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.     

A phylogenetic hypothesis for Asilidae based on a total evidence analysis of morphological and DNA sequence data (Insecta: Diptera: Brachycera: Asiloidea)

An hypothesis of phylogenetic relationships of Asilidae and its constituent taxa is presented, combining morphological and DNA sequence data in a total evidence framework. It is based on 77 robber fly species, 11 Asiloidea outgroup species, 211 morphological characters of the adult fly, and approximately 7300 bp of nuclear DNA from five genes (18S and 28S rDNA, AATS, CAD, and EF-1α protein-encoding DNA). The equally weighted, simultaneous parsimony analysis under dynamic homology in POY resulted in a single most parsimonious cladogram with a cost of 27,582 (iterative pass optimization; 27,703 under regular direct optimization). Six of the 12 included subfamily taxa are recovered as monophyletic. Trigonomiminae, previously always considered as monophyletic based on morphology, is shown to be non-monophyletic. Two of the three Trigonomiminae genera, Holcocephala Jaennicke, 1867 and Rhipidocephala Hermann, 1926, group unexpectedly as the sister taxon to all other Asilidae. Laphriinae, previously seen in the latter position, is the sister group of the remaining Asilidae. Five other subfamily taxa, i.e. Brachyrhopalinae, Dasypogoninae, Stenopogoninae, Tillobromatinae, and Willistonininae, are also shown to be non-monophyletic. The phylogenetic relationships among the higher-level taxa are partly at odds with findings of a recently published morphological study based on more extensive taxon sampling. The total evidence hypothesis is considered as the most informative one, but the respective topologies from the total-evidence, morphology-only, and molecular-only analyses are compared and contrasted in order to discuss the signals from morphological versus molecular data, and to analyze whether the molecular data outcompete the fewer morphological characters. A clade Apioceridae+Mydidae is corroborated as the sister taxon to Asilidae.     

A comparative study on mouthpart morphology of certain larvae of Chironomini (Diptera: Chironomidae) with reference to the larval feeding habits

Mouthparts of six species of chironomid larvae were compared on both an intra- and inter-specific level. The species were: Chironomus riparius, Cryptochironomus defectus gr., Endochironomus albipennis, Glyptotendipes pallens, Microtendipes pedellus and Parachironomus arcuatus . The mouthpart morphology in C. defectus gr. and P. arcuatus is considerably different from the four other species and changes little between instars. Within C. riparius, E. albipennis, G. pallens and M. pedellus there is a considerable difference in the mouthpart morphology between the first instar and the remaining instars. The morphology of the mouthparts of the first instar of C. riparius, E. albipennis, G. pallens and M. pedellus resembles in many ways that found in C. defectus gr. and P. arcuatus .     

Morphology and function of the proboscis in Bombyliidae (Diptera, Brachycera) and implications for proboscis evolution in Brachycera

Based on serial semithin sections and SEM photographs of representatives of European Bombyliinae and Anthracinae, the mouthparts of Bombyliidae are studied and compared with the relevant data from literature on other families of Diptera Brachycera. The three moving units of the proboscis (clypeo-cibarial region, haustellum-maxillary base region, and labella) and their structures and muscles are described. Functions and possible movements are inferred from the structures observed. Articulations both between the parts of the organ and to the head capsule enable the fly to retract its proboscis into a resting position. Proboscis movement from a resting to a feeding position encompasses the following submovements: rotating of the basal clypeo-cibarial region (= fulcrum) against the head capsule, folding of the haustellum-maxillary base region against the fulcrum, evagination and invagination of the labial base, and the labella movements. This is a novelty as compared to the rigid proboscis of Tabanidae and agrees largely with the conditions in the Cyclorrhapha. The evolution of these novelties and their functional significance are discussed. The fulcrum, as well as the haustellum-maxillary base, as the new moving units are deduced from the plesiomorphic state as present in Tabanidae by fusions of sclerites, shifts of musculature and formation of new articulations. Accepted: 5 April 2000     

