A new specimen of Biseridens qilianicus indicates its phylogenetic position as the most basal anomodont

A new well-preserved basal therapsid skull from the Xidagou Formation, Middle Permian of China, is identified as Biseridens qilianicus. The following synapomorphies distinguish Biseridens as an anomodont and not an eotitanosuchian as previously described: short snout; dorsally elevated zygomatic arch and septomaxilla lacking elongated posterodorsal process between nasal and maxilla. The presence of a differentiated tooth row; denticles on vomer, palatine and pterygoid; contact between tabular and opisthotic; lateral process of transverse flange of pterygoid free of posterior ramus and absence of mandibular foramen exclude it from other anomodonts. Our cladistic analysis indicates Biseridens to be the most basal anomodont, highlights separate Laurasian and Gondwanan basal anomodont clades and suggests that dicynodonts had their origins in the Gondwanan clade. The co-occurrence of the most basal anomodont (Biseridens) together with the most basal therapsid (Raranimus), basal anteosaurid dinocephalians, bolosaurids and dissorophids suggests that the earliest therapsid faunas are from China.     

Growth correlations inBryophyllum leaves and exogenous growth regulators

Growth correlations in leaves ofBryophyllum may be recognized by the development of marginal shoots varying in their particular lamina regions, the correlative inhibition increasing from the top to the base. Cytokinins extend their promoting action inB. crenatum leaves farther in the apical than in the basal direction. In the uppermost leaves ofB. daigremontianum they evoke the development of marginal shoots with flowers in the apical and of vegetative shoots in the basal region. Less suitable for this research are auxins, gibberellins, and triiodobenzoic acid which are unable to stimulate the development of marginal shoots on leaves grown out under short days if not supplemented by cytokinins. Only TIBA induces under long days formation of adventitious shoots directly from the lamina surface on leaves developed below the terminal ring fasciation exhibiting at free ends of connate leaves normal marginal shoots. Under short days a complete reduction of lateral teeth takes place inB. verticillatum both on the extremity of ring fasciation and on the uppermost leaves, the marginal shoots being formed under the subsequent long days only in the angles between the ring fasciation parts. Furthermore the correlative inhibition of the leaf causes anisophylly in marginal shoots in the same way as that of axillaries inBryophyllum.     

Anatomy and relationships of Pachyrhachis problematicus,a primitive snake with hindlimbs

The anatomy of Pachyrhachis problematicus, an elongate, limb-reduced squamate from the Upper Cretaceous of Israel, is described and evaluated in detail. Previously considered a snake-like ''lizard'' of uncertain affinities, it is here shown to be the most primitive snake, and the sister-group to all other snakes. Pachyrhachis exhibits numerous derived characters uniting it with modern snakes (scolecophidians and alethinophidians): e.g. mobile premaxilla-maxilla articulation, braincase enclosed by frontals and parietals, sagittal parietal crest, absence of tympanic recess, single postdentary bone, over 140 presacral vertebrae, and complete loss of shoulder girdle and forelimb. However, it is more primitive than all modern snakes in retaining some strikingly primitive (lizard-like) features: presence of a jugal, squamosal, normal sacral attachment, and well-developed hindlimb composed of femur, tibia, fibula, and tarsals. Pachyrhachis provides additional support for the hypothesis that snakes are most closely related to Cretaceous marine lizards (mosasauroids). Almost all of the derived characters proposed to unite snakes and mosasauroids are highly developed in Pachyrhachis: the mobile mandibular symphysis, intramandibular joint, long and recurved pterygoid teeth, quadrate suspended by the supratemporal, loosely united pelvic elements (ilium, ischium, and pubis), and separate astragalus and calcaneum.     

