HIGHER SYSTEMATICS OF SCORPIONS FROM THE CRATO FORMATION, LOWER CRETACEOUS OF BRAZIL

Abstract:  Several new specimens of Protoischnurus axelrodorum Carvalho and Lourenço and Araripescorpius ligabuei Campos from the Crato Formation, Brazil, are described. The preservation and recognition of new morphological features allows a re-diagnosis of both species and a modification of their familial placement. Protoischnurus axelrodorum is the oldest species belonging to the scorpionoid family Hemiscorpiidae Pocock (= Ischnuridae Simon; = Liochelidae Fet and Bechly) and the first Cretaceous record. It was originally placed in the extinct family Protoischnuridae Carvalho and Lourenço, which is here synonymized with Hemiscorpiidae. Araripescorpius ligabuei, now assigned to Chactidae Pocock, is the first chactoid recorded for the Cretaceous of Brazil. These findings confirm that the lineages of two modern families date back at least to the Early Cretaceous and, considering their current distribution, were probably present before the break-up of the supercontinent Gondwana. Palaeoecological inferences indicate the presence of tropical habitats in the vicinity of the Crato lake/lagoon.     

EARLY ORDOVICIAN (ARENIG) TRILOBITE PALAEOECOLOGY AND PALAEOBIOGEOGRAPHY OF THE SOUTH CHINA PLATE

Abstract:  Faunal composition of South Chinese Arenig benthic trilobite associations is investigated using the multivariate techniques of TWINSPAN, DCA and seriation. Eight fairly distinct benthic associations can be differentiated, organized primarily along a palaeobathymetrical gradient across the Yangtze Platform in southern Shaanxi and western Hubei and the Jiangnan Transitional Belt in northern Hunan: the Taihungshania , Neseuretus , Trinucleid and Asaphid- Pseudocalymene associations (shallow shelf), the Asaphid-Raphiophorid and Nileid-Asaphid associations (< 100 m outer shelf), the Nileid-Illaenid Association (deep outer shelf carbonates) and the Pseudopetigurus Association (deep outer shelf clastics). The highest levels of diversity are displayed by the Asaphid-Raphiophorid Association of western Hubei. Investigation of the biogeographical affinities of this fauna indicates that South China is biogeographically closest to the other Chinese geotectonic units, and displays strong faunal connections to other central and eastern Asian regions as well as to Tethyan and South American regions. Subdivision of the South Chinese trilobite fauna into different depth zones shows a statistically significant correlation between increasing water depth and increasing faunal cosmopolitanism, although taxa endemic to China or Gondwana also occur even in slope faunas. The results of this biogeographical analysis are consistent with a tropical or subtropical peri-Gondwanan association of South China with other Asian terranes.     

FEDEKURTZIA,A NEW CARBONIFEROUS FROND FROM GONDWANALAND AND ITS FRUCTIFICATION

Fedekurtzia gen. nov. is proposed for specimens referred to Archaeopteris argentina (Kurtz, 1921) from middle Carboniferous strata of the Paganzo Basin in Argentina. Similar materials from the same area were assigned to at least six different genera by later authors. A revision of specimens from various collections shows that they all belong to a single type of polymorph frond. A progymnospermous affinity is suggested by the small fructifications resembling spikes which are attached to main rachides, as well as by the helicoidal insertion and bilateral disposition of the pinnae on the main axis. Similar fronds and fructifications have been found in Australia. These are referred in part to the new genus which seems to have had a wide distribution over Gondwanaland. Paleoclimatic and palaeogeographic evidence is considered for the Carboniferous in the Gondwana Realm. Rather poor and primitive palaeofloristic associations are usually found.     

A first phylogenetic analysis of Giant Pill-Millipedes (Diplopoda: Sphaerotheriida), a new model Gondwanan taxon, with special emphasis on island gigantism

