The evolution of the dicynodont feeding system

The skull structure of dicynodonts may be regarded as a complex adaptation towards herbivorous feeding. The present work examines how and why this adaptation may have evolved. A cladogram of the dicynodonts is presented and from it a sequence of hypothetical ancestral forms is inferred. The jaw musculature of dicynodonts and other therapsids is described and in particular the early dicynodont Eodicynodon oosthuizeni is described in detail. This information is used to draw up a sequence of ancestral stages whose basic skull anatomy, jaw muscle organization and masticatory properties are described. Differences in masticatory properties between these stages are pinpointed and an explanation to account for the development of these differences is advanced. It is concluded that the changes in skull organization seen during the evolution of dicynodonts are consistent with the hypothesis that a propalinal jaw action was being improved by selection, and that this was required to permit dicynodonts to be efficient herbivores.     

The origin and early radiation of the therapsid mammal-like reptiles: a palaeobiological hypothesis

The replacement of the basal synapsid pelycosaurs by the more 'mammal-like' therapsids in the Permian was an important event in the history of tetrapods because it initiated the eventual transition to the mammals. It is also an example of taxon replacement in the fossil record that is unusually amenable to explanation, based on a combination of analysis of the biological significance of the inferred character changes, with the stratigraphic, palaeogeographic and palaeoecological circumstances of the time. An hypothesis is presented in which the origin of the therapsids resulted from a correlated progression of character evolution leading to higher levels of metabolic activity and homeostatic regulation of the body. It was a response to the availability of a seasonally arid, savanna-like biome. The subsequent explosive radiation of therapsids was associated with habitat expansion made possible by the Mid-Permian development of geographical continuity between that biome and the temperate biomes. The final extinction of the pelycosaurs was a case of incumbent replacement by the new therapsid lineages.     

A new basal dicynodont from the Upper Permian of South Africa

The skull of a small anomodont therapsid, from the Tapinocephalus Assemblage Zone (Abrahamskraal Formation, Beaufort Group, Upper Permian) in Northern Cape Province, South Africa, represents a new basal dicynodont and is described in detail. Colobodectes cluveri gen. et sp. nov. is distinguished from other dicynodonts by an anteroposteriorly extensive caniniform process, parietals that were broadly overlapped posterolaterally by posterodorsal processes of the postorbitals, diverging anterior palatal ridges, and a dorsoventrally low foramen magnum. A phylogenetic analysis indicates that Colobodectes is the basalmost member of a dicynodont clade that excludes Eodicynodon . This position is not particularly strong, as two additional steps are needed to make Colobodectes and Eodicynodon oosthuizeni exchange places on the most parsimonious tree. Another discovery of the phylogenetic analysis is that there is little basis for recognizing Eodicynodon oelofseni as the closest relative of E. oosthuizeni . The former species is identified as the sister taxon of a clade that includes the latter and all other dicynodonts.     

A new burnetiamorph (Therapsida: Biarmosuchia) from the Lower Beaufort Group of South Africa

The basal clade Burnetiamorpha is known from only two specimens representing two genera, Proburnetia from the Severodvinskian horizon of the Vyatka River Basin in the Kotelnich district of Russia, and Burnetia from the Dicynodon Assemblage Zone of the Beaufort Group of South Africa. Both genera are of Late Tatarian (Late Permian) age. This paper describes the cranial morphology of a new genus of burnetiamorph, Bullacephalus , from the Late Permian Tapinocephalus Assemblage Zone of the Beaufort Group of South Africa. It is known from a relatively complete skull and lower jaw and is the best preserved burnetiamorph yet discovered. Apart from being the oldest member of the clade, Bullacephalus is also morphologically the least derived and provides new evidence on the phylogeny of this poorly understood group of basal therapsids.     

A primitive anomodont therapsid from the base of the Beaufort Group (Upper Permian) of South Africa

Patranomodon nyaphulii , known from a nearly complete skull, lower jaw and partial postcranial skeleton, is morphologically the most primitive anomodont therapsid yet discovered. It is from the Eodicynodon Assemblage Zone, the lowest biozone of the Beaufort Group of South Africa, which has a primitive therapsid fauna comparable with that of the Russian Zone I. Patranaomodon is primitive with respect to other anomodonts in having short palatal exposure of the premaxilla, an unreduced tabular, a slit-like interpterygoidal vacuity, a screw-shaped jaw articulation (which precludes fore-aft sliding of the lower jaw), and only three sacral vertebrae. The poorly-known Galechirus and Galepus from the younger Cistecephalus Assemblage Zone appear to be at a comparably primitive evolutionary grade, and the three genera are tentatively united in the family Galechiridae. The taxon Dromasauria is shown to be paraphyletic and therefore should be discarded.     

