Study of the larva of Nosodendron fasciculare (Olivier 1790) (Coleoptera: Nosodendridae) with implications for the phylogeny of Bostrichiformia

Internal and external features of larvae of Nosodendron fasciculare were examined and compared to character states found in other groups of Derodontoidea and Bostrichoidea. Synapomorphic larval features indicate a sistergroup relationship between Nosodendridae and Derodontidae: tubercular surface structure, body compressed dorsoventrally, tergites with lateral projections, spiracles located on tubular processus. These families share three derived character states with Jacobsoniidae in one of two equally parsimonious trees. However, the monophyletic origin of Nosodendridae + Derodontidae + Jacobsoniidae is not sufficiently established at present. The monophyly of Bostrichoidea (Dermestidae + Bostrichidae + Anobiidae + Ptinidae) is suggested by hypognathism. Larvae of these families are characterized by the absence of the mandibular mola and a robust apical part of the mandible. The monophyly of Bostrichidae + Anobiidae + Ptinidae is indicated by a C-shaped, grub like body and the abdominal apex formed by an enlarged and rounded segment IX. Bostrychiformia are probably paraphyletic. A closer relationship between Bostrichoidea with Cucujiformia is suggested by the possession of cryptonephric malpighian tubules in adults. The specific type of cryptonephridism in Bostrichoidea is probably derived from this condition and is considered autapomorphic. The monophyly of Nosodendridae ( Nosodendron ) is supported by several autapomorphies. The assignment of the supposed larva of Nosodendron ovatum remains unclear. An inclusion of the dermestid genus Orphilus in Nosodendridae is rejected. Muscular features of larvae of Nosodendron (and Derodontus ) are largely plesiomorphic.     

Larval morphology and phylogenetic position of Micromalthus debilis LeConte (Coleoptera: Micromalthidae)

Abstract External and internal structures of the cerambycoid and triungulin larvae of Micromalthus debilis are described and compared to features found in larvae of other groups of Coleoptera. The morphological data are evaluated with respect to the systematic position of Micromalthidae. A cladistic analysis was carried out with fifty characters. Micromalthidae are not closely related to Lymexylidae (Polyphaga: Cucujiformia) but belong to Archostemata, which is confirmed as a monophyletic unit. Micromalthidae are specialized in terms of morphology and life cycle and are characterized by a considerable number of larval autapomorphies. Their sister-group relationship with Cupedidae is supported by several apomorphic features, which are probably correlated with xylobiontic habits: head transverse and strongly rounded laterally, absence of stemmata, shortened antennae, presence of sternal asperities and presence of eversible lobes of segment IX. Cupedidae is monophyletic and Priamca is the sister group of the remaining genera of Cupedidae included in the analysis. A closer relationship between Tenomerga and Rhipsideigma is supported by several larval synapomorphies. The ancestral life style of larvae of Archostemata was probably xylobiontic. This is suggested by derived groundplan features of the suborder, which are also found in larvae of non-related, wood-associated families.     

Phylogenetic analysis of Staphyliniformia (Coleoptera) based on characters of larvae and adults

Abstract. One hundred and twenty-one morphological characters of larvae and adults of the series Staphyliniformia were scored (multistate coding) and analysed to determine the family group relationships of the polyphagan groups Scarabaeoidea, Histeroidea, Hydrophiloidea and Staphylinoidea. Cladograms were rooted with exemplars of Adephaga, Archostemata, Myxophaga and the polyphagan families Dascillidae, Derodontidae, Eucinetidae and Scirtidae. Analyses of the same dataset with multistate characters re-coded as presence/absence (144 characters) produced cladograms that were similar to those produced from analyses of the original characters. Cladograms produced from partitioned larval and adult characters differed strongly, with adult-only trees more similar to those produced by combined data. The results confirm the monophyly of Hydrophiloidea + Histeroidea and of Staphylinoidea (including Hydraenidae). The Epimetopidae + Georissidae are the only strongly supported clade within Hydrophiloidea. A clade comprising Hydrochidae, Spercheidae and Hydrophilidae, and a sister-group relationship between the latter two families were confirmed in analyses of the data with presence/absence coding. Helophoridae, Epimetopidae and Georissidae are probably not a monophyletic unit, and additional evidence is needed for a reliable placement of Helophoridae. Scarabaeoidea are placed as a sister taxon of Hydrophiloidea + Histeroidea, but support for this relationship is weak. The branching pattern ((Hydraenidae + Ptiliidae) + (Leiodidae + Agyrtidae)), and a clade comprising Scydmaenidae, Silphidae and Staphylinidae (= ‘staphylinid group’) are well founded. The branching pattern (Orchymontiinae + (Prosthetopinae + (Ochthebiinae + Hydraeninae))) within Hydraenidae is confirmed. Poor resolution at the base of the trees and the placement of some nonstaphyliniform taxa (Dascillidae, Derodontidae, Scirtidae and Eucinetidae) as a sister group to a clade comprising Scarabaeoidea, Hydrophiloidea and Histeroidea suggests that Staphyliniformia may be paraphyletic. It is recommended that series names are eliminated from the classification of Polyphaga, at least for the more ‘primitive’ groups.     

