South America and the other from northwestern South America to southern North America. Reconciling the phylogeny with the available distributional data, a biogeographical analysis of the genus is proposed and discussed. The pattern found for Coenosopsia corroborated a previously proposed model of vicariance events for the Neotropical region. Although the presence of anthomyiid fauna in the region was explained previously on the basis of the North-to-South America dispersal, we suggest an alternative hypothesis, that of a Gondwanan origin for the Neotropical anthomyiids.     

A molecular phylogeny and classification of Verbenaceae

? Premise of the study: Verbenaceae consist of trees, shrubs, lianas, and herbs distributed primarily in Latin America, where they occur in a wide array of ecosystems. A second center of diversity exists in Africa. Competing morphology-based classifications that rely on different traits conflict in significant ways. A broad phylogenetic study was undertaken to assess those classifications and to examine the historical geography of the family. ? Methods: Analysis of seven chloroplast DNA regions for 109 species, representing all genera except one monotypic genus, provide inference into evolutionary relationships in Verbenaceae. ? Key results: The phylogeny shows that none of the traditional classifications reflect phylogenetic relationships very well. Eight clades are recognized as tribes (Casselieae, Citharexyleae, Duranteae, Lantaneae, Neospartoneae trib. nov., Petreeae, Priveae, and Verbeneae). Two genera, Dipyrena and Rhaphithamnus, remain unplaced in these larger clades. Petreeae, which consist of Neotropical lianas, are sister to the rest of the family. Lantaneae and Verbeneae together form a derived clade that comprises approximately two-thirds of the species in Verbenaceae. ? Conclusions: We present a new tribal classification, including one new tribe, Neospartoneae trib. nov., to accommodate three small genera of Argentine species (Diostea, Neosparton, and Lampaya). Phylogenetic inference suggests a South American origin for Verbenaceae, with approximately six colonization events having given rise to the Old World species.     

Cladistic biogeography of the Neotropical region: identifying the main events in the diversification of the terrestrial biota

A cladistic biogeographical analysis was undertaken to identify the main events in the biotic diversification of the terrestrial Neotropical biota. For the 36 animal and plant taxa analysed, a component × area matrix was constructed, associating geographical data only with informative nodes, and it was analysed under implied weights using the software TNT. The general area cladogram obtained shows that the Neotropical region constitutes a monophyletic unit, with a first split separating the Antilles and a second one dividing the continental areas into a north‐western and a south‐eastern component. Within the north‐western component the areas split following the sequence northern Amazonia, south‐western Amazonia, north‐western South America, and Mesoamerica. Within the south‐eastern component the areas split following the sequence south‐eastern Amazonia, Chaco, and Parana. The three main components are treated as subregions: Antillean, Amazonian (northern Amazonian, south‐western Amazonian, Mesoamerican, and north‐western South American dominions), and Chacoan (south‐eastern Amazonian, Chacoan, and Parana dominions). Dispersal and vicariant events postulated to explain these pattens might have occurred during the Cretaceous, when the Caribbean plate collided with the Americas, a combination of eustatic sea‐level changes and tectonic deformations of the continental platform exposed large parts of South America to episodes of marine transgressions, and the Andean uplift reconfigured the Amazonian area. Tertiary and Quaternary events are assumed to have later induced the diversification within these large biogeographical units.     

Phylogenetic Relationships and the Radiation of Sigmodontine Rodents in South America: Evidence from Cytochrome b

Phylogenetic relationships among South American sigmodontine rodents were examined based on the complete sequence for the mitochondrial cytochrome b gene [1140 base pairs (bp)] for 66 species and between 759 and 1140 bp for an additional 19 species. Thirty-eight South American genera were represented, coming from eight of nine tribes. Outgroups included the North American murid rodents Peromyscus, Reithrodontomys, Scotinomys, and Neotoma, the Old World murine rodents Mus and Rattus, and the geomyoid genera Thomomys, Geomys, Dipodomys, and Perognathus as the most distant outgroup. The South American sigmodontines were supported as a monophyletic lineage. Within this radiation several clear-cut suprageneric groupings were identified. Many of the currently recognized tribal groupings of genera were found fairly consistently, although not always with high levels of bootstrap support. The various tribes could not be linked hierarchically with any confidence. In addition, several genera stand out as unique entities, without any apparent close relatives. The overall pattern suggests a rapid radiation of the sigmodontines in South America, followed by differentiation at the tribal and generic levels.     

