Putative Autapomorphies Research Articles

Dentalion (Hymenoptera: Eulophidae: Entedoninae) a new genus from tropical America with eleven new species

Dentalion gen. nov. (Hymenoptera: Eulophidae: Entedoninae), is described from the northern part of the Neotropical region, from Costa Rica to Ecuador and Brazil, and the Caribbean (Trinidad & Tobago). It comprises eleven new species: D. alveum, D. ambonatum, D. apertum, D. apon, D. crassicornis, D. jimenezi, D. mischum, D. noyesi, D. pinguicornis, D. pnigaliae, D. quadrifer. These species are diagnosed, described and illustrated, and a key for their identification is provided. Dentalion is characterized by two putative autapomorphies and two synapomorphies. The phylogenetic implications of these apomorphies are discussed briefly.

Description of three new genera and four new species of Neanastatinae (Hymenoptera, Eupelmidae) from Baltic amber, with discussion of their relationships to extant taxa

Aspidopleura baltica gen. n. and sp. n., Brevivulva electroma gen. n. and sp. n., Neanaperiallus masneri gen. n. and sp. n., and Metapelma archetypon sp. n. are described in Neanastatinae (Chalcidoidea, Eupelmidae) from Baltic amber. New information is given on the morphology of Eopelma Gibson, Lambdobregma Gibson, Metapelma Westwood and Neanastatus Girault, the four extant genera previously classified in Neanastatinae, and the extinct and extant taxa are compared with other Chalcidoidea for hypotheses of character evolution and phylogenetic relationships. Structural features of Neanaperiallus and Aspidopleura indicate that except for a greatly enlarged acropleuron the groundplan structure of Neanastatinae was similar to male Eupelminae or some Cleonyminae (Pteromalidae). Furthermore, classification of Neanaperiallus in Neanastatinae results in no putative autapomorphies to support monophyly of the subfamily. Some features of Encyrtidae, Tanaostigmatidae and Neanastatinae suggest that the taxa may have had a common ancestor, therefore rendering Eupelmidae paraphyletic. A key to the seven recognized genera and a revised diagnosis of Neanastatinae are given.

Pachybolini - a tribe of giant Afrotropical millipedes: arguments for monophyly and the description of a new genus from Madagascar (Diplopoda:Spirobolida:Pachybolidae)

Madabolus, gen. nov. is described to accommodate M. maximus, sp. nov. (type species) from several localities in western Madagascar. A phylogenetic analysis of 26 spirobolidan taxa, based on morphological characters, places Madabolus, gen. nov. with the Afrotropical genera Pelmatojulus de Saussure, 1860; Pachybolus Cook, 1897; Hadrobolus Cook, 1897 and Epibolus Cook, 1897. The tribe Pachybolini is proposed to include Madabolus, gen. nov. and the four abovementioned genera. Putative autapomorphies of Pachybolini are: a closed incisura lateralis; setae of gnathochilarial lamella lingualis apical; vulva kidney- or crescent-shaped, with valves meeting in a sinuous fissure and crest protruding. Within Pachybolini, Pelmatojulus (West Africa) stands out by virtue of an increased number of apical antennal sensory cones, with great probability an apomorphy. Madabolus, gen. nov. and east African genera of the tribe share another potential synapomorphy, namely, an increased number of setae on the gnathochilarial lamellae linguales. Other Malagasy genera of Pachybolidae do not seem to be related to Afrotropical genera.

