Phylogenetic analysis of higher-level relationships of Odonata

Abstract. This is the most comprehensive analysis of higher‐level relationships in Odonata conducted thus far. The analysis was based on a detailed study of the skeletal morphology and wing venation of adults, complemented with a few larval characters, resulting in 122 phylogenetically informative characters. Eighty‐five genera from forty‐five currently recognized families and subfamilies were examined. In most cases, several species were chosen to serve as exemplars for a given genus. The seven fossil outgroup taxa included were exemplar genera from five successively more distant odonatoid orders and suborders: Tarsophlebiidae (the closest sister group of Odonata, previously placed as a family within ‘Anisozygoptera’), Archizygoptera, Protanisoptera, Protodonata and Geroptera. Parsimony analysis of the data, in which characters were treated both under equal weights and implied weighting, produced cladograms that were highly congruent, and in spite of considerable homoplasy in the odonate data, many groupings in the most parsimonious cladograms were well supported in all analyses, as indicated by Bremer support. The analyses supported the monophyly of both Anisoptera and Zygoptera, contrary to the well known hypothesis of zygopteran paraphyly. Within Zygoptera, two large sister clades were indicated, one comprised of the classical (Selysian) Calopterygoidea, except that Amphipterygidae, which have traditionally been placed as a calopterygoid family, nested within the other large zygopteran clade comprised of Fraser's ‘Lestinoidea’ plus ‘Coenagrionoidea’ (both of which were shown to be paraphyletic as currently defined). Philoganga alone appeared as the sister group to the rest of the Zygoptera in unweighted cladograms, whereas Philoganga + Diphlebia comprised the sister group to the remaining Zygoptera in all weighted cladograms. ‘Anisozygoptera’ was confirmed as a paraphyletic assemblage that forms a ‘grade’ towards the true Anisoptera, with Epiophlebia as the most basal taxon. Within Anisoptera, Petaluridae appeared as the sister group to other dragonflies.     

Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae)

Euptychiina is the most species‐rich subtribe of Neotropical Satyrinae, with over 450 known species in 47 genera (14 monotypic). Here, we use morphological characters to examine the phylogenetic relationships within Euptychiina. Taxonomic sampling included 105 species representing the majority of the genera, as well as five outgroups. A total of 103 characters were obtained: 45 from wing pattern, 48 from genitalia and 10 from wing venation. The data matrix was analysed using maximum parsimony under both equal and extended implied weights. Euptychiina was recovered as monophyletic with ten monophyletic genera, contrasting previous DNA sequence‐based phylogenies that did not recover the monophyly of the group. In agreement with sequence‐based hypotheses, however, three main clades were recognized: the ‘Megisto clade’ with six monophyletic and three polyphyletic genera, the ‘Taygetis clade’ with nine genera of which three were monophyletic, and the ‘Pareuptyhia clade’ with four monophyletic and two polyphyletic genera. This is the first morphology‐based phylogenetic hypothesis for Euptychiina and the results will be used to complement molecular data in a combined analysis and to provide critical synapomorphies for clades and genera in this taxonomically confused group.     

The species‐level phylogeny of archostematan beetles—where do Micromalthus debilis and Crowsoniella relicta belong?

A species‐level phylogenetic analysis comprising 37 of the 45 known extant species of archostematan beetles and a total of 110 morphological characters from adults and larvae is presented. For the first time, characters of the male genitalia are included in a phylogenetic analysis of Archostemata. The dataset is analysed with parsimony as well as with Bayesian algorithms. Analyses with differently arranged datasets, with larval characters included or excluded, and including or excluding Micromalthus debilis and Crowsoniella relicta are conducted. The resulting hypothesis of the species‐level phylogeny of Archostemata confirms Cupedidae and Ommatidae as monophyletic taxa. Within Cupedidae, the South American Paracupes and the North American Priacma together are the sister group to all remaining Cupedidae. Among the latter, the identification of a clade comprising Rhipsideigma, Cupes capitatus and Tenomerga leucophaea renders Tenomerga polyphyletic. Ascioplaga scalena comb. nov . is transferred from Adinolepis. Characters of the male genitalia support phylogenetic affinities of Micromalthus debilis with Ommatidae and of Crowsoniella relicta with Cupedidae.     

