Showing papers in "Systematic Entomology in 2004"
TL;DR: The ancestral predatory condition as generalist hunting is reconstructed, with three independent shifts to cursorial hunting and one shift to ambush hunting, associated with the largest radiation of mantid species.
Abstract: . The previously unknown phylogenetic relationships among Mantodea (praying mantids) were inferred from DNA sequence data. Five genes (16S rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase II and histone 3) were sequenced for sixty-three taxa representing major mantid lineages and outgroups. The monophyly of mantid families and subfamilies was tested under varying parameter settings using parsimony and Bayesian analyses. The analyses revealed the paraphyly of Hymenopodidae, Iridopterygidae, Mantidae, and Thespidae and the monophyly of the Amorphoscelidae subfamily Paraoxypilinae. All represented subfamilies of Iridopterygidae and Mantidae appear paraphyletic. Mantoididae is sister group to the rest of the sampled mantid taxa. Lineages congruent with current subfamilial taxonomy include Paraoxypilinae, Hoplocoryphinae, Hymenopodinae, Acromantinae and Oligonicinae. The mantid hunting strategy is defined as either generalist, cursorial or ambush predators. By mapping hunting strategy onto our phylogeny, we reconstructed the ancestral predatory condition as generalist hunting, with three independent shifts to cursorial hunting and one shift to ambush hunting, associated with the largest radiation of mantid species.
160 citations
TL;DR: P phylogenetic analysis of the heterogeneous Neuroptera remains a challenge with respect to selection of the proper genes and mutatis mutandis the morphological approach, and the hypothesis of a sister‐group relationship Raphidioptera put forward in recent morphological analyses is supported.
Abstract: In a first molecular approach specially dedicated to examining the phylogeny of the Neuropterida, two nuclear and two mitochondrial genes were tested: 18S rRNA, translation elongation factor-1a, cytochrome c oxidase subunit 3 and 16S rRNA. Molecular results are discussed in the light of a previous holomorphological cladistic analysis. The hypothesis of a sister-group relationship Raphidioptera þ (Neuroptera þ Megaloptera) put forward in recent morpho- logical analyses is supported by our data, which is in contrast to the traditional view (Raphidioptera þ Megaloptera) þ Neuroptera. Furthermore, the Nevrorthidae (constituting the suborder Nevrorthiformia) as a sister group of all other Neur- optera is confirmed. The disruption of the suborder Hemerobiiformia is the most conflicting result of the molecular analysis. Sisyridae and Osmylidae do not cluster within Hemerobiiformia, but represent two distinct and widely separated branches. The remaining Hemerobiiformia emerge as the sister group of the suborder Myrmeleontiformia, which is once more confirmed as monophyletic. Among the genes tested, cytochrome c oxidase subunit 3 proved to be most potent for resolving the phylogenetic relationships among Neuropterida. The nuclear gene for the ribosomal 18S rRNA is too conserved within the alignable regions, whereas the variable sections are too divergent to be applicable within this evolu- tionary time frame. The elongation factor-1a gene proved to exist in more than one copy in Neuropterida, and thus is not applicable in the present state of knowledge. With respect to the mitochondrial sequences (cytochrome c oxidase subunit 3, 16S rRNA), saturation impedes the unambiguous resolution of deeper nodes. Apparently, due to early diversification of the heterogeneous Neuroptera, phylogenetic analysis of this group remains a challenge with respect to selection of the proper genes and mutatis mutandis the morphological approach.
127 citations
TL;DR: The first comprehensive phylogenetic study of the Pseudococcidae based on analysis of nucleotide sequence data confirmed that the Sphaerococcinae as presently constituted is polyphyletic and the identities of the secondary endosymbionts may be useful in inferring mealybug relationships.
Abstract: . Mealybugs (Hemiptera: Pseudococcidae) are small, plant-sucking insects which comprise the second largest family of scale insects (Coccoidea). Relationships among many pseudococcid genera are poorly known and there is no stable higher level classification. Here we review previous hypotheses on relationships and classification and present the first comprehensive phylogenetic study of the Pseudococcidae based on analysis of nucleotide sequence data. We used three nuclear genes, comprising two noncontiguous fragments of elongation factor 1α (EF-1α 5′ and EF-1α 3′), fragments of the D2 and D10 expansion regions of the large subunit ribosomal DNA gene (28S), and a region of the small subunit ribosomal DNA gene (18S). We sampled sixty-four species of mealybug belonging to thirty-five genera and representing each of the five subfamilies which had been recognized previously, and included four species of Puto (Putoidae) and one species each of Aclerda (Aclerdidae) and Icerya (Margarodidae), using Icerya as the most distant outgroup. A combined analysis of all data found three major clades of mealybugs which we equate to the subfamilies Pseudococcinae, Phenacoccinae and Rhizoecinae. Within Pseudococcinae, we recognize the tribes Pseudococcini (for Pseudococcus, Dysmicoccus, Trionymus and a few smaller genera), Planococcini (for Planococcus and possibly Planococcoides) and Trabutinini (represented by a diverse range of genera, including Amonostherium, Antonina, Balanococcus, Nipaecoccus and non-African Paracoccus), as well as the Ferrisia group (for Ferrisia and Anisococcus), some ungrouped African taxa (Grewiacoccus, Paracoccus, Paraputo and Vryburgia), Chaetococcus bambusae and Maconellicoccus. The ‘legless’ mealybugs Antonina and Chaetococcus were not closely related and thus we confirmed that the Sphaerococcinae as presently constituted is polyphyletic. In our analyses, the subfamily Phenacoccinae was represented by just Phenacoccus and Heliococcus. The hypogeic mealybugs of the Rhizoecinae usually formed a monophyletic group sister to all other taxa. Our molecular data also suggest that the genera Pseudococcus, Dysmicoccus, Nipaecoccus and Paracoccus are not monophyletic (probably polyphyletic) and that Phenacoccus may be paraphyletic, but further sampling of species and genes is required. We compare our phylogenetic results with published information on the intracellular endosymbionts of mealybugs and hypothesize that the subfamily Pseudococcinae may be characterized by the possession of β-Proteobacteria (primary endosymbionts) capable of intracellular symbiosis with γ-Proteobacteria (secondary endosymbionts). Furthermore, our data suggest that the identities of the secondary endosymbionts may be useful in inferring mealybug relationships. Finally, cloning polymerase chain reaction products showed that paralogous copies of EF-1α were present in at least three taxa. Unlike the situation in Apis and Drosophila, the paralogues could not be distinguished by either the presence/absence or position of an intron.
