Mandibulate chironomids: primitive or derived? (Diptera: Chironomidae)

Abstract Mandibulate functional mouthparts are reported in males and females of the two Early Cretaceous Chironomidae (Diptera): Wadelius libanicus Veltz et al., 2007 (in Tanypodinae) and Libanochlites Brundin, 1976 (transferred from the Podonominae to the Tanypodinae). Females of Haematotanypus libanicus gen.n. et sp.n. (subfamily Tanypodinae) have mandibulate mouthparts. Although currently considered as plesiomorphic structures, the presence of such mandibulate mouthparts in these Tanypodinae and in the recent Podonominae genera Archaeochlus and Austrochlus could correspond to reversals, based on a parsimony argument after the current chironomid phylogeny. On the contrary, similar mandibulate mouthparts probably are plesiomorphic in the Early Cretaceous Cretaenne kobeyssii gen.n. et sp.n. and Cretaenne inexpectata sp.n. (Aenneinae or stem group of recent Chironomidae).     

A dated molecular phylogeny for the Chironomidae (Diptera)

We provide the first highly sampled phylogeny estimate for the dipteran family Chironomidae using molecular data from fragments of two ribosomal genes (18S and 28S), one nuclear protein‐coding gene (CAD), and one mitochondrial protein‐coding gene (COI), analysed using mixed‐model Bayesian and maximum likelihood inference methods. The most recently described subfamilies Chilenomyiinae and Usambaromyiinae proved elusive, and are unsampled. We confirm monophyly of all sampled subfamilies except Prodiamesinae, which contains Propsilocerus Kieffer, previously in Orthocladiinae. The semifamily Chironomoinae is confirmed only if Telmatogetoninae is included, which is closer to Brundin's original suggestion. Buchonomyiinae is excluded from Chironomoinae: it is a sister group to all remaining Chironomidae, conforming more to Murray and Ashe's argumentation. Semifamily Tanypodoinae is a grade and unsupported as monophyletic: the austral Aphroteniinae alone is sister to all Chironomidae (less Buchonomyiinae). Podonominae is weakly supported as the next sister group, in contrast to some estimates that place this subfamily as sister group to Tanypodinae alone. In Diamesinae, the southern African Harrisonini is confirmed as a member, but embedded within austral tribe Heptagiini, which is confirmed as sister to the undersampled Diamesini. Tribe Pentaneurini and ‘non‐Pentaneurini’ taxa are reciprocally monophyletic in Tanypodinae. Recent molecular findings concerning Podonominae are substantiated, with a monophyletic tribe Podonomini, Boreochlini forming a grade and Lasiodiamesa Kieffer placed as sister to all other Podonominae, but with uncertainty. In Orthocladiinae, a postulated two‐tribe system of Orthocladiini and Metriocnemini can be supported after exclusion of a Corynoneura group and a Brillia group, which is revealed as sister to Stictocladius Edwards. The marine Clunio Haliday and Thalassosmittia Strenzke & Remmert (given high rank in the past) are clearly embedded deep in Orthocladiinae. The finding of Shangomyia Sæther & Wang + Xyiaomyia Sæther & Wang as sister group to all other Chironominae justifies high rank, as their authors suggested. Pseudochironomini (untested by sampling shortfall) is sister to a monophyletic Tanytarsini (with a weakly supported inclusion of the enigmatic Nandeva Wiedenbrug, Reiss & Fittkau). The tribe Chironomini can be supported only by excluding Shangomyia + Xyiaomyia, and a postulated monophyletic clade comprising several taxa such as Microtendipes Kieffer, with six‐segmented larval antennae and alternate Lauterborn organs, that is sister group to Pseudochironomini + Tanytarsini. The tempo of diversification of the family, deduced by divergence time analysis (beast ), shows Permian origination with subfamily stem‐group origination from the mid–late Triassic to the early Cretaceous. Crown‐group origination ranged from Podonominae on a short stem originating in the mid Jurassic to long‐stemmed Aphroteninae from the late Cretaceous. Node dates allow inference of some vicariance via Gondwanan fragmentation, including certain nodes involving southern Africa.     