Divergence times and origin of neotropical sheath-tailed bats (tribe Diclidurini) in South America

Times of divergence and origin of sheath-tailed bats (family Emballonuridae) in the New World were approximated with a relaxed molecular clock approach using Bayesian analysis of introns from the three nuclear genetic transmission systems in mammals (autosomal, X and Y sex chromosomes). An upper constraint of 30 mya for the oldest known Neotropical emballonurid fossil and a lower constraint of 13 mya for the only pre-Pleistocene fossil of an extant genus were used as calibration points. Differentiation began in the Late Oligocene with the appearance of two subtribes as independently corroborated by each gene. Following an explosive model of evolution, the genera diversified relatively suddenly in the Early Miocene with seven of the eight genera radiating within 1.4 myr and most intrageneric speciation occurring before the Pliocene. Optimization of ancestral areas onto the phylogeny suggests that the ancestor of New World emballonurid bats has its origin in Africa and this is the third report of placental mammals colonizing South America by trans-Atlantic dispersal and subsequent speciation in allopatry.     

Phylogeny, classification, and biogeography of the cycloteline Therevinae (Insecta: Diptera: Thereavidae)

Twenty-one members of the Laurasian group of Therevinae (Diptera: Therevidae) are compared using 65 adult morphological characters. Cladistic analysis using parsimony on the 17 ingroup and 4 outgroup taxa provides a well-supported hypothesis of relationships among taxa within the Gyclotelini, tribe nov. The Cyclotelini is a monophyletic assemblage of mostly New World genera, including Anolinga , gen. nov. , Breviperna Irwin, Coleiana , gen. nov. , Crebraseta , gen. nov. , Cyclotelus Walker, Mesonana , gen.nov. , and Ozodiceromyia Bigot. In addition, three Old World genera, Ammothereva Lyneborg, Bugulaverpa , gen. nov. , and Procyclotelus Nagatomi & Lyneborg, are included in the tribe. These ten genera are divided into two monophyletic genus-groups, the Brevipema-group and the Cyclotelus-group. Keys are provided for the genera of Cyclotelini. The tribe, the two informal genus-groups, and all genera are diagnosed; five new genera and six new species are proposed. The biogeographical histories of the genera are discussed in terms of their cladistic relationships using methods of cladistic biogeography. Two major vicariant events account for the current distribution of the tribe. The first relates to the Beringian land bridge connecting western North America and eastern Asia. Second, New World cyclotelines were profoundly affected by the Early Eocene breakup of the archipelagic bridge between North and South America, and the distributions support the hypotheses favouring the continental origin of the Greater Antilles.     

Mitochondrial and nuclear phylogenies of Cervidae (Mammalia, Ruminantia): Systematics, morphology, and biogeography

