Phylogeny of Early Cretaceous spatangoids (Echinodermata: Echinoidea) and taxonomic implications

A phylogenetic analysis of 36 species provides a test for the taxonomy and the history of Early Cretaceous spatangoids. Most taxonomic units from genera to suborders are consistent with the proposed phylogenetic framework. We retain Hemiasterina, Micrasterina, Hemiasteridae, Schizasteridae, Hemiaster , Heteraster , Mecaster , and Periaster as original monophyletic groups. However, all of these clades originate without the classical apomorphies normally ascribed to them. We suggest a revision of their diagnoses and of the generic attributions of basal species. Some ill-defined, 'primitive', and paraphyletic taxa are recognised: Toxaster , Epiaster , Palhemiaster , and Toxasteridae. Even if they do not have phylogenetic meaning, they are retained here, pending a more complete revision.     

Aspects of the fossil record and phylogeny of the familyRutaceae (Zanthoxyleae,Toddalioideae)

A comparative analysis of the seed morphology and anatomy of fossil and extantRutaceae (mainlyZanthoxyleae andToddalioideae) is presented. This allows to place the most important fossil taxa in a time-table and on paleogeographical maps. A phylogenetic scheme demonstrates the postulated historical relationships ofEvodia, Zanthoxylum, Fagara, Rutaspermum, Acronychia, Toddalia, Fagaropsis, andPhellodendron.     

Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography

We present a comprehensively sampled three‐gene phylogeny of the monophyletic Forcipulatacea, one of three major lineages within the crown‐group Asteroidea. We present substantially more Southern Hemisphere and deep‐sea taxa than were sampled in previous molecular studies of this group. Morphologically distinct groups, such as the Brisingida and the Zoroasteridae, are upheld as monophyletic. Brisingida is supported as the derived sister group to the Asteriidae (restricted), rather than as a basal taxon. The Asteriidae is paraphyletic, and is broken up into the Stichasteridae and four primary asteriid clades: (1) a highly diverse boreal clade, containing members from the Arctic and sub‐Arctic in the Northern Hemisphere; (2) the genus Sclerasterias; (3) and (4) two sister clades that contain asteriids from the Antarctic and pantropical regions. The Stichasteridae, which was regarded as a synonym of the Asteriidae, is resurrected by our results, and represents the most diverse Southern Hemisphere forcipulatacean clade (although two deep‐sea stichasterid genera occur in the Northern Hemisphere). The Labidiasteridae is artificial, and should be synonymized into the Heliasteridae. The Pedicellasteridae is paraphyletic, with three separate clades containing pedicellasterid taxa emerging among the basal Forcipulatacea. Fossils and timing estimates from species‐level phylogeographic studies are consistent with prior phylogenetic hypotheses for the Forcipulatacea, suggesting diversification of basal taxa in the early Mesozoic, with some evidence for more widely distributed ranges from Cretacous taxa. Our analysis suggests a hypothesis of an older fauna present in the Antarctic during the Eocene, which was succeeded by a modern Antarctic fauna that is represented by the recently derived Antarctic Asteriidae and other forcipulatacean lineages. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 646–660.     

Seven Hox gene sequences from the asterinid starfish Patiriella exigua (Echinodermata: Asteroidea)

Partial homeobox gene fragments were amplified from Patiriella exigua genomic DNA by PCR using degenerate primers. The primers spanned a region of 82 nucleotides within the homeobox, encompassing amino acids 21–47 of the homeodomain. The deduced amino acid sequences of the seven Hox gene sequences plus one Xlox-type gene sequence from P. exigua are presented and compared with similar sequences from other organisms. This work is a preliminary step in the analysis of the evolution of development in the Patiriella species group and the roles of Hox gene expression therein.     

