Phylogenetic Interpretation Research Articles

A new marattiaceous fern from the Lower Jurassic of Patagonia (Argentina): the renaissance of Marattiopsis

We here describe Marattiopsis patagonica sp. nov. (Marattiaceae) based on vegetative and fertile foliage impressions from the Lower Jurassic of Chubut Province, Patagonia, Argentina. The new species exhibits a unique combination of vegetative and reproductive features, including a prominent basal auricle; finely undulate, denticulate, to serrulate pinnule margins; interstitial fibres (venuli recurrentes) alternating with regular veins; submarginal position and distinctive relative length of synangia (about 12–20% of the pinnule width); and relatively small number of sporangia pairs per synangium (usually 14 or 16). The new species is assigned to the morphogenus Marattiopsis, since it shows no characters that would allow an unambiguous placement in any of the modern genera, Marattia s.s., Ptisana, or Eupodium. The newly established paraphyly of the Marattia s.l. species has a profound impact on the classification of fossil species. In this context, we provide a brief assessment of the significance of the fossil Marattiopsis for phylogenetic and biogeographical interpretations. It appears that essentially modern marattioid ferns, including Marattiopsis, which are commonly considered to be typical elements of tropical areas of the northern hemisphere during the Mesozoic, may have left an important but largely ignored fossil record in subtropical Gondwana.

Three-dimensional visualization of fossil flowers, fruits, seeds, and other plant remains using synchrotron radiation X-ray tomographic microscopy (SRXTM): new insights into Cretaceous plant diversity

The application of synchrotron radiation X-ray tomographic microscopy (SRXTM) to the study of mesofossils of Cretaceous age has created new possibilities for the three-dimensional visualization and analysis of the external and internal structure of critical plant fossil material. SRXTM provides cellular and subcellular resolution of comparable or higher quality to that obtained from permineralized material using thin sections or the peel technique. SRXTM also has the advantage of being non-destructive and results in the rapid acquisition of large quantities of data in digital form. SRXTM thus refocuses the effort of the investigator from physical preparation to the digital post-processing of X-ray tomographic data, which allows great flexibility in the reconstruction, visualization, and analysis of the internal and external structure of fossil material in multiple planes and in two or three dimensions. A review of recent applications in paleobotany demonstrates that SRXTM will dramatically expand the level of information available for diverse fossil plants. Future refinement of SRXTM approaches that further increases resolution and eases digital post-processing, will transform the study of mesofossils and create new possibilities for advancing paleobotanical knowledge. We illustrate these points using a variety of Cretaceous mesofossils, highlighting in particular those cases where SRXTM has been essential for resolving critical structural details that have enhanced systematic understanding and improved phylogenetic interpretations.

Virtual reconstruction of endocast anatomy in early ray-finned fishes (Osteichthyes, Actinopterygii)

Cranial endocasts, infillings of the skeletal void that once contained the brain and associated soft tissues, represent detailed anatomical structures that have long been the focus of paleontological investigation. We applied computed tomographics (CTs) in order to generate endocast models for the Paleozoic actinopterygian fishesMimipiscisandKentuckia, which serve as key representatives of anatomically primitive, early ray fins in analyses of early vertebrate relationships. The resultant endocranial models generally corroborate existing accounts of endocranial anatomy in these genera, drawn from descriptions of the inner face of the brain cavity. However, the endocasts also provide new anatomical details, the most significant of which are the presence inMimipiscisof widely divergent olfactory tracts, small optic lobes, and anterior and posterior semicircular canals that extend dorsal to the roof of the endocranial chamber. By contrast,Kentuckiapossesses a single, straight olfactory tract, wide optic lobes, and anterior and posterior semicircular canals that do not reach the dorsal surface of the endocast. In each of these features,Kentuckiaresembles stratigraphically younger actinopterygians such asLawrenciellaandKansasiella, whereasMimipiscismore closely resembles sarcopterygians and other outgroups. This character distribution provides further support for earlier phylogenetic interpretations of these genera.