A new species of Theriosuchus (Atoposauridae, Crocodylomorpha) from the Late Jurassic (Kimmeridgian) of Guimarota, Portugal

A new species of an atoposaurid crocodilian, Theriosuchus guimarotae, is described from the Late Jurassic (Kimmeridgian) of Portugal. Theriosuchus guimarotae can be distinguished from other species of Theriosuchus by a lateral surface of squamosal bevelled ventrally; a rounded, caudally projecting and dorsally sculptured caudolateral corner of the squamosal; a premaxillomaxillary suture aligned caudomedially in dorsal aspect; a minimum space between the supratemporal foramina that comprises one third of the total width of the cranial table; a minimum width of the frontal between the orbits that comprises one third of the maximum width of the skull at the orbits; a dentition that comprises only pseudocaniniform and lanceolate-shaped teeth; the presence of an external mandibular fenestra and all vertebral bodies amphicoelous. Its osteology also sheds light on the diagnosis of Theriosuchus within Atoposauridae. The material additionally includes specimens representative of several ontogenetic stages, each of which is discussed here. With its Late Jurassic age, T. guimarotae represents the oldest well-preserved material of Theriosuchus and reveals further knowledge about the palaeobiogeography of the genus in western Europe.     

Burkholderia sp. Induces Functional Nodules on the South African Invasive Legume Dipogon lignosus (Phaseoleae) in New Zealand Soils

The South African invasive legume Dipogon lignosus (Phaseoleae) produces nodules with both determinate and indeterminate characteristics in New Zealand (NZ) soils. Ten bacterial isolates produced functional nodules on D. lignosus. The 16S ribosomal RNA (rRNA) gene sequences identified one isolate as Bradyrhizobium sp., one isolate as Rhizobium sp. and eight isolates as Burkholderia sp. The Bradyrhizobium sp. and Rhizobium sp. 16S rRNA sequences were identical to those of strains previously isolated from crop plants and may have originated from inocula used on crops. Both 16S rRNA and DNA recombinase A (recA) gene sequences placed the eight Burkholderia isolates separate from previously described Burkholderia rhizobial species. However, the isolates showed a very close relationship to Burkholderia rhizobial strains isolated from South African plants with respect to their nitrogenase iron protein (nifH), N-acyltransferase nodulation protein A (nodA) and N-acetylglucosaminyl transferase nodulation protein C (nodC) gene sequences. Gene sequences and enterobacterial repetitive intergenic consensus (ERIC) PCR and repetitive element palindromic PCR (rep-PCR) banding patterns indicated that the eight Burkholderia isolates separated into five clones of one strain and three of another. One strain was tested and shown to produce functional nodules on a range of South African plants previously reported to be nodulated by Burkholderia tuberum STM678^T which was isolated from the Cape Region. Thus, evidence is strong that the Burkholderia strains isolated here originated in South Africa and were somehow transported with the plants from their native habitat to NZ. It is possible that the strains are of a new species capable of nodulating legumes.     

The early anomodont Venjukovia and the evolution of the anomodont skull

The skull morphology and function of the anomodont mammal-like reptile Venjukovia is described and compared with that of Patranomodon, Eodicynodon, Galeops and Diictodon. Changes in osteology and functional anatomy of these forms are noted.
It is concluded that the acquisition of herbivorous adaptations in the anomodonts may have occurred in the following sequence: division of the external adductor muscles of the jaw into a medial and lateral component, propalinal jaw movement, increased Moment Arm of the external adductors by raising the level of the jaw hinge, anterior migration of the adductor insertions, reduction of the size and number of teeth and acquisition of a horn covering to the jaws, elaboration of the external adductors and reduction of the internal adductors (pterygoideus muscles), and elaboration of palatal areas for mastication of food.
The first (in the morphological sense) committed herbivorous anomodont is identified as the South African form, Galeops. As far as anomodonts are concerned, the major correlate of successful herbivorous feeding is the acquisition of propaliny.
A chronology of character acquisition in the anomodonts cannot yet be drawn up since the stratigraphic correlation of Russian and South African sequences is uncertain. However, it appears that the transition from a generalized carnivore to a committed herbivore took place very rapidly compared to the species longevity of anomodonts.     