Origin and diversification of the cryptic ant genus Stenamma Westwood (Hymenoptera: Formicidae), inferred from multilocus molecular data,biogeography and natural history

The genus Stenamma Westwood comprises a group of cryptic, cold tolerant ants that occur throughout the Holarctic and Middle American regions. Traditional approaches to taxonomy and phylogeny are confounded by multiple factors, including the conservative and often convergent morphology of workers and the rarity of reproductive castes in collections. Monophyly of Stenamma and relationships within the genus are uncertain as nearly all previous taxonomic work has been regional in scope. Furthermore, the sister group to Stenamma has not been well established. Here an extensive molecular dataset consisting of ten genes (～8 kb of data), 48 ingroup taxa (20 Nearctic, 6 Palaearctic and 22 Neotropical) and 8 outgroup taxa (6 closely related non‐Stenamma and 2 additional myrmicines) is used to investigate the broad‐scale phylogeny and evolutionary history of Stenamma. Phylogenetic analysis is performed under maximum likelihood and Bayesian frameworks on individual genes and several alternate concatenated datasets, which are used to investigate the effects of inclusion or exclusion of COI and intronic regions. The timing of Stenamma evolution is inferred in beast and ancestral areas are reconstructed using both the s‐diva and DEC methods, as implemented in the programs rasp and lagrange , respectively. Stenamma is revealed as monophyletic with high support and tentatively is sister to a group of New World species placed currently in Aphaenogaster Mayr and Messor Forel. Within Stenamma, two major clades are recovered: a ‘Holarctic clade’ (HOC) and a ‘Middle American clade’ (MAC). The HOC consists of the European S. striatulum Emery sister to two well‐supported groups, the informal ‘debile’ and ‘brevicorne’ clades. The ‘brevicorne’ clade is entirely Nearctic, whereas the ‘debile’ clade includes both Nearctic and Palaearctic representatives. The MAC occurs from the southern United States to northern South America and, with the exception of S. huachucanum Smith, is almost completely isolated geographically from the HOC. It includes a depauperate northern clade and the ‘MAC core’, which is a diverse assemblage of wet forest adapted species found throughout Central America. Divergence dating and biogeographic reconstruction show that Stenamma is most likely to have originated in the Nearctic at the Eocene–Oligocene boundary (～35 Ma) and diversified more rapidly at 16 and 8 Ma for the HOC and MAC, respectively. Potential environmental factors affecting the evolution of Stenamma include the intense global cooling of the late Eocene combined with aridification and mountain building. The phylogenetic results are discussed in relation to the current Stenamma species groups and several new morphological characters are presented to help in identification.     

Phylogeny of Callibaetis (Ephemeroptera: Baetidae) based on morphology of nymphs and adults

Callibaetis is considered to be one of the most problematic genera among mayflies by a series of taxonomic inaccuracies that have accumulated over the last two centuries. Despite these taxonomic problems, two independent hypotheses of species groups have been proposed. In the first hypothesis, three species groups for North America were proposed, and in the second, three species groups were proposed for South American. In these hypotheses, the generic delimitation and monophyly of Callibaetis have not been evaluated under a cladistic framework. Taking this into account, the objectives of this study were to verify the monophyly of Callibaetis and whether the groups of species proposed for the genus are corroborated as natural. The matrix included 128 morphological characters and one habitat character, 119 discrete characters were compiled (101 of nymphs and 18 of adults), and six imaginal characters are related to pigmentation pattern. Continuous characters were ratios and were represented by 10 characters (nine for nymphs and one for adults). The data set was analysed under implied weights. Group support was estimated with relative Bremer support and frequency differences. The results corroborated the monophyletic nature of Callibaetis and the generic status of Callibaetoides; however, the groups proposed for North and South American Callibaetis species were not corroborated. Our study indicated four groups of species for the genus, which we proposed as subgenera: Callibaetis, Abaetetuba subgen. n. Aiso subgen. n. and Cunhaporanga subgen. n.     