106 citations
TL;DR: The present analysis strongly supports a sister group relationship between the families Aphelocheiridae and Potamocoridae, a monophylum for which a new superfamily,Aphelocheiroidea, is proposed.
Abstract: . Morphological characters and molecular sequence data were for the first time analysed separately and combined for the true water bugs (Hemiptera–Heteroptera, infraorder Nepomorpha). Data from forty species representing all families were included, together with two outgroup species representing the infraorders Gerromorpha and Leptopodomorpha. The morphological data matrix consisted of sixty-five characters obtained from literature sources. Molecular data included approximately 960 bp from the mitochondrial gene 16S and the nuclear gene 28S for all forty-two terminal taxa. The morphological dataset was analysed using maximum parsimony and the combined morphological and molecular (16S + 28S rDNA) dataset was analysed using direct optimization. A sensitivity analysis of sixteen different sets of parameters (various combinations of insertion–deletion cost and transversion costs) was undertaken. Character congruence was used as an optimality criterion to choose among competing phylogenetic hypotheses. The final hypothesis was obtained from the analysis of the combined molecular and mor phological dataset with the most congruent parameter set. This hypothesis supports the monophyly of all currently recognized families of Nepomorpha, and of the superfamilies Nepoidea (Nepidae + Belostomatidae), Corixoidea (Corixidae), Ochteroidea Ochteridae + Gelastocoridae), Notonectoidea (Notonectidae), and Pleoidea (Pleidae + Helotrephidae), but not the monophyly of the Naucoroidea (Naucoridae + Aphelocheiridae + Potamocoridae). The close relationship between the Notonectidae and Pleoidea is also supported. Our hypothesis concurs with Mahner in the placement of the Corixidae as a sister group to the remaining nepomorphan superfamilies except the Nepoidea, but differs in the placement of the Ochteroidea as a sister group to the Notonectoidea + Pleoidea. The superfamily Naucoroidea should be limited to only including the family Naucoridae and not the families Aphelocheiridae and Potamocoridae. The present analysis strongly supports a sister group relationship between the families Aphelocheiridae and Potamocoridae, a monophylum for which we propose a new superfamily, Aphelocheiroidea.
91 citations
TL;DR: Comparisons of mitochondrial cytochrome oxidase I gene sequences for allopatric and sympatric Bemisia tabaci populations collected from Africa and India and from the host‐plants cassava, sweet‐potato and a common weed suggest that biologically discrete populations exist within B. tabaci.
Abstract: Reciprocal-crossing experiments were carried out and mitochondrial cytochrome oxidase I gene (mtCOI) sequences were compared for allopatric and sympatric Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) populations collected
from Africa and India, and from the host-plants cassava, sweet-potato and a common weed, Euphorbia geniculata. Three incompatible mating groups were discovered, which involved the cassava B. tabaci colonies from Africa and India, the cassava and sweet-potato B. tabaci populations from Uganda, and the cassava and E. geniculata B. tabaci from India. Successful reciprocal mating occurred between cassava-specific B. tabaci from Uganda, Tanzania and Ghana, and
between two Indian cassava B. tabaci populations. Thep arsimony and neighbourjoining analyses of 699 bp mtCOI gene sequences divided the colonies primarily into those originating from Africa and India. Further subgrouping corresponded to host-plant specialization. Cassava-specific Ugandan, Tanzanian and Ghanaian colonies formed a single group and the sympatric sweet-potato colony from Uganda grouped separately from them. The two geographically distant Indian cassava B. tabaci populations were similar and formed a single group, whereas the sympatric E. geniculata colony formed a sister clade. The clades generated by the phylogenetic analyses were maintained, with highly supported bootstrap values, when other published mtCOI gene sequences were included in the tree-building process and the divisions matched those revealed by the reciprocal-crossing experiments. These data suggest that biologically discrete populations exist within B. tabaci (sensu Russell, 1957).
85 citations
TL;DR: The phylogenetic analysis of ninety‐two adult morphological characters supports the treatment of Nirvaninae as a junior synonym of Evacanthinae and the redefinition of Evacellinae to include the tribes Nirvanini, Balbillini, Evacantini and Pagaroniini and indicates that Nirvaninee, as previously defined, is polyphyletic.
Abstract: . The phylogenetic analysis of ninety-two adult morphological characters supports the treatment of Nirvaninae as a junior synonym of Evacanthinae and the redefinition of Evacanthinae to include the tribes Nirvanini, Balbillini, Evacanthini and Pagaroniini. The analysis indicates that Nirvaninae, as previously defined, is polyphyletic. A key to tribes and Neotropical genera is provided and diagnostic features for these taxa are reviewed. Jassoqualus Kramer, Neonirvana Oman, Synogonia Melichar (=Jassopronus Nielson & Godoy, syn.n.) and Tahura Melichar are retained within Nirvanini and two new Neotropical genera of this tribe are described and illustrated: Antillonirvana, gen.n., based on two new species from the Dominican Republic and one from Cuba; and Chibchanirvana, gen.n., based on two new species from Colombia. Pentoffia Kramer and Evanirvana Hill are treated as incertae sedis within Evacanthinae. Six new species of Pentoffia, a new species of Synogonia, a new species of Jassoqualus, two new species of Neonirvana and eleven new species of Tahura, all from South America, are also described and illustrated. The following taxa included previously in Nirvaninae are excluded from Evacanthinae, sensu lato: Tungurahuala Kramer to Cicadellinae; Columbonirvana Linnavuori to Typhlocybinae; Macroceratogoniini to Coelidiinae; Occinirvanini Evans to Deltocephalinae. Omaranus Distant, placed previously in Occinirvanini, is transferred to Doraturini (Deltocephalinae).