Cranial anatomy and phylogenetic position of Suminia getmanovi, a basal anomodont (Amniota: Therapsida) from the Late Permian of Eastern Europe

Suminia getmanovi , a recently discovered basal anomodont from the Late Permian of Russia, is characterized by robust, 'leaf-shaped' teeth, and a masticatory architecture that is similar to that of the highly diverse and cosmopolitan group of Permo-Triassic herbivores, Dicynodontia (Anomodontia). Based on new material, the skull is reconstructed in three dimensions and described in detail. A cladistic analysis of the basal anomodonts, Patranomodon, Galeops, Otsheria, Ulemica , and Suminia , using 37 cranial characters, resulted in a single most parsimonious tree, in which Suminia is united with the Russian taxa, Ulemica and Otsheria. This clade, diagnosed by four unambiguous characters, is designated as Venyukovioidea. The South African anomodont, Galeops , appears as the sister taxon to Dicynodontia. Patranomodon is the most basal anomodont. The cladistic analysis suggests that a 'dicynodont-type' masticatory architecture, with an expanded adductor musculature and sliding jaw articulation, may have originated prior to the advent of the (Venyukovioidea + ( Galeops + Dicynodontia)) clade.     

Structural heterogeneity in the basal regions of the teeth of the red-backed salamander,Plethodon cinereus (Amphibia,Plethodontidae)

Ultrastructural and histochemical features of marginal (monostichous) teeth associated with the jawbones are compared with those of palatal (polystichous) teeth that compose two patches in the roof of the mouth. The apices and uncalcified regions are similar in both kinds of teeth, but the basal regions display distinctive differences. While bases (pedestals) of marginal teeth are essentially hollow cylinders that attach to the jawbones by their labial faces, bases of teeth in palatal patches are fused to form two horizontal plates which lack direct attachment to underlying bone. The plates are separated from each other by a pulp-filled space containing fibroblasts, blood vessels, and vertically oriented elements resembling bony spicules. Cylindrical pedestals like those of marginal teeth project from the ventral plate. While the identity of the material composing the basal regions remains controversial, the following evidence suggests that it is similar to “bone of attachment” (Tomes, '23): most of it, unlike dentin, does not develop in direct association with an enamel organ; alcian blue stains the bases of developing teeth but stains dentin, developing dentin, enamel, or mature bone very weakly (if at all); bases of teeth in palatal patches develop in isolation from the parasphenoid bone and thus cannot be considered extensions of it; and marginal teeth attach directly to the jawbones, but the material composing their bases does not blend with the bone. Structural heterogeneity of the basal regions appears to be linked to functional differences exhibited by these two types of teeth.     

First procolophonid (Reptilia,Parareptilia) from the Lower Triassic of Madagascar

Lasasaurus beltanae nov. gen. nov. sp., a new procolophonid (Reptilia, Parareptilia) from the Lower Triassic of Madagascar, is represented by a single partial skeleton preserved in a ferro-calcareous nodule from the Middle Sakamena Formation, in the North of the island. This new taxon is unique in possessing peculiar, fine and dendritic crests running along the posterolateral side of the squamosal, widely spaced maxillary teeth, subparallel mesiodistal ridges connecting maxillary teeth to the tooth row, and a strongly acute anterior margin of the copula (hyoid bone). This well-preserved specimen belongs to a juvenile individual. The inclusion of L. beltanae nov. gen. nov. sp. in a phylogenetic analysis suggests that it is close to Theledectinae, Procolophoninae, and Leptopleurinae, though their respective relationships are uncertain. This specimen is the first procolophonid described from Madagascar and represents a minor terrestrial component of a coastal vertebrate assemblage dominated by amphibious to fully-aquatic taxa.     

A Finding of the Red Squirrel (Rodentia,Sciuridae, <Emphasis Type="Italic">Sciurus vulgaris</Emphasis> Linnaeus, 1758) and Forest Dormouse (Rodentia,Gliridae, <Emphasis Type="Italic">Dryomys nitedula</Emphasis> Linnaeus, 1778) in the Pleistocene of the Southern Urals

The study deals with a new sample of the mammalian fossils from the Pleistocene deposits of the Ignatievskaya Cave in Southern Urals (54°53' N, 57°46' E). Among the rodent fossils, the teeth of red squirrel (Sciurus vulgaris) and forest dormouse (Dryomys nitedula) have been identified. The enamel differentiation quotient (SDQ) of the water vole teeth matches the Arvicola terrestris species from locations of the end of Middle and Late Pleistocene (oxygen isotope stages, OIS 6 and OIS 5–2, respectively). The high SDQ value for the water vole and the presence of the squirrel and dormouse in the fauna allow dating the finding near the end of the Middle Pleistocene and beginning of the Late Pleistocene (OIS 6 and OIS 5e, respectively). The widespread open and forest landscapes were inhabited by this fauna.     