Systematics and biogeography of Australian rhopalosomatid wasps (Hymenoptera: Rhopalosomatidae) with a global synopsis of the enigmatic genus Olixon Cameron

Abstract The wasp family Rhopalosomatidae is represented in Australia only by the genus Olixon Cameron. Species of this genus have previously been considered rare based on material in collections and have rarely been observed in the field. All known species of Olixon are brachypterous, solitary ectoparasitoids of crickets (Orthoptera: Gryllidae). Prior to our study the world fauna comprised 11 species of which two were endemic to north‐eastern Australia. Based on specimens collected during recent intensive surveys in Australia we record 17 species for the continent. Fifteen of these species are described as new, which more than doubles the world fauna of Olixon: Olixon abrahami, O. danggari, O. ferrugineum, O. guyim, O. harveyi, O. helgae, O. jandakotae, O. jawoyn, O. jenningsi, O. kakadui, O. pilbara, O. wajuk, O. waldockae, O. wuthathi and O. zonale spp.n . Females of Olixon australiae Perkins and Olixon flavibase Townes are redescribed, and the male of O. flavibase is described for the first time. A key is provided for all 17 Australian species and their distribution is discussed. Results of a cladistic analysis of the world species of Olixon based on 41 morphological characters for 24 ingroup species are presented. The results obtained from equal and implied weighting parsimony analyses indicate that: (i) the Australian species of Olixon are not monophyletic, (ii) about three‐quarters of the Australian species form a monophyletic group, and (iii) a Central/South American Olixon is sister to all other species. The implications of these results for the biogeography of world Olixon species are briefly discussed.     

Beyond the wasp‐waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera)

A comprehensive data set of hymenopteran mesosomal anatomy is presented and analysed. Eighty‐nine taxa, including three outgroups, were scored for 273 characters. Analyses were carried out under different weighting conditions (equal and implied weights). Topologies retrieved for the non‐apocritan Hymenoptera were highly congruent with previously published results. Apocrita were always retrieved as monophyletic, as were most superfamilies. Relationships amongst apocritan superfamilies were mostly weakly corroborated. Stephanoidea were almost always the sister group to the remaining Apocrita. Evaniomorpha were usually retrieved, Ceraphronoidea being the sister group to Megalyroidea, and Evanioidea to Trigonaloidea. Aculeata did not always come out as monophyletic, and of the aculeate superfamilies, only Apoidea was retrieved. Ichneumonoidea were always monophyletic and often the sister group of Aculeata. Maamingidae and Mymarommatoidea were usually sister groups; together, they often form the sister group of Chalcidoidea. A large clade comprising Cynipoidea, Platygastroidea, and Proctotrupoidea was usually retrieved, the two former superfamilies being nested within Proctotrupoidea. Cynipoidea were usually closely related to some of the Diapriidae. Platygastroidea were usually the sister group of a clade comprising Heloridae, Pelecinidae, Proctotrupidae, and Vanhorniidae. The mesosomal region proved to be a very substantial source of phylogenetically relevant information. The results of the present analyses indicate that a reclassification, especially of Proctotrupoidea, is required, but this should be carried out after thorough analyses of more comprehensive combined data sets. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 22–194.     

Phylogenetics and evolutionary morphology of the Neotropical true bug genus Epipolops (Hemiptera: Heteroptera: Geocoridae)

Species of Epipolops Herrich‐Schaeffer (Hemiptera: Geocoridae), comprising the largest genus of Pamphantinae, are among the most bizarre true bugs because of their striking morphology. To elucidate evolutionary morphology in Epipolops, a phylogenetic analysis was performed using 17 species and 36 adult morphological characters. Two cladograms were obtained under equal and implied weight analyses, showing slight differences between them. Two new species, E. stridulatus sp.n . and E. univallensis sp.n ., are described, and E. meridionalis Pirán is resurrected from synonymy with E. frondosus Herrich‐Schaeffer. A key to the known species of Epipolops is provided. The systematic relevance of the unique characters of the genus is discussed and the sequence of character state transformations for both the anterior and posterior lateral processes of the pronotum are optimized on the cladogram obtained under implied weights. Species of Epipolops are found in the Neotropical region and the Mexican Transition Zone, with some clades and species restricted to certain South American subregions. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:CB72D5DA‐D86B‐4B91‐93A2‐88894F7120C9 .     