The family Cervidae includes 40 species of deer distributed throughout the northern hemisphere, as well as in South America and Southeast Asia. Here, we examine the phylogeny of this family by analyzing two mitochondrial protein-coding genes and two nuclear introns for 25 species of deer representing most of the taxonomic diversity of the family. Our results provide strong support for intergeneric relationships. To reconcile taxonomy and phylogeny, we propose a new classification where the family Cervidae is divided in two subfamilies and five tribes. The subfamily Cervinae is composed of two tribes: the tribe Cervini groups the genera Cervus, Axis, Dama, and Rucervus, with the Père David's deer (Elaphurus davidianus) included in the genus Cervus, and the swamp deer (Cervus duvauceli) placed in the genus Rucervus; the tribe Muntiacini contains Muntiacus and Elaphodus. The subfamily Capreolinae consists of the tribes Capreolini (Capreolus and Hydropotes), Alceini (Alces), and Odocoileini (Rangifer + American genera). Deer endemic to the New World fall in two biogeographic lineages: the first one groups Odocoileus and Mazama americana and is distributed in North, Central, and South America, whereas the second one is composed of South American species only and includes Mazama gouazoubira. This implies that the genus Mazama is not a valid taxon. Molecular dating suggests that the family originated and radiated in central Asia during the Late Miocene, and that Odocoileini dispersed to North America during the Miocene/Pliocene boundary, and underwent an adaptive radiation in South America after their Pliocene dispersal across the Isthmus of Panama. Our phylogenetic inferences show that the evolution of secondary sexual characters (antlers, tusk-like upper canines, and body size) has been strongly influenced by changes in habitat and behaviour.     

New fossil paussids from Dominican amber with notes on the phylogenetic systematics of the paussine complex (Coleoptera: Carabidae)

The first fossil representative of the Protopaussinae and the second fossil Eohomopterus (Paussinae) are described from amber inclusions of Tertiary age collected in the Dominican Republic on the West Indian island of Hispaniola ( Protopaussus pristinus sp.n., Eohomopterus poinari sp.n.). These finds increase the total number of paussids present in Dominican amber to four species in three genera ( Protopaussus , Eohomopterus , Homopterus ). A key is provided for the identification of the New World genera of Protopaussinae and Paussinae and for the species of Eohomopterus . Based on synapomorphic character states, the paussid beetles are split into two subfamilies, the monobasic Protopaussinae and the more inclusive Paussinae (the twenty-three genera are placed into two supertribes: the Carabidomemnitae comprising two tribes and the Paussitae comprising eight tribes. The extant congeners of Eohomopterus and Homopterus live in the Neotropical Region, while the extant members of Protopaussus are known only from the Oriental Region. This record of New World Tertiary Protopaussus adds another example of the Old World relationships of taxa from Dominican amber. The temporal and geographical distribution of Protopaussus indicates that its present zoogeographical range may represent the relict of a wide Laurasian distribution in the past. It does not support the previously suspected Oriental centre of origin for the ancestral stock of {Protopaussinae + Paussinae}.     

Mitochondrial DNA phylogeny of the woodpecker genus Veniliornis (Picidae, Picinae) and related genera implies convergent evolution of plumage patterns

The woodpecker genus Veniliornis comprises 12 species, all restricted to the New World tropics. The seemingly distantly related genus Picoides is broadly distributed in Eurasia and North America with two putative species, P. lignarius and P. mixtus , occurring in South America. The two genera are clearly distinct with respect to general plumage colouration and patterning as well as habitat utilization and thus traditionally have been placed in different tribes. Phylogenetic analyses of mtDNA sequences from the COI and cyt b genes indicated that both genera are reciprocally paraphyletic. The two South American species of Picoides belong to a clade comprising most species of Veniliornis , but V. fumigatus of Central and north-western South America belongs to a clade comprising species of Picoides . The mtDNA tree also indicated that Veniliornis is not closely related to the genus Piculus, as is implicit in current classifications. Misclassifications involving Veniliornis at both the generic and tribal levels appear to result from convergent evolution of plumage traits in specific forest types. We infer that the common ancestor of Veniliornis entered South America approximately at the time the Isthmus of Panama was formed, and diversification within South America was rapid.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 611–624.     