Mass mortality of an asteriid starfish (Forcipulatida,Asteroidea, Echinodermata) from the late Maastrichtian (Late Cretaceous) of Morocco

Abstract: A mass occurrence, numbering tens of thousands of individuals in a single lens, belonging to a single species of asteroid, is described from the late Maastrichtian (Late Cetaceous) of Morocco. The lens of partially silicified asteroidal limestone is made up largely of fully articulated specimens of similar size and probably represents the mass mortality of a single recruitment. By comparison with mass strandings of the present‐day species Asterias rubens (Linnaeus), it can be inferred that a feeding swarm of individuals was swept into a submarine channel by either a storm or an exceptionally strong tidal current, and permanently buried. The genus and species are herein described as Cretasterias reticulatus gen. et sp. nov. The exceptional preservation of the material enables the identification of wreath organs (clusters of crossed pedicellariae set in a dermal pad around spines) for the first time in the fossil record. Comparison between extant Asteriidae, putative fossil asteriids and C. reticulatus provides ambiguous evidence of its affinities; it appears to display a combination of plesiomorphic and derived characters. It is shown that all Mesozoic forcipulatid asteroids described so far share a very simple arm construction (single row of adradial ossicles) unknown in adult extant Forcipulatida.     

The fossil pollen record of Araceae

The fossil record of Araceae pollen beginning in the late Early Cretaceous and peaking in the Paleocene/Eocene is very sparse up to now, consisting of three highly distinctive types: zona-aperturate pollen of the Monstera or Gonatopus type (very similar to Proxapertites operculatus), an ulcerate-spiny type typical for Limnobiophyllum, and a polyplicate, omniaperturate pollen type (an ephedroid pollen with non-gnetalean affinities) which was recently reported from the late Early Cretaceous (Mayoa portugallica). An extensive literature search has shown that some distinctive Ephedripites forms (the Paleogene Ephedripites vanegensis, and the Late Cretaceous Ephedripites elsikii) are very similar to pollen of Spathiphyllum and both species are here transferred from Ephedripites to Spathiphyllum (as comb. nov.). We also add new fossil findings to the Araceae record. The new findings include a zona-aperturate, microperforate to microreticulate pollen type from the Palaeocene of Colombia, highly similar to extant Gonatopus or Zamioculcas or Monstera pollen (Araceae) and to fossil Proxapertites operculatus, which is currently seen as a fossil equivalent; and, an ulcerate, spiny pollen from the Eocene of Stolzenbach, Germany, extending the range of Limnobiophyllum (Pandaniidites), which is thought to be an extinct member of extant Araceae. The three pollen types add considerably to the reliable fossil record of the family that now contains more than 20 records of these three pollen types: with the zona-aperturate type recorded from the tropical or subtropical regions of Northern and Southern America, Central Africa, Southern and Central Europe, from the Indian subcontinent and the Malayan Archipelago; the ulcerate type occurring in North America and Europe; and the polyplicate type mainly occurring in South America and South-West Europe. Now we have good evidence that some of the aroid subfamilies were already in existence in the Cretaceous, increasing in diversity and worldwide distribution in the Paleogene. Dedicated to Prof. Dr. Stefan Vogel on the occasion of his 80th birthday.     

Glandulocalyx upatoiensis, a fossil flower of Ericales (Actinidiaceae/Clethraceae) from the Late Cretaceous (Santonian) of Georgia, USA

Background and Aims

Ericales are a major group of extant asterid angiosperms that are well represented in the Late Cretaceous fossil record, mainly by flowers, fruits and seeds. Exceptionally well preserved fossil flowers, here described as Glandulocalyx upatoiensis gen. & sp. nov., from the Santonian of Georgia, USA, yield new detailed evidence of floral structure in one of these early members of Ericales and provide a secure basis for comparison with extant taxa.

Methods

The floral structure of several fossil specimens was studied by scanning electron microscopy (SEM), light microscopy of microtome thin sections and synchrotron-radiation X-ray tomographic microscopy (SRXTM). For direct comparisons with flowers of extant Ericales, selected floral features of Actinidiaceae and Clethraceae were studied with SEM.

Key Results

Flowers of G. upatoiensis have five sepals with quincuncial aestivation, five free petals with quincuncial aestivation, 20–28 stamens arranged in a single series, extrorse anther orientation in the bud, ventral anther attachment and a tricarpellate, syncarpous ovary with three free styles and numerous small ovules on axile, protruding-diffuse and pendant placentae. The calyx is characterized by a conspicuous indumentum of large, densely arranged, multicellular and possibly glandular trichomes.

