Sister Taxon Relationship Research Articles

Cranial Osteology and Ontogeny ofSaurolophus angustirostrisfrom the Late Cretaceous of Mongolia with Comments onSaurolophus osbornifrom Canada

Reanalysis of the skull of the crested Asian hadrosaurine Saurolophus angustirostris confirms its status as a distinct species from its North American relative, Saurolophus osborni. In addition to its greater absolute size, S. angustirostris is differentiated from Saurolophus osborni by an upturned premaxillary body, a more strongly reflected oral margin of the premaxilla, the absence of an anterior notch in the prenarial fossa, a sigmoidal contour of the ventral half of the anterior process of the jugal, a shallow quadratojugal notch on the quadrate, and by a strongly bowed quadrate in lateral view. Phylogenetic analysis corroborates a sister taxon relationship between S. angustirostris and S. osborni. Saurolophus itself is characterised by a solid, rod-like crest composed of the nasals, frontals, and prefrontals; secondary elongation of the frontal and prefrontal resulting in the backwards extension of the frontal platform; a frontal platform that extends dorsal to the anterior portion of the supratempo...

Podargiform affinities of the enigmatic Fluvioviridavis platyrhamphus and the early diversification of Strisores ("Caprimulgiformes" + Apodiformes).

BackgroundThe early Eocene Green River Formation avifauna preserves exceptional exemplars of the earliest unambiguous stem representatives of many extant avian clades. We identify the basal-most member of Podargiformes (extant and fossil stem lineage frogmouths) based on a new specimen of Fluvioviridavis platyrhamphus, a unique neoavian bird from the Fossil Butte Member of the Green River Formation of Wyoming. Extant frogmouths (Podargidae) comprise approximately 13 nocturnal species with an exclusively Australasian distribution.Methodology/Principal FindingsThe new specimen was included in a combined phylogenetic analysis of morphological (osteology and soft tissue) and molecular sequence (cytochrome b, c-myc exon 3, and RAG) data sampling species-level taxa from both extant and extinct members of Steatornithidae, Podargidae, Caprimulgidae, Nyctibiidae, Aegothelidae, and Apodiformes ( = Strisores). New data from F. platyrhamphus help resolve phylogenetic relationships within Strisores, supporting placement of F. platyrhamphus and the European fossil form Masillapodargus longipes as basal parts of Podargiformes and also supporting a sister taxon relationship between Podargiformes and Steatornithiformes (oilbirds) within Strisores. This relationship is recovered only when fossil taxa are included, reaffirming the potential impact of stem fossil taxa on inferences of phylogenetic relationships. The well-preserved mandible and palate of the new specimen demonstrate that many of the unique characteristics of the skull that characterize the crown frogmouth clade Podargidae arose early in the evolutionary history of the clade, over 50 million years ago. Comparisons with the new specimen also indicate that Eurofluvioviridavis and Fluvioviridavis are not closely related.Conclusions/SignificanceTogether with the European fossil frogmouth Masillapodargus, Fluvioviridavis shows that Podargiformes had a much wider geographic distribution in the past, whereas extant species are restricted to Australasia. The Eocene record of Strisores from the Green River Formation and Messel Formation indicates most major subclade divergences had already occurred by the early-middle Eocene.

Phylogenomics reveals deep molluscan relationships

Evolutionary relationships among the eight major lineages of Mollusca have remained unresolved despite their diversity and importance. Previous investigations of molluscan phylogeny, based primarily on nuclear ribosomal gene sequences or morphological data, have been unsuccessful at elucidating these relationships. Recently, phylogenomic studies using dozens to hundreds of genes have greatly improved our understanding of deep animal relationships. However, limited genomic resources spanning molluscan diversity has prevented use of a phylogenomic approach. Here we use transcriptome and genome data from all major lineages (except Monoplacophora) and recover a well-supported topology for Mollusca. Our results strongly support the Aculifera hypothesis placing Polyplacophora (chitons) in a clade with a monophyletic Aplacophora (worm-like molluscs). Additionally, within Conchifera, a sister-taxon relationship between Gastropoda and Bivalvia is supported. This grouping has received little consideration and contains most (>95%) molluscan species. Thus we propose the node-based name Pleistomollusca. In light of these results, we examined the evolution of morphological characters and found support for advanced cephalization and shells as possibly having multiple origins within Mollusca.