A phylogenetic analysis of Coelopidae (Diptera) based on morphological and DNA sequence data

The phylogenetic relationships of 22 species of Coelopidae are reconstructed based on a data matrix consisting of morphological and DNA sequence characters (16S rDNA, EF-1alpha). Optimal gap and transversion costs are determined via a sensitivity analysis and both equal weighting and a transversion cost of 2 are found to perform best based on taxonomic congruence, character incongruence, and tree support. The preferred phylogenetic hypothesis is fully resolved and well-supported by jackknife, bootstrap, and Bremer support values, but it is in conflict with the cladogram based on morphological characters alone. Most notably, the Coelopidae and the genus Coelopa are not monophyletic. However, partitioned Bremer Support and an analysis of node stability under different gap and transversion costs reveal that the critical clades rendering these taxa non-monophyletic are poorly supported. Furthermore, the monophyly of Coelopidae and Coelopa is not rejected in analyses using 16S rDNA that was manually aligned. The resolution of the tree based on this reduced data sets is, however, lower than for the tree based on the full data sets. Partitioned Bremer support values reveal that 16S rDNA characters provide the largest amount of tree support, but the support values are heavily dependent on analysis conditions. Problems with direct comparison of branch support values for trees derived using fixed alignments with those obtained under optimization alignment are discussed. Biogeographic history and available behavioral and genetic data are also discussed in light of this first cladogram for Coelopidae based on a quantitative phylogenetic analysis.     

The adult head structures of Tipulomorpha (Diptera,Insecta) and their phylogenetic implications

Schneeberg, K. and Beutel, R.G. 2011. The adult head structures of Tipulomorpha (Diptera, Insecta) and their phylogenetic implications. —Acta Zoologica (Stockholm) 92 : 316–343. Head structures of adults of Tipula paludosa, Limonia sp. and Trichocera saltator were examined and described. The results are compared with conditions found in other dipterans and other antliophoran groups, notably Nannochoristidae. Several potential synapomorphies of a dipteran–nannomecopteran–siphonapteran clade are present in Tipuloidea and Trichocera, the labro‐epipharyngeal food channel, the loss of the galea and the postpharyngeal pumping apparatus. The sensorial field of the maxillary palpomere 3, a potential dipteran–nannomecopteran synapomorphy, is also present but modified. The presence of M. clypeolabralis, labellae and mandibular stylets are groundplan apomorphies of Diptera, with secondary loss of the mandibles in Tipuloidea, Trichoceridae and many other groups. Tipuloidea is supported by the origin of M. tentorioscapalis anterior on the head capsule, the reduction of M. frontobuccalis anterior and the loss of the ocelli. The reduced tentorium, the origin of two further antennal muscles on the head capsule, the maxillary sensorial field with sensilla in individual pits, the lacking dorsal prelabial concavity and the unpaired salivary channel entering the head are apomorphies of Tipulidae. Closer affinities of Tipulidae and Cylindrotomidae are suggested by pseudotracheae of the advanced type, which have evolved independently in this lineage. The results do neither support a basal placement of Tipuloidea nor close affinities with Brachycera.     

Molecular phylogenetic analysis of nycteribiid and streblid bat flies (Diptera: Brachycera, Calyptratae): implications for host associations and phylogeographic origins