The head morphology of Clambidae and its implications for the phylogeny of Scirtoidea (Coleoptera: Polyphaga)

External and internal structures of the head of adults of Clambus are described and illustrated in detail. The results are compared with structural features found in the clambid genus Calyptomerus, in representatives of other scirtoid families, and also in species of other coleopteran suborders, notably Myxophaga. The results tentatively support the monophyly of Scirtoidea and a close relationship between Clambidae and Eucinetidae is suggested by one shared derived feature of the mandible, a long and slender apical tooth with a serrate edge. The monophyly of Clambidae is very strongly supported and Acalyptomerus is probably the sistergroup of a clade Calyptomerus + Clambinae. Potential scirtoid autapomorphies are the loss of the dorsal tentorial arms, a bulging gula, a strongly transverse labrum, and a ridge separating the mediostipes from the lacinia. However, all these features are homoplasious. The monophyly of Clambidae is supported by modifications of the head capsule which is strongly flattened and broadened, by a deep clypeofrontal incision enabling vertical antennal movements, and a series of antennal features. Synapomorphies of Clambinae + Calyptomerus (Clambidae excluding Acalyptomerus) are the conglobate body form with the ventral side of the head capsule in contact with the mesocoxae, and compound eyes integrated in the contour of the head. The completely subdivided eye is an autapomorphy of Clambus. An entire series of features is shared by Clambidae (or Scirtoidea) and Myxophaga. Most of them are apomorphies that apparently evolved independently in both groups. However, the presence of well‐developed maxillary and labial glands is arguably a retained groundplan feature of Coleoptera, with parallel loss in Archostemata, Adephaga and various groups of Polyphaga. J. Morphol. 277:615–633, 2016. © 2016 Wiley Periodicals, Inc.     

Building the Coleoptera tree‐of‐life for >8000 species: composition of public DNA data and fit with Linnaean classification

The species representation of public databases is growing rapidly and permits increasingly detailed phylogenetic inferences. We present a supermatrix based on all gene sequences of Coleoptera available in Genbank for two nuclear (18S and 28S rRNA) and two mitochondrial (rrnL and cox1) genes. After filtering for unique species names and the addition of ?2000 unpublished sequences for cox1 and 18S rRNA, the resulting data matrix included 8441 species‐level terminals and 6600 aligned nucleotide positions. The concatenated matrix represents the equivalent of 2.17% of the 390 000 described species of Coleoptera and includes 152 beetle families. The remaining 29 families constitute small lineages with ?250 known species in total. Taxonomic coverage remains low for several major lineages, including Buprestidae (0.16% of described species), Staphylinidae (1.03%), Tenebrionidae (0.90%) and Cerambycidae (0.58%). The current taxon sampling was strongly biased towards the Northern Hemisphere. Phylogenetic trees obtained from the supermatrix were in very good agreement with the Linnaean classification, in particular at the family level, but lower for the subfamily and lowest for the genus level. The topology supports the basal split of Derodontidae and Scirtoidea from the remaining Polyphaga, and the broad paraphyly of Cucujoidea. The data extraction pipeline and detailed tree provide a framework for placement of any new sequences, including environmental samples, into a DNA‐based classification system of Coleoptera.     