Phylogeny of western Mediterranean Leptodirini, with an emphasis on genital characters (Coleoptera: Leiodidae: Cholevinae)

Abstract The tribe Leptodirini (Leiodidae: Cholevinae) is one of the largest radiations of Coleoptera in the subterranean environment. Although subjected to systematic and evolutionary studies, the phylogeny remains poorly understood. We assessed the phylogeny of the western Mediterranean lineages (Iberian Peninsula, Pyrenees and Sardinia) based on a cladistic analysis of fourteen characters of external morphology and twenty characters of the male and female genitalia, studied in 182 species belonging to thirty‐nine genera. We tested the monophyly of the traditional two main divisions of the group (infraflagellates and supraflagellates), as well as that of some ‘phyletic series’. The final matrix contained fifty‐eight terminal taxa, twenty‐four of which had different character state combinations. The strict consensus of the sixty most parsimonious trees recovered a monophyletic Leptodirini, but not their separation into infraflagellates and supraflagellates. The supraflagellates formed a paraphyletic group with respect to the infraflagellates (corresponding to our sampled ‘Speonomus’ series), with Notidocharis sister to all other included Leptodirini, and Speonomidius sister to Leptodirini excluding Notidocharis. The series ‘Spelaeochlamys’, including the Sardinian genera but excluding Pseudochlamys, was recovered as monophyletic with weak support. The ‘Quaestus’ series formed a polytomy with Pseudochlamys plus the ‘Speonomus’ series (including Bathysciola), which was recovered as monophyletic with strong support. Speonomus, Bathysciola, Quaestus and Troglophyes were para‐ or polyphyletic. Our results suggested the respective monophyletic origin of the Leptodirini from the Pyrenees (Pseudochlamys plus the ‘Speonomus’ series) and the Mediterranean coast plus Sardinia (series ‘Spelaeochlamys’). On the contrary, the Leptodirini of the Atlantic north coast of the Iberian Peninsula (series ‘Quaestus’ and ‘Speonomidius’) were not monophyletic.     

Evidence from chloroplast DNA restriction site analysis on the relationships of Scalesia (Asteraceae: Heliantheae)

Chloroplast DNA restriction site variation provided data with which to compare the Galápagos Island endemic Scalesia to potential sister groups within subtribe Helianthinae. Pappobolus is suggested by these data to be the most likely sister group to Scalesia. It is an Andean endemic genus that includes the South American species once regarded as a subgenus of Helianthus and later assigned to Helianthopsis. Two other groups considered as potential sister groups based on their geographic distribution in South America were not placed near Scalesia in the most parsimonious tree. Viguiera sect. Diplostichis appears to be relatively basal within subtribe Helianthinae, and the South American species of Viguiera, although previously classified in more than one subgenus, appear to form a single, monophyletic group that is not the sister group to Scalesia. The minimum of ten restriction site differences between Scalesia and Pappobolus of approximately 525 sites surveyed yielded an estimated sequence divergence of 0.19%, and an estimated time of divergence of approximately 1.9–6.2 million years.     

Phylogeny of the death's head hawkmoths, Acherontia[Laspeyres], and related genera (Lepidoptera: Sphingidae: Sphinginae: Acherontiini)

Abstract. Adult death's head hawkmoths (Acherontia species) have a unique feeding biology as cleptoparasites of honeybees, stealing honey from the combs, rather than imbibing nectar from flowers. The moths have a range of features, both morphological and behavioural, that enable them to successfully enter, feed and escape from the colonies. These adaptations may vary among the three Acherontia species and allow them each to target different species of honeybee. A cladistic analysis is presented of the hawkmoths of tribe Acherontiini. The study aims to resolve the relationships of the genera and species of Acherontiini, with a particular focus on the three species of Acherontia. The dataset comprises sixty‐five characters derived from adult, larval and pupal morphology, and larva host‐plant biology. These data are analysed using equal weighting and implied weighting. Acherontiini and each constituent genus are recovered as monophyletic. However, within Coelonia, there is ambiguity in that the sister‐species relationships C. brevis+C. fulvinotata and C. fulvinotata + C. solani are equally parsimonious under both weighting schemes. Furthermore, under equal weighting Agrius is placed equally parsimoniously as the sister group of either Acherontia + Coelonia or Callosphingia. Under implied weighting, however, only the latter relationship is most parsimonious (fit). Within Acherontia, A. atropos and A. styx are always recovered as sister species to the exclusion of A. lachesis. The results of the phylogenetic analysis provide an objective basis for future studies of the unique cleptoparasitic association of these moths.     