A New Basal Lystrosaurid Dicynodont from the Upper Permian of South Africa

A new genus and species of late Permian dicynodont, Kwazulusaurus shakai , is described on the basis of a complete skull from the late Permian Dicynodon Assemblage Zone of the South African Beaufort Group. It is an advanced form which shows characters, such as the shape of the snout and the loss of the ectopterygoid, that link it to the early Triassic genus Lystrosaurus. Kwazulusaurus represents the most basal member of the lystrosaurian lineage. The phylogeny of progressive pristerodontian dicynodonts is discussed. It appears possible that the Kannemeyeriiformes and Lystrosauridae do not form a monophylum, as previously assumed. Instead a sister-group relationship between lystrosaurids and dicynodontids plus kannemeyeriiforms is suggested.     

Virtual reconstruction and description of the cranial endocast of Pristerodon mackayi (Therapsida,Anomodontia)

The cranial endocast of Pristerodon mackayi is described, which has been virtually reconstructed on the basis of neutron tomographic data. The gross anatomy of the cranial endocast of Pristerodon resembles other nonmammalian synapsids such as Thrinaxodon liorhinus in having a narrow, tubular forebrain, well developed olfactory bulbs, a large parietal foramen and unossified zone. As it is the case in cynodonts the hindbrain of Pristerodon is broader than the mid‐ and forebrain. Large paraflocculi are developed. The medulla oblongata can be well distinguished from the pons. The pons is divided by a median ridge into two portions. There is no evidence for a neocortex, which seems to be also reflected in the low encephalization quotient of 0.18 estimated according to the method of Eisenberg. J. Morphol. 276:1089–1099, 2015. © 2015 Wiley Periodicals, Inc.     

Vertebrate assemblages from the north‐central Main Karoo Basin,South Africa,and their implications for mid‐Permian biogeography

The rich fossil vertebrate record from the Beaufort Group, Main Karoo Basin, provides a global standard for mid‐Permian to Mid‐Triassic continental faunas. However, recent studies have demonstrated variability in the composition of contemporaneous faunas across Gondwana. This raises the question of how much the vertebrate faunas differ within the Karoo, where the taxonomic composition of vertebrate assemblage zones (AZs) is mostly considered to be uniform. Although fossil material is known from across the outcrop of the Beaufort Group, the lowest Beaufort strata have received little attention, particularly north of S31°10′. Here, we report two fossil tetrapod assemblages from the lowest Beaufort Group in the southern Free State Province, which represent the northernmost point at which the lowest Beaufort has been targeted for collecting. The lower assemblage is characterized by an abundance of the small dicynodont Eosimops and can thus be attributed to the Tapinocephalus AZ (Guadalupian), but the absence of dinocephalian or pareiasaurian material is unlike contemporaneous assemblages found further south. This suggests that the Tapinocephalus AZ was not uniform across the entire basin and highlights that the abundance, distribution and taxonomic composition of Karoo biozones may vary more than currently appreciated. The upper assemblage, characterized by the dicynodonts Oudenodon, Aulacephalodon and Dinanomodon, is attributable to the upper Cistecephalus AZ to lower Daptocephalus AZ. The juxtaposition of the lower Tapinocephalus AZ and upper Cistecephalus\lower Daptocephalus AZ in the southern Free State implies a stratigraphic gap from the Middle to Late Permian of up to 6 million years.     

Morphology of the palate and braincase of Dimetrodon milleri

The palate and partial braincase of the holotype of Dimetrodon milleri (MCZ 1365) are preserved in three dimensions, but have yet to be described in detail. Here, we describe these structures for the first time for this species, and compare them with the better-known specimens of D. limbatus. Interesting characteristics of the morphology include the patterns of articulation of the palatal elements, including the palatine and vomer, and anatomy of the pterygoid in the posterior region of the palatal vacuities. Dimetrodon milleri is found to differ from D. limbatus in the lack of teeth on the ectopterygoid, the shape of the basal process of the epipterygoid, and the anterior extent of the palatine and pterygoid. The two species are similar in the relative position of the basicranial articulation, but differ significantly from that in other sphenacodontids, including Secodontosaurus and Sphenacodon. The evolution of these cranial features will be the subject of future phylogenetic analyses of sphenacodontids.     

