Permian bryophytes of Western Gondwanaland from the Paraná Basin in Brazil

Abstract: This study involves bryophyte fossils from the Teresina Formation (Permian–Guadalupian) collected in the Rio Preto Quarry in the state of Paraná, southern Brazil. Two new genera are proposed, with two new species: Capimirinus riopretensis sp. nov. and Yguajemanus yucapirus sp. nov. C. riopretensis sp. nov. included a lateral sporophyte that was organically attached to a gametophyte and is thus one of the oldest pleurocarpous fossils known today. This discovery provides clear evidence of the evolution of mosses during the Permian, because an acrocarpous moss from this period was also found in India. The preservation of such a fragile structure demonstrates that no prefossilization displacement took place and that the sediments accumulated in a low‐energy, fresh‐water environment, similar to a lake fed by rain, and isolated from the direct influence of the sea. Campimirinus riopretensis may be placed within the earlier pleurocarps, and Yguajemanus yucapirus sp. nov. may belong to the Bryalean families. However, we were not able to place neither of these species in any existing family today.     

The presence of Compsocerops (Brachyopoidea: Chigutisauridae) (Late Triassic) in southern Brazil with comments on chigutisaurid palaeobiogeography

Abstract: Chigutisauridae is the longest‐lived trematosaurian clade (from early Triassic to early Cretaceous). They were reported in Argentina, Australia, India and South Africa. This contribution reports a putative chigutisaurid specimen in the Carnian of southern Brazil (Santa Maria Formation, Paraná Basin). The material comprises two skull fragments, a mandibular fragment, a clavicular blade and a humerus. Ontogenetic features point to an early development stage of the specimen. The presence of a long, straight and pointed tabular horn, which runs parallel to the skull midline towards its tip, and a distinctive projection in the posterior border of the postparietal indicates a close relationship of the Brazilian chigutisaurid with the Indian Compsocerops cosgriffi. Three distinctive and combined characters suggest that the Brazilian chigutisaurid is a distinctive specimen: the presence of an alar process of the jugal in the ventral margin of the orbit; jugal does not extend well beyond the anterior margin of the orbit; and tabular does not contact the parietal. These characters could justify the erection of a new taxon; however, they might reflect its immature ontogenetic stage as well. Accordingly, we attribute this new specimen to Compsocerops sp. Argentinean and Indian occurrences are dated as Norian, so the presence of a Carnian chigutisaurid in southern Brazil indicates that western Gondwana chigutisaurids have first occupied the Paraná Basin and later migrated towards west (to Argentina) and east (India). However, the presence of ghost chigutisaurid taxa cannot be dismissed, because their long temporal range contrasts with their still short (in comparison with other temnospondyl groups) geographic distribution. Hence, they might have been more geographically widespread than their fossil record suggests.     

Late Carboniferous palynostratigraphy of the Itararé Subgroup, northeastern Paraná Basin, Brazil

A new biozonation is proposed for the Late Carboniferous strata of the northeastern Paraná Basin (São Paulo and Paraná States), based on well-preserved palynological assemblages recovered from outcrops and core samples retrieved from the Itararé Subgroup. Assemblages include forty-nine species of trilete spores and twenty-five pollen species. The oldest biozone, the Ahrensisporites cristatus Interval Zone (AcZ), in the basal portion of the Itararé Subgroup, includes eleven exclusive spore species, and is assigned a Pennsylvanian (late Bashkirian to Kasimovian) age. The overlying proposed biozone, the Crucisaccites monoletus Interval Zone (CmZ), dated late Pennsylvanian (Kasimovian to Gzhelian), ranges approximately from the top of the lower portion to the middle portion of the Itararé Subgroup, bears only one exclusive species of pollen. Both zones are characterized by the dominance of trilete spores and monosaccate pollen grains, including also few taeniate pollen grains. Furthermore, the basal Protohaploxypinus goraiensis Subzone of the Vittatina costabilis Interval Zone (VcZ) is broadly recognized for the first time in the upper portion of the Itararé Subgroup in the northeastern Paraná Basin. This subzone is attributed to the Early Permian (Early Cisuralian), and is characterized by the dominance of monosaccate pollen grains, a significant increase of taeniate grains (mainly Protohaploxypinus spp.), apart from the appearance of polyplicate pollen (Vittatina spp.). Reference sections and the main characteristics of the two new zones (AcZ and CmZ) are presented, along with a preliminary palynological correlation and discussion on their ages.     

