Phylogeny of Schisandraceae based on morphological data: evidence from modern plants and the fossil record

Schisandraceae are traditionally subdivided in two genera, Schisandra and Kadsura, based on differences in the organisation of the floral receptacle, the carpels, and the presence or absence of a ``pseudostigma'. Recently, phylogenetic analyses utilizing ITS sequence data and morphological data resulted in incongruent tree topologies, with the morphological trees suggesting monophyly of the two genera, whereas ITS trees did not resolve Schisandra and Kadsura as monophyletic clades. In the present paper we study seed morphology and leaf epidermal features of 22 species of Schisandraceae in order to provide additional data for a morphological data matrix. Seed morphological characters are highly homoplastic and do not yield further evidence for monophyly of the two genera. Instead, a number of characters appear to support sister group relationships between taxa within the genera, such as, for instance, for K. coccinea and K. scandens, both of which have large seeds along with a multi-layered mesotesta. Considering leaf epidermal characteristics, species of Kadsura were found to be consistently amphistomatic, whereas species of Schisandra are always hypostomatic. Phylogenetic analysis using the extended data matrix resulted in weakly supported Kadsura and Schisandra clades with five and four synapomorphies indicating monophyly of Kadsura and Schisandra, respectively. Fossils ascribed to Schisandraceae date back to the Late Cretaceous. These are tri-and hexacolpate pollen types displaying a combination of features found in modern Schisandraceae and partly also in Illiciaceae. Leaf remains from this period are poorly preserved and difficult to ascribe to Schisandraceae because of the lack of synapomorphies for the family. In the Early Cainozoic, leaf and seed remains from North America and Europe unambiguously belong to the family. Seeds from the Eocene of North America show some similarities to the modern Schisandra glabra from North America, while fossils from Europe show more similarities to modern Asian species.     

The mosasaur fossil record through the lens of fossil completeness

The quality of the fossil record affects our understanding of macroevolutionary patterns. Palaeodiversity is filtered through geological and human processes; efforts to correct for these biases are part of a debate concerning the role of sampling proxies and standardization in biodiversity models. We analyse the fossil record of mosasaurs in terms of fossil completeness as a measure of fossil quality, using three novel, correlating metrics of fossil completeness and 4083 specimens. A new qualitative measure of character completeness (QCM) correlates with the phylogenetic character completeness metric. Mean completeness by species decreases with specimen count; average completeness by substage varies significantly. Mean specimen completeness is higher for species‐named fossils than those identified to genus and family. We consider the effect of tooth‐only specimens. Importantly, we find that completeness of species does not correlate with completeness of specimens. Completeness varies by palaeogeography: North American specimens show higher completeness than those from Eurasia and Gondwana. These metrics can be used to identify exceptional preservation; specimen completeness varies significantly by both formation and lithology. The Belgian Ciply Formation displays the highest completeness; clay lithologies show higher completeness values. Neither species diversity nor sea level correlates significantly with fossil completeness. A generalized least squares (GLS) analysis using multiple variables agrees with this result, but reveals two variables with significant predictive value for modelling averaged diversity: sea level, and mosasaur and plesiosaur‐bearing formations (the latter is redundant with diversity). Mosasaur completeness is not driven by sea level, nor does completeness limit the mosasaur diversity signal.     

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.     

First record of a pterosaur landing trackway

The terrestrial progression of pterosaurs, the flying reptiles of the Mesozoic Era, has been debated for over two centuries. The recent discovery of quadrupedal pterodactyloid pterosaur tracks from Late Jurassic sediments near Crayssac, France, shows that the hindlimbs moved parasagittally, as in mammals, birds and other dinosaurs, and the hypertrophied forelimbs could make tracks both close to the body wall and far outside it. Their manus tracks are unique in form, position and kinematics, which would be expected because the forelimbs were used for flight. Here, we report the first record of a pterosaur landing track, which differs substantially from typical walking trackways. The individual landed on both hind feet in parallel fashion, dragged its toes slightly as it left the track, landed again almost immediately and placed the hindfeet parallel again, then placed its forelimbs on the ground, took another short step with both hindlimbs and adjusted its forelimbs, and then began to walk off normally. The trackway shows that pterosaurs stalled to land, a reflection of their highly developed capacity for flight control and manoeuverability.     