Phylogeny and historical biogeography of leafhopper subfamily Iassinae (Hemiptera: Cicadellidae) with a revised tribal classification based on morphological and molecular data

Phylogenetic relationships among major lineages of the leafhopper subfamily Iassinae were explored by analysing a dataset of 91 discrete morphological characters and DNA sequence data from nuclear 28S rDNA and histone H3 genes and mitochondrial 12S rDNA. Bayesian, maximum‐likelihood and maximum parsimony analyses yielded similar tree topologies that were well resolved with strong branch support except at the base of the tree, resulting in equivocal support for inclusion of Bythoniini as a tribe of Iassinae but strong support for the monophyly of Iassinae (excluding Bythoniini) and most previously recognized iassine tribes. Divergence times for recovered nodes were estimated using a Bayesian relaxed clock method with two fossil calibration points. The results suggest that the deepest divergences coincided with Gondwanan vicariant events but that more recent divergences resulted from long‐range dispersal and colonization. Biogeographical analyses suggest that the group most likely has a Neotropical origin. The following changes to the taxonomic classification are proposed: establishment of three new tribes, Batracomorphini trib.n. (based on type genus Batracomorphus Lewis), Hoplojassini trib.n. (based on type genus Hoplojassus Dietrich and including one other South American genus), Lipokrisnini trib.n. (based on type genus Lipokrisna Freytag and including two other endemic Caribbean genera); Krisnini is redefined to include only the Old World genera Krisna and Gessius; Iassini is redefined to include only the type genus and four endemic Afrotropical genera; Bascarrhinus Fowler and Platyhynna Berg, recently treated as genera incertae sedis, are placed in Hyalojassini; Thalattoscopus Kirkaldy is added to the previously monobasic tribe Trocnadini. Iassinae now includes 12 tribes, all of which appear to be monophyletic. Revised morphological diagnoses of the subfamily and each of the included tribes are provided and a key to tribes is also given. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:41295B68‐2DAB‐4C4F‐B260‐F7C054922173 .     

Phylogeny of the lizard subfamily Lygosominae (Reptilia: Scincidae), with special reference to the origin of the new world taxa

Phylogenetic relationships of the three lygosomine skink genera occurring both in the Old World and the New World (Mabuya, Scincella and Sphenomorphus) were inferred from mitochondrial DNA sequence of 12S and 16S rRNA genes. Results strongly suggested the non-monophyly for any of these three genera. Within the Mabuya group, Asian members appear to have diverged first, leaving the Neotropical and the Afro-Malagasy Mabuya as sister groups. These relationships, together with the absence of extant or fossil representatives of the Mabuya group from North America, strongly suggest the trans-Atlantic dispersals of Mabuya from Africa to Neotropics. Our results also indicated a closer affinity of the New World Scincella with the New World Sphenomorphus than with the Old World Scincella. Such relationships suggest the trans-Beringian dispersal of the common ancestor from Asia and its subsequent divergence into the North American Scincella and the Neotropical Sphenomorphus.     

Dating an impressive Neotropical radiation: Molecular time estimates for the Sigmodontinae (Rodentia) provide insights into its historical biogeography

With about 400 living species and 82 genera, rodents of the subfamily Sigmodontinae comprise one of the most diverse and more broadly distributed Neotropical mammalian clades. There has been much debate on the origin of the lineage or the lineages of sigmodontines that entered South America, the timing of entrance and different aspects of further diversification within South America. The ages of divergence of the main lineages and the crown age of the subfamily were estimated by using sequences of the interphotoreceptor retinoid binding protein and cytochrome b genes for a dense sigmodontine and muroid sampling. Bayesian inference using three fossil calibration points and a relaxed molecular clock estimated a middle Miocene origin for Sigmodontinae (～12 Ma), with most tribes diversifying throughout the Late Miocene (6.9–9.4 Ma). These estimates together results of analyses of ancestral area reconstructions suggest a distribution for the most recent common ancestor of Sigmodontinae in Central-South America and a South American distribution for the most recent common ancestor of Oryzomyalia.     