The Late Permian herbivore Suminia and the early evolution of arboreality in terrestrial vertebrate ecosystems

Vertebrates have repeatedly filled and partitioned the terrestrial ecosystem, and have been able to occupy new, previously unexplored habitats throughout their history on land. The arboreal ecospace is particularly important in vertebrate evolution because it provides new food resources and protection from large ground-dwelling predators. We investigated the skeletal anatomy of the Late Permian (approx. 260 Ma) herbivorous synapsid Suminia getmanovi and performed a morphometric analysis of the phalangeal proportions of a great variety of extant and extinct terrestrial and arboreal tetrapods to discern locomotor function and habitat preference in fossil taxa, with special reference to Suminia. The postcranial anatomy of Suminia provides the earliest skeletal evidence for prehensile abilities and arboreality in vertebrates, as indicated by its elongate limbs, intrinsic phalangeal proportions, a divergent first digit and potentially prehensile tail. The morphometric analysis further suggests a differentiation between grasping and clinging morphotypes among arboreal vertebrates, the former displaying elongated proximal phalanges and the latter showing an elongation of the penultimate phalanges. The fossil assemblage that includes Suminia demonstrates that arboreality and resource partitioning occurred shortly after the initial establishment of the modern type of terrestrial vertebrate ecosystems, with a large number of primary consumers and few top predators.     

Deforestation Trends of Tropical Dry Forests in Central Brazil

Tropical dry forests are the most threatened forest type in the world yet a paucity of research about them stymies development of appropriate conservation actions. The Paranã River Basin has the most significant dry forest formations in the Cerrado biome of central Brazil and is threatened by intense land conversion to pastures and agriculture. We examined changes in Paranã River Basin deforestation rates and fragmentation across three time intervals that covered 31 yr using Landsat imagery. Our results indicated a 66.3 percent decrease in forest extent between 1977 and 2008, with an annual rate of forest cover change of 3.5 percent. Landscape metrics further indicated severe forest loss and fragmentation, resulting in an increase in the number of fragments and reduction in patch sizes. Forest fragments in flatlands have virtually disappeared and the only significant forest remnants are mostly found over limestone outcrops in the eastern part of the basin. If current patterns persist, we project that these forests will likely disappear within 25 yr. These patterns may be reversed with creation of protected areas and involvement of local people to preserve small fragments that can be managed for restoration.     

A Late carboniferous palynoflora from the Itararé Subgroup (Paraná Basin) in Campinas, São Paulo State, Brazil

Upper Paleozoic sedimentary sequences from several localities of the northeastern Paraná Basin were sampled for paleobotanical and palynological studies. Well-preserved terrestrial assemblages from samples of Campinas, São Paulo State, from an outcrop at the 97 km mark on the Bandeirantes Highway, located at the Municipality of Campinas, São Paulo State were recorded where fertile bryophytes and megaspores were previously also found. The Campinas palynoflora is composed of 23 species of bryophytic and pteridophytic spores, 16 species of pollen grains, and one algae (Botryococcus braunii Kützing). Among the sporomorphs, Reticulatisporites asperidictyus Playford and Helby is described for the first time from the Paraná Basin. Assemblages bear diagnostic species of the Ahrensisporites cristatus Interval Zone, such as Anapiculatisporites argentinensis Azcuy, Cristatisporites inordinatus (Menéndez and Azcuy) Playford, and Psomospora detecta Playford and Helby, of Late Carboniferous age (Pennsylvanian). Palynological content suggests a glacio-deltaic/lacustrine environment of deposition. This is supported by lithological data and by the absence of marine palynomorphs.     

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.     

Distribution of non‐native suckermouth armoured catfish Pterygoplichthys ambrosettii in the upper Paraná River basin,Brazil

This study describes the invasion of the upper Paraná River basin by Pterygoplichthys ambrosettii based on a literature review and field samples. Pterygoplichthys ambrosettii has been reported in 42 localities throughout the upper Paraná River basin, including the Tietê, Paranapanema, Paraná, Grande and Aguapeí rivers. The ascent of P. ambrosettii after the inundation of the Sete Quedas Falls on the Paraná River and the release of individuals by aquarium hobbyists were the primary drivers of this invasion.     