Bat flies are a small but diverse group of highly specialized ectoparasitic, obligatory bloodsucking Diptera. For the first time, the phylogenetic relationships of 26 species and five subfamilies were investigated using four genes (18S rDNA, 16S rDNA, COII, and cytB) under three optimality criteria (maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference). Tree topology tests of previous hypotheses were conducted under likelihood (Shimodaira-Hasegawa test). Major findings include the non-monophyly of the Streblidae and the recovery of an Old World- and a New World-Clade of bat flies. These data ambiguously resolve basal relationships between Hippoboscidae, Glossinidae, and bat flies. Recovered phylogenies resulted in either monophyly (Bayesian approach) or paraphyly (MP/ML topologies) of the bat flies, thus obscuring the potential number of possible associations with bats throughout the history of this group. Dispersal-vicariance analysis suggested the Neotropical region as the possible ancestral distribution area of the New World Streblidae and the Oriental region for the Old World bat flies. The genes examined show conflicting support across the nodes of the tree, particularly in the basal positions. Additionally, there is poor character support among all genes for the nodes associated with early hippoboscoid diversification. This results in extremely short basal branches, adding support to the idea of a rapid radiation among the four major groups of Hippoboscoidea.     

A phylogenetic analysis of relationships among genera of Conopidae (Diptera) based on molecular and morphological data

Members of the family Conopidae (Diptera) have been the focus of little targeted phylogenetic research. The most comprehensive test of phylogenetic support for the present subfamily classification of Conopidae is presented here using 66 specimens, including 59 species of Conopidae and seven outgroup taxa. Relationships among subfamily clades are also explored. A total of 6824 bp of DNA sequence data from five gene regions (12S ribosomal DNA, cytochrome c oxidase subunit I, cytochrome b, 28S ribosomal DNA and alanyl‐tRNA synthetase) are combined with 111 morphological characters in a combined analysis using both parsimony and Bayesian methods. Parsimony analysis recovers three shortest trees. Bayesian analysis recovers a nearly identical tree. Five monophyletic subfamilies of Conopidae are recovered. The rarely acknowledged Zodioninae is restored, including the genera Zodion and Parazodion. The genus Sicus is removed from Myopinae. Morphological synapomorphies are discussed for each subfamily and inter‐subfamily clade, including a comprehensive review of the character interpretaions of previous authors. Included are detailed comparative illustrations of male and female genitalia of representatives of all five subfamilies with new morphological interpretation.     

A phylogenetic analysis of Sciomyzidae (Diptera) and some related genera

Sciomyzidae is a family of acalyptrate flies with 546 species in 61 genera that is among the most extensively studied groups of higher Diptera. Most of the known larvae are obligate enemies of Gastropoda. Hundreds of studies published over the past 50 years have resulted in detailed information concerning morphology of adults and immature stages, biology, development, behaviour, phenology and distribution. However, studies of phylogenetic relationships are based almost exclusively on morphological characters of adults, and no comprehensive molecular analysis across the family has been published. Here we fill this void by generating and analysing molecular data for 54 species of Sciomyzidae (22 genera), including Phaeomyiidae (one genus), and seven representative species of five other families of Sciomyzoidea (Coelopidae, Dryomyzidae, Helcomyzidae, Heteromyzidae and Huttoninidae) as outgroups. The reconstruction is based on morphological characters as well as nucleotide sequences for genes from the mitochondrial (12S, 16S, COI, COII, Cytb) and nuclear genome (28S, EF1α). The results are compared with recent morphological analyses. Our analyses support the monophyly of Sciomyzidae + Phaeomyiidae, and place Phaeomyiinae as a unique lineage within Sciomyzidae. A modified classification comprising three subfamilies is proposed. The major subfamily, Sciomyzinae, consists of two monophyletic and well separated groups, the tribes Sciomyzini and Tetanocerini.     