74 citations
TL;DR: The phylogenetic relationships of twenty‐five species of Cotesia commonly used in field and laboratory research are investigated, using the genes 16S, ND1, 28S and LW opsin and employing a range of phylogeny estimation methods including maximum parsimony, maximum likelihood, minimum evolution and Bayesian inference.
Abstract: . The braconid wasp genus Cotesia Cameron (Braconidae: Microgastrinae) is one of the largest genera of parasitoid wasps, and its species are employed frequently as biological control agents against pest insects. Several Cotesia species are also used as model organisms in physiology, ecology and population genetics studies. The genus thus has considerable importance from both applied and basic science perspectives. We investigated the phylogenetic relationships of twenty-five species of Cotesia commonly used in field and laboratory research, using the genes 16S, ND1, 28S and LW opsin and employing a range of phylogeny estimation methods including maximum parsimony, maximum likelihood, minimum evolution and Bayesian inference. Phylogenetic estimates differed little among methods, especially for the combined data analysis. The combined analysis of four genes identified four well-supported clades within Cotesia: the melanoscela group (containing C. melanoscela, the C. flavipes species complex and probably also C. ruficrus), the kariyai group (containing C. kariyai, C. kazak, C. cyaniridis, C. flaviconchae and probably also C. anisotae and C. griffini), the rubecula group (containing C. congregata, C.electrae, C.euchaetis, C. marginiventris, C. obsuricornis and C. schizurae), and the glomerata group (consisting of C. glomerata, C. melitaearum and C. plutellae), plus a basal unresolved complex including C. hyphantriae, C. diacrisiae and C. empretiae. These groups correspond poorly with previous broad subgroups of Cotesia defined by Papp based on morphology. The current work constitutes the first real framework for comparative studies in systematics, ecology, physiology and population genetics of Cotesia. A preliminary analysis of the evolution of gregarious development from solitary is presented, in which it is apparent that solitary development is ancestral, and gregariousness has arisen several times independently within separate groups.
74 citations
TL;DR: This issue is raised in an editorial because editors of Systematic Entomology regularly receive discussions of ‘basal’ clades or taxa and try regularly to convince the authors individually that this doesn't make sense.
72 citations
TL;DR: The molecular phylogeny contradicts previous morphological hypotheses, in which Gollumiella and Anorasema are within Eucharitini, and suggests that very distinct traits such as the fusion of the pronotum and prepectus in adults and the Fusion of the first two tergites of the planidia are convergent apomorphies.
Abstract: Gollumiellinae is proposed as a new subfamily for the Indo-Pacific genera Gollumiella Hedqvist and Anorasema Bouc˘ ek based on analyses of three ribosomal transcript gene regions (28S-D2 and -D3, and 18S-E23; 1262 aligned base pairs) sequenced for twenty-eight genera and fifty-four species of Euchariti- dae, and twelve genera and nineteen species of Pteromalidae (Pteromalinae) and Perilampidae (Chrysolampinae and Perilampinae). Gollumiella and Anorasema have been treated as either a monophyletic or paraphyletic group within the Eucharitini (Hymenoptera: Eucharitidae). The monophyly of the Eucharitidae and a sister-group relationship between Gollumiellinae and Oraseminae þ Eucharitinae are supported strongly in parsimony and Bayesian analyses. The molecular phylogeny contradicts previous morphological hypotheses, in which Gollumiella and Anorasema are within Eucharitini. The strength of the molecular hypothesis is explored through evaluations of data alignments that are eye optim- ized, CLUSTAL-X aligned, eye optimized but with gaps coded as a fifth base, and eye optimized and combined with morphological data. The oviposition behaviour and host associations of G. longipetiolata on tree ferns (Cyathea latebrosa, Cyathea- ceae), and the morphology of the planidium and pupa are newly described. Eggs are deposited vertically, with the base anchored into the leaf or petiolar surface. Unlike other Eucharitidae, the eggs are not stalked, but rather tipped with a secretion that may act as an attractant for their ant host, Paratrechina sp. (For- micidae: Formicinae). Various morphological character state optimizations and their implications for convergent morphology, behaviour and host associations are discussed. All results using gene regions treated separately or combined with the morphological data reach the same conclusion: Anorasema þ Gollumiella form the sister group of Oraseminae þ Eucharitinae, and thus deserve subfamily status. This suggests that very distinct traits such as the fusion of the pronotum and prepectus in adults and the fusion of the first two tergites of the planidia are convergent apomorphies. Molecular data, when strongly supported, can provide new information to unravel convergent from synapomorphic changes, resulting in more robust hypotheses of relationship.
71 citations
Journal Article•
TL;DR: In this article, the authors argue that there is no such a thing as themost basal clade, and that there are always two most basal clades (if the most basal branching is completely resolved, Fig. 1).
Abstract: When a tree is rooted, it has a base. Everything near that base is basal. A clade branching off near the base is a basal clade, isn’t it? Does this make sense? No, it doesn’t. As editors of Systematic Entomology we regularly receive discussions of ‘basal’ clades or taxa. We find them in a great proportion of submitted papers and even in a widely used textbook (Forey et al., 1992: 134; eliminated from the second edition). Google found 100 hits for ‘the most basal clade’ and seventy-five for ‘the most basal taxon’, including a few from Systematic Entomology (search on 14 January 2004). Although we try regularly to convince the authors individually that this doesn’t make sense, we have decided to raise this issue in an editorial. Why one clade cannot be most basal Every branching in a (phylogenetic) tree is rotatable (see Fig. 1). Of course, the tree has a base, and there is a most basal branching and a next most basal branching, but there is no such thing as themost basal clade. Because branchings are rotatable, there are always two most basal clades (if the most basal branching is completely resolved, Fig. 1) or even more most basal clades (if the most basal branching is not completely resolved, e.g. in Polyneoptera in Fig. 1). Both branches originating from a node (i.e. the two sister groups) are of equal age and have undergone equivalent evolutionary change. Whether a group has branched off early (basal)
70 citations
TL;DR: Data indicate a paraphyletic Erotylidae and Languriidae, which has experienced at least one evolutionary transition from mycophagy to phytophagy, three transitions from Aphyllophorales hosts to Euagarics, and one transition from EUagarics hosts to Mucorales.