Conclusions

Comparison with extant taxa provides clear evidence for a relationship with core Ericales comprised of the extant families Actinidiaceae, Roridulaceae, Sarraceniaceae, Clethraceae, Cyrillaceae and Ericaceae. Within this group, the most marked similarities are with extant Actinidiaceae and, to a lesser degree, with Clethraceae. More detailed analyses of the relationships of Glandulocalyx and other Ericales from the Late Cretaceous will require an improved understanding of the morphological features that diagnose particular extant groups defined on the basis of molecular data.     

Molecular phylogeny of the Valvatacea (Asteroidea: Echinodermata)

The Valvatacea is one the most ecologically important, taxonomically diverse, and widespread groups of post‐Palaeozoic (i.e. modern) Asteroidea. Classification within the group has been historically problematic. We present a comprehensively sampled, three‐gene (12S, 16S, early‐stage histone H3) molecular phylogenetic analysis of the Valvatacea. We include five of the six families within the Paxillosida, the monotypic Notomyotida, and 13 of the 16 families of the living Valvatida. The Solasteridae is removed from the Velatida (Spinulosacea) and joins the Ganeriidae and the Leilasteridae as members of the clade containing the Asterinidae. The Poraniidae is supported as the sister group to the large cluster of Valvatacea. Asteropseids and poraniids are phylogenetically distant, contrary to morphological evidence. Several goniasterid‐like ophidiasterids, such as Fromia and Neoferdina are supported as derived goniasterids rather than as Ophidiasteridae. The Benthopectinidae (Notomyotida) are supported as members of the Paxillosida as are two members of the Pseudarchasterinae that have traditionally been considered members of the Goniasteridae. Our data suggest that Antarctic valvataceans may be derived from sister taxa in adjacent regions. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 769–788.     

Coulonia parva n. sp., a new species of Astropectinidae (Asteroidea; Echinodermata) from the Santonian (Upper Cretaceous) of Sierra de Montsech (southern Pyrenees, Spain)

A new astropectinid sea star,Coulonia parva n. sp., is described from the Santonian of the Sierra de Montsech, northern Spain. The single specimen shows part of a rather weathered ventral side. It is characterized by comparatively long arms (R/r approx. 3/1), with one row of actinals reaching far into the arms; inferomarginals on the interbrachial arc have deep fasciolar grooves that are less developed on the arms; there are no larger spines on adambulacrals and probably also on inferomarginals. Based on presence of deep marginal fascioles,C. parva n. sp. is inferred to have shared the capability for self-burial with the relatedC. platyspina Hess &Blake.      

Hettangian Asteriidae (Echinodermata: Asteroidea) from southern Germany: taxonomy,phylogeny and life habits

Germanasterias amplipapularia andHystrixasterias hettangiurnus are new genera and species of Asteriidae (Asteroidea: Forcipulatida) described from the Hettangian (lowest Jurassic) of southern Germany. They are among the oldest known relatively complete asteroids assignable to surviving families, yet they are remarkably similar to certain living species. Modern asteriids are efficient predators of sessile and slow-moving stoutly-armored mullusks, however much asteriid morphology is suggested to have originated with suspension-feeding habits. The lineage leading to the modern Forcipulatacea might have separated from that of the remainder of the modern asteroids as early as the Carboniferous.     

Preface: The phylogeny, fossil record and ecological diversity of ostracod crustaceans

After more than two centuries of research, more than 65,000 living and fossil ostracod species have been described and studied, yet much remains to be learned about this ancient, widespread and diverse group of bivalved arthropods. Their higher classification and phylogeny are subjects of vigorous debate, as is their position in the broader picture of crustacean phylogeny. At the same time, major advances in our understanding of ostracod lineages and their relationships are resulting from the application of innovative approaches and techniques. This preface provides a contextual overview of the 15 contributions to this volume, which resulted from the 14th International Symposium on Ostracoda (ISO2001) held in 2001at Shizuoka, Japan. As such it provides a cross-section of topics at the forefront of research on the evolution and diversity of Ostracoda, and indicates directions for future work.     