Dated historical biogeography of the temperate Loliinae (Poaceae, Pooideae) grasses in the northern and southern hemispheres

Divergence times and biogeographical analyses have been conducted within the Loliinae, one of the largest subtribes of temperate grasses. New sequence data from representatives of the almost unexplored New World, New Zealand, and Eastern Asian centres were added to those of the panMediterranean region and used to reconstruct the phylogeny of the group and to calculate the times of lineage-splitting using Bayesian approaches. The traditional separation between broad-leaved and fine-leaved Festuca species was still maintained, though several new broad-leaved lineages fell within the fine-leaved clade or were placed in an unsupported intermediate position. A strong biogeographical signal was detected for several Asian-American, American, Neozeylandic, and Macaronesian clades with different affinities to both the broad and the fine-leaved Festuca. Bayesian estimates of divergence and dispersal-vicariance analyses indicate that the broad-leaved and fine-leaved Loliinae likely originated in the Miocene (13My) in the panMediterranean-SW Asian region and then expanded towards C and E Asia from where they colonized the New World. Further expansions in America (10-3.8My) showed a predominant migratory route from North to South (N America<-->the Andes<-->Patagonia). This late Tertiary scenario of successive colonizations and secondary polyploid radiations in the southern hemisphere from the northern hemisphere was accompanied by occasional transcontinental long-distance dispersal events between South America and New Zealand. Multiple Pliocene dispersal events (3.6-2.5My) from the near SW European and NW African continents gave rise to the Macaronesian Loliinae flora, while a more recent Pleistocene origin (2-1My) is hypothesized for the high polyploid lineages that successfully colonized newly deglaciated areas in both hemispheres.     

Molecular phylogeny of the Athetini–Lomechusini–Ecitocharini clade of aleocharine rove beetles (Insecta)

Elven, E., Bachmann, L. & Gusarov V. I. (2012) Molecular phylogeny of the Athetini–Lomechusini–Ecitocharini clade of aleocharine rove beetles (Insecta). —Zoologica Scripta, 41, 617–636. It has previously been shown that the Aleocharinae tribes Athetini and Lomechusini form a well‐supported clade, which also includes the small Neotropical tribe Ecitocharini. However, neither Athetini nor Lomechusini were recovered as monophyletic. In this study, we addressed the basal phylogenetic relationships among the three tribes using sequence data from (i) a mitochondrial fragment covering the COI, Leu2 and COII genes; (ii) a mitochondrial fragment covering part of the 16S gene, the Leu1 gene and part of the NADH 1 gene; and (iii) a part of the nuclear 18S gene, for 68 Athetini, 33 Lomechusini and 2 Ecitocharini species, plus representatives from 10 other tribes. The athetine subtribe Geostibina was recovered as sister group to the ‘true Lomechusini’, which included the type genus Lomechusa. The two clades formed a sister group to the main Athetini clade, which also included Ecitocharini and the ‘false Lomechusini’, a group of New World genera normally placed in Lomechusini. The following changes in classification are proposed: (i) Geostibina Seevers, 1978 is raised to tribal rank, and 13 Athetini genera are placed in Geostibini; (ii) Ecitodonia Seevers, 1965; Ecitopora Wasmann, 1887, and Tetradonia Wasmann, 1894 are moved from Lomechusini to Athetini; (iii) Ecitocharini Seevers, 1965 is placed in synonymy with Athetini; (iv) Discerota Mulsant & Rey, 1874 is tentatively included in Oxypodini; (v) Actocharina Bernhauer, 1907 is placed in synonymy with Hydrosmecta Thomson, 1858.     

A preliminary phylogenetic analysis of the New World Helopini (Coleoptera,Tenebrionidae, Tenebrioninae) indicates the need for profound rearrangements of the classification