The larval head anatomy of Rhyacophila (Rhyacophilidae) with discussion on mouthpart homology and the groundplan of Trichoptera

The external and internal features of the larval head of Rhyacophila fasciata (Trichoptera: Rhyacophilidae) were described in detail. Anatomical examinations were carried out using a multimethod approach including histology, scanning electron microscopy, confocal laser‐scanning microscopy, microcomputed tomography, and computer‐based three‐dimensional reconstructions. Additionally, the information on the larval head of Limnephilus flavicornis (Limnephilidae) and Hydropsyche angustipennis (Hydropsychidae) available in the literature were reinvestigated. These anatomical data were used to address major questions of homology and terminology, that is, the ventral closure of the head capsule, the sclerites, and appendages of labium and maxilla and their muscles. These topics were discussed by summarizing the main hypotheses present in the literature and a critical inclusion of new findings. Consequently, the inner lobe of the maxilla very likely represents the galea. The distal maxillary sclerite (palpifer) is an anatomical composite formation at least including dististipes and lacinia. Based on these homology hypotheses several potential groundplan features of the larval head of Trichoptera were reconstructed. The head of Rhyacophila shows several presumably plesiomorphic features as for instance the prognath orientation of the mouthparts, the well‐developed hypocranial bridge, the triangular submentum and eyes composed of seven stemmata. Derived features of Rhyacophila are the reduced antennae, the anterior directing of three stemmata and the shift of the tentorio‐stipital muscle to the mentum. J. Morphol. 276:1505–1524, 2015. © 2015 Wiley Periodicals, Inc.     

Relationships of extant lower Brachycera (Diptera): a quantitative synthesis of morphological characters

With over 80 000 described species, Brachycera represent one of the most diverse clades of organisms with a Mesozoic origin. Larvae of the majority of early lineages are detritivores or carnivores. However, Brachycera are ecologically innovative and they now employ a diverse range of feeding strategies. Brachyceran relationships have been the subject of numerous qualitative analyses using morphological characters. These analyses are often based on characters from one or a few character systems and general agreement on relationships has been elusive. In order to understand the evolution of basal brachyceran lineages, 101 discrete morphological characters were scored and compiled into a single data set. Terminals were scored at the family level, and the data set includes characters from larvae, pupae and adults, internal and external morphology, and male and female terminalia. The results show that all infraorders of Brachycera are monophyletic, but there is little evidence for relationships between the infraorders. Stratiomyomorpha, Tabanomorpha, and Xylophagomorpha together form the sister group to Muscomorpha. Xylophagomorpha and Tabanomorpha are sister groups. Within Muscomorpha, the paraphyletic Nemestrinoidea form the two most basal lineages. There is weak evidence for the monophyly of Asiloidea, and Hilarimorphidae appear to be more closely related to Eremoneura than other muscomorphs. Apsilocephalidae, Scenopinidae and Therevidae form a clade of Asiloidea. This phylogenetic evidence is consistent with the contemporaneous differentiation of the main brachyceran lineages in the early Jurassic. The first major radiation of Muscomorpha were asiloids and they may have diversified in response to the radiation of angiosperms in the early Cretaceous.     

The phylogeny and systematics of European predacious Syrphidae (Diptera) based on larval and puparial stages

The external larval morphology of predacious Syrphidae is examined and compared with that of other groups of syrphids. Predacious larvae form a distinctive, monophyletic group. Relationships among 29 of the 34 European genera and subgenera are analysed using cladistic methods based on larval and puparial characters. Interpretation of the cladogram suggests that, within Syrphinae, a system of five tribes can be recognized and that a major trend has been a shift from ground layer to arboreal larval habitats, with an increasing degree of obligate dependence on soft-bodied Homoptera as prey. Generic diagnoses with biological and taxonomic notes are given and a key to genera based on larvae and puparia is provided.     

Molecular Evolution and Phylogenetic Utility of the Internal Transcribed Spacer 2 (ITS2) in Calyptratae (Diptera: Brachycera)

The resolution potential of internal transcribed spacer 2 (ITS2) at deeper levels remains controversial. In this study, 105 ITS2 sequences of 55 species in Calyptratae were analyzed to examine the phylogenetic utility of the spacer above the subfamily level and to further understand its evolutionary characteristics. We predicted the secondary structure of each sequence using the minimum-energy algorithm and constructed two data matrixes for phylogenetic analysis. The ITS2 regions of Calyptratae display strong A-T bias and slight variation in length. The tandem and dispersed repeats embedded in the spacers possibly resulted from replication slippage or transposition. Most foldings conformed to the four-domain model. Sequence comparison in combination with the secondary structures revealed six conserved motifs. Covariation analysis from the conserved motifs indicated that the secondary structure restrains the sequence evolution of the spacer. The deep-level phylogeny derived from the ITS2 data largely agreed with the phylogenetic hypotheses from morphologic and other molecular evidence. Our analyses suggest that the accordant resolutions generated from different analyses can be used to infer deep-level phylogenetic relations.     