Abstract: Phylogenetic relationships of Erotylidae (pleasing fungus beetles) were inferred based on DNA sequence data. Relationships of clades within Erotylidae were examined, as was the relationship of the entire family to Languriidae (lizard beetles). 18S and 28S ribosomal DNA were sequenced for sixty-one taxa repre- senting major erotylid lineages and outgroups. Phylogenetic analyses under vary- ing parameter settings using standard parsimony and likelihood techniques were performed. These data indicate a paraphyletic Erotylidae and Languriidae. Encaustinae (including Coptengis), Megalodacninae and Erotylinae are supported as monophyletic, whereas Dacninae and Tritominae are paraphyletic. Taxonomic and biological implications are discussed. Gregariousness has arisen at least three times in Erotylidae. The erotylid clade has experienced at least one evolutionary transition from mycophagy (on Aphyllophorales) to phytophagy, three transitions from Aphyllophorales hosts to Euagarics, and one transition from Euagarics hosts to Mucorales (Zygomycetes). There are no recognizable phylogenetic trends in coloration across higher-level erotylid lineages.
TL;DR: In this paper, the authors re-examine the wing re-evolution hypothesis of Whiting et al. and conclude that there are no grounds for overturning the traditional view of stick insect evolution.
Abstract: In a recent issue of Nature, Whiting et al. (2003) report a surprising and novel result: the re-evolution of insect wings from wingless ancestors in Phasmatodea (stick insects). If true, this represents an important advance in our understanding of insect and gene evolution. It overturns a long-held notion that wings evolved just once in insects and have been lost many times. The allegedly re-evolved wings have exactly the same detailed structure as other insect wings (Whiting et al., 2003, their Fig. 1b) and if this feature truly has been regained in stick insects it would demonstrate a remarkable maintenance of ‘wing genes’ despite inactivation of the developmental pathway over long periods of evolutionary time. Whiting et al.’s conclusions have already been promulgated in the popular literature (New Scientist, Scientific American, The New York Times, Anon, 2003a, b, c) but before this extraordinary evolutionary scenario reaches the entomology textbooks a re-examination is in order. Two lines of evidence are essential to Whiting et al.’s wing re-evolution hypothesis: (1) the topology in their stick insect phylogenetic tree, and (2) the distribution of wing/wingless transformations on that tree. The latter issue, character optimization, is addressed here. Our reappraisal of the evidence convinces us that Whiting et al. have overstated significantly the probability of wing re-evolution in stick insects. When this is taken into account we see no grounds for overturning the traditional view of stick insect evolution.
TL;DR: A scenario in which species from temporary habitats evade drought by entering the hyporheic zone, and may be driven to a subterranean existence by desertification of their ancestral habitats, as can be observed in Australia is suggested.
Abstract: Michael Balke, Chris H. S. Watts, Steven J. B. Cooper, William F. Humphreys and Alfried P. Vogler
TL;DR: The results of the combined data analyses grouped membracids into two well‐supported lineages, consistent with the following hypotheses: (1) the posterior pronotal process was derived and lost multiple times during the evolution of Membracidae; (3) maternal care evolved independently multiple times and may or may not have been preceded by the acquisition of ant mutualism.
Abstract: . Recent independent phylogenetic analyses of membracid relationships based on molecular and morphological data have identified monophyletic lineages within the family. However, the results of these studies have not fully resolved treehopper phylogeny, and relationships among some higher membracid lineages remain in doubt. Portions of three datasets (958 aligned nucleotides from elongation factor-1α, 2363 aligned nucleotides from 28S ribosomal DNA, and eighty-three morphological features of adults and nymphs) introduced in recent studies were reanalysed separately and in combination with two new molecular datasets (321 aligned nucleotides from wingless and 1829 aligned nucleotides from 18S ribosomal DNA). The results of the combined data analyses, contrary to previous analyses of morphological data alone, grouped membracids into two well-supported lineages, one comprising Stegaspidinae and Centrotinae, the other comprising Membracinae, Darninae and Smiliinae. The analyses recovered Centrotinae, Membracinae and Darninae as monophyletic groups, but Stegaspidinae was paraphyletic with respect to Centrotinae, and Smiliinae was polyphyletic with Micrutalini placed as a sister group to the clade comprising Membracinae, Darninae and Smiliinae. These results are consistent with the following hypotheses, proposed previously based on an analysis of morphological data: (1) the posterior pronotal process was derived and lost multiple times during the evolution of Membracidae; (2) Membracidae originated in the New World and reached the Old World subsequently via dispersal; (3) maternal care evolved independently multiple times and may or may not have been preceded by the acquisition of ant mutualism.
TL;DR: Cladistic analysis of adult morphological characters corroborates Engel's hypotheses of monophyly of Augochlorella phylogenetic relationships with related genera as follow: (Augochlorlla ((Ceratalictus, Pereirapis) Augochlora)).
Abstract: . The New World halictid bee genus Augochlorella (Augochlorini) is revised. Sixteen species are recognized, with five described as new: Augochlorella acarinata sp. n., A. una sp. n., A. meridionalis sp. n., A. stenothoracica sp. n. from South America, and A. karankawa sp. n. from U.S.A. The following new synonymies are proposed: Augochlorella michaelis (Vachal) with A. urania (Smith), A. edendata Michener with A. comis (Vachal); A. striata (Provancher) with Augochlorella aurata (Smith) and A. neglectula maritima Ordway with A. neglectula (Cockerell). The female of A. tredecim (Vachal) and the male of A. iopoecila Moure are described for the first time. Keys to the species are provided. Cladistic analysis of adult morphological characters corroborates Engel's hypotheses of monophyly of Augochlorella phylogenetic relationships with related genera as follow: (Augochlorella ((Ceratalictus, Pereirapis) Augochlora)). Vicariant events shown in the cladistic analyses are discussed, and an account of distribution is presented.