Human calcanei from the Middle Pleistocene site of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain)

The existence of calcanei in the fossil record prior to modern humans and Neandertals is very scarce. This skeletal element is fundamental to understanding the evolution of the morphology of the foot in human evolution. Here we present and metrically and comparatively describe 29 calcaneus remains from the Middle Pleistocene site of Sima de los Huesos (SH) (Sierra de Atapuerca, Burgos, Spain). These calcanei belong to 15 individuals (nine adults, two adolescents and four immature individuals). The metric and morphological differences in the calcanei among Middle and Late Pleistocene hominins tend to be subtle. However, the calcanei from SH are broad and robust with large articular surfaces and most significantly, exhibit a very projected sustentaculum tali. A biomechanical and phylogenetic interpretation is proffered to explain the observed morphology of these calcanei. It has been possible to propose tentative sex assignments for the SH calcanei based on size, using methods similar to those used to establish sex from the talus bones from SH. The estimation of stature based on the calcaneus provides a mean of 175.3 cm for males and 160.6 for females, which is similar to that obtained using other skeletal parts from the site. In sum, the SH calcanei are robust with a proportionally long tubercle and a projected sustentaculum tali, which are traits shared by Neandertals.

Three-dimensional microCT analysis of the Ediacara fossil Pteridinium simplex sheds new light on its ecology and phylogenetic affinity

Ediacara fossils often exhibit enigmatic taphonomy that complicates morphological characterization and ecological and phylogenetic interpretation; such is the case with Pteridinium simplex from the late Ediacaran Aar Member in southern Namibia. P. simplex is often preserved as three-dimensional (3D) casts and molds in coarse-grained quartzites, making detailed morphological characterization difficult. In addition, P. simplex is often transported, distorted, and embedded in gutter fills or channel deposits, further obscuring its morphologies. By utilizing microfocus X-ray computed tomography (microCT) techniques, we are able to trace individual specimens and their vanes in order to digitally restore the 3D morphology of this enigmatic fossil. Our analysis shows that P. simplex has a very flexible integument that can be bent, folded, twisted, stretched, and torn, indicating a certain degree of elasticity. In the analyzed specimens, we find no evidence for vane identity change or penetrative growth that were previously used as evidence to support a fully endobenthic lifestyle of P. simplex; instead, evidence is consistent with the traditional interpretation of a semi-endobenthic or epibenthic lifestyle. This interpretation needs to be further tested through microCT analysis of P. simplex specimens preserved in situ rather than transported ones. The elastic integument of P. simplex is inconsistent with a phylogenetic affinity with xenophyophore protists; instead, its physical property is consistent with the presence of collagen, chitin, and cellulose, an inference that would provide constraints on the phylogenetic affinity of P. simplex.

Hypothesized evolutionary trends in zoospore ultrastructural characters in Chytridiales (Chytridiomycota)

Chytridiales is an order of zoosporic fungi currently comprising species representing 19 genera. Although morphologically and genetically diverse, these taxa have in common a zoospore with a suite of ultrastructural characters unique among Chytridiomycota. However, multiple states have been reported for almost every character that defines the Chytridiales zoospore. Two zoospore types have been recognized, each corresponding to a family. Here we examine zoospore ultrastructure of 52 isolates in Chytridiales and assess states for six characters to hypothesize evolutionary trends, using parsimony ancestral state reconstruction for evolutionary analysis. Based on suites of character states, we describe four additional zoospore types in Chytridiales. Five of the six characters ([i] location of the nucleus, [ii] morphology of the kinetosome-associated structure, [iii] complexity of the microtubular root, [iv] microbody-lipid globule complex cisterna structure and [v] thickness of the flagellar plug) revealed ancestral and derived states. The sixth character, structure of the paracrystalline inclusion, did not resolve ancestral and derived states. In each of the lineages within Chytridiales, the evolutionary trend appears to have been from a more complex zoospore to a less complex zoospore with reduced features. As we isolate and analyze additional taxa, we discover new ultrastructural character states that assist in taxon delineation and phylogenetic interpretation.