The phylogeny of the Giant Pill-Millipedes, order Sphaerotheriida, is investigated using a new morphological character matrix comprising 89 characters. The majority of these characters are employed for the first time in millipedes. All trees obtained agree on the monophyletic status of the Sphaerotheriida and several of its tribes, each restricted to a modern land mass. The species from Madagascar displaying island gigantism do not form a monophyletic group. The classic division of Giant Pill-Millipedes into two families, Sphaerotheriidae and Zephronidae, was not reflected in the analysis. The genus Procyliosoma is the sister-group to all other Sphaerotheriida, rendering the family Sphaerotheriidae paraphyletic. A new family-level classification of Giant Pill-Millipedes, based on the current phylogeny, is introduced. The new family Procyliosomatidae contains only the genus Procyliosoma , distributed in Australia and New Zealand. The family Zephronidae remains unchanged, while the family Sphaerotheriidae now incorporates only the African Giant Pill-Millipede genera. All genera from southern India and Madagascar form a monophyletic group and are placed in the new family Arthrosphaeridae. The Malagasy genus Sphaeromimus is more closely related to the Indian Arthrosphaera species than to other genera from Madagascar. A biogeographical analysis identifies the group as a Gondwana taxon (with a notable absence from South America). The current phylogeny of Giant Pill-Millipede families mirrors perfectly the suggested break-up of Gondwana fragments 160–90 Ma. No evidence for a dispersal event could be found, highlighting the importance of Giant Pill-Millipedes as a potential model taxon.     

Modern hydrophilid clades present and widespread in the Late Jurassic and Early Cretaceous (Coleoptera: Hydrophiloidea: Hydrophilidae)

We present a summary of the fossil evidence documenting the worldwide occurrence of the family Hydrophilidae (Insecta: Coleoptera: Polyphaga: Hydrophiloidea) in the Late Jurassic and Early Cretaceous. We present the first known fossils of the family from the Mesozoic, being c. 100 Myr older than the fossil record available until now. Two Late Jurassic fossils are documented: P rotochares brevipalpis gen. nov., sp. nov. from the Talbragar Fish Bed (New South Wales, Australia) and ‘Mesosperchus’ schultzi Ponomarenko, 1985 from Solnhofen (Bavaria, Germany). The occurrence of the Hydrophilidae in the Early Cretaceous is documented by six species, all of which may be already assigned to modern subfamilies/tribes: B aissalarva hydrobioides gen. nov., sp. nov. from the Baissa outcrops (Buryat Republic, Russia) and C retoxenus australis gen. nov., sp. nov. from Koonwarra outcrops (Victoria, Australia) are both assigned to the tribe Hydrobiusini (Hydrophilinae); A legorius yixianus gen. nov., sp. nov. and Alegorius sp. from the Yixian Formation (Liaoning, China) may represent the Acidocerinae or Enochrinae, H ydroyixia elongata gen. nov., sp. nov. and H . latissima sp. nov. from the same locality are assigned to the Acidocerinae. The phylogenetic position of Baissalarva hydrobioides is also tested by a phylogenetic analysis. The presence of extant clades (Hydrophilinae: Hydrobiusini, Acidocerinae) in the Early Cretaceous and the wide distribution of the Hydrobiusini in both Gondwana and Laurasia at the same time suggests that the principal extant clades of the Hydrophilidae are at least of Early–Middle Jurassic origin. © 2014 The Linnean Society of London     

The oldest Gondwanan fossil of Leiinae (Diptera,Mycetophilidae): Phylogenetic and evolutionary implications

A fossil Mycetophilidae from the Aptian Crato Formation—Cretomanota gondwanica gen. nov., sp. nov.—is described, which is the first mycetophilid from the Crato Formation and corresponds to the oldest known fossil leiine and only the second Gondwanan fossil mycetophilid described so far. Cretomanota gondwanica and both species of Alavamanota Blagoderov and Arillo were added as terminals to the data matrix of a general phylogenetic analysis of the Mycetophilidae, and both fit into the Leiinae. Alavamanota is monophyletic, sister to the clade composed by Cretomanota and the extant genus Manota Williston. The biology of the extant members of this fungivorous family corroborates the reconstruction of the Crato palaeoenvironment as including woodlands with humid habitats and microhabitats. The presence of a Cretaceous member of the tribe Manotini at low latitudes in South America reinforces the hypothesis that the clade with all manotines except Leiella Edwards corresponds to a Lower Cretaceous offshoot from a group in southern Gondwana expanding its distribution to more northern areas into the Gondwana and into Laurasia.     

Biogeography of Mesozoic leptosporangiate ferns related to extant ferns

The paleobiogeography of the extant leptosporangiate fern families present in the Mesozoic is reviewed based mainly on data from megafossils that possess enough characters to be assigned to a family. Times of first appearances are discussed for each family, followed by their distribution throughout the Mesozoic. Paleolatitudes and climatic conditions indicate that first appearances of many fern families occur about or above 30° and in conditions of higher moisture. Much of fern distribution in the past was likely due to long-distance dispersal.     