A reappraisal of the monophyly of the genus Pseudomonocelis Meixner, 1943 (Platyhelminthes: Proseriata), with the description of a new species from the Mediterranean

Pseudomonocelis paupercula nov. sp. is described from brackish-water habitats of the Mediterranean. It is distinguished from other members of the genus by the copulatory organ provided with a stylet, combined with lack of vagina and presence of a muscular organ close to the female pore. Its phylogenetic relationships have been investigated sequencing complete 18S rRNA gene and partial 28S rRNA gene, spanning variable domains D1-D6. Both BI and ML suggest a sistertaxon relationships of P. paupercula nov. sp. with the east African P. cf cavernicola. However, statistical support is low. Conversely, MP indicates P. paupercula nov. sp. as sister-taxon to all the remaining Pseudomonocelis and Minona ilenae. Overall, results of the combined analysis do not support the monophyly of the genus Pseudomonocelis. The need for wider molecular and taxonomic sampling is stressed.

Two species ofHesslandona(Phosphatocopida, Crustacea) from the Upper Cambrian of western Hunan, South China and the phylogeny of Phosphatocopida

The phosphatocopidsHesslandona necopinaMüller, 1964 andHesslandona longispinosa(Kozur, 1974) new combination, recovered from the Upper Cambrian in western Hunan, South China, are described. The ontogenetic stages ofH. necopinaare revised, with newly defined second and third ontogenetic stages. The second stage is characterized by a bipartite mandibular limb stem consisting of separate coxa and basipod. The third stage is characterized by the partial fusion of the mandibular coxa and basipod. These two parts may become completely fused possibly in a later ontogenetic stage.Vestrogothia longispinosaKozur, 1974 is reassigned toHesslandonabecause of the presence of an interdorsum and a relatively narrow doublure. The new data reported here, plus earlier reports of phosphatocopids from the same section and horizon in western Hunan since 2005, are included in an updated phylogenetic analysis of the Phosphatocopida. Autapomorphies that define several monophyletic groups (e.g., the Crustacea sensu lato, Labrophora, Eucrustacea, Phosphatocopida, Euphosphatocopida, Vestrogothiina, Hesslandonina, and Dorsospinata) are discussed. The present analysis confirms results from earlier phylogenetic analyses in showing the Phosphatocopida-Eucrustacea sister-taxon relationship, but differs from them in supporting the paraphyly of the Hesslandonidae and the monophyly of the Vestrogothiidae.

Biogeographical Consequences of Cenozoic Tectonic Events within East Asian Margins: A Case Study of Hynobius Biogeography

Few studies have explored the role of Cenozoic tectonic evolution in shaping patterns and processes of extant animal distributions within East Asian margins. We select Hynobius salamanders (Amphibia: Hynobiidae) as a model to examine biogeographical consequences of Cenozoic tectonic events within East Asian margins. First, we use GenBank molecular data to reconstruct phylogenetic interrelationships of Hynobius by Bayesian and maximum likelihood analyses. Second, we estimate the divergence time using the Bayesian relaxed clock approach and infer dispersal/vicariance histories under the ‘dispersal–extinction–cladogenesis’ model. Finally, we test whether evolutionary history and biogeographical processes of Hynobius should coincide with the predictions of two major hypotheses (the ‘vicariance’/‘out of southwestern Japan’ hypothesis). The resulting phylogeny confirmed Hynobius as a monophyletic group, which could be divided into nine major clades associated with six geographical areas. Our results show that: (1) the most recent common ancestor of Hynobius was distributed in southwestern Japan and Hokkaido Island, (2) a sister taxon relationship between Hynobius retardatus and all remaining species was the results of a vicariance event between Hokkaido Island and southwestern Japan in the Middle Eocene, (3) ancestral Hynobius in southwestern Japan dispersed into the Taiwan Island, central China, ‘Korean Peninsula and northeastern China’ as well as northeastern Honshu during the Late Eocene–Late Miocene. Our findings suggest that Cenozoic tectonic evolution plays an important role in shaping disjunctive distributions of extant Hynobius within East Asian margins.