The gas bladder of puffers and porcupinefishes (Acanthomorpha: Tetraodontiformes): Phylogenetic interpretations

The anatomy of the gas bladder of Diodontidae (porcupinefishes) and Tetraodontidae (pufferfishes) was studied on the basis of dissections and magnetic resonance imaging. Among the examined taxa of Tetraodontiformes, only puffers and porcupinefishes possess a thick walled and dorsally U-shaped or crescent-moon-shaped gas bladder. In the tetraodontid genus Lagocephalus the gas bladder is reduced to a rudiment. The species belonging to the genera Canthigaster, Arothron, and some species of Tetraodon differ in the positioning of their crescent-moon-shaped gas bladder. These observations confirm the close relationship of: (i) Diodontidae and Tetraodontidae and (ii) Canthigaster, Arothron, and some species of Tetraodon. The heterogeneity of the genus Tetraodon is supported by the gas bladder morphology, as previously suggested by molecular studies.

Were There Miocene Meridiolestidans? Assessing the Phylogenetic Placement of Necrolestes patagonensis and the Presence of a 40 Million Year Meridiolestidan Ghost Lineage

The enigmatic mammal Necrolestes patagonensis from the Miocene of Patagonia possesses a highly apomorphic osteological form that has confounded phylogenetic interpretation for over a century. In this time it has been affiliated with both eutherians and metatherians; however, a recent study by Rougier et al. (Proc Natl Acad Sci USA 109:19871–19872, 2012) raises the intriguing possibility that Necrolestes is a relictual member of a clade of South American non-therian dryolestoids, the Meridiolestida. This group is known chiefly from the Cretaceous of South America and assignment of Necrolestes to Meridiolestida implies a ghost lineage of about 40 million years. Such a lengthy ghost lineage requires strong evidence, which minimizes potentially circular assumptions of anatomical homology. Here, we vary the coding of cusp homologies in Necrolestes, previously assumed to diverge from the metatherian pattern, and add zalambdodont and incipiently zalambdodont metatherian taxa to the analyses, in order to assess the effects of non-independent characters and taxon sampling on the original topology. The results of both maximum parsimony and Bayesian analysis using the Mk model show that these possible sources of bias have little effect on the topology and ultimately increase confidence in the placement of Necrolestes in Meridiolestida and its concomitant 40 million year ghost lineage. Additionally, our Bayesian analysis resolves Australosphenida in a trichotomy with Peramus and Vincelestes + Boreosphenida. This contrasts with the majority of existing topologies, and raises interesting questions regarding both the evolution of tribospheny and the use of the Mk model with paleontological datasets.

New information on Bobosaurus forojuliensis (Reptilia: Sauropterygia): implications for plesiosaurian evolution

Preparation of the holotype specimen of Bobosaurus forojuliensis, a large sauropterygian from the lower Carnian of northeastern Italy, revealed new morphological data relevant in establishing its phylogenetic affinities among pistosauroid taxa and its relationships with plesiosaurians. Inclusion of B. forojuliensis in two phylogenetic analyses focusing, respectively, on sauropterygians and pistosauroids agreed in placing the Italian taxon as closer to plesiosaurians than to other pistosauroids. The phylogenetic interpretation of Bobosaurus was not biased by assumptions on character weighting, is consistent with its relatively younger age compared to most pistosauroids, extends the fossil record of the plesiosaurian basal lineage back to the Carnian and supports the earliest diversification of the clade during the Late Triassic in agreement with the record of several distinct lineages of rhomaleosaurids, plesiosauroids and pliosauroids in the lowermost Jurassic. Bobosaurus shows that the evolution of the plesiosaurian body plan from the ancestral pistosauroid grade was a step-wise process, and that some of the vertebral and appendicular specialisations of Jurassic and Cretaceous plesiosaurians had already developed in the earliest Late Triassic.