New basal procolophonid reptile from the Katberg formation (Lower Triassic) of the South African Karoo

A new procolophonid reptile, Kitchingnathus untabeni n. gen. et n. sp., is described from the uppermost strata of the Lystrosaurus Assemblage Zone of the Karoo Basin, South Africa. The new taxon co-occurs with the well-known Procolophon trigoniceps. The most distinctive feature of the new taxon is the presence of numerous small bicuspid molariforms in both the maxilla and the dentary. A phylogenetic analysis indicates that Kitchingnathus occupies a basal position among procolophonids. Character optimisation suggests that bicuspid teeth were acquired independently by the new taxon, and originated twice in procolophonid evolution.     

A preliminary revision of sordidus-like species of Ilyocryptus Sars, 1862 (Anomopoda, Branchiopoda) in North America, with description of I. bernerae n. sp.

Presently the inter-generic structure of the Ilyocryptus Sars, 1862 is not clear, and we regard as `sordidus-like' all the species with (1) incomplete moulting, (2) valves without lateral horns or other remarkable structures, (3) non-distal position of anus on postabdomen, (4) long lateral setae on the postabdomen, which reaches the anus; (5) single or doubled teeth on preanal margin of postabdomen. Available material on sordidus-like forms of Ilyocryptus Sars, 1862 (Anomopoda, Branchiopoda) from North America was investigated. We also analyzed a limited amount of material similar to I. sordidus s. str., but not enough to be presented now. Three other conventional species were found and redescribed: the exclusively North American I. gouldeni Williams, 1978, as well as I. cuneatus?tifter, 1988 and I. spinosus?tifter, 1988, both described earlier from Europe. An original analysis of the differences between all species was performed. A new sordidus-like species, I. bernerae n. sp., from a single locality in South Carolina, U.S.A., is described. The main diagnostic features of this new species within sordidus-like members of the Ilyocryptusare: thin and sharp dorsal keel; exclusively single preanal teeth, greatly increasing in size in basal direction; lateral setae on postabdomen, not decreasing in size basad; absence of denticles on base of postabdomen; complete absence of any denticles on distal and middle portion of ventral margin of postabdominal claws; distalmost spine on claw base longer than basalmost; bases of antennules compressed against each other (although not touching); slightly different armature of two sides of lateral swimming setae; presence of small hooks on tips of lateral swimming setae; distal segments of setae on trunk of limb I setulated only in basal portion; sensillum on gnathobase II curved.     

Craniodental morphology and systematics of a new family of hystricognathous rodents (Gaudeamuridae) from the late eocene and early oligocene of Egypt

Background

Gaudeamus is an enigmatic hystricognathous rodent that was, until recently, known solely from fragmentary material from early Oligocene sites in Egypt, Oman, and Libya. Gaudeamus'' molars are similar to those of the extant cane rat Thryonomys, and multiple authorities have aligned Gaudeamus with Thryonomys to the exclusion of other living and extinct African hystricognaths; recent phylogenetic analyses have, however, also suggested affinities with South American caviomorphs or Old World porcupines (Hystricidae).

Methodology/Principal Findings

Here we describe the oldest known remains of Gaudeamus, including largely complete but crushed crania and complete upper and lower dentitions. Unlike younger Gaudeamus species, the primitive species described here have relatively complex occlusal patterns, and retain a number of plesiomorphic features. Unconstrained parsimony analysis nests Gaudeamus and Hystrix within the South American caviomorph radiation, implying what we consider to be an implausible back-dispersal across the Atlantic Ocean to account for Gaudeamus'' presence in the late Eocene of Africa. An analysis that was constrained to recover the biogeographically more plausible hypothesis of caviomorph monophyly does not place Gaudeamus as a stem caviomorph, but rather as a sister taxon of hystricids.

Conclusions/Significance

We place Gaudeamus species in a new family, Gaudeamuridae, and consider it likely that the group originated, diversified, and then went extinct over a geologically brief period of time during the latest Eocene and early Oligocene in Afro-Arabia. Gaudeamurids are the only known crown hystricognaths from Afro-Arabia that are likely to be aligned with non-phiomorph members of that clade, and as such provide additional support for an Afro-Arabian origin of advanced stem and basal crown members of Hystricognathi.     