The Infrabranchial Musculature and Its Bearing on the Phylogeny of Percomorph Fishes (Osteichthyes: Teleostei)

The muscles serving the ventral portion of the gill arches ( = infrabranchial musculature) are poorly known in bony fishes. A comparative analysis of the infrabranchial muscles in the major percomorph lineages reveals a large amount of phylogenetically-relevant information. Characters derived from this anatomical system are identified and discussed in light of current hypotheses of phylogenetic relationships among percomorphs. New evidence supports a sister-group relationship between the Batrachoidiformes and Lophiiformes and between the Callionymoidei and Gobiesocoidei. Investigated data also corroborate the existence of two monophyletic groups, one including the Pristolepididae, Badidae, and Nandidae, and a second clade consisting of all non-amarsipid stromateiforms. New synapomorphies are proposed for the Atherinomorphae, Blenniiformes, Lophiiformes, Scombroidei (including Sphyraenidae), and Gobiiformes. Within the latter order, the Rhyacichthyidae and Odontobutidae are supported as the successive sister families of all remaining gobiiforms. The present analysis further confirms the validity of infrabranchial musculature characters previously proposed to support the grouping of the Mugiliformes with the Atherinomorphae and the monophyly of the Labriformes with the possible inclusion of the Pholidichthyiformes. Interestingly, most hypotheses of relationships supported by the infrabranchial musculature have been advanced by preceding anatomists on the basis of distinct data sources, but were never recovered in recent molecular phylogenies. These conflicts clearly indicate the current unsatisfactory resolution of the higher-level phylogeny of percomorphs.     

The structure of the skull and jaw adductor musculature in the Gekkota, with comments on the phylogenetic relationships of the Xantusiidae (Reptilia: Lacertilia)

The skull and trigeminal jaw adductor musculature of the lizard families Gekkonidae, Pygopodidae and Xantusiidae are described. The external jaw adductor shows a different structure in the Gekkonidae and Pygopodidae than is observed in other lizards, approached only by the Xantusiidae and Feyliniidae. Paedomorphosis seems to be involved in the differentiation of the jaw adductor musculature in the Gekkonidae. The Gekkonidae and Pygopodidae may be hypothesized to form a monophyletic group, the Gekkota, on the basis of numerous synapomorphies. Within the Gekkota, the Pygopodidae are the sister-group of the Gekkonidae and retain some plesiomorphous features which are absent in the latter. The Xantusiidae share few synapomorphies with the Gekkota on the one hand, and some with scincomorph lizards on the other, especially with the Lacertidae.     

A phylogenetic study of Dermestidae (Coleoptera) based on larval morphology

Abstract.  The results of a phylogenetic analysis of Dermestidae (Coleoptera) based on the morphology of immature stages are presented and implications for the classification of the family are discussed. The monophyly of (( Orphilus ) + (Dermestidae) + (Endecatomidae (Bostrichidae + Anobiidae))) is strongly supported. Dermestidae, excluding Orphilus Erichson, forms a monophyletic group. The phylogenetic position of the Orphilinae remains enigmatic. The hypothesized relationships of the remaining dermestid subfamilies are (Thorictinae (Dermestinae (Attageninae (Megatominae + Trinodinae)))). Phylogenetically informative morphological characters of larvae and pupae are illustrated and described. Character state transformations implied by the phylogenetic hypothesis are discussed. New morphological observations are provided, including the first records of the presence of gastric caeca in the larva of Dermestes L., and the first report of the number of Malpighian tubules and the number of abdominal ganglia in Dermestidae, Bostrichidae, Anobiidae, Nosodendridae and Derodontidae. The form of the mandibular base is described for the first time for Thylodriini Beal and Trinodini Beal. The fossil record and distribution of extant Dermestidae are discussed in the light of the phylogenetic hypothesis. The origin of the family is hypothesized to be in the Late Cretaceous.     