Environmental History and the Desertification of the Karoo,South Africa

Abstract

This paper reviews the desertification debate in South Africa and emphasizes the methods that have been used to understand the environmental history of the semi–arid rangelands in the eastern Karoo during the last 500 years. These mixed grass/dwarf shrub rangelands, with mean annual rainfall totals typically between 300–400 mm, are described and the main driving variables discussed. A brief historical account of the European settlement of the region is also presented. The desertification debate has focussed on three main issues. Firstly, because arid and semi–arid lands appear to be heavily influenced by climate, the changing climatic regime during the Holocene and particularly during the last two hundred years has been the subject of intensive investigation. While all studies conclude that there is no evidence in the historical record to support the popular perception that rainfall totals have decreased this century, the question of changing rainfall seasonality has not been adequately explored. Although mean annual temperatures over the last 50 years have not changed there have been significant increases in mean monthly maximum temperatures and significant decreases in mean monthly minimum temperatures for some of the stations investigated. The second issue of great interest in the South African desertification literature is that of the nature of pre–colonial environments. A wide variety of archaeological, historical and ecological techniques have been used, including analyses of fossil mammal bones in owl pellets, fossil pollen in hyrax middens, notes from traveller's accounts of the region and stable carbon isotopes in soil as well as fossil mammal bones. While all authors agree that the eastern Karoo was more grassy at some stage in the past there is disagreement as to both the timing and cause of the changes to a more shrubby vegetation. The final issue of great concern to the desertification debate in South Africa concerns the rate of change during the last 100 years. Satellite imagery, matched ground and aerial photography, survey data and an analysis of historical stock records cannot agree as to whether the Karoo is degrading or not. Certainly, the classic view of an annually expanding desert margin has been replaced in recent years by a more realistic understanding of the seasonal dynamics of the vegetation. The recent trend to detailed modelling of the demographic process in key species holds much promise for our understanding of the degradation process. The vibrant community of researchers, employing a range of archaeological, historical and ecological techniques, will make important contributions to South Africa's National Action Plan to Combat Desertification.     

The distribution and biology of nomadic birds in the Karoo, South Africa

Dryland nomadic bird species, as a proportion of all bird species in a biome in southern Africa, are highest in the arid grassland and arid and semi-arid Karoo in South Africa. Nomadic birds, of which the most widespread species is the greybacked finchlark Eremopterix verticalis (Smith), are most frequently observed in the north-central and north western Nama Karoo. The species richness of nomadic species is inversely correlated with species richness of all bird species in the Karoo. Since the distribution of nomadic birds is in areas where rainfall is patchy, low (<250 mm per year) and aseasonal, this supports the idea that fewer species are able to cope with resources that are patchy in time and space, and that there has been selection for nomadism in the species that are able to use patchy environments. Species richness and abundance of nomadic birds is negatively correlated with rainfall amount but positively correlated with the coefficient of variation of the rainfall and with rainfall in autumn. The frequency of nomadic birds is inversely correlated with altitude range; nomadic species are most often recorded in structurally simple habitats (shrubland and grassland) on open plains. Most nomadic bird species in the Karoo are granivorous. Perennial desert grasses are important components of the habitat and diet of small nomadic granivores, and also provide nest sites and nest material. Nomadic birds can breed throughout the year, without a clearly defined ‘season’ in both the Succulent and Nama Karoo. Average clutch sizes do not differ significantly between resident and all nomadic species in the arid and semi-arid Karoo. Nomadism is an evolutionary stable strategy for individual species only when extremes in environmental conditions are frequent enough, and unpredictable enough, to maintain movements to high resource patches or to maintain dispersal away from low resource patches. If high rainfall years are too regular or infrequent, or peaks in fluctuations of resources in the environment too low, or rainfall patches are randomly distributed, nomadism would not be maintained as part of the individual behaviour pattern.     