Egarapithecus narcisoi, a new genus of Pliopithecidae (primates, catarrhini) from the Late Miocene of Spain

Pliopithecid remains from the Spanish locality of Torrent de Febulines (Late Vallesian, MN 10), consisting of right and left mandibular fragments with partial tooth rows and an isolated P(3) probably belonging to the same individual, are described and assigned to Egarapithecus narcisoi gen. et sp. nov. (Pliopithecidae, Crouzeliinae). This is a highly derived species dated at around 9 Ma (Ma = 10(6) years), representing the latest appearance of the family in the European continent. Morphologically it is the most distant member from the inferred primitive pliopithecid morphotype, displaying many autapomorphies that notably accentuate those of the remaining Crouzeliinae. A cladistic analysis based on lower cheek teeth, performed in order to tentatively assess the phylogenetic relationships of Egarapithecus within the Crouzeliinae, indicates that several equally parsimonious cladograms are possible in the light of current evidence. This is due to uncertainties regarding the position of Plesiopliopithecus and Crouzelia (here considered distinct genera), as a result of missing characters and the significant degree of homoplasy apparently involved in crouzeliine dental evolution. Whether Egarapithecus is more closely related to Crouzelia or to Anapithecus (the latter hypothesis tentatively favored here) cannot be definitively resolved with the currently available material and deserves further investigation. It is clear, however, that Egarapithecus is one of the more derived and specialized members of the Pliopithecidae.     

Skeletal anatomy of the Late Cretaceous shark,Squalicorax (Neoselachii: Anacoracidae)

The paleobiology of the Cretaceous neoselachian shark,Squalicorax, has largely been based on isolated teeth. We examined partial and nearly complete skeletons of three species ofSqualicorax, S. falcatus (Aoassiz),S. kaupi (Agassiz), andS. pristodontus (Agassiz), that were collected from the U.S.A. These specimens suggest that the total body length (TL) ofS. falcatus typically measured 1.8–2.0 m, and probably did not exceed 3 m. Moderatesized individuals ofS. kaupi andS. pristodontus perhaps measured about 3 m TL. AlthoughS. pristodontus was the largest form among the three species examined, this taxon possessed a set of large jaws (with large but fewer teeth) relative to its body size compared toS. falcatus orS. kaupi. This suggests that tooth size is not an accurate indicator of the TL if one compares oneSqualicorax species to another. Neurocranial features suggest that the vision ofSqualicorax was not as acute as that of a contemporaneous macrophagous lamniform shark,Cretoxyrhina mantelli (Agassiz) , but olfaction ofSqualicorax may have been better thanC. mantelli. The morphology of placoid scales suggests thatSqualicorax was capable of fast swimming. New skeletal data support the view that the feeding dynamics ofSqualicorax was similar to the modern tiger shark (Galeocerdo Müller & Henle). The present data do not allow for exact ordinal placement, but, contrary to some previous interpretations,Squalicorax can be excluded from the Hexanchiformes and Orectolobiformes. The taxon should more appropriately be placed within the Lamniformes or Carcharhiniformes.      

The fossil record and limb disparity of enantiornithines, the dominant flying birds of the Cretaceous

Morphometric and stratigraphic analyses that encompass the known fossil record of enantiornithine birds (Enantiornithes) are presented. These predominantly flighted taxa were the dominant birds of the second half of the Mesozoic; the enantiornithine lineage is known to have lasted for at least 60 million years (Ma), up until the end of the Cretaceous. Analyses of fossil record dynamics show that enantiornithine 'collectorship' since the 1980s approaches an exponential distribution, indicating that an asymptote in proportion of specimens has yet to be achieved. Data demonstrate that the fossil record of enantiornithines is complete enough for the extraction of biological patterns. Comparison of the available fossil specimens with a large data set of modern bird (Neornithes) limb proportions also illustrates that the known forelimb proportions of enantiornithines fall within the range of extant taxa; thus these birds likely encompassed the range of flight styles of extant birds. In contrast, most enantiornithines had hindlimb proportions that differ from any extant taxa. To explore this, ternary diagrams are used to graph enantiornithine limb variation and to identify some morphological oddities ( Otogornis , Gobipteryx ); taxa not directly comparable to modern birds. These exceptions are interesting – although anatomically uniform, and similar to extant avians in their wing proportions, some fossil enantiornithines likely had flight styles not seen among their living counterparts.     