Fish‐based multimetric index for evaluating land use effects on large neotropical reservoirs

Because of their importance to human well‐being and ecosystem services, it is important to have cost‐effective methods for assessing reservoir ecological condition or health. To do so, we constructed a preliminary reservoir fish assemblage index (RFAI) to assess the effect of predominantly agricultural land use on two oligotrophic neotropical reservoirs that differed in disturbance level. We first determined the disturbance levels around each reservoir through use of an integrated disturbance index (IDI). The IDI was calculated from the combination of land use metrics at local (Local Disturbance Index ‐ LDI) and buffer scales (Buffer Disturbance Index ‐ BDI). Afterwards, we employed the IDI to assess the response of candidate metrics to environmental disturbance in the RFAI construction. The final RFAI included three metrics selected by range, responsiveness, and redundancy tests. Metrics were scored continuously, and the final RFAI was divided in four quality classes: very poor, poor, acceptable and good. The results showed a clear disturbance gradient in both reservoirs with the IDI values varying between 0.07 and 1.33. However, the IDI values were greater around Volta Grande reservoir because of the predominance of agriculture land use (at local and buffer scales). The RFAI scores in Nova Ponte reservoir were significantly higher than in Volta Grande. All the sites scored as very poor or poor were in Volta Grande and all the sites scored as good were in Nova Ponte. In addition, RFAI scores in the reservoir river arms did not differ from those in the reservoir main body, indicating that it was applicable to both habitat types. Finally, our RFAI scores indicated no significant seasonal difference in the two reservoirs; however, the rainy season produced a greater range and more low scores, particularly in Volta Grande. We related this result to the harmful effect of agriculture and the surface runoff of fertilizers, herbicides, and pesticides. Thus, the RFAI was effective in detecting agricultural impacts at the site scale, even in oligotrophic reservoirs.     

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.     

Chromosome characterization and biogeographic relations among three populations of the driftwood catfish Parauchenipterus galeatus (Linnaeus, 1766) (Siluriformes: Auchenipteridae) in Brazil

The present study employed basic and molecular cytogenetic methods to characterize three populations of Parauchenipterus galeatus from the basins of the Paraná and São Francisco Rivers, and a region of connection between the two basins. Although the diploid number was equal to 58 chromosomes, variations in karyotype formula were detected among the populations. B chromosomes were detected only in the population from the São Francisco River. Heterochromatin was located in the terminal position in almost all the chromosomes and in the pericentromeric position in some acrocentric chromosomes in the three populations. A single nucleolus organizer region was detected by silver nitrate and 18S rDNA‐fluorescent in situ hybridization in the short arm of one subtelocentric pair in the three populations, varying only in the chromosome pair bearing this site. The 5S rDNA sites were located in two submetacentric chromosome pairs in the three populations, varying only in the chromosome pairs bearing these sequences. Classic and molecular chromosome markers, along with the context of the natural history of the formation of hydrographic basins, ecological aspects, and the geographic isolation of populations between hydrographic basins and within the same basin, were important contributions to the discussion on possible biogeographic relations among the populations of Parauchenipterus galeatus. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 648–656.     

The jaw function and adaptive radiation of the dicynodont mammal-like reptiles of the Karoo basin of South Africa