Afrochlus Freeman: an African gondwanan midge and the phylogeny of the Podonominae (Diptera: Chironomidae)

The immature stages and male adult of Afrochlus harrisoni Freeman are described for the first time, from the type locality, a granite outcrop in Zimbabwe. A phylogenetic position within the subfamily Podonominae as sister to South African-western/central Australian genus Archaeochlus Brundin, suggested in 1966 by Brundin from the adult female alone, is confirmed by cladistic analysis of all life history stages combined. Within the Podonominae, the tribes Boreochlini and Podonomini erected by Brundin are monophyletic only if Trichotanypus Kieffer is transferred from the former to the latter. Little or none of the structure provided by the combined evidence from all stages is retrieved by cladistic analyses of characters derived solely from the adult stage. The implications of erection of monotypic genera for adult-only taxa are discussed.     

A preliminary survey of the Chironomidae (Diptera) of Costa Rica,with emphasis on the lotic fauna

Extensive collections of Chironomidae were made in Costa Rica, Central America, during 1986 and 1987. Fifty-five genera and at least 148 species belonging to the subfamilies Podonominae, Tanypodinae, Orthocladiinae and Chironomidae were found. Chironominae and Orthocladiinae predominated. Only one species of Podonominae was collected. Tanypodinae was represented by many genera, but species richness was low.Cricotopus was the most widespread and diverse genus of Orthocladiinae. Among the Chironominae the generaPolypedilum, Pseudochironomus, Tanytarsus andRheotanytarsus showed high species richness. Several species were collected that could not be assigned to genus. A number of range extensions were recorded for taxa found in the Neotropical region for the first time and for Neotropical taxa recorded outside of South America for the first time. The Costa Rican chironomid fauna consists of cosmopolitan, holarctic and neotropical components. There is probably an endemic Central American chironomid fauna at the species level.     

Phylogenetic relationships of nonbiting midges in the subfamily Tanypodinae (Diptera: Chironomidae) inferred from morphology

The nonbiting midge subfamily Tanypodinae represents one of the most diverse lineages of Chironomidae. Despite the wide distribution and high diversity of tanypodine chironomids, the evolutionary history of the subfamily remains poorly understood. Here, we present the first phylogenetic analysis of the subfamily Tanypodinae based on morphological data. Cladistic analyses were conducted using 86 morphological characters from 115 species belonging to 54 tanypodine genera, including the eight currently recognised tribes: Anatopyniini, Clinotanypodini, Coelopyniini, Macropelopiini, Natarsiini, Pentaneurini, Procladiini and Tanypodini. We use characters from fourth‐instar larvae, pupae and adults of both sexes. We examine the effects of implied weighting by reanalysing the data with varying values of concavity constant (k). Our analysis supports the monophyly of Tanypodinae with Podonominae as its sister group. All previously proposed tribes are recovered as monophyletic assemblages under a wide range of weighting factors. Under these conditions, the genus Fittkauimyia is the sister group of the remaining Macropelopiini and is erected as a new monobasic tribe, Fittkauimyiini trib.n . The tribe Pentaneurini is recovered as monophyletic with some internal relationships resolved. The genus Paramerina, recovered as sister of Reomyia + Zavrelimyia, is formally synonymised with Zavrelimyia syn.n. , based on morphological similarity in all three life stages and treated as a subgenus of the latter. Finally, the recently suggested synonymies of Gressittius and Guassutanypus with Alotanypus and the establishment of the subgenera Conchapelopia (Helopelopia), Macropelopia (Bethbilbeckia), Monopelopia (Cantopelopia), Thienemannimyia (Hayesomyia) and Zavrelimyia (Reomyia and Schineriella) are investigated. Our results support all proposed changes, except for the subgenus‐level status of Helopelopia and Cantopelopia. We suggest re‐establishment of Helopelopia as a genus, but refrain from promoting genus‐level status of Cantopelopia at present because the apparent sister‐relationship between Monopelopia + Nilotanypus likely is due to wing vein reduction caused by miniaturisation. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:DF012C17‐AFB3‐4904‐83DC‐30DD94D0B376 .     

Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules

Aim The sequential break‐up of Gondwana is thought to be a dominant process in the establishment of shared biota across landmasses of the Southern Hemisphere. Yet similar distributions are shared by taxa whose radiations clearly post‐date the Gondwanan break‐up. Thus, determining the contribution of vicariance versus dispersal to seemingly Gondwanan biota is complex. The southern freshwater crayfishes (family Parastacidae) are distributed on Australia and New Guinea, South America, Madagascar and New Zealand and are unlikely to have dispersed via oceans, owing to strict freshwater limitations. We test the hypotheses that the break‐up of Gondwana has led to (1) a predominately east–west (((Australia, New Zealand: 80 Ma) Madagascar: 160–121 Ma) South America: 165–140 Ma), or (2) a southern (((Australia, South America: 52–35 Ma) New Zealand: 80 Ma) Madagascar: 160–121 Ma) pattern for parastacid crayfish. Further, we examine the evidence for a complete drowning of New Zealand and subsequent colonization by freshwater crayfish. Location Southern Hemisphere. Methods The evolutionary relationships among the 15 genera of Parastacidae were reconstructed using mitochondrial [16S, cytochrome c oxidase subunit I (COI)] and nuclear (18S, 28S) sequence data and maximum likelihood and Bayesian methods of phylogenetic reconstruction. A Bayesian (multidivtime ) molecular dating method using six fossil calibrations and phylogenetic inference was used to estimate divergence time among crayfish clades on Gondwanan landmasses. Results The South American crayfish are monophyletic and a sister group to all other southern crayfish. Australian crayfish are not monophyletic, with two Tasmanian genera, Spinastacoides and Ombrastacoides, forming a clade with New Zealand and Malagasy crayfish (both monophyletic). Divergence of crayfish among southern landmasses is estimated to have occurred around the Late Jurassic to Early Cretaceous (109–178 Ma). Main conclusions The estimated phylogenetic relationships and time of divergence among the Southern Hemisphere crayfishes were consistent with an east–west pattern of Gondwanan divergence. The divergence between Australia and New Zealand (109–160 Ma) pre‐dated the rifting at around 80 Ma, suggesting that these lineages were established prior to the break‐up. Owing to the age of the New Zealand crayfish, we reject the hypothesis that there was a complete drowning of New Zealand crayfish habitat.     

The 'Ice Worm' - the Immature Stages,Phylogeny and Biology of the Glacier Midge Zelandochlus (Diptera: Chironomidae)

The previously unknown female and larva for the New Zealand glacier midge, Zealandochlus latipalpis Brundin are described for the first time, and the pupa described more fully than previously. Unlike the male, which is brachypterous, the female has large wings possessing traces of a vein between R₁ and R₄₊₅ uniquely for the subfamily Podonominae. The larvae, known locally as ice-worms, live in meltwater pools and ice caves of the Franz Joseph and Fox glaciers, New Zealand. Cladistic analysis of this highly autapomorphic species results in an unresolved trichotomy Zelandochlus + Parochlus + Podonomus, which is no advance on the suggestion made by Brundin (1966) concerning the relationships. Information is too scanty on many species of the putative related genera, but additional morphological features indicate that there may be a sister group relationship with part of Parochlus.     

Tri-locus sequence data reject a “Gondwanan origin hypothesis” for the African/South Pacific crab genus Hymenosoma

Crabs of the family Hymenosomatidae are common in coastal and shelf regions throughout much of the southern hemisphere. One of the genera in the family, Hymenosoma, is represented in Africa and the South Pacific (Australia and New Zealand). This distribution can be explained either by vicariance (presence of the genus on the Gondwanan supercontinent and divergence following its break-up) or more recent transoceanic dispersal from one region to the other. We tested these hypotheses by reconstructing phylogenetic relationships among the seven presently-accepted species in the genus, as well as examining their placement among other hymenosomatid crabs, using sequence data from two nuclear markers (Adenine Nucleotide Transporter [ANT] exon 2 and 18S rDNA) and three mitochondrial markers (COI, 12S and 16S rDNA). The five southern African representatives of the genus were recovered as a monophyletic lineage, and another southern African species, Neorhynchoplax bovis, was identified as their sister taxon. The two species of Hymenosoma from the South Pacific neither clustered with their African congeners, nor with each other, and should therefore both be placed into different genera. Molecular dating supports a post-Gondwanan origin of the Hymenosomatidae. While long-distance dispersal cannot be ruled out to explain the presence of the family Hymenosomatidae on the former Gondwanan land-masses and beyond, the evolutionary history of the African species of Hymenosoma indicates that a third means of speciation may be important in this group: gradual along-coast dispersal from tropical towards temperate regions, with range expansions into formerly inhospitable habitat during warm climatic phases, followed by adaptation and speciation during subsequent cooler phases.     