Head anatomy of Xyelidae (Hexapoda: Hymenoptera) and phylogenetic implications

External and internal head structures of Macroxyela ferruginea (Say) and Xyela julii (Brébisson) were examined. A detailed description is provided for Macroxyela. The results are compared to the conditions found in other basal hymenopterans and representatives of other groups of endopterygote insects. Hitherto unnoticed autapomorphies of Hymenoptera are the concavity of the posterior head capsule, the very dense, regular vestiture of hairs, the collar-like, strongly developed posterior tentorium, and a large epipharyngopharyngeal lobe. Microphagous habits and associated features (asymmetric mandibular molae, epipharyngeal brush, infrabuccal pouch) are possibly groundplan features of Hymenoptera and Endopterygota. A switch to more or less liquefied food took place early in the evolution of Hymenoptera. The sitophore plate and a constricted, elongated prepharyngeal tube are likely synapomorphies of Hymenoptera and Mecopterida. Monophyly of Hymenoptera excluding Xyelidae is supported by the reduction of the mandibular molae and epipharyngeal brush. These changes are likely related to modified feeding habits. Widely separated mandibular bases, the loss of the median labral retractor (parallel loss in Xyelidae), and the presence of a hypostomal bridge are potential apomorphies of Hymenoptera excluding Xyelidae and Tenthredinoidea. Monophyly of Xyelinae and Macroxyelinae, respectively, is well supported by the results of our study. There is conflicting evidence as to whether Xyelidae is monophyletic. The presence of a subdivided galea is a putative autapomorphy of the family. The presence of unsclerotised paraglossae with dense fringes of thin hairs and the presence of a muscle connecting the anterior tentorial arm with the posterior edge of the sitophore plate are features shared by Xyelinae and members of non-xyelid families.

The affinities of mites and ticks: a review

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The genital morphology of Zorotypus hubbardi Caudell, 1918 (Insecta: Zoraptera: Zorotypidae)

The internal and external genitalia of both sexes of Zorotypus hubbardi Caudell 1918 were examined. A detailed description is provided, especially for the male genitalia. The results were compared with the conditions in potentially related neopteran groups. Most features of the internal genitalia of both sexes are plesiomorphic. Putative autapomorphies are: fusion of the distal parts of the vasa deferentia, division of the ductus ejaculatorius (de) into two sections by a distinct circular furrow, dorso-median processes on tergites VIII–X, an asymmetric aedoeagus with a cryptic slit-like phallotrema, reduction of the right outer paramere, the specific asymmetric shape of the endophallus with three conspicuous pouches, the endophallic sclerites scl1–scl6 and their specific arrangement, the specific arrangement of the endophallic muscularis and the unpaired genital muscles m2, m4, m5, m6 and m7. These features cannot be regarded as autapomorphies of the whole order Zoraptera with reliability, as the morphology of the external male genitalia seems to be highly variable within the genus Zorotypus and the intra-generic relationships are poorly resolved. Potential synapomorphies were found with several neopteran groups under comparison: with Embiidina a strengthened muscularis surrounding the de, reduction of the ovipositor and a shortened tergite IX, with Psocoptera the occurance of dense sperm bundles and with Hemiptera a strengthened muscularis of the de. No synapomorphies were found with Dictyoptera. Homologies of the muscles m1 and m3 found in Z. hubbardi with some genital muscles in Dictyoptera and Psocoptera are highly ambiguous.

Phylogenetic implications of the mesosomal skeleton in Chalcidoidea (Hymenoptera, Apocrita) – tree searches in a jungle of homoplasy

Results from a comparative anatomical study of the mesosomal skeleton of Chalcidoidea are presented. External and internal features are described and illustrated for 39 chalcidoid taxa, representing 16 families and 29 subfamilies. This is the most comprehensive morphological study ever conducted for the superfamily. The mesosoma was dissected, macerated and investigated using scanning electron microscopy. The mesothorax and metathorax contributed most of the phylogenetically relevant information. The metafurca is highly variable within Chalcidoidea but seems to be relatively constant at the subfamily level. One hundred and fifty-four morphological characters were scored and analysed cladistically. Outgroup species were chosen from six apocritan superfamilies: Stephanoidea, Ceraphronoidea, Cynipoidea, Platygastroidea, Proctotrupoidea and Mymarommatoidea. Some previously suggested chalcidoid relationships were retrieved: (1) Pteromalidae: Pteromalinae + Miscogasterinae + Panstenoninae; (2) Perilampidae + Eucharitidae; (3) Chalcididae + Leucospidae + Eurytomidae; (4) Eulophidae: Eulophinae + Tetrastichinae + Entedoninae; and (5) Eupelmidae + Encyrtidae. Mymarommatoidea renders Chalcidoidea paraphyletic in our analyses; however, the taxon sample is too restricted to provide a robust hypothesis. Three previously unreported putative autapomorphies of Chalcidoidea were revealed: (1) presence of an exposed, triangular or diamond-shaped prosternum; (2) presence of a percurrent mesopleural sulcus anteriorly terminating in the acropleuron; and (3) presence of paired metapectal plates lateral to the metafurca.