Musculature of the male genitalia of Xylophagus cinctus (De Geer) and relationships within the superfamily Xylophagoidea (Diptera, Brachycera)

There is no general agreement on relationships within Xylophagoidea (Diptera, Brachycera). The musculature of the male genitalia of Xylophagus cinctus (De Geer) (Xylophagidae, the most primitive family of Brachycera) is described and compared with that of some other Xylophagoidea: Exeretonevridae (Exeretonevra angustifrons Hardy), Coenomyiidae (Anacanthaspis biafasciata Röder), and Rhagionidae (Rhagio montanus Becker, Chrysopilus dives Loew, and Ch. helvolus Meigen) discussed earlier (Ovtshinnikova, 1989, 1998; Palmer et al., 2000). In spite of the differences in the structure of the genital sclerites, Xylophagidae possess all the muscles found in Coenomyiidae and Rhagionidae. The musculature of the male genitalia of Xylophagus cinctus includes two muscle pairs of the aedeagus sheath (M1 and M2); three muscle pairs of the ejaculatory complex (M30, M31, and M32); one muscle pair of the gonocoxites (M33); two muscle pairs of the gonostyli (M27 and M28); one muscle pair of the proctiger (M21), one muscle pair of the cerci (M29); two pairs of the tergosternal muscles (M5 ¹ and M5 ²); and two pairs of the pregenital muscles (M18 and M19). Muscles of the family Exeretonevridae are mostly the same, except for the muscles of the cerci M29, proctiger M29, and pregenital muscles M18 and M19, that are subdivided into two parts. This fact and also a different degree of the development of muscles M32 and M5 ² clearly distinguish Exeretonevridae from closely related families. The attachment places of the muscles of the aedeagus sheath M2 and of the gonostyli M28, as well as the split character of the tergosternal muscle pair M5 ¹ makes it possible to distinguish two sister groups, Xylophagidae plus Exeretonevridae, versus Coenomyiidae plus Rhagionidae. It should be noted that the muscles of the male genitalia of Xylophagidae, Exeretonevridae, Coenomyiidae, and Rhagionidae possess similar plesiomorphic characters, and these families should be united into the superfamily Xylophagoidea. This superfamily is the most primitive superfamily of Brachycera Orthorrhapha and possesses the most stable set and arrangement of male genital muscles within the entire suborder. An improved dendrogram of the phylogenetic relationships between the known groups of Xylophagoidea is proposed on the basis of the structure of male genital muscles.     

Differing strategies of patterning of follicular cells in higher and lower brachycerans (Diptera: Brachycera)