Novel Arenavirus Sequences in Hylomyscus sp. and Mus (Nannomys) setulosus from Côte d'Ivoire: Implications for Evolution of Arenaviruses in Africa

This study aimed to identify new arenaviruses and gather insights in the evolution of arenaviruses in Africa. During 2003 through 2005, 1,228 small mammals representing 14 different genera were trapped in 9 villages in south, east, and middle west of Côte d'Ivoire. Specimens were screened by pan-Old World arenavirus RT-PCRs targeting S and L RNA segments as well as immunofluorescence assay. Sequences of two novel tentative species of the family Arenaviridae, Menekre and Gbagroube virus, were detected in Hylomyscus sp. and Mus (Nannomys) setulosus, respectively. Arenavirus infection of Mus (Nannomys) setulosus was also demonstrated by serological testing. Lassa virus was not found, although 60% of the captured animals were Mastomys natalensis. Complete S RNA and partial L RNA sequences of the novel viruses were recovered from the rodent specimens and subjected to phylogenetic analysis. Gbagroube virus is a closely related sister taxon of Lassa virus, while Menekre virus clusters with the Ippy/Mobala/Mopeia virus complex. Reconstruction of possible virus–host co-phylogeny scenarios suggests that, within the African continent, signatures of co-evolution might have been obliterated by multiple host-switching events.

Molecular phylogeny and taxonomic revision of the Philippine endemic Villaria Rolfe (Rubiaceae)

The little known Rubiaceae genus Villaria is endemic mostly to the coastal forests of the Philippines. Traditionally, it has been placed in the tribe Gardenieae. Later it was transferred to Octotropideae sensu Robbrecht and Puff. Villaria was placed among the “primitive” genera of the tribe, which are essentially characterized by large fruits, horizontal ovules and numerous seeds. Parsimony and Bayesian analyses of the combined plastid (rps16 and trnT-F) dataset strongly support the inclusion of Villaria in Octotropideae as well as monophyly of the genus. However, our molecular results do not conform to the current informal groups of the tribe delimited by fruit size, ovule position, number of seeds and exotesta thickenings. Instead, a close relationship between Villaria and two “central genera” (Hypobathrum and Pouchetia) is revealed for the first time. This clade is sister to a group comprising “primitive” (Fernelia), “advanced” (Kraussia and Polysphaeria) and “central” (Feretia) representatives. In addition, our combined tree strongly supports a sister taxa relationship between Canephora and Paragenipa. Villaria is characterized by unilocular ovaries, parietal placentation and strictly horizontal ovules. These features are unique within the Octotropideae. We recognize a total of five Villaria species, one new species (V. leytensis) is described here, and two species (V. philippinensis and V. rolfei) are transferred into synonymy with V. odorata. Each species is fully described, and a key to the species, a distribution map and illustrations are provided.

A novel growth-promoting microbe, Methylobacterium funariae sp. nov., isolated from the leaf surface of a common moss

Land plants (embryophytes) evolved in the presence of prokaryotic microbes. As a result, numerous mutually beneficial associations (symbioses) developed that can be analyzed using a variety of methods. Here we describe the isolation and characterization of a new pink-pigmented facultatively methylotrophic symbiotic bacterium of the genus Methylobacterium (laboratory strain F3.2) that was isolated from the gametophytic phylloids of the common cord moss Funaria hygrometrica Hedw. Plantlets were collected in the field and analyzed in the laboratory. Colonies of methylobacteria were obtained by the agar-impression-method. Based on its unique phenotype (the bacterial cells are characterized by fimbriae-like appendages), a comparative 16S rRNA gene (DNA) sequence analysis, and an average DNA-DNA hybridization value of 8,4 %, compared with its most closely related sister taxon, this isolate is described as a new species, Methylobacterium funariae sp. nov. (type strain F3.2). This new epiphytic bacterium inhabits the leaf surface of “primitive” land plants such as mosses and interacts with its host organism via the secretion of phytohormones (cytokinines, auxins). These external signals are perceived by the plant cells that divide and grow more rapidly than in the absence of their prokaryotic phytosymbionts. We suggest that M. funariae sp. nov. uses methanol emitted from the stomatal pores as principal carbon source for cell metabolism. However, our novel data indicate that, in this unique symbiotic plant-microbe interaction, the uptake of amino acids leached from the surface of the epidermal cells of the green host organism may be of importance as microbial carbon- and nitrogen-source.

Intercontinental dispersal by a microendemic burrowing reptile (Dibamidae).