A new genus of mantidflies discovered in the Oriental region,with a higher‐level phylogeny of Mantispidae (Neuroptera) using DNA sequences and morphology

A remarkable new genus and two new species of Mantispidae (Neuroptera) are described from the Oriental region. Allomantispa Liu, Wu, Winterton & Ohl gen.n. , currently including A. tibetana Liu, Wu & Winterton sp.n. and A. mirimaculata Liu & Ohl sp.n. The new genus is placed in the subfamily Drepanicinae based on a series of morphological characteristics and on the results of total evidence phylogenetic analyses. Bayesian and Parsimony analyses were undertaken using three gene loci (CAD, 16S rDNA and COI) combined with 74 morphological characters from living and fossil exemplars of Mantispidae (17 genera), Rhachiberothidae (two genera) and Berothidae (five genera), with outgroup taxa from Dilaridae and Osmylidae. The resultant phylogeny presented here recovered a monophyletic Mantispidae with ?Mesomantispinae sister to the rest of the family. Relationships among Mantispidae, Rhachiberothidae and Berothidae support Rhachiberothidae as a separate family sister to Mantispidae. Within Mantispidae, Drepanicinae are a monophyletic clade sister to Calomantispinae and Mantispinae. In a combined analysis, Allomantispa gen.n. was recovered in a clade comprising Ditaxis McLachlan from Australia, and two fossil genera from the Palaearctic, ?Promantispa Panfilov (Kazakhstan; late Jurassic) and ?Liassochrysa Ansorge & Schlüter (Germany; Jurassic), suggesting a highly disjunct and relictual distribution for the family. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:464B06E8‐47E6‐482E‐8136‐83FE3B2E9D6B .     

Phylogeny, character evolution, and classification of Sapotaceae (Ericales)

We present the first cladistic study of the largely tropical family Sapotaceae based on both morphological and molecular data. The data were analyzed with standard parsimony and parsimony jackknife algorithms using equally and successive weighted characters. Sapotaceae are confirmed to constitute two main evolutionary lineages corresponding to the tribes Isonandreae‐Mimusopeae‐Sideroxyleae and Chrysophylleae‐Omphalocarpeae. The Sideroxyleae are monophyletic, Isonandreae are polyphyletic as presently circumscribed, and as suggested by the analyses, the subtribe Mimusopeae‐Mimusopinae has evolved within the Mimusopeae‐Manilkarinae, which hence is also paraphyletic. Generic limits must be altered within Sideroxyleae with the current members Argania, Nesoluma and Sideroxylon. Argania cannot be maintained at a generic level unless a narrower generic concept is adopted for Sideroxylon. Nesoluma cannot be upheld in a narrow or broad generic concept of Sideroxylon. The large tribe Chrysophylleae circumscribes genera such as Chrysophyllum, Pouteria, Synsepalum, and Xantolis, but the tribe is monophyletic only if the taxa from Omphalocarpeae are also included. Neither Chrysophyllum nor Pouteria are monophyletic in their current definitions. The results indicate that the African taxa of Pouteria are monophyletic and distinguishable from the South American taxa. Resurrection of Planchonella, corresponding to Pouteria section Oligotheca, is proposed. The African genera Synsepalum and Englerophytum form a monophyletic group, but their generic limits are uncertain. Classification of the Asian genus Xantolis is particularly interesting. Morphology alone is indecisive regarding Xantolis relationships, the combined unweighted data of molecules and morphology indicates a sister position to Isonandreae‐Mimusopeae‐Sideroxyleae, whereas molecular data alone, as well as successive weighted combined data point to a sister position to Chrysophylleae‐Omphalocarpeae. An amended subfamily classification is proposed corresponding to the monophyletic groups: Sarcospermatoideae (Sarcosperma), Sapotoideae (Isonandreae‐Mimusopeae‐Sideroxyleae) and Chrysophylloideae (Chrysophylleae‐Omphalocarpeae), where Sapotoideae circumscribes the tribes Sapoteae and Sideroxyleae as well as two or three as yet unnamed lineages. Morphological characters are often highly homoplasious and unambiguous synapomorphies cannot be identified for subfamilies or tribes, which we believe are the reason for the variations seen between different classifications of Sapotaceae. © The Willi Hennig Society 2005.     