In all higher dipterans (Brachycera), including the fruitfly, Drosophila melanogaster, each egg chamber (ovarian follicle) consists of a group (clone) of germ cells (one oocyte and 15 accompanying nurse cells) that is surrounded by a layer of somatic mesodermal follicular cells (FCs). As oogenesis progresses the initially uniform FCs diversify into several morphologically and functionally distinct subpopulations. In D. melanogaster some of these subpopulations, e.g., border, centripetal, and dorsolateral cells, undertake coordinated migration or rearrangement over the surface of the germ cells. During the final stages of oogenesis these subpopulations participate in the formation of a complex, regionally specialized eggshell. In representatives of lower brachycerans (Orthorrhapha), only FCs that undertake active, directed migration are the border cells. These cells originate at the anterior pole of the ovarian follicle and migrate between the nurse cells to the anterior pole of the oocyte. Reduced motility of FCs in lower brachycerans results in the absence of certain FC subpopulations in their egg chambers and subsequent simplicity of their eggshells. We found that the lack of some FC subpopulations coincided with the appearance of lamellipodium-like protrusions of the oocyte. These protrusions penetrated between the apposing membranes of nurse and FCs and partially enveloped the nurse cell compartment. Analysis of whole-mount preparations stained with rhodamine-conjugated phalloidin revealed that the protrusions contained microfilaments and that their tips were equipped with actin-rich filopodium-like processes. We also found that in some lower brachycerans (representatives of the family Rhagionidae), the FCs located at the posterior pole of the oocyte, became enlarged and morphologically similar to the anterior border cells. These findings indicate that in higher dipterans the processes leading to the formation of a functional egg are variable and often markedly different from those in the model organism, D. melanogaster.     

Analysis of the taxonomic structure of the crane fly family Limoniidae (Diptera) based on the larval characters

A morphological study of the larval characters in the genera Elephantomyia, Helius, Microlimonia, Lipsothrix, and Teucholabis was conducted. Significant differences between Elephantomyia and Helius were discovered; Elephantomyia should be placed in the subfamily Hexatominae, while Helius cannot be assigned to any known subfamily. The genus Microlimonia has nothing in common with other representatives of the tribe Limoniini, and the genera Lipsothrix and Teucholabis, with those of Gonomyini; these genera cannot be included in any of the existing tribes. The presently accepted subdivision of the Limoniidae into 4 subfamilies does not reflect its actual diversity.     

Structure and phylogenetic interpretation of diplopod eyes (Diplopoda)

Summary The structural organization of the ocelli of several diplopod species has been studied by means of electron microscopy. The results provide evidence that diplopodan ocelli are derived from typical mandibulate ommatidia, which consequently had been present in diplopod ancestors. The recent representatives of the two sister groups, Pselaphognatha and Chilognatha are characterized by two essentially different types of eye morphology: The eyes of the Pselaphognatha comprise a bilayered rhabdom (built up by 3+4 retinular cells), a few corneagenous cells, a corneal lens, and two vitreous bodies. The latter probably represent relics of a former crystalline cone. On the contrary, the ocelli of the Chilognatha consist of a multilayered rhabdom (built up by a large number of retinular cells), numerous corneagenous cells, and a corneal lens. The dioptric apparatus lacks a crystalline cone. Further structural elements, the distribution of which varies, are the covering cells and processes of hypodermal cells which contain screening pigments. Whereas the eye of the Pselaphognatha can be traced back to a single ommatidium, the ocellus of the Chilognatha can only be interpreted as a merging product of several associated ommatidia or as the result of multiplication and rearrangement of former ommatidial elements. This concept is substantiated by analogous phenomena which occur within other arthropod groups and thus serve as models for the phylogeny of the diplopodan eyes. The comparison of the morphology and the ecology of palaeozoic and recent diplopods demonstrates that the disintegration of former facetted eyes and the modification of ommatidia were induced by the adaptation to cryptic modes of life.     

Morphological deformations of hard-chitinized mouthpart structures in larvae of the genus Chironomus (Diptera,Chironomidae) as the index of organic pollution in freshwater ecosystems

Deformations of hard-chitinized mouthpart structures (mentum and mandibles) in larvae of the genus Chironomus affected by organic pollution have been studied. A system for the quantitative assessment of these structures has been developed. Indices for the severity of mental and mandibular deformation have been calculated. They allow one to determine the pollution of freshwater ecosystems by toxic organic substances.     