Molecular phylogenetics of Elateriformia (Coleoptera): evolution of bioluminescence and neoteny

Phylogenetic relationships in the coleopteran Series Elateriformia (click beetles, jewel beetles, fireflies and allies) were investigated using > 3800 nucleotides of partial nuclear (small and large subunit rRNA genes) and mitochondrial (large subunit rRNA and cytochrome oxidase subunit I) gene sequences. The Elateriformia includes several soft‐bodied lineages, some of which retain larviform features in the adult stage (neoteny), and several major bioluminescent groups, including the families Lampyridae (fireflies), Phengodidae and Rhagophthalmidae whose relationships have been contentious. All recognized superfamilies (Elateroidea, Cantharoidea, Byrrhoidea, Buprestoidea, Dascilloidea, Scirtoidea) and 28 of the 37 families, represented in 112 individuals, were included in the analysis. Sequence alignment was based on static and dynamic homology assignments and partial removal of sequences of uncertain homology. Alignment variable regions caused a great deal of uncertainty but also contributed much of the phylogenetic signal that was insufficient to resolve deep relationships when these were removed. The main features of most analyses were the monophyly of Elateroidea + Cantharoidea (= Elateroidea sensu lato), with Omethidae + Telegeusidae frequently occupying the basal node in this group; the affinities of Dascilloidea, Buprestoidea and a (broadly paraphyletic) Byrrhoidea, with unclear relationships among them; and the monophyly of Scirtoidea (including Decliniidae) as a rather distant outgroup to all others. When mapped on the resulting trees, soft‐bodied lineages were polyphyletic, contradicting the former Cantharoidea that had been united by this trait. Transitions to neoteny were either simultaneous with, or subsequent to, the origin of soft‐bodiedness in a minimum of seven lineages. The bioluminescent groups Lampyridae (including the enigmatic genus Drilaster) and the tightly allied Phengodidae + Rhagophthalmidae were never monophyletic. The former showed close relationship to the species‐rich, soft‐bodied families Lycidae and Cantharidae, while the latter grouped with poorly resolved lineages at the base of Elateridae (click beetles). Hence, although key features as soft‐bodiedness, neoteny and bioluminescence in Coleoptera are largely confined to the Elateriformia, they appear to result from multiple origins, showing the propensity of closely related lineages to acquire similar features independently. © The Willi Hennig Society 2007.     

Facial and gill musculature of polynemid fishes,with notes on their possible relationships with sciaenids (Percomorphacea: Perciformes)

The Polynemidae is a family of primarily marine fishes with eight genera and 42 extant species. Many aspects of their morphology are largely unknown, with few reports about their osteology and barely any information on their myology. This paper describes and illustrates in detail all facial and branchial muscles of representative species of polynemids. Our analysis demonstrates the existence of several remarkable and previously unknown specializations in the polynemid musculature. The aponeurotic and completely independent origin of the pars promalaris of the adductor mandibulae is apparently unique among percomorphs. The differentiation of this section into lateral and medial subsections; the total separation of the promalaris from the retromalaris; the differentiation of the pars primordialis of the levator arcus palatini into external and internal subsections are also uncommon features of polynemids that are shared by sciaenids, thus supporting the hypothesis of a closer relationship between these families.     

The musculature of Squatinella rostrum (Milne, 1886) (Rotifera: Lepadellidae) as revealed by confocal laser scanning microscopy with additional new data on its trophi and overall morphology

Wilts, E.F., Wulfken, D., Ahlrichs, W.H. and Martínez Arbizu, P. 2012. The musculature of Squatinella rostrum (Milne, 1886) (Rotifera: Lepadellidae) as revealed by confocal laser scanning microscopy with additional new data on its trophi and overall morphology.—Acta Zoologica (Stockholm) 93 : 14–27. The monogonont rotifer Squatinella rostrum was investigated with light, scanning electron and confocal laser scanning microscopy to reveal new morphological data on its inner and outer anatomy. In total, the visualized somatic musculature displays five paired longitudinal muscles (musculi longitudinales I–V) and nine circular muscles (musculi circulares I–IX). Compared to other species, S. rostrum is characterized by the absence of several longitudinal and circular muscles (e.g. musculus longitudinalis capitis, corona sphincter and pars coronalis). A reconstruction of the mastax musculature revealed a total number of seven paired and two unpaired mastax muscles. Possibly homologous somatic and mastax muscles in other, thus far investigated rotifers are discussed. Moreover, we provide a phylogenetic evaluation of the revealed morphological characters and suggest possible autapomorphic characters supporting Squatinella and Lepadellidae. Finally, we refer to some striking similarities in the morphology, ecology and way of movement of Squatinella and Bryceella that may indicate a closer relationship of both taxa.     