Helopini is a diverse tribe in the subfamily Tenebrioninae with a worldwide distribution. The New World helopine species have not been reviewed recently and several doubts emerge regarding their generic assignment as well as the naturalness of the tribe and subordinate taxa. To assess these questions, a preliminary cladistic analysis was conducted with emphasis on sampling the genera distributed in the New World, but including representatives from other regions. The parsimony analysis includes 30 ingroup species from America, Europe and Asia of the subtribes Helopina and Cylindrinotina, plus three outgroups, and 67 morphological characters. Construction of the matrix resulted in the discovery of morphological character states not previously reported for the tribe, particularly from the genitalia of New World species. A consensus of the 12 most parsimonious trees supports the monophyly of the tribe based on a unique combination of characters, including one synapomorphy. None of the subtribes or the genera of the New World represented by more than one species (Helops Fabricius, Nautes Pascoe and Tarpela Bates) were recovered as monophyletic. Helopina was recovered as paraphyletic in relation to Cylindrinotina. One Nearctic species of Helops and one Palearctic species of Tarpela (subtribe Helopina) were more closely related to species of Cylindrinotina. A relatively derived clade, mainly composed by Neotropical species, was found; it includes seven species of Tarpela, seven species of Nautes, and three species of Helops, two Nearctic and one Neotropical. Our results reveal the need to deeply re-evaluate the current classification of the tribe and subordinated taxa, but a broader taxon sampling and further character exploration is needed in order to fully recognize monophyletic groups at different taxonomic levels (from subtribes to genera).     

Phylogeny, biogeography, and host-plant association in the subfamily Apaturinae (Insecta: Lepidoptera: Nymphalidae) inferred from eight nuclear and seven mitochondrial genes

The subfamily Apaturinae consists of 20 genera and shows disjunct distributions and unique host-plant associations. Most genera of this subfamily are distributed in Eurasia South-East Asia and Africa, whereas the genera Doxocopa and Asterocampa are distributed mainly in South America and North America, respectively. Although the Apaturinae larvae mainly feed on the Cannabaceae, those of the genus Apatura are associated with Salix and Populus (Salicaceae), which are distantly related to the Cannabaceae. Here, we infer the phylogeny of Apaturinae and reconstruct the history of host shifting and of colonization in the New World. We analyzed 9761 bp of nuclear and mitochondrial DNA sequence data, including the genes encoding EF1a, Wg, ArgK, CAD, GAPDH, IDH, MDH, RpS5, COI, COII, ATPase8, ATPase6, COIII, ND3, and ND5 for 12 apaturine genera. We also inferred the phylogeny with six additional genera using mitochondrial sequence data alone. Within the Apaturinae, two major clades are recovered in all the datasets. These clades separate the New World genera, Doxocopa and Asterocampa, indicating that dispersal to the New World occurred at least twice. According to our divergence time estimates, these genera originated during the Early Oligocene to the Early Miocene, implying that they migrated across the Bering Land Bridge rather than the Atlantic Land Bridge. The temporal estimates also show that host shifting to Salix or Populus in Apatura occurred more than 15 million years after the divergence of their host plants. Our phylogenetic results are inconsistent with the previously accepted apaturine genus groups and indicate that their higher classification should be reconsidered.     

Synchronous intercontinental splits between assemblages of woodpeckers suggested by molecular data

The woodpeckers (Piciformes: Picinae) comprise a widely distributed and species-rich clade of birds that is strongly associated with trees for feeding, nesting, or both. Because of this association, woodpeckers provide a useful model for evaluating the impact of climatic and tectonic events on the diversification of forest birds during the Tertiary. In order to resolve the biogeographical history of the woodpeckers, we have analysed sequences from two nuclear introns and one mitochondrial gene using likelihood and Bayesian approaches. Our analyses favour a tropical Eurasian origin; divergences between African, Indo-Malayan and New World clades with subsequent colonizations of Africa and the New World occurred synchronously during the Middle Miocene, a period corresponding to the expansion of the C4 grasses and the uplift of the Himalayan-Tibetan plateau. The taxonomic diversification of woodpeckers at this time may be attributed to the fragmentation of forests in response to the drier climate, which in turn prevented gene flow between tropical stocks in Africa, Indo-Malaya and the New World. Our estimates of colonization times of South America predate the closure of the Panama Isthmus and support the hypothesis of a short-lived, terrestrial corridor at the end of the Miocene, 5.7 Myr BP.