The phylogeny of varanoid lizards and the affinities of snakes

Evidence that platynotan squamates (living varanoid lizards, snakes and their fossil relatives) are monophyletic is presented. Evolutionary relationships within this group are then ascertained through a cladistic analysis of 144 osteological characters. Mosasauroids (aigialosaurs and mosasaurs), a group of large marine lizards, are identified as the nearest relatives of snakes, thus resolving the long-standing problem of snake affinities. The mosasauroid–snake clade (Pythonomorpha) is corroborated by 40 derived characters, including recumbent replacement teeth, thecodonty, four or fewer premaxillary teeth, supratemporal–prootic contact, free mandibular tips, crista circumfenestralis, straight vertical splenio-angular joint, loss of posterior ramus of the coronoid, reduced basipterygoid processes, reduced interpterygoid vacuity, zygosphene–zygantral articulations, and absence of epiphyses on the axial skeleton and skull. After mosasauroids, the next closest relatives of snakes are varanids (Varanus, Saniwa and Saniwides) and lanthanotids (Lanthanotus and Cherminotus). Derived features uniting varanids and lanthanotids include nine cervical vertebrae and three or fewer pairs of sternal ribs. The varanid–lanthanotid–pythonomorph clade, here termed Thecoglossa, is supported by features such as the anteriorly positioned basal tubera, and the loss of the second epibranchial. Successive outgroups to thecoglossans are Telmasaurus, an unresolved polytomy (Estesia, Gobidermatidae and Helodermatidae), Paravaranus and Proplatynota. The ''necrosaurs'' are demonstrated to be an artificial (polyphyletic) assemblage of primitive platynotans that are not particularly closely related to each other.Snakes are presumed to have evolved from small, limbless, burrowing lizards and the inability of previous analyses to resolve the affinities of snakes has been attributed to extensive convergence among the numerous lineages of such lizards. The present study contradicts this claim, demonstrating that the problem is due instead to omission of critical fossil taxa. No modern phylogenetic analysis of squamate relationships has simultaneously included both mosasauroids and snakes: previous studies have therefore failed to identify the mosasauroid–snake association and the suite of derived characters supporting it. Mosasauroids are large aquatic animals with well-developed appendages, and none of the derived characters uniting mosasauroids and snakes is obviously correlated with miniaturization, limb reduction or fossoriality. Recognition that mosasauroids, followed by varanids and lanthanotids, are the nearest relatives of snakes will also facilitate studies of relationships within snakes, which until now have been hampered by uncertainty over the most appropriate (closely-related) lizard outgroups.     

The last hadrosaurid dinosaurs of Europe: A new lambeosaurine from the Uppermost Cretaceous of Aren (Huesca, Spain)

A new hadrosaurid dinosaur, Arenysaurus ardevoli gen. et sp. nov., from the Late Maastrichtian of Aren (Huesca, South-central Pyrenees) is described on the basis of a partial, articulated skull, mandibular remains and postcranial elements, including vertebrae, girdle and limb bones. Arenysaurus is characterized by having a very prominent frontal dome; nearly vertical prequadratic (squamosal) and jugal (postorbital) processes, and deltopectoral crest of the humerus oriented anteriorly. Moreover, it possesses a unique combination of characters: short frontal (length/width approximately 0.5); midline ridge of parietal at level of the postorbital-squamosal bar; parietal excluded from the occiput; squamosal low above the cotyloid cavity. A phylogenetical analysis indicates that Arenysaurus is a rather basal member of Lambeosaurinae and the sister-taxon to Amurosaurus and the Corythosaurini-Parasaurolophini clade. The phylogenetic and biogeographical relationships of Arenysaurus and other lambeosaurines suggest a palaeogeographical connection between Asia and Europe during the Late Cretaceous.     