Molecular phylogeny of the South American land slug Phyllocaulis (Mollusca,Soleolifera, Veronicellidae)

Gomes, S. R., Britto da Silva, F., Mendes, I. L. V., Thomé, J. W. & Bonatto, S. L. (2009). Molecular phylogeny of the South American land slug Phyllocaulis (Mollusca, Soleolifera, Veronicellidae). —Zoologica Scripta, 39, 177–186. Our main objectives were investigate the phylogenetic relationships between the species of the land slug Phyllocaulis and the monophyly of the genus based on mitochondrial (16S and COI) and nuclear (ITS2) DNA sequences from multiple individuals from each species. Evolutionary trees were constructed using Bayesian inference, maximum likelihood, maximum parsimony and neighbor‐joining methods. All species accepted in the current taxonomy based on penial characteristics and radular measurements were reciprocally monophyletic. Five species out of six formed a clade with the following highly supported relationship: Phyllocaulis gayi, Phyllocaulis soleiformis, Phyllocaulis renschi, Phyllocaulis variegatus, Phyllocaulis boraceiensis. The position of Vaginulus taunaisii changed according to the analysis, appearing as sister‐group of Phyllocaulis or as sister‐group of Phyllocaulis tuberculosus. Divergence times estimated from the 16S tree indicated that the extant species of Phyllocaulis shared a common ancestor around 1.3 Ma and that most species originated between 0.8 and 0.6 Ma.     

Phylogeny,taxonomy, and evolution of Neotropical cichlids (Teleostei: Cichlidae: Cichlinae)

Despite recent progress on the higher‐level relationships of Cichlidae and its Indian, Malagasy, and Greater Antillean components, conflict and uncertainty remain within the species‐rich African, South American, and Middle American assemblages. Herein, we combine morphological and nucleotide characters from the mitochondrial large ribosomal subunit, cytochrome c oxidase subunit I, NADH dehydrogenase four, and cytochrome b genes and from the nuclear histone H3, recombination activating gene two, Tmo‐4C4, Tmo‐M27, and ribosomal S7 loci to analyse relationships within the Neotropical cichlid subfamily Cichlinae. The simultaneous analysis of 6309 characters for 90 terminals, including representatives of all major cichlid lineages and all Neotropical genera, resulted in the first well‐supported and resolved generic‐level phylogeny for Neotropical cichlids. The Neotropical subfamily Cichlinae was recovered as monophyletic and partitioned into seven tribes: Astronotini, Chaetobranchini, Cichlasomatini, Cichlini, Geophagini, Heroini, and Retroculini. Chaetobranchini + Geophagini (including the “crenicichlines”) was resolved as the sister group of Heroini + Cichlasomatini (including Acaronia). The monogeneric Astronotini was recovered as the sister group of these four tribes. Finally, a clade composed of Cichlini + Retroculini was resolved as the sister group to all other cichlines. The analysis included the recently described ?Proterocara argentina, the oldest known cichlid fossil (Eocene), which was placed in an apical position within Geophagini, further supporting a Gondwanan origin for Cichlidae. These phylogenetic results were used as the basis for generating a monophyletic cichline taxonomy. © The Willi Hennig Society 2008.     

Non‐monophyly of Bostrychia simpliciuscula (Ceramiales,Rhodophyta): Multiple species with very similar morphologies,a revised taxonomy of cryptic species