The structure and functioning of the dicynodont jaw system are described. A pivoting action of the lower jaw around the palate posteromedian to the caniniform processes is as basic to the jaw movement as is the well-known double-convex jaw articulation. The sequence of origin of the characteristic features of the dicynodonts is analysed cladistically; this demonstrates the patterns of association of these characters into functional character-complexes. The structures of the palate and lower jaw, and their functional integration in feeding, are described and illustrated in standard format. Five different lineages of dicynodont can be identified: Eodicynodon; the robertoids (including Diictodon ); the dicynodontoids (including Dicynodon and the majority of the large dicynodont genera of the Permian and Triassic); the endothiodontoids (including Prodicynodon [='Chelydontops'] and Pristerodon ), and the emydopoids (including Cistecephalus, Myosaurus and Kingoria ). Eodicynodon or a similar form could have been ancestral to the other four lineages. The robertoids probably fed upon the stems and rhizomes of equisetaleans, while the varied dicynodontoids probably fed upon the varied glossopterid seed-ferns. The endothiodontoids, too, were probably herbivorous, but many, perhaps all, of the small emydopoids were burrowing and may have been omnivorous. The dicynodonts were probably ectothermal, and the dicynodontoids may have migrated to warmer latitudes in the winter. Only c. 20 genera of Karoo dicynodont are now recognized as valid, and it is suggested that this fauna is now almost completely known. Their distribution in the Karoo biozones is reviewed and correlated with environmental changes. The Permian ancestors of the Triassic dicynodonts, including Lystrosaurus , probably lived on higher, drier ground, and were therefore already adapted to the more fibrous food that spread into the basins as the climate became drier in the Triassic.     

Revision of Late Permian tetrapod tracks from the Dolomites (Trentino-Alto Adige,Italy)

The Val Gardena Formation of the Dolomites region in northern Italy preserves the most significant assemblage of Late Permian tetrapod footprints in the world. More than 120 years of collecting resulted in about 900 publicly accessible specimens from the study area. This huge amount of data is comprehensively revised in the light of recent advances in the study of Late Palaeozoic – Early Mesozoic tetrapod ichnofossils. According to our analyses, the Val Gardena Sandstone Formation includes tracks that can be assigned to cf. Batrachichnus isp. (temnospondyl amphibian), Capitosauroides isp. (amphibian), Dicynodontipus isp. (cynodont therapsid), Dolomitipes accordii n. igen. n. comb. (dicynodont therapsid), cf. Dromopus isp. (neodiapsid), Pachypes dolomiticus (pareiasaurian parareptile), Paradoxichnium problematicum (archosauromorph neodiapsid), Procolophonichnium tirolensis n. comb. (procolophonoid parareptile), cf. Protochirotherium isp. (archosauriform neodiapsid) and Rhynchosauroides pallinii (neodiapsid). The ichnoassociation is dominated by tracks of neodiapsid and parareptilian tetrapods, whereas synapsid and anamniote tracks are rather minor components. It includes 10 out of 12 tetrapod ichnogenera known from Lopingian deposits and thus it constitutes a reference for the Paradoxichnium biochron. It shows striking similarities with other low-latitude non-aeolian contemporaneous ichnoassociations of Europe and North Africa, differences may be linked to the palaeoenvironment. Moreover, it shows a clear Triassic affinity.

The new ichnogenus Dolomitipes was registered in Zoobank.org. urn:lsid:zoobank.org:act:5B4D871C-D16A-4E93-8211-CEE08019BA60     

Pennsylvanian palynofloras from the Itu rhythmites (Itararé Subgroup, Paraná Basin) in São Paulo State, Brazil

The rhythmites of Itu present the best exposures of glacial varvites known in the Paraná Basin, and constitute a classic geological monument related to the Late Palaeozoic Gondwanan glaciation. Palynological results in this paper are based on rhythmites from two quarries in Itu area, central-eastern State of São Paulo, Brazil, including correlate levels of the “Itu Varvite Park”, as well as based on samples from the borehole IT-IG-85 (at 171 and 228 m) drilled also in the Itu City. Well-preserved indigenous miospores and microphytoplankton elements have been recorded. The former comprise 15 spore species and 19 pollen species, of which three are recorded for the first time in the Brazilian part of the Paraná Basin (Verrucosisporites cf. V. andersonii, Convolutispora archangelskyi and Caheniasaccites verrucosus). Microalgae include prasinophyceans (Leiosphaeridia sp., Tasmanites sp., Deusilites tenuistriatus), chlorophyceans (Botryococcus braunii) and zignemataceans (Tetraporina). The palynological content confirms a late Pennsylvanian (Kasimovian/Gzhelian) age for these rhythmites, which are assigned to the Crucisaccites monoletus interval Zone, based on the record of the eponymous pollen species and Scheuringipollenites maximus. Although very scarce, prasinophycean algae suggest low salinity marine conditions.     