Phylogenetic analysis of the New World Ptininae (Coleoptera: Bostrichoidea)

A phylogenetic analysis of the New World Ptininae (Anobiidae) was conducted with representatives of nine of ten New World genera, several Old World genera and seven more of the ten subfamilies of Anobiidae. One hundred and two characters (forty‐three multistate) from thirty‐four taxa were used. The single cladogram shows Ptininae as monophyletic and the sister group of the remaining Anobiidae, supporting their placement as subfamilies of a monophyletic Bostrichidae. Genus Niptus Boieldieu is polyphyletic supporting recognition of Pseudeurostus Heyden and the creation of a new genus to encompass the remaining New World species of Niptus. Flightlessness has evolved a minimum of three times within Ptininae and myrmecophily has probably evolved three times within just the New World taxa. The classifications of Ptininae and the remaining Anobiidae are examined and the evolution of feeding habits, myrmecophily and wing loss are discussed.     

Molecules,morphology and minute hooded beetles: a phylogenetic study with implications for the evolution and classification of Corylophidae (Coleoptera: Cucujoidea)

Eight genes (nuclear: 18S, 28S, H3, CAD; mitochondrial: 12S, 16S, COI, COII) and morphology were used to infer the evolutionary history of Corylophidae, some of the smallest free‐living insects. The study included 36 corylophid exemplars, representing approximately 60% of the known generic diversity of the family and 16 cucujoid outgroup taxa. Multiple partitioning strategies, molecular datasets, combined datasets and different taxon sampling regimes using maximum likelihood and mixed‐model Bayesian inference were utilized to analyse these data. Most results were highly concordant across analyses. There was strong agreement across (i) partitioning strategies, (ii) maximum likelihood and Bayesian inference analyses of the molecular data, and (iii) Bayesian inference of the molecular data alone and Bayesian inference of the combined morphological and molecular data when all terminal taxa were included. When a strict taxon sampling protocol was employed so that only single generic exemplars were included, deep relationships were affected in the resulting phylogenetic hypotheses. Under such narrow sampling strategies, deep phylogenetic relationships were also sensitive to the choice of generic exemplars. Although it is often challenging to obtain single representatives for many taxa in higher‐level phylogenetic analyses, these results indicate the importance of using denser taxon sampling approaches even at the specific level for genera included in such studies. Molecular data alone support Anamorphinae (Endomychidae) strongly as the sister group of Corylophidae. In combined data analyses, Coccinellidae is recovered as the sister group to Corylophidae. In all analyses, Corylophidae and the subfamily Corylophinae are recovered as monophyletic. The monophyly of Periptyctinae was untested, as only a single species was included. All included corylophine tribes were recovered as monophyletic with the exception of Aenigmaticini; Aenigmaticum Matthews forms the sister group to Orthoperus Stephens and Stanus?lipiński et al. is recovered as the sister group of Sericoderus Stephens. Stanus tasmanicus?lipiński et al. is transferred to a new genus, Pseudostanus Robertson, ?lipiński & McHugh gen.n. incertae sedis. We propose a new tribe, Stanini Robertson, ?lipiński & McHugh trib.n. for Stanus bowesteadi?lipiński et al. and a new concept of Aenigmaticini sensu.n. to include only the nominate genus. Anatomical transitions associated with corylophid miniaturization are highlighted. Key phenotypic modifications and elevated rates of substitution in nuclear rRNA genes are evident in a subgroup of Corylophinae that includes the most diminutive species. Other taxonomic and evolutionary implications are discussed in light of the results.     