The discovery of a plethora of cryptic species in many algal groups has led to speculation as to the causes of this observation and has affected taxonomy, with reluctance to give names to species that look identical. While this is defensible for monophyletic cryptic species complexes, both our understanding of similar morphologies (crypsis) and nomenclature is challenged when we encounter non‐monophyletic ‘cryptic’ species. Bostrychia simpliciuscula is a wide‐ranging species in which multiple cryptic species are known. Our increased sampling shows that this species consists of four lineages that do not form a clade, but lineages are sister to species with different morphologies. Careful morphological examination shows that characters, especially branched monosiphonous laterals and rhizoid morphology in haptera, are able to distinguish these four lineages into two groups, that are still not monophyletic. The similar morphologies in these lineages could be due to convergence, but not developmental constraints or lack of time to diverge morphologically; or possibly maintenance of a generalized body plan. These lineages appear to have specific biogeographic patterns and these will be used to propose a new taxonomy. B. simpliciuscula is now confined to the tropics. Another of these lineages matches a previously described species, B. tenuissima, that was synonymized with B. simpliciuscula and is from cold temperate Australasia, and is resurrected. Another lineage is found in Japan in which a previous name is also available, B. hamana‐tokidae; the last lineage is found in central New South Wales, morphologically it resembles B. tenuissima, with which it overlaps in distribution around Sydney, and is named as a new species, B. kingii sp. nov.     

The biogeography of Gunnera L.: vicariance and dispersal

Aim The genus Gunnera is distributed in South America, Africa and the Australasian region, a few species reaching Hawaii and southern Mexico in the North. A cladogram was used to (1) discuss the biogeography of Gunnera and (2) subsequently compare this biogeographical pattern with the geological history of continents and the patterns reported for other Southern Hemisphere organisms. Location Africa, northern South America, southern South America, Tasmania, New Zealand, New Guinea/Malaya, Hawaii, North America, Antarctica. Methods A phylogenetic analysis of twenty‐six species of Gunnera combining morphological characters and new as well as published sequences of the ITS region, rbcL and the rps16 intron, was used to interpret the biogeographical patterns in Gunnera. Vicariance was applied in the first place and dispersal was only assumed as a second best explanation. Results The Uruguayan/Brazilian Gunnera herteri Osten (subgenus Ostenigunnera Mattfeld) is sister to the rest of the genus, followed sequentially upwards by the African G. perpensa L. (subgenus Gunnera), in turn sister to all other, American and Australasian, species. These are divided into two clades, one containing American/Hawaiian species, the other containing all Australasian species. Within the Australasian clade, G. macrophylla Blume (subgenus Pseudogunnera Schindler), occurring in New Guinea and Malaya, is sister to a clade including the species from New Zealand and Tasmania (subgenus Milligania Schindler). The southern South American subgenus Misandra Schindler is sister to a clade containing the remaining American, as well as the Hawaiian species (subgenus Panke Schindler). Within subgenus Panke, G. mexicana Brandegee, the only North American species in the genus, is sister to a clade wherein the Hawaiian species are basal to all south and central American taxa. Main conclusions According to the cladogram, South America appears in two places, suggesting an historical explanation for northern South America to be separate from southern South America. Following a well‐known biogeographical pattern of vicariance, Africa is the sister area to the combined southern South America/Australasian clade. Within the Australasian clade, New Zealand is more closely related to New Guinea/Malaya than to southern South America, a pattern found in other plant cladograms, contradictory to some of the patterns supported by animal clades and by the geological hypothesis, respectively. The position of the Tasmanian G. cordifolia, nested within the New Zealand clade indicates dispersal of this species to Tasmania. The position of G. mexicana, the only North American species, as sister to the remaining species of subgenus Panke together with the subsequent sister relation between Hawaii and southern South America, may reflect a North American origin of Panke and a recolonization of South America from the north. This is in agreement with the early North American fossil record of Gunnera and the apparent young age of the South American clade.     

Monophyly of Euaesthetinae (Coleoptera: Staphylinidae): phylogenetic evidence from adults and larvae,review of austral genera,and new larval descriptions