Phylogenetic relationships and the larval head of the lower Cyclorrhapha (Diptera)

We examined final‐stage larvae of all currently recognized lower cyclorrhaphan (= Aschiza) families, except Ironomyiidae and Sciadoceridae, and those of the higher cyclorrhaphan (= Schizophora) families Calliphoridae, Conopidae, Lonchaeidae, Muscidae, and Ulidiidae, and compared them with larvae of two out‐group families, Rhagionidae and Dolichopodidae, paying particular attention to structures of the head. A set of 86 morphological characters were analysed phylogenetically. The results show that the lower Cyclorrhapha is paraphyletic in relation to the higher Cyclorrhapha. The monophyly of the Cyclorrhapha is strongly supported. The lower Cyclorrhapha is resolved into two clades, based on the Lonchopteridae. Within the Syrphidae the traditional three‐subfamily system is supported, based on the Microdontinae. Within the lower Cyclorrhapha, the larval head is variable in form and arrangement of components. In Lonchopteridae, the mouth lies at the back of an open trough or furrow, comprising ventrally an elongate labium and laterally the maxilla. This arrangement of components appears to facilitate scooping food in water films. In Platypezoidea there is no furrow, and the dorsolateral lobes bearing the antennae are connected by a dorsal extension of the pseudocephalon. The main food‐gathering structure is the hooked apex of the labium, but in Phoridae the mandibles may also be important. In Eumuscomorpha the mandibles are at the apex of the head skeleton. The pseudocephalon is extended and infolded dorsally to form an oral pocket over the mouth. In the Pipunculidae, and the Microdontinae and Syrphinae of the Syrphidae, ventrally it forms a V‐shaped groove or guide along which the mandibles project. The labium is sclerotized apically, and forms a plate or tapered projection. This arrangement of components facilitates holding, piercing and extracting prey tissues. In Eristalinae the pseudocephalon is attached to the mandibles and is formed into a pair of cirri bearing mandibular lobes that lie either side of the mouth. Furthermore, the epipharynx is produced anteriorly in relation to the hypopharynx, and the labium is attached to the anterior part of the epipharynx to form a cavity or atrium. This arrangement is suited to fragmenting and imbibing solid food in Eristalinae with hooked mandibles, and when the mandibles are reduced and the mandibular lobes are inverted and sclerotized, these structures form a filter for separating fluid‐suspended particulate food. In higher Cyclorrhapha an atrium is present as in Eristalinae, but a connection between the pseudocephalon and the mandibles is absent. Instead, the pseudocephalon is bifurcate dorsally and forms a pair of cephalic lobes that ventrally ensheath each mandible. The surface of the sheath may be coated in cirri and other food‐gathering structures. The cephalic lobes, mandibular sheaths and the head skeleton are maneuverable and retractile to a higher degree than in lower Cyclorrhapha. This arrangement of components facilitates feeding on both solid food, in which the mouthooks may extend from the sheath to break the food up, and particulate and suspended food, in which the food‐gathering structures of the sheath scoop up the food. In many higher Cyclorrhapha, maneuverability is enhanced by a break between the labium and the basal sclerite, to which it is fused in all lower Cyclorrhapha. Intermediate characters and states for the structures of the higher cyclorrhaphan larval head are present in out‐groups, and lower Cyclorrhapha and homologies are discussed. Liquidity of the food is an important factor explaining the structure of the larval head in Cyclorrhapha. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 287–323.     

A functional and phylogenetic interpretation of the skull of the Erycinae (Reptilia, Serpentes)

Based on skull structure, the genus Gongylophis Wagler is again placed in synonomy with the genus Eryx Daudin. Lichanura is considered to be structurally close to the early erycine stock from which Charina evolved as an early, separate off-shoot.
Skull characters which adapt erycines to burrowing habits show clinal variation: they involve shortening of the skull and streamlining of its contours through snout depression, the development of a spatulate transverse process of the pre-maxilla reinforced by the nasals and a specialization of the naso-frontal joint which supports the snout. The jaws retain their mobility. Burrowing habits are a specialized feature of the Erycinae.