Molecular phylogeny of Nitidulidae: assessment of subfamilial and tribal classification and formalization of the family Cybocephalidae (Coleoptera: Cucujoidea)

We present a molecular phylogeny of Nitidulidae based on thirty ingroup taxa representing eight of the ten currently recognized subfamilies. Approximately 10 K base pairs from seven loci (12S, 16S, 18S, 28S, COI, COII and H3) were used for the phylogenetic reconstruction. The phylogeny supports the following main conclusions: (i) Cybocephalidae are formally recognized as a distinct family not closely related to Nitidulidae and its constituent taxa are defined; (ii) Kateretidae are sister to Nitidulidae; (iii) Cryptarchinae are monophyletic and sister to the remaining nitidulid subfamilies; (iv) subfamily Prometopinae stat. res. is reinstated and defined, to accommodate taxa allied to Axyra Erichson, Prometopia Erichson and Megauchenia MacLeay; (v) Amphicrossinae, Carpophilinae and Epuraeinae are shown to be closely related taxa within a well‐supported monophyletic clade; (vi) tribal affinities and respective monophyly within Nitidulinae are poorly resolved by our data and must be more rigorously tested as there was little or no support for prior morphologically based tribes or genus‐level complexes; (vii) Nitidulinae are found to be paraphyletic with respect to Cillaeinae and Meligethinae, suggesting that they should either be subsumed as tribes, or Nitidulinae should be divided into several subfamilies to preserve the status of Cillaeinae and Meligethinae; (viii) Teichostethus Sharp stat. res. is not a synonym of Hebascus Erichson and the former is reinstated as a valid genus. These conclusions and emendations are discussed in detail and presented within a morphological framework.     

Molecular phylogeny of telenomine egg parasitoids (Hymenoptera: Platygastridae s.l.: Telenominae): evolution of host shifts and implications for classification

Parasitoid wasps of the subfamily Telenominae (Hymenoptera: Platygastroidea, Platygastridae) develop as immatures within the eggs of other insects (Lepidoptera, Hemiptera, Diptera and Neuroptera). Rearing records indicate that individual species are restricted to attack hosts within only one of these four main groups. We conducted a phylogenetic analysis of the group using sequence data from multiple genes (18S, 28S, COI, EF‐1α) to assess the pattern of shifts among host groups and to test the monophyly of and relationships among genera and species‐groups. Telenominae sensu Masner—that is, including only the nominate tribe Telenomini—is not monophyletic. Representatives of the Psix group of genera (Psix Kozlov & Lê and Paratelenomus Dodd) form a monophyletic group that is sister to Gryon Haliday (Scelioninae: Gryonini) and are excluded from the subfamily. The remaining telenomines are monophyletic. The genus Phanuromyia Dodd and the crassiclava group of Telenomus Haliday, both recorded as parasitoids of planthopper eggs (Hemiptera: Auchenorrhyncha, Fulgoroidea), form a monophyletic group that is sister to all other telenomines exclusive of the Psix group. Twenty‐nine species of the crassiclava and aradi groups of Telenomus are transferred to Phanuromyia as new combinations. Basal elements of the remaining species are all in groups reared from the eggs of true bugs (Heteroptera), primarily the stink bugs (Pentatomoidea) and seed bugs (Lygaeoidea). A shift to parasitism of lepidopteran eggs evolved within a single clade, occurring either one or two times. From this clade a small group of species, the Telenomus tabanivorus group, subsequently shifted to parasitism of egg masses of true flies (Tabanidae and Stratiomyiidae). Aholcus Kieffer and Platytelenomus Dodd both belong to the clade of lepidopteran parasitoids and are considered as junior synonyms of Telenomus (new synonymy for Aholcus). The monophyletic status of the two core genera, Telenomus and Trissolcus could not be resolved using these data. The phylogenetic pattern of host shifts suggests comparisons among taxa that may be fruitful in elucidating mechanisms by which parasitoids locate their hosts, the proximate factors that determine the host range, and the changes in these factors that influence host changes.     