Abstract We develop a morphological dataset for the rove beetle subfamily Euaesthetinae comprising 167 morphological characters (135 adult and 32 larval) scored from 30 terminal taxa including 25 ingroup terminals (from subfamilies Euaesthetinae and Steninae) and five outgroups. Four maximum parsimony analyses using different sets of terminals and character sets were run to test the monophyly of (1) Euaesthetinae, (2) Steninae, (3) Euaesthetinae + Steninae, (4) euaesthetine tribes Austroesthetini, Alzadaesthetini, Euaesthetini, Fenderiini and Stenaesthetini, and (5) the ten currently known austral endemic genera together. Analyses of adult and larval character sets separately and in combination recovered the monophyly of Euaesthetinae, Steninae, and both subfamilies together, with strong support. Analysis of 13 ingroup terminals for which complete data were available suggests that monophyly of Euaesthetinae is supported by 19 synapomorphies (13 adult, six larval), of Steninae by 23 synapomorphies (14 adult, nine larval), and of both subfamilies together by 24 synapomorphies (21 adult, three larval). Within Euaesthetinae, only the tribe Stenaesthetini was recovered as monophyletic based on adult characters, and in no analyses were the ten austral endemic genera recovered as a monophyletic group. Phylogenetic relationships among euaesthetine genera were weakly supported, although analyses including adult characters supported monophyly of Octavius and Protopristus separately, and of Octavius + Protopristus, Austroesthetus + Chilioesthetus and Edaphus + Euaesthetus. Steninae may include a third genus comprising two undescribed species probably possessing a ‘stick–capture’ method of prey capture, similar to that in Stenus. These two species formed a strongly supported clade recovered as the sister group of Stenus based on adult characters. Diagnoses and a key to adults are provided for the 15 euaesthetine genera currently known from the austral region (Australia, New Zealand, South Africa and southern South America). Euaesthetine larvae previously were known only for Euaesthetus, and we describe the larvae of nine more genera and provide the first larval identification key for genera of Euaesthetinae.     

The phylogenetic relationships of Morgan's Sphinx, Xanthopan morganii (Walker), the tribe Acherontiini, and allied long-tongued hawkmoths (Lepidoptera: Sphingidae, Sphinginae)

A cladistic analysis is presented of the hawkmoths of the tribe Acherontiini, Morgan's Sphinx (Xanthopan morganii (Walker)), and related genera. The study aims to test the monophyly of tribe Acherontiini; the hypothesis that all taxa with extremely long probosces (some Acherontiini, Meganoton rubescens, Neococytius, Xanthopan) form a monophyletic group, or at least fall within a single reasonably compact clade; and, within this group, to determine whether Xanthopan is more closely related to Acherontiini or to Cocytius and Neococytius. The data set comprises 109 characters derived from adult and immature stage morphology, biology and behaviour. These data were analysed using equal weighting, successive approximations character weighting (SACW) and implied weighting. All weighting schemes agreed on the monophyly of Acherontiini and of a group of genera comprising Amphimoea, Cocytius and Neococytius (the Cocytius group). Several other generic and suprageneric clades were also consistently recovered. However, those hawkmoths with extremely long probosces were never recovered as a monophyletic group. The relationships of Xanthopan were also ambiguous. Equal weighting and SACW placed Xanthopan + Meganoton rubescens (Butler) as sister to the Cocytius group, while implied weighting placed Xanthopan as sister to Acherontiini. This latter relationship is based primarily on shared possession of a pilifer/palp hearing organ. Further analyses suggested the two components of this organ were not biologically independent. Downweighting this feature accordingly resulted in all weighting schemes converging on the topology found by equal weighting. Exclusion of the incomplete subset of immature stage data had no effect under implied weighting but equal weighting and SACW now recovered a Neotropical clade comprising Manduca and the Cocytius group, while Xanthopan was placed with M. rubescens and Panogena. Downweighting the pilifer/palp hearing organ under implied weighting again caused convergence with the equal weighting/SACW results. Thus, the relationships of Xanthopan remain equivocal and further data, particularly from the immature stages, will be required to elucidate its phylogenetic position further. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society, 2002, 135 , 471–527.     

A new Veturius (Veturius) of the “cephalotes” group from southern Colombia,Putumayo (Coleoptera: Passalidae)

A new species of Veturius (Veturius) Kaup from southern Colombia, Putumayo, lowland forests, is described and illustrated. V. paya n. sp. belongs to the South American “cephalotes” species group and is sister species of V. cephalotes (Le Peletier & Serville, 1825). Both species are distinguished by few characters of the head and thorax. They seem to be allopatric close to the western distributional limits of V. cephalotes. The endemism of V. paya n. sp. corresponds to a geographic vicariance in the axis of the Amazon Basin. An overview is given on the phylogeny and the chorology of the completed “cephalotes” species group.     

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.