Intercontinental dispersal via land bridge connections has been important in the biogeographic history of many Holarctic plant and animal groups. Likewise, some groups appear to have accomplished trans-oceanic dispersal via rafting. Dibamid lizards are a clade of poorly known fossorial, essentially limbless species traditionally split into two geographically disjunct genera: Dibamus comprises approximately 20 Southeast Asian species, many of which have very limited geographical distributions, and the monotypic genus Anelytropsis occupies a small area of northeastern Mexico. Although no formal phylogeny of the group exists, a sister-taxon relationship between the two genera has been assumed based on biogeographic considerations. We used DNA sequence data from one mitochondrial and six nuclear protein-coding genes to construct a phylogeny of Dibamidae and to estimate divergence times within the group. Surprisingly, sampled Dibamus species form two deeply divergent, morphologically conserved and geographically concordant clades, one of which is the sister taxon of Anelytropsis papillosus. Our analyses indicate Palaearctic to Nearctic Beringian dispersal in the Late Palaeocene to Eocene. Alternatively, a trans-Pacific rafting scenario would extend the upper limit on dispersal to the Late Cretaceous. Either scenario constitutes a remarkable long-distance dispersal in what would seem an unlikely candidate.

A new taxon of phytosaur (Archosauria: Pseudosuchia) from the Late Triassic (Norian) Sonsela Member (Chinle Formation) in Arizona, and a critical reevaluation of Leptosuchus Case, 1922

Abstract: Leptosuchus Case, 1922 (Reptilia: Phytosauria) from the Late Triassic of the American West is represented by many specimens. Here, I present complete morphological descriptions of the skull material of a new taxon from the Sonsela Member (Chinle Formation) of Petrified Forest National Park, Arizona, with the first rigorous phylogenetic analysis focused on the interrelationships of Leptosuchus. The new taxon is recovered as the sister taxon to Pseudopalatinae. It possesses one unambiguous synapomorphy (the ‘septomaxillae’ form part of the lateral borders of the nares) and shares the presence of a subsidiary opisthotic process with Pseudopalatinae. The new taxon does not fall within the restricted clade Leptosuchus. In my analysis, the previously proposed, but undemonstrated, sister taxon relationship between Angistorhinus and Rutiodon is not supported, Paleorhinus is recovered as paraphyletic, and a subset of taxa traditionally included within Leptosuchus are found to be more closely related to Pseudopalatinae, rendering Leptosuchus paraphyletic. ‘Leptosuchus’adamanensis emerges as sister taxon to Smilosuchus gregorii and is here referred to as Smilosuchus adamanensis nov. comb., and ‘Machaeroprosopus’lithodendrorum is also transferred to Smilosuchus lithodendrorum nov. comb. Documentation of the variation present within Phytosauria, and specifically within Leptosuchus sensu lato, demonstrates higher diversity within Phytosauria than previously appreciated and places the character states previously proposed for Pseudopalatinae into a broader context of shared characters.

Gene and species trees of a Neotropical group of treefrogs: Genetic diversification in the Brazilian Atlantic Forest and the origin of a polyploid species

The Neotropical Phyllomedusa burmeisteri treefrog group includes four diploid ( P. bahiana, P. burmeisteri, P. distincta and P. iheringii) and one tetraploid ( P. tetraploidea) forms. Here we use mitochondrial and nuclear sequence variation from across its range to verify if recognized morphospecies correspond to phylogenetic clades, examine the origin of the polyploid P. tetraploidea, and compare range wide patterns of diversification to those of other BAF organisms. We compared single gene trees with one Bayesian multi-gene tree, and one Bayesian species tree inferred under a coalescent framework. Our mtDNA phylogenetic analyses showed that P. bahiana, P. burmeisteri and P. iheringii correspond to monophyletic clades, while P. distincta and P. tetraploidea were paraphyletic. The nuclear gene trees were concordant in revealing two moderately supported groups including (i) P. bahiana and P. burmeisteri (northern species) and (ii) P. distincta, P. tetraploidea and P. iheringii (southern species). The multi-gene tree and the species tree retrieved similar topologies, giving high support to the northern and southern clades, and to the sister-taxa relationship between P. tetraploidea and P. distincta. Estimates of t MRCA suggest a major split within the P. burmeisteri group at ≈5 Myr (between northern and southern groups), while the main clades were originated between ≈0.4 and 2.5 Myr, spanning the late Pliocene and Pleistocene. Patterns of geographic and temporal diversification within the group were congruent with those uncovered for other co-distributed organisms. Independent paleoecological and geological data suggest that vicariance associated with climatic oscillations and neotectonic activity may have driven lineage divergence within the P. burmeisteri group. P. tetraploidea probably originated from polyploidization of P. distincta or from a common ancestor.