Molecular phylogeny of flat‐footed flies (Diptera: Platypezidae): main clades supported by new morphological evidence

The molecular phylogeny of flat‐footed flies is inferred from analysis of DNA sequence data from the five mitochondrial genes 12S, 16S, COI, COII and CytB, and the nuclear gene 28S and discussed with the recent systematics based on morphological features. The Bayesian inference, maximum likelihood and maximum parsimony analyses included 42 species of 18 genera, representing all four extant subfamilies (Microsaniinae, Melanderomyiinae, Callomyiinae and Platypezinae) and all known genera except one (Metaclythia). Representatives of the brachycerous taxa Lonchopteridae, Phoridae, Sciadocerinae (Phoridae) and Opetiidae are used as outgroups, and Lonchoptera was used to root the trees. Our results show Platypezidae consisting of two well‐supported clades, the first with the subfamilies Melanderomyiinae + Callomyiinae and the second formed by subfamily Platypezinae. Genus Microsania was resolved as a separate lineage distant from Platypezidae which clustered with Opetiidae as its sister group, both together forming a sister group to Platypezidae. At the generic level, the genus Agathomyia proved not to be monophyletic in any of the analyses. The species Chydaeopeza tibialis is sister to Agathomyia sexmaculata, and consequently, the genus Chydaeopeza Shatalkin, 1992 is a new junior synonym of Agathomyia Verrall, 1901. Bifurcated setae on legs of adult Platypezidae are documented as a new synapomorphy of the family, exclusive of Microsania. Outstretched wings and only a small overlap of their surfaces at resting position are considered a new synapomorphy for the subfamily Platypezinae. Other phylogenetically important characters defining main clades are documented, and their relevance/validity in phylogenetic studies is discussed. The current systematic concept of Platypezidae is discussed, and new phylogenetic hypotheses are proposed.     

Orthoglymma wangapeka gen.n., sp.n. (Coleoptera: Carabidae: Broscini): a newly discovered relict from the Buller Terrane,north‐western South Island,New Zealand,corroborates a general pattern of Gondwanan endemism

Orthoglymma Liebherr, Marris, Emberson, Syrett & Roig‐Juñent gen.n. (Coleoptera: Carabidae: Broscini) is described to accommodate the single type species Orthoglymma wangapeka Liebherr, Marris, Emberson, Syrett & Roig‐Juñent sp.n., known from the Wangapeka Track, Kahurangi National Park, north‐western South Island, New Zealand. Orthoglymma wangapeka sp.n. is analysed cladistically along with a comprehensive array of 42 other broscine generic terminals and four out‐group taxa, using information obtained from 73 morphological characters, and placed as adelphotaxon to the remainder of subtribe Nothobroscina, a clade distributed in New Zealand, southern South America and Australia. Based on fossil evidence for Carabidae, the occurrence of Orthoglymma wangapeka sp.n. on the Buller Terrane, a geological feature once situated on the eastern margin of Gondwana, and early cladistic divergence of Orthoglymma from the remaining Nothobroscina, Orthoglymma wangapeka sp.n. is interpreted as a Gondwanan relict. The New Zealand arthropod fauna is reviewed to identify other taxa in existence at the time of Cretaceous vicariance of New Zealand and Australia. These candidate Gondwanan taxa, all of which are specified using fossil data or molecular divergence‐based estimates, are analysed biogeographically. Where phylogenetic hypotheses are available, primordial distributions are optimized using event‐based, dispersal‐vicariance (DIVA) analysis. The hypothesized Gondwanan‐aged taxa demonstrate inordinate fidelity to the Gondwanan‐aged geological terranes that constitute the western portions of New Zealand, especially in the South Island. Persistence of these relicts through a hypothesized ‘Oligocene drowning’ event is the most parsimonious explanation for the concentration of Gondwanan relicts in the Nelson, Buller and Fiordland districts of the South Island. Geographic patterns of Gondwanan‐aged taxa are compared with distributions of taxa hypothesized to have colonized New Zealand across the Tasman Sea from Australia and New Caledonia, subsequent to Cretaceous vicariance. These post‐Gondwanan taxa exhibit very different patterns of distribution and diversification in New Zealand, including: (i) abundant endemism in Northland, and the islands and peninsulas of the North Island; (ii) species geographically restricted to areas underlain by the youngest Rakaia and Pahau geological terranes; and (iii) species exhibiting exceedingly widespread geographic distributions spanning geological terranes of disparate ages.