Trichomycterus maracaya, a new catfish from the upper rio Paraná, southeastern Brazil (Siluriformes: Trichomycteridae), with notes on the T. brasiliensis species-complex

Trichomycterus maracaya, a new species of Trichomycteridae, is described from a streamlet in the upper rio Paraná, Poços de Caldas, State of Minas Gerais, southeastern Brazil. The following putative autapomorphies distinguishes T. maracaya from congeneric species: 1) row of lateral blotches not forming a stripe at any phase during ontogeny; 2) superficial layer of pigmentation of juveniles and large (presumably adults) specimens consisting solely of scattered chromatophores. Furthermore, the new species is characterized by a combination of yellow ground color in life and mottled pattern formed by small to medium-sized, brown, irregularly-coalescent, well-defined deeper-lying blotches, and more superficial dots on the body. Trichomycterus maracaya is assigned to the T. brasiliensis species-complex (which includes T. brasiliensis, T. iheringi, T. mimonha, T. potschi, T. vermiculatus, and several undescribed species apparently endemic to the main river basins draining the Brazilian Shield) based on the presence of: 1) blotches in four longitudinal rows of deeper-lying pigmentation on the trunk large, horizontally-elongated, and well-defined; 2) pectoral fin with I+5-6 rays; 3) separation between the anterior and posterior cranial fontanels by the primordial epiphyseal cartilaginous bar being present only in larger specimens; and 4) pelvic-fin bases very close to each other, sometimes in contact.

The ovipositor apparatus of basal Hymenoptera (Insecta): phylogenetic implications and functional morphology

The skeleto‐musculature of the ovipositor apparatus and the external sculpture of the 1st and 2nd valvulae was studied in representatives from all ‘symphytan’ families. Nineteen informative characters were coded and scored. The distribution of character states are discussed with reference to recent cladistic treatments of the Hymenoptera. Putative autapomorphies of the Hymenoptera are the presence of cordate apodemes on T9 and basal articulations and associated musculature between the 2nd valvifers and the 2nd valvulae. It is a ground plan feature of the order to have the gonocoxites of abdominal segment 8 fused with the gonangula. The configuration of the musculature of the ovipositor apparatus did not display much variation among the taxa examined, except within the Pamphilioidea. There is considerable variation in the external ovipositor sculpture within the Tenthredinoidea. Putative synapomorphies for the tenthredinoid families except the Blasticotomidae are the presence of alternating strongly and weakly sclerotized zones on the first and/or second valvulae and the presence of serrulae on the sawteeth. The presence of transverse rows of large ctenidia on the 1st valvulae is an autapomorphy of the Diprionidae. Fusion of the 2nd valvifers and the 3rd valvulae is a synapomorphy for the Argidae + Pergidae. The ovipositor apparatus of the Pamphilioidea is highly derived, putative autapomorphies being the close association between T9 and the first valvifers, the reduction of the distal parts of the 1st valvulae, and the fusion of the 2nd valvulae for their entire length. The changes in the ovipositor apparatus of Pamphilioidea are associated with a decrease in the amount of work it has to perform during ovipositing, as the eggs are placed predominantly externally on the substrate. The ovipositor apparatus of the ‘Siricoidea’ is enlarged and modified for ovipositing into wood. Putative synapomorphies of the ‘Siricoidea’ + Orussidae + Apocrita are the presence of sawteeth only distally on the ovipositor and elongation of the cordate apodemes of T9.