Intergeneric relationships ofLolium,Festuca, andVulpia (Poaceae) and their phylogeny

Morphological and seed protein analyses of 26 species of the generaLolium, Festuca andVulpia confirmed their close systematic affinities. Six inflorescence characters readily differentiatedFestuca fromLolium. Protein similarities betweenFestuca of sect.Bovinae and cross-pollinated species ofLolium, coupled with cytogenetic and crossability data, substantiate that they should be united into one genus.Vulpia had phenetic similarities with sect.Scariosae, Montanae andOvinae ofFestuca. Lolium, Festuca, andVulpia are most likely derived from a common ancestral form which was close toFestuca pratensis andLolium perenne.     

A new fossil bird from the Early Cretaceous of Gansu Province,northwestern China

We report on the discovery of an Early Cretaceous bird from northwestern Gansu Province, in northwestern China. Represented by a nearly complete left wing and shoulder girdle the size of a rock dove, the new bird was quarried from laminated yellowish mudstones of the Xiagou Formation (Xinminpu Group) near Changma, in the Jiuquan area. These deposits have previously yielded the only known specimen of Gansus yumenensis, a basal ornithuromorph represented by the distal half of a hind limb with long and slender digits. Several derived characters of the new occurrence supports its allocation within Enantiornithes: (1) a convex lateral margin of the coracoid, (2) a minor metacarpal that projects distally more than the major metacarpal and (3) a proximal phalanx of the major digit longer than the intermediate (second) phalanx. The general proportions of the wing suggest it was a flier comparable to most other known enantiornithine birds. Although, direct comparisons between the new fossil and Gansus are not possible, phylogenetic based inferences supports their placement into two different clades. While the new fossil falls definitively within the enantiornithines, G. Yumenensis falls within the ornithuromorphs. The new occurrence thus adds to the taxonomic diversity of Early Cretaceous birds from Gansu Province in particular and central Asia in general.     

New crested specimens of the Late Cretaceous pterosaurNyctosaurus

Two recently collected specimens of the Late Cretaceous pterosaurNyctosaurus differ from all previously known specimens in the possession of a large branching cranial crest. The crest extends upward and backward from the posterior skull roof and is nearly three times the length of the skull proper. Despite the large crest, the specimens do not differ significantly in morphology from previously known specimens ofNyctosaurus, and do not seem to represent a new species ofNyctosaurus. The specimens suggest that the cranial crest was developed late in ontogeny, which is consistent with the interpretation of pterosaur cranial crests as intraspecific display structures.      

Formation binning: a new method for increased temporal resolution in regional studies,applied to the Late Cretaceous dinosaur fossil record of North America

The advent of palaeontological occurrence databases has allowed for detailed reconstruction and analyses of species richness through deep time. While a substantial literature has evolved ensuring that taxa are fairly counted within and between different time periods, how time itself is divided has received less attention. Stage-level or equal-interval age bins have frequently been used for regional and global studies in vertebrate palaeontology. However, when assessing diversity at a regional scale, these resolutions can prove inappropriate with the available data. Herein, we propose a new method of binning geological time for regional studies that intrinsically incorporates the chronostratigraphic heterogeneity of different rock formations to generate unique stratigraphic bins. We use this method to investigate the diversity dynamics of dinosaurs from the Late Cretaceous of the Western Interior of North America prior to the Cretaceous–Palaeogene mass extinction. Increased resolution through formation binning pinpoints the Maastrichtian diversity decline to between 68 and 66 Ma, coinciding with the retreat of the Western Interior Seaway. Diversity curves are shown to exhibit volatile patterns using different binning methods, supporting claims that heterogeneous biases in this time-frame affect the pre-extinction palaeobiological record. We also show that the apparent high endemicity of dinosaurs in the Campanian is a result of non-contemporaneous geological units within large time bins. This study helps to illustrate the utility of high-resolution, regional studies to supplement our understanding of factors governing global diversity in deep time and ultimately how geology is inherently tied to our understanding of past changes in species richness.