Study on the systematic position of Metriini based on characters of the larval head (Coleoptera: Carabidae)

Abstract. Characters of the head of larvae of Metrius contractus Eschscholtz, Ozaenini and Paussini are interpreted phylogenetically. The monophyly of Metriini + Ozaenini + Paussini is substantiated by several synapomorphies such as hyperprognathism and strong constriction of the neck. Ozaenini and Paussini together form the sister-group of Metriini. Ozaenini are paraphyletic. The monophyly of Paussini + Ozaenini excluding Pachyteles is indicated by two possible synapomorphies. Several synapomorphies are shared by Physea + Paussini. Secondary prognathism, large membranous submento-mental area and other derived features are considered autapomorphies of Paussini. Paussini excluding Platyrhopalopsis are characterized by the loss of the palpifer. The monophyly of a group which comprises Geadephaga excluding Trachypachini is suggested by several synapomorphic features. A very basal position of the metriine—paussine lineage within Carabidae is indicated by several plesiomorphic features. A hypopharyngeal filter apparatus with a dense fringe of well-arranged, long hairs is a possible autapomorphy of Anisochaeta. The results of this study do not indicate a close relationship between the metriine—paussine lineage and the tribes Brachinini and Crepidogastrini as has been suggested in recent works.     

Molecular Phylogenetics at the Felsenstein Zone: Approaching the Strepsiptera Problem Using 5.8S and 28S rDNA Sequences

Recent efforts to reconstruct the phylogenetic position of the insect order Strepsiptera have elicited a major controversy in molecular phylogenetics. We sequenced the 5.8S rDNA and major parts of the 28S rDNA 5′ region of the strepsipteran speciesStylops melittae.Their evolutionary dynamics were analyzed together with previously published insect rDNA sequences to identify tree estimation bias risks and to explore additional sources of phylogenetic information. Several major secondary structure changes were found as being autapomorphic for the Diptera, the Strepsiptera, or the Archaeognatha. Besides elevated substitution rates a significant AT bias was present in dipteran and strepsipteran 28S rDNA which, however, was restricted to stem sites in the Diptera while also affecting single-stranded sites in the Strepsiptera. When dipteran taxa were excluded from tree estimation all methods consistently supported the placement of Strepsiptera to within the Holometabola. When dipteran taxa were included maximum likelihood continued to favor a sister-group relationship of Strepsiptera with Mecoptera while remaining methods strongly supported a sister-group relationship with Diptera. Parametric bootstrap analysis revealed maximum likelihood as a consistent estimator if rate heterogeneity across sites was taken into account. Though the position of Strepsiptera within Holometabola remains elusive, we conclude that the evolution of dipteran and strepsipteran rDNA involved similar yet independent changes of substitution parameters.     

The larval head of Raphidia (Raphidioptera, Insecta) and its phylogenetic significance

External and internal head structures of larval representatives of Raphidiidae are described. The obtained data were compared to characters of other neuropterid larvae and to larval characters of representatives of other endopterygote lineages. Characters potentially relevant for phylogenetic reconstruction are listed and discussed. The larvae of Raphidioptera differ distinctly from other neuropterid larvae in their morphology. They are mainly characterised by autapomorphic and plesiomorphic character states and few features indicate systematic affinities with other groups. Endopterygote groundplan features maintained in Raphidioptera are the complete tentorium, the free labrum, the full set of labral muscles, the presence of four extrinsic antennal muscles, the three-segmented labial palpi, the presence of a full set of extrinsic maxillary and labial muscles, the presence of a salivarium, and possibly the high number of stemmata. Apomorphies likely correlated with predaceous habits are the long gula, the protracted maxillae, the longitudinal arrangement of extrinsic maxillary muscles, and the elongated prepharyngeal tube. Highly unusual, potentially autapomorphic features are the presence of a dorsal ligament of the tentorium and paired gland-like structures below the pharynx. A prognathous or very slightly inclined head and slender mandibles without mola are features shared by larvae of all orders of Neuropterida. The parallel-sided head is a potential synapomorphy of Raphidioptera and Megaloptera. A fully prognathous head with anteriorly shifted posterior tentorial grooves and the presence of a parietal ridge and a distinct neck region are features shared with Corydalidae. Characters of the larval head are not sufficient for a reliable placement of Raphidioptera.     