Classification and phylogeny of the diapsid reptiles

Reptiles with two temporal openings in the skull are generally divided into two groups–the Lepidosauria (lizards, snakes, Sphenodon , 'eosuchians') and the Archosauria (crocodiles, thecodontians, dinosaurs, pterosaurs). Recent suggestions that these two are not sister-groups are shown to be unproven, whereas there is strong evidence that they form a monophyletic group, the Diapsida, on the basis of several synapomorphies of living and fossil forms. A cladistic analysis of skull and skeletal characters of all described Permo-Triassic diapsid reptiles suggests some significant rearrangements to commonly held views. The genus Petrolacosaurus is the sister-group of all later diapsids which fall into two large groups–the Archosauromorpha (Pterosauria, Rhynchosauria, Prolacertiformes, Archosauria) and the Lepidosauromorpha (Younginiformes, Sphenodontia, Squamata). The pterosaurs are not archosaurs, but they are the sister-group of all other archosauromorphs. There is no close relationship between rhynchosaurs and sphenodontids, nor between Prolacerta or Tanystropheus and lizards. The terms 'Eosuchia', 'Rhynchocephalia' and 'Protorosauria' have become too wide in application and they are not used. A cladistic classification of the Diapsida is given, as well as a phylogenetic tree which uses cladistic and stratigraphic data.     

Evaluation of Restoration Techniques for the Succulent Karoo, South Africa

Abstract Possible constraints on the passive recovery of bare areas in the Karoo, a semiarid region in South Africa, include inadequate supply of seed, availability of suitable microsites for plant establishment, altered soil properties, and the truncation of key soil biotic processes. Here we investigate the possibility of initiating the restoration of bare areas by soil surface treatments with gypsum (CaSO₄) and/or organic mulch. We also apply an exogenous seed source to test the hypothesis that seed availability limits autogenic recovery. Both gypsum and mulch improved rain water infiltration, gypsum more so than mulch, and both treatments resulted in significantly higher numbers of reseeded seedlings compared with controls. Gypsum also improved the survival of the cohorts of seedlings of the larger seeded Tripteris sinuata. Tripteris showed the highest number of seedlings (maximum count of 150 seedlings/1,000 viable seeds sown) and surviving plants of the three reseeded species, which included two small‐seeded species, Ruschia spinosa and Chaetobromus dregeanus. Throughout the study period significantly higher plant volumes of naturally seeded annuals and perennials were recorded in the gypsum and/or mulch treatments compared with the controls. Germination and emergence of reseeded and naturally seeded plants appears to be determined by the availability of cool season (autumn to spring ) soil moisture, whereas follow‐up rainfall during this time is important for plant survival. Mulching of bare areas in the Succulent Karoo has the potential to re‐create vegetated areas that will further capture and conserve water, soil, and nutrients. Gypsum also showed positive results but might not be a cost‐effective option because of transport costs to these remote arid areas.     

New data on the jaw apparatus of fossil cephalopods

A newly discovered fossil cephalopod jaw apparatus that may belong to Permian representatives of the Endocochlia is described. Permorhynchus dentatus n. gen. n. sp. is established on the basis of this apparatus. The asymmetry of jaws in the Ectocochlia may be connected with the double function of the ventral jaw apparatus, and the well-developed, relatively large frontal plate of the ventral jaw should be regarded as a feature common to all representatives of ectocochlian cephalopods evolved from early Palaeozoic stock. Distinct features seen in the jaw apparatus of Upper Permian endocochlians include the pronounced beak form of both jaws and the presence of oblong wings on the ventral mandible.     

Fungal abundance spike and the Permian-Triassic boundary in the Karoo Supergroup (South Africa)

The most severe mass extinction of marine species and terrestrial vertebrates and plants is associated with the Permian-Triassic boundary (∼251 Ma). The extinction interval is also marked by the disappearance of most Late Permian gymnosperm palynomorphs at a layer containing solely the abundant remains of fungi. This ‘fungal spike’ apparently represents widespread devastation of arboreous vegetation. Stratigraphic and palynological study of the Carlton Heights section in the southern Karoo Basin of South Africa revealed a 1-m-thick fungal spike zone that occurs simultaneously with the last appearance of typically Late Permian gymnosperm pollen. The plant extinction and fungal spike zone are found above the last occurrence of Late Permian mammal-like reptiles of the Dicynodont Zone at other Karoo sections. Using the fungal event as a time line in marine and non-marine sections allows placement of the marine extinctions and the extinction of terrestrial plants and reptiles within a brief crisis interval of less than about 40?000 years at the end of the Permian.     