The dentition of Amygdalodon patagonicus (Dinosauria: Sauropoda) and the dental evolution in basal sauropods

Amygdalodon patagonicus is the oldest record of Sauropoda from South America and is known from several teeth and fragmentary postcranial remains. Here we describe in detail its dental morphology, characterized by the presence of broad spatulated teeth (with low SI values) and the absence of denticles in their crowns. The enamel bears a particular wrinkling pattern composed only of apicobasally-aligned pits, which are frequently joined to each other by a continuous sulcus. Some worn teeth have large wear-facets that extend over only one of the crown's edges. This unique combination of characters provides a proper diagnosis for Amygdalodon. Despite the fragmentary available material, Amygdalodon is here interpreted as a non-eusauropod sauropod based on the results of a phylogenetic analysis. The presence of derived dental characters in Amygdalodon, such as the presence of tooth-tooth occlusion, shows several features previously thought to diagnose Eusauropoda or Gravisauria appeared earlier during the early evolution of sauropods.     

Pseudogenization of the tooth gene enamelysin (MMP20) in the common ancestor of extant baleen whales

Whales in the suborder Mysticeti are filter feeders that use baleen to sift zooplankton and small fish from ocean waters. Adult mysticetes lack teeth, although tooth buds are present in foetal stages. Cladistic analyses suggest that functional teeth were lost in the common ancestor of crown-group Mysticeti. DNA sequences for the tooth-specific genes, ameloblastin (AMBN), enamelin (ENAM) and amelogenin (AMEL), have frameshift mutations and/or stop codons in this taxon, but none of these molecular cavities are shared by all extant mysticetes. Here, we provide the first evidence for pseudogenization of a tooth gene, enamelysin (MMP20), in the common ancestor of living baleen whales. Specifically, pseudogenization resulted from the insertion of a CHR-2 SINE retroposon in exon 2 of MMP20. Genomic and palaeontological data now provide congruent support for the loss of enamel-capped teeth on the common ancestral branch of crown-group mysticetes. The new data for MMP20 also document a polymorphic stop codon in exon 2 of the pygmy sperm whale (Kogia breviceps), which has enamel-less teeth. These results, in conjunction with the evidence for pseudogenization of MMP20 in Hoffmann''s two-toed sloth (Choloepus hoffmanni), another enamel-less species, support the hypothesis that the only unique, non-overlapping function of the MMP20 gene is in enamel formation.     

Changing status of the parasites of potato tuber mothPhthorimaea operculella [Lepidoptera: Gelechiidae] in Australia

Three introduced parasites ofPhthorimaea operculella (Zeller) are now widespread, and a fourth established in a limited area, in Australia.Apanteles subandinus Blanchard, which occurs from Tasmania to Queensland, is the most effective species and the outstanding parasite in Victoria. The extent of parasitism and the status of the various parasites were monitored from the time of their release until 1971. In many areas whereCopidosoma koehleri Blanchard was the only parasite found in 1966, a substantial increase in parasitism was recorded in subsequent years, following the liberation of other species. In the 1969–70 season there was an explosive increase inOrgilus lepidus Muesebeck, which became the dominant species in some coastal areas of New South Wales, displacingA. subandinus. Its change of status to a dominant position was associated with a change of sex ratio from 1.6 ♀: 1♂ to 1.0 ♀: 1 ♂.     

Niche Partitioning in Oligocene Toothed Mysticetes (Mysticeti: Aetiocetidae)

Niche partitioning has long been considered to be a fundamental part of speciation and difference in body size is a straightforward proxy for ecology and niche partitioning. Here we report a new aetiocetid specimen from the upper part of Morawan Formation (Oligocene; late Chattian, ~26–24 Ma), Ashoro, Hokkaido, Japan. This specimen comprises an isolated left squamosal and an isolated left periotic; the morphology of the squamosal is distinguishable from all aetiocetids, but shows a close match with the aetiocetid Morawanocetus yabukii in that: the dorsal margin of the zygomatic process of the squamosal is twisted dorsomedially; the zygomatic process expands dorsoventrally; the zygomatic process is transversely thin; and the anterior tip of the zygomatic process is acutely tapered. The estimated body size of this Morawanocetus-like specimen is ca. 8 m, almost twice the size of all known aetiocetids (3–4 m). Moreover, the preserved periotic reveals structures that are otherwise poorly known in aetiocetids. This large Morawanocetus-like aetiocetid sheds new light on niche partitioning of Oligocene toothed mysticetes and evolution of body size in Mysticeti.