The evolutionary history of HSA7/16 synteny in vertebrates: a critical interpretation of comparative cytogenetic and genome sequence data

The current work is an in silico study of data from previous publications and genome browsers, on the origin of the human synteny HSA7a/16p. The molecular composition of the chromosomal segments identified as HSA7a/16 and 7b (free or differently associated) is not yet clear. This means that a syntenic association 7/16, which can be detected by an in situ hybridization (FISH) method in different taxa, may not necessarily correspond to those of the same association in different lineages. In silico research, together with comparative cytogenetics, have been applied in order to define the composition of the 7/16 syntenic association. These results allow a confident reconstruction of the syntenic associations HSA7/16 in diverse vertebrate lineages, indicating various levels of homology, but also considerable levels of non-homology, which should elicit caution with phylogenetic interpretations. The 7/16 association represents a paradigmatic example of the complexity involved in the interpretation of comparative...

Bioinformatic progress and applications in metaproteogenomics for bridging the gap between genomic sequences and metabolic functions in microbial communities

Metaproteomics of microbial communities promises to add functional information to the blueprint of genes derived from metagenomics. Right from its beginning, the achievements and developments in metaproteomics were closely interlinked with metagenomics. In addition, the evaluation, visualization, and interpretation of metaproteome data demanded for the developments in bioinformatics. This review will give an overview about recent strategies to use genomic data either from public databases or organismal specific genomes/metagenomes to increase the number of identified proteins obtained by mass spectrometric measurements. We will review different published metaproteogenomic approaches in respect to the used MS pipeline and to the used protein identification workflow. Furthermore, different approaches of data visualization and strategies for phylogenetic interpretation of metaproteome data are discussed as well as approaches for functional mapping of the results to the investigated biological systems. This information will in the end allow a comprehensive analysis of interactions and interdependencies within microbial communities.

Hemichordata (Pterobranchia, Enteropneusta) and the fossil record

The Hemichordata are generally interpreted as early deuterostomes, closely related to the chordates, a notion important for modern analyses of the origin of the deuterostomes. Because their fossil record is quite scanty, modern phylogenetic interpretations largely rely on the analysis of DNA of the available extant taxa. The tripartite body plan of the group of worm-like hemichordates, the Enteropneusta, may be traced back in deep time to a few poorly known Middle Cambrian (Series 3, Stage 5) taxa from the Burgess Shale biota. The derived small, colonial or pseudocolonial Pterobranchia (Cephalodiscida and Graptolithina) have a more complete fossil record due to their preservable housing construction, the tubarium. The relationships of fossil taxa, putatively identified as early deuterostomes and possible hemichordates or even as pterobranchs of Lower to Middle Cambrian age (e.g. Galeaplumosus, Herpetogaster), cannot be substantiated. The Pterobranchia and their housing construction are first seen in the Middle Cambrian Series 3, Stage 5 but a clonal, colonial organization of the tubaria can only be recognized in the basal Drumian. The fossil enteropneust Mazoglossus ramsdelliBardack, 1997 from the Carboniferous Mazon Creek Biota is re-described, its lectotype designated and illustrated for the first time.

A unique DNA repair and recombination gene (recN) sequence for identification and intraspecific molecular typing of bacterial wilt pathogen Ralstonia solanacearum and its comparative analysis with ribosomal DNA sequences

Ribosomal gene sequences are a popular choice for identification of bacterial species and, often, for making phylogenetic interpretations. Although very popular, the sequences of 16S rDNA and 16-23S intergenic sequences often fail to differentiate closely related species of bacteria. The availability of complete genome sequences of bacteria, in the recent years, has accelerated the search for new genome targets for phylogenetic interpretations. The recently published full genome data of nine strains of R. solanacearum, which causes bacterial wilt of crop plants, has provided enormous genomic choices for phylogenetic analysis in this globally important plant pathogen. We have compared a gene candidate recN, which codes for DNA repair and recombination function, with 16S rDNA/16-23S intergenic ribosomal gene sequences for identification and intraspecific phylogenetic interpretations in R. solanacearum. recN gene sequence analysis of R. solanacearum revealed subgroups within phylotypes (or newly proposed species within plant pathogenic genus, Ralstonia), indicating its usefulness for intraspecific genotyping. The taxonomic discriminatory power of recN gene sequence was found to be superior to ribosomal DNA sequences. In all, the recN-sequence-based phylogenetic tree generated with the Bayesian model depicted 21 haplotypes against 15 and 13 haplotypes obtained with 16S rDNA and 16-23S rDNA intergenic sequences, respectively. Besides this, we have observed high percentage of polymorphic sites (S 23.04%), high rate of mutations (Eta 276) and high codon bias index (CBI 0.60), which makes the recN an ideal gene candidate for intraspecific molecular typing of this important plant pathogen.