TL;DR: Character analysis reveals that in Panacanthus the ancestral condition of calling song resonance has given rise to a more nonresonant (transient) stridulation, and a correlation between the production of more complex sound waves and spinous protection of the body (especially the pronotum) is noted.
Abstract: A cladistic analysis and systematic revision of the genus Panacanthus accompanies the description of three new species, with calling songs reported for four species. The evolutionary origin of spines is considered as a defensive mechanism in Panacanthus; both morphological and behavioural (i.e. acoustic) traits allow inferences about relationships. Phylogenetic analysis produced one most parsimonious cladogram eighty-two steps long, with the ensemble consist- ency index ¼ 0.84. Panacanthus cuspidatus and P. pallicornis (formerly Storniza Walker 1869, Martinezia Bolivar 1881) are properly incorporated in Panacanthus. On morphology, Panacanthus is more related to the Neotropical Copiphora and Lirometopum than to the Old World Lesina. Character analysis reveals that in Panacanthus the ancestral condition of calling song resonance (the production of musical sounds) has given rise to a more nonresonant (transient) stridulation. A correlation between the production of more complex sound waves and spinous protection of the body (especially the pronotum) is noted. Because early workers grouped Panacanthus with other spiny genera, based on pronotal morphology, we present a critique of the evolutionary and ecological implications of the develop- ment of defensive spines in this genus. This approach may be applied to other taxa using a similar protective mechanism. We advise against arrangement of the pronotal, cephalic and femoral armature as a homologous characteristic across subfamilies. Several pronotal processes and modifications evolved independently in other genera of Conocephalinae, Hetrodinae, Pseudophyllinae and Phanerop- terinae. The pronotal structure of Panacanthus is unique and may be taken as a synapomorphic characteristic of all its species and as an autapomorphic feature of the genus.
TL;DR: The phylogenetic trees show that one group composed of ssp.
Abstract: . The phylogeny of the butterflies Parides (Byasa) alcinous caught at various localities in Japan and the Ryukyu Islands and in the eastern part of the Eurasian Continent was analysed using mitochondrial DNA sequences coding for NADH dehydrogenase subunit 5 (778 bp). The same phylogenetic relationship among P. (B.) alcinous subspecies was obtained with all analytical methods used, and was supported by high bootstrap values. The female butterfly wing pattern that characterizes each subspecies was unrelated to the phylogenetic relationship among the subspecies. The phylogenetic trees show that one group composed of ssp. alcinous and yakushimanus, which are distributed in the main area of the Japan Archipelago, the Korean Peninsula and southern Primorski of Russia, and the other group comprised of ssp. loochooanus, bradanus and miyakoensis, all of which are distributed in the Ryukyu Islands, diverged from a common ancestor. The ssp. loochooanus distributed in Amami and Okinawa Islands then diverged, and ssp. bradanus and miyakoensis distributed in Yaeyama and the Miyako Islands, respectively, finally diverged. This divergence order nearly agrees with the palaeogeography of the Ryukyu Islands that has been established in Pliocene and Pleistocene (0.2–2 MYA), suggesting that P. (B.) alcinous has been isolated in the Ryukyu Islands since the establishment of the islands.
TL;DR: Both coprophagy and male–female co‐operation in nesting appear as derived traits in Geotrupinae, and Odonteus species (Bolboceratinae s.str.) appear to constitute the closest sister group to Geotrupes.
Abstract: . Thirty-eight characters derived from the larvae of Geotrupidae (Scarabaeoidea, Coleoptera) were analysed using parsimony and Bayesian inference. Trees were rooted with two Trogidae species and one species of Pleocomidae as outgroups. The monophyly of Geotrupidae (including Bolboceratinae) is supported by four autapomorphies: abdominal segments 3–7 with two dorsal annulets, chaetoparia and acanthoparia of the epipharynx not prominent, glossa and hypopharynx fused and without sclerome, trochanter and femur without fossorial setae. Bolboceratinae showed notable differences with Pleocomidae, being more related to Geotrupinae than to other groups. Odonteus species (Bolboceratinae s.str.) appear to constitute the closest sister group to Geotrupinae. Polyphyly of Bolboceratinae is implied by the following apomorphic characters observed in the ‘Odonteus lineage’: anterior and posterior epitormae of epipharynx developed, tormae of epipharynx fused, oncyli of hypopharynx developed, tarsal claws reduced or absent, plectrum and pars stridens of legs well developed and apex of antennal segment 2 with a unique sensorium. A ‘Bolbelasmus lineage’ is supported by the autapomorphic presence of various sensoria on the apex of the antennal segment, and the subtriangular labrum (except Eucanthus). This group constituted by Bolbelasmus, Bolbocerosoma and Eucanthus is the first evidence for a close relationship among genera, but more characters should be analysed to test the support for the clade. A preliminary classification at tribe level of Geotrupinae is suggested as follows: Chromogeotrupini (type genus Chromogeotrupes), Lethrini (type genus Lethrus), Taurocerastini (type genus Taurocerastes) and Geotrupini (type genus Geotrupes). Some ecological facts of Geotrupidae evolution could also be explained by the present results, such as those related to diet and nesting behaviour. Both coprophagy and male–female co-operation in nesting appear as derived traits.
TL;DR: Phylogenetic relationships among species groups of Trichadenotecnum were inferred based on morphology and the partial sequences of five gene regions and the short ventral valve of gonapophyses independently evolved at least three or four times, although this character was used initially to diagnose the spiniserrulum group.