DNA barcoding of freshwater ichthyoplankton in the Neotropics as a tool for ecological monitoring

Quantifying and classifying ichthyoplankton is one of the most effective ways of monitoring the recruitment process in fishes. However, correctly identifying the fish based on morphological characters is extremely difficult, especially in the early stages of development. We examined ichthyoplankton from tributaries and reservoirs along the middle stretch of the Paranapanema River, one of the areas most impacted by hydroelectric projects in the Neotropics. Matching DNA sequences of the COI gene (628–648 bp) allowed us to identify 99.25% of 536 samples of eggs (293) and larvae (243) subjected to BOLD‐IDS similarity analysis with a species‐level threshold of 1.3%. The results revealed 37 species in 27 genera, 15 families and four orders, some 23.8% of documented fish species in the Paranapanema River. Molecular identification meant that we could include data from egg samples that accounted for about 30% of the species richness observed. The results in this study confirm the efficacy of DNA barcoding in identifying Neotropical ichthyoplankton and show how the data produced provide valuable information for preparing plans for conserving and managing inland waters.     

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 dicynodont (Anomodontia, Eotherapsida) from the Upper Permian of Tatarstan

A new dicynodont genus and species, Idelesaurus tataricus sp. nov. (Cryptodontidae), from the Semin Ovrag locality (Tatarstan, Tetyushinskii District; Upper Permian, Upper Tatarian Substage, Severodvinian Horizon) is described. The skull patterns of the East European and South African Cryptodontidae and Aulocephalodontidae are compared from the morphofunctional point of view.     

High‐latitude Hirnantian (latest Ordovician) brachiopods from the Eusebio Ayala Formation of Paraguay,Paraná Basin

Abstract: Examination of newly collected brachiopods from the Eusebio Ayala Formation of Paraguay reveals the occurrence of Arenorthis paranaensis sp. nov., Plectothyrella? itacurubiensis sp. nov., Hindella sp. and Eostropheodonta conradii (Harrington). Associated graptolites of the N. persculptus Zone indicate that the age of the fossiliferous beds is Hirnantian. The overall generic composition of the fauna is similar to that of the atypical Hirnantia Fauna of the Bani Province. The record in Paraguay of Arenorthis, hitherto only known from North Africa, together with species of Plectothyrella? and Eostropheodonta different from those recorded in the Kosov Province, emphasizing the affinities between the Paraguayan fauna and the low‐diversity African assemblages. Stratigraphic and faunal evidence indicates that biogeographical links between South America and Africa already existed by the end of the Ordovician when most of the intra‐cratonic basins of Gondwana were flooded during the postglacial sea level rise.     

Functional aspects of the postcranial anatomy of the Permian dicynodont Diictodon and their ecological implications

The postcranial skeleton of the Permian dicynodont Diictodon is described, major skeletal muscles restored and functional aspects of the skeleton analyzed. The humerus was caudolaterally oriented and articulated in a near sagittal plane. This reduced the transverse component of thrust during locomotion. Throughout the stride, the femur, which is relatively longer than the humerus, was held in a parasagittal position. Diictodon probably had a hindlimb gait similar to the high walk of crocodilians. Comparisons between several dicynodont genera, including Diictodon, reveal considerable variation in the limb orientations, articulations, posture and gait among these taxa. Expansion of the preacetabular iliac process and the overall iliac length with respect to its height among these dicynodonts may be correlated with the increase in the number of the sacral vertebrae. Diictodon does not show the cranial and forelimb modifications suitable for digging as seen in Cistecephalus, Kawingasaurus and extant burrowers such as a broad skull, rounded occiput for considerable neck musculature, a robust humerus and a prominent olecranon process. However, a cylindrical body, short limbs with well–developed limb elevators and depressors and a long wide manus with long, blunt claws probably helped in digging. In addition, the hindlimb with well–developed retractors and short, blunt claws participated in soil removal. The caudolaterally oriented humerus and a laterally flexible vertebral column facilitated movement in the narrow confines of the burrows. Forelimb articulation and morphology indicate that its method of digging was probably rotation–thrust, where humeral excursion and not forearm extension played a dominant role. The close associations of articulated paired skeletons of adult individuals suggest that Diictodon was gregarious. A large number of unconnected burrow–casts in a small area of about 500 m², indicates that although the animals lived close together, they did not share a single large communal structure.