Belichnus Traces Produced on Shells of the Bivalve Lutraria lutraria by Gulls

In February 2011, after a storm, thousands of adult, articulated, and still-living common otter shells Lutraria lutraria (L. 1758) were stranded on the North Sea beach of the Island Texel (NL). These 9 to12 cm long bivalves were rapidly found and consumed by both herring- and lesser black-backed gulls. Holes, irregular in outline, were observed in some 10% of the articulated shells of these bivalves. These holes were always smaller on the outside of the valves than on the inside and varied in size from 1 to 20 mm (outside) to 4 to 22 mm (inside). Often the other valve was crushed indicating consumption by gulls. We concluded that these holes were made by the gulls probing the shells; in a few cases, we observed that valves were broken starting from such a hole. Such traces are described in the literature as the ichnogenus Belichnus and were until now attributed to Stomatopoda only. We also suggest that a separate ichnospecies name for two Belichnus holes in one shell should not be used, as we see them simply as a double injury due to two blows. Our findings stress once more the importance of avoiding premature phylogenetic interpretation of traces and the use of a separate ichnotaxonomy.

The phylogenetic significance of the carpophore in Apiaceae

Fruit structural characters have traditionally been important in the taxonomy of the family Apiaceae. Previous investigations using a limited number of taxa have shown that the carpophore may be especially useful in helping to circumscribe subfamily Azorelloideae. The present study examines, for the first time, carpophore structure in 92 species from 43 genera, representing all subfamilies of Apiaceae, and including all genera assigned to subfamily Azorelloideae. Phylogenetic interpretations are made for the first time, using all available information, and a standard terminology is proposed to describe the various character states found in carpophores. Carpophore structure was studied in detail using light microscopy. Carpophores, when present, may be categorized into two main groups (B and C) based mainly on the arrangement of the vascular bundles in transverse section, and further divided into six sub-types according to the length of the carpophore (short in B1 and C1) and whether they are entire (B1-B3 and C1) or bifurcate (B4 and C2). Free carpophores are absent in subfamily Mackinlayoideae, and in tribes Lichtensteinieae and Phlyctidocarpeae, which have two opposite vascular bundles (Group A). Entire carpophores with one or two vascular bundles, or bifurcate carpophores with lateral vascular bundles (arranged side by side within the commissural plane), are the main types characterizing Azorelloideae. The short, hygroscopic carpophores found in Choritaenia are unique in Apiaceae and provide additional evidence for the exclusion of this genus from Azorelloideae. Carpophore type C2 is typical for most Apioideae sensu lato (exceptions are, for example, Arctopus and Alepidea, which have type B2). A single carpophore and ventral vascular bundles not forming free carpophores are proposed to be the ancestral conditions in Apiaceae, while bifurcate carpophores with opposite vascular bundles are the derived state, present in most Apioideae. Secondary reductions seem to have occurred in several unrelated lineages in all major groups, e.g. many Azorelloideae, several protoapioids (including nearly all members of the tribe Saniculeae) and 29 euapioid genera (e.g. some Oenantheae).