Abstract: Phylogenetic relationships among species groups of Trichadenotecnum were inferred based on morphology and the partial sequences of five gene regions (mitochondrial 12S rDNA, 16S rDNA, cytochrome oxidase I, NADH dehydro- genase subunit 5 and nuclear 18S rDNA). All analyses supported the monophyly of Trichadenotecnum and all previously proposed species groups, except that T. circularoides was excluded from the spiniserrulum group. To examine the phylogenetic usefulness of morphological data, the morphological characters used in the construction of an earlier taxonomic system for Trichadenotecnum were mapped parsimoniously on the molecular tree. As a result: (1) commonly used forewing marking features (sparsely or extensively spotted) are considered to be very homoplastic and less informative of higher-level phylogenetic relation- ships; (2) a broadly expanded epiproct lobe is considered to be independently evolved at least two or three times, and a detailed morphological re-examination allows recognition of these convergent structures; (3) the short ventral valve of gonapophyses independently evolved at least three or four times, although this character was used initially to diagnose the spiniserrulum group.
TL;DR: According to the present study, Hydrophilinae is monophyletic, and except for the tribe hydrophilini which appears as polyphyletic (it includes the subtribes HydrophILina, Hydrobiina, and Acidocerina), the remaining tribes are monophy letic.
Abstract: Fil: Archangelsky, Miguel. Consejo Nacional de Investigaciones Cientificas y Tecnicas; Argentina. Universidad Nacional de la Patagonia; Argentina
TL;DR: An alternative hypothesis is suggested, that of a Gondwanan origin for the Neotropical anthomyiids, that is distributed mainly in southeastern South America and the other from northwestern South America to southern North America.
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.
TL;DR: The erection of the new subfamily Succinoraphidiinee as the sister group of the Raphidiinae (comprising all other known members of the family) triggers reconsiderations concerning reliable autapomorphies of Raphidioptera, which are discussed.
Abstract: Succinoraphidia exhibens genn, spn (Raphidiidae: Succinoraphidiinae subfamn) and Succinofibla aperta genn, spn (Inocelliidae) from Baltic Amber are described and figured Both male holotypes are significant for giving access to their genital sclerites, allowing comparison with relevant structures of extant species The erection of the new subfamily Succinoraphidiinae as the sister group of the Raphidiinae (comprising all other known members of the family), triggers reconsiderations concerning reliable autapomorphies of Raphidioptera, which are discussed As a consequence, the Jurassic Hondelagia and Priscaenigma have to be dismissed from the order
TL;DR: The association of the included Larentiinae taxa with the Cosymbiini-Rhodometrini-Timandrini lineage questions the monophyly of Sterrhinae.
Abstract: The phylogenetic relationships of tribes of the geometrid subfamily Sterrhinae (Lepidoptera) were studied, with special emphasis on finding delimiting characters for the tribe Scopulini Two cladistic analyses were conducted for fifty-nine species representing all previously recognized Sterrhinae tribes and covering the geographical range of the subfamily In the first analysis, twelve putative synapomorphies of Scopulini, taken from the literature, were coded for actual specimens in order to test their ability to support the monophyly of the group The resulting strict consensus cladogram was totally unresolved In the second analysis, the twelve characters were combined with additional information from the morphology and ecology of adults and immature stages Analysis of these ninety-six characters resulted in a well-resolved cladogram The tribes were found to be monophyletic, except Cosymbiini and Rhodostrophiini There are two main lineages within Sterrhinae: Cosymbiini + Rhodometrini + Timandrini and Rhodostrophiini + Cyllopodini + Sterrhini + Scopulini Aletini and Problepsini lay within the concept of Scopulini The association of the included Larentiinae taxa with the Cosymbiini + Rhodometrini + Timandrini lineage questions the monophyly of Sterrhinae A majority of the recovered synapomorphic characters had been recognized previously, but several new phylogenetically informative characters were found, especially from the thorax No unique characters diagnosing the tribe Scopulini were found, but many homoplastic synapomorphic features were found which diagnose parts of it All recognized Sterrhinae genera are assigned tentatively to tribes and problematic cases are discussed
TL;DR: It appears that in the present state of the analytical strategies, hypotheses concerning arthropod phylogenies obtained from morphological and developmental criteria and combined analyses involving molecular and morphological data provide more reliable results than those generated by molecular information alone.
Abstract: . A review of different studies on the phylogenetic relationships of the early Hexapoda lineages shows that analyses based on molecular sequence data have led to labile and sometimes incongruous results, introducing doubt as to the reliability of the cladograms as a whole. In a recent analysis using molecular data, the Collembola, usually considered as early branching hexapods, appear to occupy a position outside the assemblage of Crustacea and Insecta, leading to the rejection of the traditional view of hexapod monophyly. However, many morphological features, as well as the results of cladistic analyses based on morphological and developmental information, contradict these conclusions. More generally, it appears that in the present state of the analytical strategies, hypotheses concerning arthropod phylogenies obtained from morphological and developmental criteria and combined analyses involving molecular and morphological data provide more reliable results than those generated by molecular information alone.
TL;DR: The authors' analysis suggests that the family Hybosoridae is paraphyletic with respect to Ceratocanthidae, and the clade comprising the hybosorid genera Hybosorus and Phaeochrous is the sister group of the remaining HybosOridae plus CeratOCanthidae.
Abstract: Larvae of the scarabaeoid genera Germarostes Paulian, Cyphopisthes Gestro, Paulianostes Ballerio, Ceratocanthus White, Pterorthochaetes Gestro, Madrasostes Paulian, Astaenomoechus Martinez & Pereira (Ceratocanthidae) and Hybosorus Macleay, Phaeochrous Castelnau, and Anaides Westwood (Hybo- soridae) are described, keyed and illustrated with fifty-seven drawings. A phylo- genetic analysis of these two families based on larval morphology is presented. Fifty-four larval morphological and three biological characters from twenty-seven taxa revealed nineteen equally parsimonious cladograms. The monophyly of (Ceratocanthidae þ Hybosoridae) is supported by four unambiguous unique syna- pomorphies: dorsal medial endocarina on cranium extended anteriorly into fron- tal sclerite; presence of large membranous spot on apical antennomere; labium dorsally with four pores in centre (secondarily reduced to two pores in some groups); and presence of stridulatory organ on fore- and middle legs (secondarily reduced in some groups). Our analysis suggests that the family Hybosoridae is paraphyletic with respect to Ceratocanthidae. The clade comprising the hybosorid genera Hybosorus and Phaeochrous is the sister group of the remaining Hybo- soridae plus Ceratocanthidae. It is supported by two unambiguous synapomorphies: two apical antennomeres completely joined and the stridulatory organ represented by seven to nine large teeth anteriorly on the middle leg. The hybosorid genus Anaides is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus and Phaeochrous) and the ceratocanthid genus Germarostes is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus, Phaeochrous and Anaides). The ceratocanthid genera Cyphopisthes, Astaenomoechus, Paulianostes, Pterorthochaetes, and Madrasostes constitute a sister group to the hybosorid genus Cryptogenius and are supported by the presence of two reversions: two dorsal pores on labium and completely reduced stridulatory organs on fore- and middle legs.