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When phylogenetics met biogeography: Willi Hennig,Lars Brundin and the roots of phylogenetic and cladistic biogeography

Willi Hennig's (Beitr. Ent. 1960, 10, 15) Die Dipteren-Fauna von Neuseeland als systematisches und tiergeographisches Problem applied a phylogenetic approach to examine the distributional patterns exhibited by the Diptera of New Zealand. Hennig showed how phylogenetic trees may be used to infer dispersal, based on the progression and deviation rules, and also discussed the existence of vicariance patterns. The most important author who applied Hennig's phylogenetic biogeography was Lars Brundin, when analysing the phylogenetic relationships of two taxa of Chironomidae (Diptera) and using them to examine the biogeographic relationships of Australia, New Zealand, South America and South Africa. The relevance of Brundin's contribution was noted by several authors, as it began the cladistic or vicariance approach to biogeography, that implies the discovery of vicariance events shared by different monophyletic groups. Both phylogenetic and cladistic biogeography have a place in contemporary biogeography, the former for analysing taxon biogeography and the latter when addressing Earth or biota biogeography. The recent use of the term “phylogenetic biogeography” to refer to a posteriori methods of cladistic biogeography is erroneous and should be avoided.     

The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae)

Abstract. To reconstruct the phylogeny of the Sericini and their systematic position among the scarabaeid beetles, cladistic analyses were performed using 107 morphological characters from the adults and larvae of forty‐nine extant scarabaeid genera. Taxa represent most ‘traditional’ subfamilies of coprophagous and phytophagous Scarabaeidae, with emphasis on the Sericini and other melolonthine lineages. Several poorly studied exoskeletal features have been examined, including the elytral base, posterior wing venation, mouth parts, endosternites, coxal articulation, and genitalia. The results of the analysis strongly support the monophyly of the ‘orphnine group’ + ‘melolonthine group’ including phytophagous scarabs such as Dynastinae, Hopliinae, Melolonthinae, Rutelinae, and Cetoniinae. This clade was identified as the sister group to the ‘dung beetle line’ represented by Aphodius + Copris. The ‘melolonthine group’ is comprised in the strict consensus tree by two major clades and two minor lineages, with the included taxa of Euchirinae, Rutelinae, and Dynastinae nested together in one of the major clades (‘melolonthine group I’). Melolonthini, Cetoniinae, and Rutelinae are strongly supported, whereas Melolonthinae and Pachydemini appear to be paraphyletic. Sericini + Ablaberini were identified to be sister taxa nested within the second major melolonthine clade (‘melolonthine group II’). As this clade is distributed primarily in the southern continents, one could assume that Sericini + Ablaberini are derived from a southern lineage. Plausibly, ancestors of Sericini + Ablaberini and Athlia were separated by a vicariance event, such as the separation of the African plate from the rest of Gondwana, whereas Sericini and Ablaberini probably diversified during the early Tertiary, with dispersal of some basal Sericini to South America.     

PHYLOGENY OF THE EUGLENOID LORICATE GENERA TRACHELOMONAS AND STROMBOMONAS (EUGLENOPHYTA) INFERRED FROM NUCLEAR SSU AND LSU rDNA1

Previous studies using the nuclear SSU rDNA and partial LSU rDNA have demonstrated that the euglenoid loricate taxa form a monophyletic clade within the photosynthetic euglenoid lineage. It was unclear, however, whether the loricate genera Trachelomonas and Strombomonas were monophyletic. In order to determine the relationships among the loricate taxa, SSU and LSU nuclear rDNA sequences were obtained for eight Strombomonas and 25 Trachelomonas strains and combined in a multigene phylogenetic analysis. Conserved regions of the aligned data set were used to generate maximum‐likelihood (ML) and Bayesian phylogenies. Both methods recovered a strongly supported monophyletic loricate clade with Strombomonas and Trachelomonas species separated into two sister clades. Taxa in the genus Strombomonas sorted into three subclades. Within the genus Trachelomonas, five strongly supported subclades were recovered in all analyses. Key morphological features could be attributed to each of the subclades, with the major separation being that all of the spine‐bearing taxa were located in two sister subclades, while the more rounded, spineless taxa formed the remaining three subclades. The separation of genera and subclades was supported by 42 distinct molecular signatures (33 in Trachelomonas and nine in Strombomonas). The morphological and molecular data supported the retention of Trachelomonas and Strombomonas as separate loricate genera.