Vulnerability of birds to contaminated water sources in the Karoo region of South Africa

The Karoo is a unique region in South Africa in terms of its ecological processes and endemic species. Large areas are needed to maintain viable populations of nomadic birds that follow erratic rainfall events and subsequent food and nesting resources, as well as ephemeral standing water. Whereas many species are adapted to arid conditions, our trait-based analysis found that an unusually large percentage (almost 45%) of 315 bird species in the semi-arid Karoo region rely on water to some degree. Indeed, some birds may have benefited from human activities to date, such as through the provision of water for livestock. However, this reliance on water makes birds vulnerable to changes in water quality stemming from various industrial developments. Given the large areas of the Karoo under consideration for concessions, the most noteworthy of these is hydraulic fracturing for shale gas, which results in a large quantity of waste water (‘produced water’) that contains a wide variety of chemicals, including petroleum byproducts. Given the negative impacts of secondary waste water on wildlife in other parts of the world where shale-gas exploration is being undertaken, careful attention must be given to preventing access to such produced water by an estimated 60 to 141 species of birds that make use of the water in the Karoo.     

Sciuridae (Rodentia,Mammalia) from the Late Miocene of Venta del Moro (Cabriel Basin,Spain): biostratigraphical,phylogenetic and palaeoecological inferences

Abstract

Fossil squirrels are relatively abundant in Aragonian and Ramblian assemblages, but very scarce in the Late Turolian and Ruscinian of the Iberian Peninsula. Until now, the locality of Venta del Moro has yielded over 4500 micromammal dental remains, but just nine of them are sciurids. In this assemblage, we have identified the taxa Pliopetaurista pliocaenica, Heteroxerus cf. mariatheresae and Atlantoxerus cf. margaritae. This is the first co-occurrence of the genera Pliopetaurista, Heteroxerus and Atlantoxerus in the same locality, being also one of the oldest records of P. pliocaenica, the oldest record of a form that could be related to A. margaritae, and the youngest record of the genus Heteroxerus. In addition, we hypothesize about the phylogeny of Pliopetaurista in the light of recent discoveries. Finally, the presence of the genera Heteroxerus and Atlantoxerus suggests an open environment and relatively dry environmental conditions, while Pliopetaurista is purportedly a flying squirrel, dweller of forested habitats and wet climates. Based on this, we infer an open environment of grassland type, bordered by forested habitats, agreeing with the conditions supposed for Venta del Moro according to its macromammal and floral assemblages.     

The gliding reptiles of the Upper Permian

The skeleton of a long-ribbed reptile from the Upper Permian Marl Slate of north-east England is described. The animal is assigned to the genus Weigeltisaurus , previously recorded from the Kupferschiefer of West Germany. Long-ribbed reptiles from the Upper Permian of Madagascar are also considered. Daedalosaurus Carroll, 1978 is a junior synonym of Coelurosauravus Piveteau, 1926. The European and Madagascan genera can be accommodated within a single family Coelurosauravidae, Infraclass Eosuchia. The skull is diapsid with an incomplete lower temporal arcade. Comparison with the modern Draco and the Upper Triassic kuehneosaurids supports the conclusion that the coelurosauravids were effective gliders.     

Effect of grazing on vegetation and soil of the heuweltjieveld in the Succulent Karoo,South Africa

We asked how historical and recent grazing intensity affect the patchy landscape of the heuweltjieveld in the semi-arid biodiversity hotspot Succulent Karoo. The study was carried out on a communal farmland 80 km south-west of Springbok, in Namaqualand. Heuweltjies are roughly circular earth mounds that are regularly distributed in this landscape. We sampled plant species and life-form composition, diversity measures, habitat and soil variables in 100 m² plots, placed in three visually distinguishable heuweltjie zones (centre, fringe, and matrix) and distributed across grazing camps with different recent and historic grazing intensities. Differences between heuweltjie zones were assessed with ANOVAs and multiple linear regressions. The effect of past and recent grazing intensity on soil and plant variables was analysed by Generalized Linear Models for each heuweltjie zone separately. The three zones constituted clearly distinguishable units in terms of vegetation and soil characteristics. Soil pH and cover of annual plants increased from matrix to centres, while total vegetation cover, species richness and perennial plant cover decreased in the same direction. Historic (pre-2000) grazing patterns had the strongest effects on fringes, showing the strongest soil and vegetation-related signs of overutilization with increased stocking density. Centres showed signs of overutilization irrespective of the stocking density. The much shorter exposure to recent grazing pattern (post-2000), which was nearly inverse to the historic grazing pattern, showed increase of vegetation cover (centres) and species richness (matrix) with recent grazing intensity. We interpret these effects as still visible responses of the lower grazing intensity in these camps during the historic period. No recovery under recent grazing was observed at any of the zones. We conclude that irrespective of their conducive growing conditions, once transformed to a disturbed state, heuweltjie centres recover slowly, whereas the less impacted soil and vegetation of fringes are more responsive than centres and matrix.