TL;DR: The ancestor of Balacha was probably distributed in southeast South America before the uplift of the mountain ranges, during the late Eocene or Oligocene, and this event may have triggered speciation of some lineages of the genus by vicariance.
Abstract: . Balacha currently includes six valid described species and B. caparao sp. nov., here described from Minas Gerais State, Brazil. The new taxon differs from other Balacha species in the ventrally inflated preapical aedeagal area and oblique bases of the ovipositor first valvulae. New records are as follows: B. decorata from Paraguay and Minas Gerais State (Brazil); B. distincta from Brazil; B. melanocephala from Montevideo Department (Uruguay) and Buenos Aires and Distrito Federal provinces (Argentina); and B. similis from Rio de Janeiro and Minas Gerais states (Brazil). Colombia is considered a dubious record for the genus. Phylogenetic relationships among Balacha species are examined based on a matrix of eighteen terminal taxa and sixty-seven morphological characters. Balacha may be defined by the following synapomorphies: crown–frons transition approximately acute, crown anteriorly produced and with anterior margin subangulate, flattened pronotum continuing contour of head and mesonotum in lateral view, and teeth of second valvulae of ovipositor with anterior dorsal projection. The genus comprises two major lineages: the red Balacha clade (B. lepida(B. distincta + B. rubripennis)), and the black Balacha clade (B. caparao(B. decorata(B. melanocephala + B. similis))). Despite the scarcity of data on host plant usage of outgroup taxa (probably generalists on Lamiaceae, Lauraceae, and Asteraceae), we believe that the shift to feeding on Eryngium(Apiaceae) occurred in the ancestor of all recent Balacha species. Their small size and depressed body appear to be adaptations to living inside the rosette-disposed leaves, and they seem to be restriced to this microhabitat. Balacha and their Eryngium hosts occur in grasslands in temperate South America, but at lower latitudes in Brazil they are isolated in alpine meadows on peaks of the southeast highlands. Dispersal between such areas through the lowland humid Atlantic forest in recent times would be difficult, thus the ancestor of Balacha was probably distributed in southeast South America before the uplift of the mountain ranges, during the late Eocene or Oligocene. This event may have triggered speciation of some lineages of the genus by vicariance.
TL;DR: Trueman et al. as mentioned in this paper showed that the ancestral phasmid is reconstructed unambiguously as wingless, with wings being reacquired later in phylogenetic evolution by using multiple molecular markers and employing several methods of tree reconstruction and character optimization.
Abstract: By using multiple molecular markers and employing several methods of tree reconstruction and character optimization, we demonstrated that the ancestral phasmid is reconstructed unambiguously as wingless, with wings being reacquired later in phasmid evolution (Whiting et al., 2003). We presented this as a compelling example of recurrence in which a complex character, once lost to evolution, is regained subsequently in a descendant lineage (West-Eberhard, 2003). Our hypothesis is refutable via additional phylogenetic analyses including a larger selection of taxa, additional molecular markers, morphological data, or by examining patterns of development of wing expression in phasmids. We are currently performing research in each of these areas to add greater precision to this hypothesis. Trueman et al. (2004) have not presented a formal test of our hypothesis, nor contributed additional data to refute our findings. Wing recurrence is a hypothesis of character transformation, requiring a phylogenetic topology for interpretation. The ‘traditional view’ of phasmid wing evolution that these authors embrace was conjured in phylogenetic ignorance, since we presented the first formal analysis of phasmid phylogeny. Clearly the current data support a basal placement of apterous taxa, and multiple researchers who have reanalysed our data, including Trueman et al., have been unable to find a topology which rejects this hypothesis, regardless of analytical methodology. Thus, Trueman et al. quibble over methods of character optimization by launching into confused, non-phylogenetic, and mutually contradictory arguments to unravel our hypothesis ‘before this extraordinary evolutionary scenario reaches the entomology textbooks’. When the cost of wing gain is set extremely high (parsimony), or the rate of transformation from wingless to winged is set extremely low (likelihood), any method of character optimization will bias against reconstructing wing recurrence. More generally, values always can be selected to make it impossible to detect character recurrence by forbidding its transformation on a phylogenetic topology (1⁄4 Dollo’s law). Because we believe that phylogenetic topologies should establish patterns for inferring evolutionary processes, the issue becomes how much evidence is required before recurrence is a well-supported hypothesis. Our analyses and those of Trueman et al. agree on one critical point: under both parsimony and likelihood methods of character optimization the ancestral stick insect is supported unambiguously as wingless, with wings gained on multiple occasions. Trueman et al. (2004) premise their argument with the curious statement that ‘reconstruction of the phasmid ancestor is not the relevant issue.’ We argue it is the only relevant issue. If an ancestral node is reconstructed as wingless, and a descendant node is reconstructed as winged, then there must be a transformation from wingless to winged. Gains and losses are not observations: they are inferences based on hypotheses of character transformations given a topology and method of character optimization. To discard ancestral reconstruction is to discard the very reason why phylogeny is critical for investigating character evolution. Trueman et al. fail to follow their own dictum in discussing the relative merits of hypotheses using values obtained directly from character optimization (e.g. four gains and three losses for the wing ‘re-evolved’ hypothesis). Moreover, if all that matters is how well the ‘model fits the data’ then we can dispense with their parsimony arguments outright, as most systematists would acknowledge that parsimony is not attempting to quantify the fit of data to specific evolutionary models. Trueman et al.’s criticisms can be summarized as three points: (1) the ‘probability’ of wing loss to gain should be 2.5 under parsimony; (2) the ‘probability’ of wing loss to gain should be 6 : 1 under parsimony; and (3) the ‘probability’ of wing loss to gain should be 13 : 1 given a likelihood analysis.