Molecular evidence suggests an ancient radiation for the fairy shrimp genus Streptocephalus (Branchiopoda: Anostraca)

Phylogenetic relationships among assumed Gondwanan aquatic inland invertebrate fauna are generally largely neglected, and biogeographical hypotheses for these organisms are generally inferred from historic (palaeogeographical) and contemporary distribution patterns. The distribution of the monogeneric thermophilic freshwater fairy shrimp family Streptocephalidae ( Streptocephalus ) provides a particularly useful framework to test the three contrasting biogeographical scenarios proposed for the evolution of this group: (1) the genus evolved in Laurasia and subsequently dispersed into Africa and North America; (2) the genus evolved and dispersed out of Africa and (3) the current distribution of the genus is the result of vicariance following the fragmentation of Gondwana. In the present study, the phylogenetic relationships of species in this genus are examined with the use of two mitochondrial genes (12S rRNA and COI mtDNA), while the phylogenetic relationships among the North American species and selected African taxa was investigated using the nuclear fragment (5.8S-ITS-1-18S). Phylogenetic results indicate that Streptocephalus probably evolved in Gondwana and that the current distribution patterns are a consequence of a combination of vicariance and limited dispersal. The implications for the evolution of continental freshwater crustaceans are discussed.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 313–327.     

The family prosthogonimidae (Trematoda) in Australia

The family Prosthogonimidae (Trematoda) in Australia. International Journal for Parasitology 3: 853–862. The family Prosthogonimidae has previously been known in Australia by three species of Prosthogonimus Lühe, 1899 (from birds) and by Coelomotrema Angel, 1970 (from marsupials). Two new genera are now proposed: Cylindrotrema cygni from Cygnus atratus (Latham) (caecum), and Mawsonotrema eudyptulae from Eudyptula minor (Forster) (liver). Prosthogonimus vitellatus Nicoll, 1914 is recorded from three new hosts, Gymnorhina tibicen leuconota (Latham), Phalacrocorax melanoleucos (Vieillot) and Chlidonias hybrida (Pallas), and the species is redescribed from 11 specimens from the first of these. The life history, not known for any Australian species, is discussed.

The only plump-bodied genera in the family are Coelomotrema, Cylindrotrema and Mawsonotrema, all of which are confined to Australia.

Mawsonotrema differs from all other members of the family in the position of the genital pore, which is away from the anterior end of the body, lying against the anterior border of the acetabulum.     

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.     

Phylogeny and historical biogeography of Gondwanan moss-bugs (Insecta: Hemiptera: Coleorrhyncha: Peloridiidae)

The moss bugs of the Peloridiidae, a small group of cryptic and mostly flightless insects, is the only living family in Coleorrhyncha (Insecta: Hemiptera). Today 37 species in 17 genera are known from eastern Australia, New Zealand, New Caledonia and Patagonia, and the peloridiids are thereby a group with a classical southern Gondwanan distribution. To explicitly test whether the present-day distribution of the Peloridiidae actually results from the sequential breakup of southern Gondwana, we provide the first total-evidence phylogenetic study based on morphological and molecular characters sampled from about 75% of recognized species representing 13 genera. The results largely confirm the established morphological phylogenetic context except that South American Peloridium hammoniorum constitutes the sister group to the remaining peloridiids. A timescale analysis indicates that the Peloridiidae began to diversify in the land mass that is today's Patagonia in the late Jurassic (153 Ma, 95% highest posterior density: 78–231 Ma), and that splitting into the three extant well-supported biogeographical clades (i.e. Australia, Patagonia and New Zealand/New Caledonia) is consistent with the sequential breakup of southern Gondwana in the late Cretaceous, indicating that the current transoceanic disjunct distributions of the Peloridiidae are best explained by a Gondwanan vicariance hypothesis.     