Primates in the Eocene

The mammalian order Primates made its first appearance in the fossil record during the Paleocene–Eocene thermal maximum (PETM), the global greenhouse warming event that marks the beginning of the Eocene. Two primate superfamilies, Tarsioidea and Adapoidea, dominate early and middle Eocene primate faunas. Warm climates enabled primates to thrive, and warming events within the Eocene facilitated cosmopolitan dispersal. Declining diversity at the end of the Eocene reflects environmental cooling. Fossils of earliest Tarsioidea and Adapoidea are similar dentally, often confused, and appear closely related as stem or crown Haplorhini. The superfamily Tarsioidea is represented by a single genus, Tarsius, living today, while Adapoidea appear to be ancestral to living Anthropoidea. Little is known of the Eocene history of strepsirrhine Lemuroidea and Lorisoidea. Temporal scaling of molecular clock ages suggests that Strepsirrhini appeared before Haplorhini in the Paleocene or possibly with Haplorhini at the beginning of the Eocene. Substantial skeletons of Eocene primates like those of adapoid Darwinius and Europolemur from Messel in Germany and Notharctus and Smilodectes from western North America constrain phylogenetic interpretation of primate relationships much more than dental remains ever can. A specialised grasping foot distinguishes early primates from other mammals. Traits associated in a functional complex include replacement of claws by nails on all digits; movement of the pedal fulcrum from the metatarsals to the tarsals; elongation of digit IV relative to digit III, with reduction of digit II and sometimes III; and then secondary development of a grooming claw or claws on digits II and sometimes III. The specialised grasping foot of early primates was later moderated in the emergence of anthropoid primates.

Application of numerical cladistic analyses to the Carnian–Norian conodonts: a new approach for phylogenetic interpretations

The high intraspecific variability of conodont platform elements in the upper Carnian–lower Norian interval and the proliferation of numerous species in this relatively short time have generated many problems for the understanding of Late Triassic conodont phylogeny, systematics and taxonomy. Since Late Triassic natural assemblages are still unknown, we have applied cladistic methodologies to investigate the evolution of Carnian–Norian conodont platforms and to infer more precise phylogenetic relationships among taxa. Numerical cladistic analysis was undertaken of species belonging to the five Late Triassic genera Paragondolella, Carnepigondolella, Metapolygnathus, Epigondolella and Norigondolella. A taxon–character data matrix describing the distribution of 64 characters amongst two outgroups and 31 ingroup taxa was compiled and processed using PAUP* 4.1. Our analyses show the evolutionary and systematic value of certain morphological characters, and lead to a reinterpretation of the phylogenetic position of the genera considered: Metapolygnathus, Epigondolella and Norigondolella are monphyletic taxa, Paragondolella represents a polyphyletic assemblage of basal members of the ingroup, and Carnepigondolella a paraphyletic group including primitive forms of the Metapolygnathus and Epigondolella lineages. Our analyses also clarify the systematic position of ‘Metapolygnathus communisti B’ and confirm the existence of four new species: Carnepigondolella angulata sp. nov., Epigondolella heinzi sp. nov., Epigondolella miettoi sp. nov. and Norigondolella trinacriae sp. nov.

The nasal and paranasal architecture of the Middle Miocene ape Pierolapithecus catalaunicus (primates: Hominidae): Phylogenetic implications

The internal (nasal and paranasal) cranial anatomy of the Middle Miocene (11.9 Ma [millions of years ago]) great ape Pierolapithecus catalaunicus (Hominidae: Dryopithecini) is described on the basis of computed-tomography scans of the holotype specimen (IPS21350), with particular emphasis on its phylogenetic implications. Pierolapithecus displays the following characters: an anteriorly-restricted maxillary sinus that posteriorly spreads towards the ethmoidal area (thus resembling the pongine condition), although being situated well above the molar roots (as in kenyapithecins, other dryopithecins and pongines); lack of frontal sinus (a synapomorphy of derived pongines, independently acquired by both cercopithecoids and hylobatids); posteriorly-situated turbinals (as in Pongo); anteriorly-projecting nasolacrimal canal (as in Pongo); and probably stepped nasal floor with non-overlapping premaxillary–maxillary contact (as in dryopithecines and stem hominoids, although it cannot be conclusively shown due to bone damage). Overall, Pierolapithecus displays a mosaic of primitive hominid and derived pongine features that are inconsistent with this taxon being a hominine (as previously suggested). Two alternative phylogenetic interpretations are possible: Pierolapithecus may be a stem member of the Hominidae as previously suggested in its original description, or alternatively this taxon may be a stem member of the Ponginae s.l. (with the European dryopithecines being the sister taxon to the Asian pongines).