Before the wasp-waist: comparative anatomy and phylogenetic implications of the skeleto-musculature of the thoraco-abdominal boundary region in basal Hymenoptera (Insecta)

The skeleto-musculature of the metathorax and first abdominal segment was studied in representatives from all ’symphytan’ families. Forty-three informative characters were coded and scored. The distribution of character states are discussed with reference to recent cladistic treatments of the Hymenoptera. Previously unreported autapomorphies for the Hymenoptera are the separation of the metathoracic trochantins from the metepisterna and metacoxae, the position of the metafurca anteriorly on the discrimenal lamella of the metathorax and the presence of second abdominal sternum (S2)-metacoxal muscles. The absence of metapleuro-S2 muscles is an autapomorphy for the non-xyelid Hymenoptera. Putative autapomorphies of the Tenthredinoidea are: (1) the presence of transverse metanotal muscles, (2) the subdivision of the second phragmo-third phragmal muscles, part of which arises from the metalaterophragmal lobes, (3) the posterior thoracic spiracle occlusor muscles arising from the mesepisterna, (4) the absence of trochantins and metanoto-trochantinal muscles and (5) the presence of elongate lateral metafurcal arms. Having the paracoxal sulci extending along the anterior margins of the metepisterna and the anterior metafurcal arms reduced are synapomorphies for all tenthredinoid families excluding Blasticotomidae. The presence of transversely extended cenchri with hooks on their entire surface is a putative synapomorphy for Diprionidae + Cimbicidae + Argidae + Pergidae. The clade Cimbicidae + Argidae + Pergidae is supported by the absence of metanoto- metabasalar muscles, the fusion of the first abdominal tergite (T1) with the metepimera and the absence of posterior metapleuro-metafurcal muscles. Autapomorphies of the Cimbicidae are the absence of the metalaterophragmal lobes and the metalaterophragmal-metafurcal muscles. Having the mesoscutello-metanotal muscle inserting on a projection from the anterior margin of the metanotum, surrounding the tendon with sclerotised cuticle, is a synapomorphy for the Argidae and Pergidae. Autapomorphies of the Cephoidea are the absence of cenchri, the presence of distinct articulations between T1 and the metepimera, and having the paracoxal sulci extending subparallel with the metafurcal discrimen. The monophyly of the Siricidae is supported by the absence of the anapleural clefts and the presence of an elongate mesospina projecting posteriorly between the anterior metafurcal arms. The presence of a membranous pouch ventrally of T1 and of large T1-metafurcal muscles is unique to Xiphydria camelus among the taxa examined. The absence of hind wing tegulae, posterior metapleuro-metafurcal, metanoto-trochantinal and anterior metanoto-metacoxal muscles, and the presence of elongate lateral metafurcal arms are synapomorphies for Xiphydriidae + Orussidae + Apocrita. The Orussidae greatly resembles the Apocrita in the region studied, a synapomorphy for the two taxa being the presence of metepisternal depressions. An autapomorphy for the Apocrita is the fusion of T1 with the metapleural arms; these structures closely abut in Orussidae. The fusion of T1 with the metepimera was preceded by the reduction of the posterior parts of the metepimera, as observed in Anaxyelidae, Xiphydriidae, and Orussidae. This makes the lines of fusion between T1 and the metepimera confluent with the metapleural sulci in the Apocrita. There is no compelling evidence for considering the configuration of T1 and the metepimera in Cephoidea to be incipient in the formation of the propodeum in Apocrita. The close association between the meso- and metathorax and the integration of T1 in the metathorax evolved gradually twice within the basal hymenopteran lineages, culminating in the Apocrita and the Cimbicidae + Argidae + Pergidae clade.

The Australian moth family Lophocoronidae and the basal phylogeny of the Lepidoptera – Glossata