SYSTEMATIC REVISION OF PSEUDOBEACONIA BORDAS, 1944, AND MENDOCINICHTHYS WHITLEY, 1953 (ACTINOPTERYGII: ‘PERLEIDIFORMES’) FROM THE TRIASSIC OF ARGENTINA

Abstract: The ‘perleidiform’Mendocinichthys and Pseudobeaconia from the Potrerillos and Santa Clara Abajo formations (Upper Triassic; Argentina) are reviewed. Mendocinichthys has been known from a review of this species that is not based on the type material, but on referred specimens in the American Museum of Natural History. However, those specimens are found here to represent a new species of Pseudobeaconia, P. celestae sp. nov. Consequently, Mendocinichthys is restricted here to the type material and, within it, to the only specimen that clearly represents a distinct taxon and is thus designated lectotype. We further performed a cladistic analysis and propose a new family, Pseudobeaconiidae for these two South American Triassic genera, which is mainly characterized by the presence of an incomplete dorsal ridge of spine‐like scales between the skull and the dorsal fin, and scales with straight posterior border, an elevated central region and marginal concentric ridges of ganoine. The cladistic analysis further indicates the existence of some lineages endemic to certain areas of Gondwana and Europe. Pseudobeaconia celestae sp. nov. represents the first record of Pseudobeaconia in the Cacheuta sub‐basin of the Cuyana Basin. The genus was previously known from the Santa Clara sub‐basin of the Cuyana Basin only, and the new record confirms the previous hypothesis of correlation between the sedimentary infilling of these sub‐basins.     

Turtle remains from the Middle Jurassic Xintiangou Formation of Yunyang,Sichuan Basin,China

Three turtle shells from the Middle Jurassic Xintiangou Formation of Yunyang (Chongqing, China) are described and assigned to Xinjiangchelyidae (Testudines: Eucryptodira). This is the first report of turtle remains from the Xintiangou Formation, Sichuan Basin and represents the oldest known Xinjiangchelyidae. The assemblage includes two taxa, Protoxinjiangchelys sp. and Xinjiangchelyidae indet. This discovery extends the stratigraphical distribution of Xinjiangchelyidae and improves our knowledge about the early evolution of that family. It demonstrates that by the Middle Jurassic, at the time of deposition of the Xintiangou Formation, the group was already diversified in the Sichuan Basin.     

A Megaraptor-like theropod (Dinosauria: Tetanurae) in Australia: support for faunal exchange across eastern and western Gondwana in the Mid-Cretaceous

The fossil record of Australian dinosaurs in general, and theropods in particular, is extremely sparse. Here we describe an ulna from the Early Cretaceous Eumeralla Formation of Australia that shares unique autapomorphies with the South American theropod Megaraptor. We also present evidence for the spinosauroid affinities of Megaraptor. This ulna represents the first Australian non-avian theropod with unquestionable affinities to taxa from other Gondwanan landmasses, suggesting faunal interchange between eastern and western Gondwana during the Mid-Cretaceous. This evidence counters claims of Laurasian affinities for Early Cretaceous Australian dinosaur faunas, and for the existence of a geographical or climatic barrier isolating Australia from the other Gondwanan continents during this time. The temporal and geographical distribution of Megaraptor and the Eumeralla ulna is also inconsistent with traditional palaeogeographic models for the fragmentation of Gondwana, but compatible with several alternative models positing connections between South America and Antarctica in the Mid-Cretaceous.     

The oldest mammals from Antarctica,early Eocene of the La Meseta Formation,Seymour Island

New fossil mammals found at the base of Acantilados II Allomember of the La Meseta Formation, from the early Eocene (Ypresian) of Seymour Island, represent the oldest evidence of this group in Antarctica. Two specimens are here described; the first belongs to a talonid portion of a lower right molar assigned to the sparnotheriodontid litoptern Notiolofos sp. cf. N. arquinotiensis. Sparnotheriodontid were medium‐ to large‐sized ungulates, with a wide distribution in the Eocene of South America and Antarctica. The second specimen is an intermediate phalanx referred to an indeterminate Eutheria, probably a South American native ungulate. These Antarctic findings in sediments of 55.3 Ma query the minimum age needed for terrestrial mammals to spread from South America to Antarctica, which should have occurred before the final break‐up of Gondwana. This event involves the disappearance of the land bridge formed by the Weddellian Isthmus, which connected West Antarctica and southern South America from the Late Cretaceous until sometime in the earliest Palaeogene.     

FOSSIL WOODS IN INTERGLACIAL SEDIMENTS FROM THE CARBONIFEROUS HOYADA VERDE FORMATION, SAN JUAN PROVINCE, ARGENTINA

Abstract:  Fossil woods are described and assigned to Cuyoxylon multipunctatus gen. et sp. nov. The woods were deposited in shales from an interglacial period, and are the first to be described in detail from the Carboniferous glaciation of western Gondwana. They have the primitive characters of Early Carboniferous woods, such as mesarch protoxylem and numerous contiguous pits in an alternate arrangement. They are the oldest fossil woods to have been described from South America. The growth ring characteristics are consistent with a seasonal cool climate.     