TL;DR: Results from both individual and combined analyses of the smaller taxon sets provided support for the monophyly of Thricops and a complete division of the genus into two monophyletic subgroups.
Abstract: . The muscid genus Thricops Rondani comprises forty-four species and two subspecies restricted to the northern hemisphere. A species-level phylogenetic analysis of Thricops was conducted using forty-four morphological characters, 426 bp of the nuclear gene white and 523 bp spanning the 5′ end of the cytochrome c oxidase subunit I (COI), the tRNA leucine gene (L2 region) and the 3′ end of the cytochrome c oxidase subunit II (COII). Thirty-nine species and two subspecies of Thricops were included in the analysis. Two species of Azelia Robineau-Desvoidy and one species of Hydrotaea Robineau-Desvoidy were used as outgroups. Morphological characters were coded for all included species, the mitochondrial gene fragment (COI + II) was sequenced for a subset of seventeen species of Thricops and three outgroup species, and white for twelve of those seventeen Thricops species and two outgroup species. Six separate maximum parsimony analyses were performed on three taxon sets of different sizes (n = 14, n = 20, n = 44). Results from the partition homogeneity test indicated no significant incongruence between data partitions, and four combined maximum parsimony analyses were conducted (DNA + morphology for n = 14; COI + II + morphology for n = 20; DNA + morphology for n = 20; DNA + morphology for n = 44). The relative contribution of each data partition to individual nodes was assessed using partitioned Bremer support. Strict consensus trees resulting from the unweighted analyses of each dataset are presented. Combination of datasets increased resolution for the small taxon set (n = 14), but not for the larger ones (n = 20, n = 44), most probably due to increasing amounts of missing data in the larger taxon sets. Results from both individual and combined analyses of the smaller taxon sets (n = 14, n = 20) provided support for the monophyly of Thricops and a complete division of the genus into two monophyletic subgroups. The strict consensus cladograms resulting from the analysis of the morphological data alone and the combined data for the large taxa set (n = 44) both supported the monophyly of the genus, but placed the species Thricops foveolatus (Zetterstedt) and Thricops bukowskii (Ringdahl) at the base of the ingroup, in a polytomy with a relatively well-resolved branch comprising all remaining species of the genus. The basal position of these two species, included in the morphological taxon set but absent in the others, illustrates the potential pitfalls of taxon sampling and missing data in phylogenetic analyses. The synonymy of Alloeostylus with Thricops as proposed by previous authors was supported by our results. Relative contributions of different data partitions is discussed, with the mitochondrial sequence generally providing finer resolution and better branch support than white.
TL;DR: The phylogenetic value of pupal characters is assessed at intra‐ and intergeneric levels within the tribe and a summary of antipredator structures of all known Coelometopini pupae is given.
Abstract: . The pupal stage of ten Coelometopini species occurring in Australia, New Guinea, Southeast Asia and the Pacific region are described and a key for their identification is provided. The species are Chrysopeplus expolitus Broun, Derosphaerus hirtipes Kaszab, Hypaulax crenata (Boisduval), Leprocaulus borneensis Kaszab, Metisopus purpureipennis Bates, Promethis carteri Kaszab, P. nigra (Blessig), P. quadraticollis (Gebien), P. quadricollis Pascoe and P. sulcigera (Boisduval). The gin trap structures of D. hirtipes and P. quadraticollis are described in detail using scanning electron micrographs. A summary of antipredator structures of all known Coelometopini pupae is given. The phylogenetic value of pupal characters is assessed at intra- and intergeneric levels within the tribe.
TL;DR: The phylogeny presented here confirms the monophyly of the centaurus, eumolphus, camdeo and epimuta groups and the amphimuta subgroup and confirms close relationships between species within the agelastus group, that together with the amphicuta sub group, centaurus and camdee groups form a monophyletic group.
Abstract: We present a phylogeny for a selection of species of the butterfly genus Arhopala Boisduval, 1832 based on molecular characters. We sequenced 1778 bases of the mitochondrial genes Cytochrome Oxidase 1 and 2 including tRNALeu, and a 393-bp fragment of the nuclear wingless gene for a total of 42 specimens of 33 species, representing all major species groups. Analyses of mtDNA and wingless genes show congruent phylogenetic signal. The phylogeny presented here confirms the monophyly of the centaurus, eumolphus, camdeo and epimuta groups and the amphimuta subgroup. It confirms close relationships between species within the agelastus group, that together with the amphimuta subgroup, centaurus and camdeo groups form a monophyletic group. However, incongruencies with previous taxonomic studies also occur; the amphimuta and silhetensis groups are not monophyletic, as is the genus Arhopala itself. One enigmatic species, A. kinabala, was evaluated further for topology and the support for basal placement of this species is due mainly to the wingless gene. However, in the Parsimony analysis, and subsequent Maximum Likelihood evaluations, certain nodes could not be resolved due to insufficient support. The mtDNA shows extreme AT bias with compositional heterogeneity at 3rd codon positions, which may result in saturation. By contrast, the wingless gene does not show compositional bias, suggesting that poor support is not due solely to saturation. The evaluation of morphological characters used in previous studies on Arhopala systematics on the molecular tree indicates that the macular pattern and the absence of tails at the hind wings show extensive homoplasy. A significant phylogenetic signal (as indicated by T-PTP tests) is present in several of these morphological characters, which are nevertheless of limited use in phylogenetic studies due to their labile nature