First μ‐CT‐based 3D reconstruction of a dipteran larva—the head morphology of protanyderus (tanyderidae) and its phylogenetic implications

The larval head of Protanyderus was examined and documented using innovative techniques, with emphasis on internal structures. A chart listing all head muscles of dipteran larvae and other holometabolan groups is presented in the Supporting Information. The results are compared to conditions found in other nematoceran lineages. The larval head of Protanyderus is characterized mainly by plesiomorphic character states such as the complete and largely exposed head capsule, the long coronal suture, V-shaped frontal sutures, lateral antennal insertion areas, a transverse labrum, a nearly horizontal plane of mandibular movements, mandibles lacking a movable distal part, a mesal hook and mesal or distal combs, separated maxillary endite lobes, a comparatively complete array of muscles, and a brain only partly located within the head capsule. An anteriorly toothed hypostomal plate and dense labral brushes of microtrichiae are also likely groundplan features of Diptera. The pharyngeal filter is a possible apomorphy of Diptera excl. Deuterophlebiidae (or Deuterophlebiidae + Nymphomyiidae). The messors have also likely evolved early in the dipteran crown group but are absent in the groundplan. The phylogenetic interpretation of externolateral plates with growth lines is ambiguous. Autapomorphies of Tanyderidae are differences between the third and fourth instar larvae, the roof-like extension above the antennal insertion area, the dorsal endocarina, and the posterodorsal internal ridge. The phylogenetic position of Tanyderidae is controversial, but features of the larval head do not support a proposed sistergroup relationship between Tanyderidae and Psychodidae. Both groups differ in many features of the larval head, and we did not identify a single potential synapomorphy. Larval characters alone are insufficient for a reliable phylogenetic reconstruction, though they vary greatly and apparently contain phylogenetic information. The evaluation of these features in the context of robust molecular phylogenies will be a sound basis for the reconstruction of complex evolutionary scenarios for the megadiverse Diptera.

Reassessment of Dolichopithecus (Kanagawapithecus) leptopostorbitalis, a colobine monkey from the Late Pliocene of Japan

In the original description of Dolichopithecus (Kanagawapithecus) leptopostorbitalis, Iwamoto, Hasegawa and Koizumi, 2005, a moderately large-sized colobine monkey from the Late Pliocene of central Japan, affinities to the European Dolichopithecus rather than to the Transbaikalian Parapresbytis were noted based on the similarities in cranial morphology. Computed tomography scans confirm the presence of the maxillary sinus in the holotype, whereas it is probably absent in specimens of the European Dolichopithecus ruscinensis, the type species of this genus. This feature is either present or absent homogeneously in any given genus of living anthropoids. Its presence or absence is unknown in Parapresbytis, but the distinct morphology of the maxillary incisors in this taxon suggests that this form had different feeding habits from the Japanese colobines. These findings suggest that the Japanese colobine should be referred to henceforth as Kanagawapithecus leptopostorbitalis. Kanagawapithecus shares many important facial and dental features with Dolichopithecus rather than with Parapresbytis, but this association depends largely on the limited availability of comparable materials for the latter. Among colobines, the presence of the maxillary sinus is recorded only in Libypithecus and Cercopithecoides. The maxillary sinus is absent in all modern Asian colobines, implying that Kanagawapithecus is an isolated form without any relationship to living forms. Nevertheless, such phylogenetic interpretations are largely dependent on the restricted fossil evidence from the Pliocene and Pleistocene of eastern Eurasia and will be reexamined when new findings are made.