A new Alokistocaridae Resser, 1939 (Trilobita) from the middle Cambrian of Spain

A new trilobite species, Schopfaspis? graciai, from the middle Cambrian of Spain is the first member of Alokistocaridae reported from west Gondwana. A cladistic analysis of this trilobite and other Gondwanan trilobites of possible alokistocarid affinities (Schopfaspis granulosa, Chelidonocephalus anatolicus, Derikaspis toluni, Kounamkites multiformis) suggests that this family can be divided into two subfamilies: Alokistocarinae and Altiocculinae. Schopfaspis? graciai nov. sp. and Schopfaspis granulosa are assigned to the subfamily Altiocculinae laying in a more basal position than Altiocculus species. The cladistic analysis also demonstrates a possible relationship of Chelidonocephalus anatolicus and Derikaspis toluni with Alokistocaridae. Enrolment is analysed in this new trilobite describing novelties in the ventral surface of the cephalon which allowed interlocking of the trunk and cephalon in a discoidal enrolment-type.     

Middle Ordovician Aporthophyla brachiopod fauna from the roof of the World,southern Tibet

A Darriwilian (late Middle Ordovician) brachiopod fauna from the Lower Formation of the Chiatsun Group at Jiacun, northern Nyalam, southern Tibet, consists of ten brachiopod species, forming a distinct Aporthophyla–Paralenorthis Association. Its taxonomic composition is typical of the Aporthophyla Fauna that occupied lower BA2 to upper BA3 benthic environments on sandy lime mud substrates. The occurrence of Paralenorthis in southern Tibet is confirmed for the first time, represented by P. costata sp. nov. Numerical analyses (PCA and CA) of 18 Darriwilian brachiopod faunas from ten palaeoplates or terranes indicate that: (1) the Aporthophyla Fauna was confined to a specific latitudinal belt although it had a wide lateral distribution from the large palaeocontinents of Gondwana to Laurentia; (2) the Saucrorthis Fauna, a typical late Middle Ordovician regional fauna, is limited to a much smaller area, marginal to the Gondwana supercontinent; (3) the strong provincialism persistent in the late Middle Ordovician contributed to increased gamma biodiversity during the Great Ordovician Biodiversification Event.     

Grasses through space and time: An overview of the biogeographical and macroevolutionary history of Poaceae

Grasses are widespread on every continent and are found in all terrestrial biomes. The dominance and spread of grasses and grassland ecosystems have led to significant changes in Earth′s climate, geochemistry, and biodiversity. The abundance of DNA sequence data, particularly chloroplast sequences, and advances in placing grass fossils within the family allows for a reappraisal of the family′s origins, timing, and geographic spread and the factors that have promoted diversification. We reconstructed a time-calibrated grass phylogeny and inferred ancestral areas using chloroplast DNA sequences from nearly 90% of extant grass genera. With a few notable exceptions, the phylogeny is well resolved to the subtribal level. The family began to diversify in the Early–Late Cretaceous (crown age of 98.54 Ma) on West Gondwana before the complete split between Africa and South America. Vicariance from the splitting of Gondwana may be responsible for the initial divergence in the family. However, Africa clearly served as the center of origin for much of the early diversification of the family. With this phylogenetic, temporal, and spatial framework, we review the evolution and biogeography of the family with the aim to facilitate the testing of biogeographical hypotheses about its origins, evolutionary tempo, and diversification. The current classification of the family is discussed with an extensive review of the extant diversity and distribution of species, molecular and morphological evidence supporting the current classification scheme, and the evidence informing our understanding of the biogeographical history of the family.     

Compartments in the abdomen of Drosophila and the role of the engrailed locus

A clonal analysis has shown that the dorsal surface of the first abdominal segment of Drosophila melanogaster is subdivided into anterior and posterior compartments. Cells of the posterior compartment grow up to but not beyond the anterior-posterior compartment border within the first abdominal segment and the intersegmental border that defines the boundary between the first and second abdominal segments. Growing within these boundaries, a narrow band of tissue clonally isolated from the adjoining tissue is formed. When these posterior cells are deficient for the engrailed locus, however, neither the compartment nor the segment border is maintained. The implications, that compartmentalization is essential for segmentation, and that all insect segments are subdivided by anterior and posterior compartments, are discussed.