Development of the ethmoidal structures of the endocranium in Discoglossus pictus (Anura: Discoglossidae)

We use histological techniques and computer-aided three-dimensional reconstructions made from serial histological sections to describe the ontogeny of the ethmoidal endocranium of discoglossid frog Discoglossus pictus. We identify a pattern of development for the suprarostral cartilage that differs from previous findings and probably represents the ancestral anuran pattern. The nasal cartilages, including the inferior prenasal cartilage, are de novo adult structures. The only larva-derived structures of the adult nasal capsules are the posterior aspects of the solum nasi and septum nasi. We also identify patterns of development for the ethmoid plate and postnasal wall that occur during early in ontogenesis. These patterns are associated with development events during metamorphic climax. The pattern of timing of chondrification of the anterior nasal cartilages more closely coincides with that of the neobatrachian species than that recorded for the pelobatid frog Spea. In addition, this study supports a sister taxon relationship between Discoglossus and Alytes.

Molecular inference of phylogenetic relationships among Decapodiformes (Mollusca: Cephalopoda) with special focus on the squid Order Oegopsida

Squids, cuttlefish and bobtail squids comprise the molluscan superorder Decapodiformes (Mollusca: Cephalopoda). Although these animals exemplify the morphological and ecological diversity seen in Cephalopoda, no previous study has focused resolving decapodiform relationships, particularly within Oegopsida, a large order comprised of pelagic squid. To further clarify the phylogenetic history of Decapodiformes, and Oegopsida in particular, molecular data for five genes (18S rRNA, 28S rRNA, Histone H3, 16S rRNA, COI) was collected for 90 taxa representing all major lineages and families and evaluated using parsimony, likelihood, and Bayesian analysis. Although ordinal relationships were sensitive to analytical method, several conclusions can be inferred: the pelagic order Myopsida is closely related to the benthic sepioids, whose relationships were ambiguous, and Bathyteuthoidea is distinct from Oegopsida. Within Oegopsida several clades are consistently recovered, some with previous morphological support (e.g. chiroteuthid, lepidoteuthid, histioteuthid families) while others suggest novel relationships (e.g. Architeuthidae + Neoteuthidae). This study, with its broad coverage of taxa, provides the first in-depth analysis of Decapodiformes with special focus on the morphologically and biogeographically diverse Oegopsida, confirms several sister-taxon relationships, and provides new hypotheses of cephalopod evolution in the open ocean.

A new dromaeosaurid (Dinosauria: Theropoda) from the Upper Cretaceous Wulansuhai Formation of Inner Mongolia, China

We describe a new dromaeosaurid theropod from the Upper Cretaceous Wulansuhai Formation of Bayan Mandahu, Inner Mongolia. The new taxon, Linheraptor exquisitus gen. et sp. nov., is based on an exceptionally well-preserved, nearly complete skeleton. This specimen represents the fifth dromaeosaurid taxon recovered from the Upper Cretaceous Djadokhta Formation and its laterally equivalent strata, which include the Wulansuhai Formation, and adds to the known diversity of Late Cretaceous dromaeosaurids. Linheraptor exquisitus closely resembles the recently reported Tsaagan mangas. Uniquely among dromaeosaurids, the two taxa share a large, anteriorly located maxillary fenestra and a contact between the jugal and the squamosal that excludes the postorbital from the infratemporal fenestra. These features suggest a sister-taxon relationship between L. exquisitus and T. mangas, which indicates the presence of a unique dromaeosaurid lineage in the Late Cretaceous of Asia. A number of cranial and dental features seen in L. exquisitus and T. mangas, and particularly some postcranial features of L. exquisitus, suggest that these two taxa are probably intermediate in systematic position between known basal and derived dromaeosaurids. The discovery of Linheraptor exquisitus is thus important for understanding the evolution of some salient features seen in the derived dromaeosaurids.

Are mimics monophyletic? The necessity of phylogenetic hypothesis tests in character evolution

BackgroundThe processes governing the origin and maintenance of mimetic phenotypes can only be understood in a phylogenetic framework. Phylogenetic estimates of evolutionary relationships can provide a context for analyses of character evolution; however, when phylogenetic estimates conflict, rigorous analyses of alternative evolutionary histories are necessary to determine the likelihood of a specific history giving rise to the observed pattern of diversity. The polyphenic butterfly Limenitis arthemis provides a case in point. This species is comprised of three lineages, two of which are mimetic and one of which is non-mimetic. Conflicting estimates of the relationships among these three lineages requires direct evaluation of the alternative hypotheses of mimicry evolution.ResultsUsing a coalescent framework, we found support for a sister-taxon relationship between the non-mimetic L. a. arthemis and the mimetic L. a. astyanax, congruent with the previous hypothesis that the non-mimetic form of L. a. arthemis was derived from a mimetic ancestor. We found no support for a mimetic clade (L. a. astyanax + L. a. arizonensis) despite analyzing numerous models of population structure.ConclusionsThese results provide the foundation for future studies of mimicry, which should integrate phylogenetic and developmental analyses of wing pattern formation. We propose future analyses of character evolution accommodate conflicting phylogenetic estimates by explicitly testing alternative evolutionary hypotheses.