The endemic Australian primitive moth family Lophocoronidae is reviewed. The family was previously known from three species represented by male 'museum' specimens only. The family now consists of one genus, Lophocorona Common, with six species of which three (L. robinsoni, L. commoni and L. flavicosta) are here described as new. L. robinsoni differs markedly from the remaining species in wing pattern and phenology, but all species have very similar male genitalia. Females of two species (L. robinsoni and L. commoni) are described. All species and parts of their male genitalia are illustrated; a key to all species is given. All new distribution records are listed and the known Australian range of the family now extends from east of Perth to south of Sydney. Lophocoronid structure is surveyed, including information on aspects of the soft anatomy of L. pediasia Common: cephalic, spiracular, abdominal base and male genital musculature, male internal genitalia, alimentary canal, gross structure of the central nervous system (CNS) and thoracic aorta. The most significant findings include the following: extrinsic labral muscles are absent; the relatively well-developed mandibles have no musculature, hence the (unknown) lophocoronid pupa must be adecticous; there is no intrinsic proboscis musculature; the posterolateral comer of the laterocervicale covers the anepisternal tooth; an anterior pronotal plate is present; the mesobasistemum is markedly produced anteriorly; wingsurface scales are largely hollow; a sizeable metapostphragma is present; the female has a piercing oviscapt similar to that of Eriocraniidae and Acanthopteroctetidae; a stomodaeal crop is well developed, extending into the abdomen, and followed by a narrow tubular portion in front of the mesenteron; there are four malpighian tubules, each opening into the gut; the deutocerebral lobes meet in front of the posterionnost pharyngeal sucking pump dilator (forming a 'deutocerebral loop'); the abdominal nerve cord has five ganglionic masses and thick connective tissue on top; the metathoracic aorta touches the dorsal pulsatile diaphragm. Six basal clades are recognised within the Lepidoptera-Glossata: (1) Eriocraniidae, (2) Acanthopteroctetidae (including Catapterix), (3) Lophocoronidae, (4) Neopseustidae, (5) Exoporia and (6) Heteroneura. Putative autapomorphies are listed and discussed for each. Several structural traits are compared throughout the six clades, and 47 potentially phylogenetically informative characters are identified (Appendices 1 and 2). Analysis of these characters with Hennig86, by using a hypothetical ancestor (reconstructed on the basis of character state distribution within the non-glossatan moth grade), yields a single shortest tree: Eriocraniidae + (Acanthopteroctetidae + (Lophocoronidae + (Neopseustidae + (Exoporia + Heteroneura)))). This tree is compared with a number of competing trees; it is concluded to be the most biologically meaningful one. The formal classification of the Glossata is discussed. The Acanthopteroctetidae are assigned to a superfamily of their own. Redundant taxon names above familygroup (Dacnonypha, Lophocoronina and Neopseustina) are discarded. The new name Coelolepida is introduced for the high-rank taxon comprising all Glossata except the Eriocraniidae; it is characterised primarily by the acquisition of hollow wing-surface scales and an apomorphic configuration of the first thoracic spiracle. Some ecological and conservation-related implications of the new insights in glossatan phylogeny are outlined.

CENTROSPERMAE REVISITED, PART I

SummaryIn preparation for continued phenetic and cladistic studies of taxonomic relationships of Centrospermae, 16 families are recognized in the order, and putative autapomorphies are designated for each where known, in order to support their monophyletic status. Polyphyly in Phytolaccaceae s.l. and paraphyly in Caryophyllaceae, Chenopodiaceae, and Portulacaceae are discussed. Pollen and floral characters used in an early study are revised and recoded. Recent molecular data bearing on centrosperm phylogeny are summarized.

The Carboniferous amphibianProterogyrinus ScheeleiRomer, and the early evolution of tetrapods

Proterogyrinus scheeleiRomer, from the Upper Mississippian of Greer, West Virginia, is one of the earliest known members of the am phibian infraorder Embolomeri. The primitive, eogyrinid-like skull conforms to Panchen’s allometric plot of the British eogyrinids. It has no identifiable autapomorphies, but shares with two other, as yet undescribed genera attributed to the same family, a unique skull table with a raised pineal foramenrim and an acuminate median ridge on the skull table posteriorly, flanked on each side by a deep depression. The kinetic junction extends from the otic notch anteriorly to the posterodorsal corner of the orbit. The braincase, bearing no ossified roof, is composed of separate otic-occipital and sphenethmoid units, the latter bearing no sagittal or parasagittal septa. The vertebrae are similar to those of other embolomeres, except that the pleurocentrum retains a dorsal suture in adults, and the intercentrum , unossified dorsally in adults, is a ventral cresent. The presacral count is 32. The atlas-axis is reptiliomorph. The limbs and girdles are similar to those ofArcheria,except that they are slightly stouter. Humerus structure is primitive. Range of limb movement, at least in the pectoral limb, is very restricted. The possession of few autapomorphies indicates that Proterogyrinusis a member of the stem family (Proterogyrinidae) of the infraorder Embolomeri. Although most of the putative autapom orphies of anthracosaurs that were formerly thought to preclude them from reptile ancestry are not considered to be valid, there are no undisputed synapomorphies with reptiles either, making it impossible to support close relationship between the two groups.