Remarks on the family-level phylogeny of butterflies (Insecta, Lepidoptera, Rhopalocera)

1. Available evidence on butterfly family-level relationships is re-examined according to the principles of phylogenetic (cladistic) systematics. 2. The assumption of a sister-group relationship between the Hesperioidea and Papilionoidea seems a reasonably substantiated working hypothesis. 3. The Papilionoid families Papilionidae, Pieridae and Lycaenidae sensu Ehrlich (1958) are definable as monophyletic entities; of Ehrlich 's two remaining families, Nymphalidae and Libytheidae, the former is paraphyletic in terms of the latter. 4. The interrelationships between the Papilionoid families may be presented as Papilionidae + (Pieridae + [Lycanidae + Nymphalidae]). 5. In a phylogenetic system any given arrangement of taxa is either correct or not: Contrary to the pheneticists' view (Ehrlich and Ehrlich 1967) phylogenetic systematists cannot accept the existence of a multitude of valid classifications.     

Phylogenetic position of Apostasia and Neuwiedia (Orchidaceae)

JUDD, W. S., STERN, W. L. & CHEADLE, V. I. 1993. Phylogenetic position of Apostasia and Neuwiedia (Orchidaceae). Cladistic analyses of the phylogenetic relationships of selected orchid taxa were conducted in order to assess the phylogenetic position of Apostasia and Neuwiedia (Orchidaceae: Apostasioideae). These analyses employed newly available anatomical characters, along with several morphological features that had been used in recent phylogenetic analyses of Orchidaceae. Our analyses indicate that Apostasia is more closely related to Neuwiedia than it is to Cypripedioideae. The two genera comprise an apostasiad clade; this clade is the sister-group to a clade including Cypripedioideae and monandrous orchids. The apostasiad clade is diagnosed by the derived features of operculate pollen colpi, Apostasia-type seeds, and vessel members with simple perforation plates. Of these, the presence of simple perforation plates is considered to be the most significant phylogenetically. Therefore, the apostasiads should not be considered ancestral to the remaining orchid groups. Vessel members of the monandrous orchids, as well as the cypripediads, are multiperforate–the hypothesized ancestral state based on the condition in Hypoxidaceae.     

Floral anatomy of Thomandersia (Lamiales), with particular reference to the nature of the retinaculum and extranuptial nectaries

Thomandersia is an isolated genus that is currently unplaced in Lamiales. In the past it has been classified in Acanthaceae or Pedaliaceae, on the basis of intuitive assessments of a limited number of morphological characters. Recent molecular sequenced-based phylogenies have suggested a relationship to Bignoniaceae, Schlegeliaceae or Verbenaceae. Here we present new observations of the floral anatomy of Thomandersia , with particular emphasis on the structure of the retinaculum, a character shared with Acanthaceae, and calyx nectaries, which may be shared with other families in Lamiales. The morphological and anatomical characters of Thomandersia are discussed in the context of recent phylogenetic hypotheses for Lamiales, with the aim of identifying potential primary and secondary homologies between Thomandersia and related families in Lamiales. We find that Thomandersia shares a range of characters with each of the families to which it might be sister-group, and that some of these primary homologies must therefore be homoplastic. In particular, if the topology based on molecular sequence data is correct, the retinacula of Thomandersia and Acanthaceae are homoplastic and represent an example of parallel morphological evolution.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 469–482.     

Phylogenetic position of the monogeneans Sundanonchus, Thaparocleidus, and Cichlidogyrus inferred from 28S rDNA sequences

A molecular phylogeny was inferred from newly obtained partial 28S rRNA gene sequences of Sundanonchus micropeltis (Sundanonchidae), Thaparocleidus siamensis and Cichlidogyrus sp. (Ancyrocephalidae), and other already available sequences. Although sequences are lacking for several families, the following phylogenetic relationships could be inferred. The Diplectanidae were the sister-group to a clade including Sundanonchus and the Ancyrocephalidae; Sundanonchus was the sister-group to the Ancyrocephalidae, therefore suggesting validity of the Sundanonchidae, which include this single genus; within the Ancyrocephalidae, Thaparocleidus (Ancylodiscoidinae) was the sister-group to the four other taxa, though with relatively low support, suggesting that the Ancylodiscoidinae are the sister-group to the Ancyrocephalinae.