Co-estimation of phylogeny and divergence times of Argonautoidea using relaxed phylogenetics

This is the first study to investigate molecular phylogenetic relationships among all four genera of the superfamily Argonautoidea, a clade of diverse pelagic cephalopods with extraordinary characters such as ovoviviparity, dwarf males and secondary "shell" development. Phylogenetic relationships and divergence times within the superfamily were co-estimated using relaxed phylogenetic techniques. A sister-taxon relationship was recovered between Argonauta and Ocythoe and between Tremoctopus and Haliphron. The most recent common ancestor of Argonautoidea was estimated to date from the early Tertiary under the scenario that a lack of a "shell" in the ocythoid lineage is a primary characteristic. In contrast, a later Tertiary most recent common ancestor was estimated under the scenario that a "shell" was present in the early ocythoid lineage and was subsequently lost.

Skeletal morphology and the phylogeny of skuas (Aves: Charadriiformes, Stercorariidae)

The skuas (Aves: Charadriiformes, Stercorariidae) consist of two assemblages. On the basis of size, plumage, and distributional similarities, each of the two assemblages has long been considered monophyletic, and this traditional hypothesis has commonly been manifested in the recognition of two genera, Stercorarius and Catharacta; conversely, more recently collected molecular and ectoparasite evidence yields an alternative hypothesis, in which one member of Stercorarius, Stercorarius pomarinus, is more closely related to the forms in Catharacta than to the other Stercorarius sp. In this study we used skeletal morphology to test the competing hypotheses of skua phylogeny. Cladistic analysis of 141 osteological characters provided strong support for the molecular/ectoparasite hypothesis. However, those skeletal data did not support a sister-taxon relationship between S. pomarinus and Catharacta skua, as inferred from mitochondrial DNA sequence data; instead, they resolved pomarinus as the sister of a monophyletic Catharacta. Additionally, our skeletal evidence did not support a sister-group relationship between skuas and auks, as constraining skua/auk monophyly increased the tree length by nearly 5%. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157, 612–621.

Evolution of seahorses' upright posture was linked to Oligocene expansion of seagrass habitats

Seahorses (Syngnathidae: Hippocampus) are iconic marine teleosts that are readily identifiable by their upright posture. The fossil record is inadequate to shed light on the evolution of this trait because it lacks transitional forms. There are, however, extant syngnathid species (the pygmy pipehorses) that look like horizontally swimming seahorses and that might represent a surviving evolutionary link between the benthic seahorses and other, free-swimming members of the family Syngnathidae. Using sequence data from five nuclear loci, we confirm the sister taxon relationship between seahorses and pygmy pipehorses. Molecular dating indicates that the two taxa diverged during the Late Oligocene. During this time, tectonic events in the Indo-West Pacific resulted in the formation of vast amounts of new shallow-water areas and associated expansion of seagrass habitats that would have favoured the seahorses' upright posture by improving their camouflage while not affecting their manoeuvrability negatively. The molecular techniques employed here provide new insights into the evolution of a taxon whose fossil record is incomplete, but whose evolutionary history is so recent that the major stages of morphological evolution are still represented in extant species.

Neurocranial osteology and systematic relationships ofVaranus(Megalania)priscaOwen, 1859 (Squamata: Varanidae)

The neurocranial osteology of the giant monitor lizard Varanus (Megalania) prisca Owen, 1859 is described in detail for the first time. Optimization of neurocranial characters onto phylogenetic topologies for varanoids, including Lanthanotus, Heloderma, and Varanus species nests V. prisca within an Indo-Australian clade of Varanus on the basis of characters of the otic capsule. A sister-taxon relationship between V. prisca and Varanus komodoensis Ouwens, 1912 is proposed based on apomorphies of the crista prootica, fenestra vestibuli, occipital recess, and supraoccipital. These results support a monophyletic clade of giant monitors among Indo-Australian species, and unambiguously synonymize Megalania with Varanus at both generic and subgeneric levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155, 445–457.