Body size of insular carnivores: evidence from the fossil record

Aim Our goals here are to: (1) assess the generality of one aspect of the island rule – the progressive trend towards decrease in size in larger species – for fossil carnivores on islands; (2) offer causal explanations for this pattern and deviations from it – as far as fossil carnivores are concerned; and (3) estimate the speed of this trend. Location Oceanic and oceanic‐like islands world‐wide. Methods Body size estimates of fossil insular carnivores and of their phylogenetically closest mainland relative were obtained from our own data and the published literature. Our dataset consisted of 18 species from nine islands world‐wide. These data were used to test whether the body size of fossil insular carnivores varies as a function of body size of the mainland species in combination with characteristics of the island ecosystem. Results Dwarfism was observed in two canid species. Moderate decrease in body mass was observed in one hyena species. Gigantism was observed in one otter species. Moderate body mass increase was observed in two otter species, one galictine mustelid and perhaps one canid. Negligible or no change in body mass at all was observed in five otter species, three galictine mustelids and one genet. Size changes in teeth do not lag behind in comparison to skeletal elements in the dwarfed canids. The evolutionary speed of dwarfism in a canid lineage is low. Main conclusions Size change in fossil terrestrial insular carnivores was constrained by certain ecological conditions, especially the availability of prey of appropriate body size. When such alternative prey was not available, the carnivores retained their mainland size. The impact of competitive carnivores seems negligible. The case of (semi‐)aquatic carnivores is much less clear. The species that maintained their ancestral body mass may have changed their diet, as is evidenced by their dentition. Among the otters, one case of significant size increase was observed, perhaps best explained as being due to it entering the niche of an obligate aquatic otter. Dwarfism was not observed in otters. The island rule seems to apply to fossil carnivores, but with exceptions. The dependency of the island rule on resource availability is emphasized by the present study.     

Vertebrate cranial evolution: Contributions and conflict from the fossil record

In modern vertebrates, the craniofacial skeleton is complex, comprising cartilage and bone of the neurocranium, dermatocranium and splanchnocranium (and their derivatives), housing a range of sensory structures such as eyes, nasal and vestibulo-acoustic capsules, with the splanchnocranium including branchial arches, used in respiration and feeding. It is well understood that the skeleton derives from neural crest and mesoderm, while the sensory elements derive from ectodermal thickenings known as placodes. Recent research demonstrates that neural crest and placodes have an evolutionary history outside of vertebrates, while the vertebrate fossil record allows the sequence of the evolution of these various features to be understood. Stem-group vertebrates such as Metaspriggina walcotti (Burgess Shale, Middle Cambrian) possess eyes, paired nasal capsules and well-developed branchial arches, the latter derived from cranial neural crest in extant vertebrates, indicating that placodes and neural crest evolved over 500 million years ago. Since that time the vertebrate craniofacial skeleton has evolved, including different types of bone, of potential neural crest or mesodermal origin. One problematic part of the craniofacial skeleton concerns the evolution of the nasal organs, with evidence for both paired and unpaired nasal sacs being the primitive state for vertebrates.     

Braincase evolution in suchian archosaurs (Reptilia: Diapsida): evidence from the rauisuchian Batrachotomus kupferzellensis

The osteology of an almost complete braincase of the rauisuchian archosaur Batrachotomus kupferzellensis Gower from the Middle Triassic of Germany is described. There is a possibly discrete epiotic ossification, the metotic fissure is undivided by bone (i.e. there is a metotic foramen), the medial wall of the otic capsule is mostly ossified, the cerebral branch of the internal carotid artery entered the lateral surface of the parabasisphenoid, the ventral ramus of the opisthotic is more prominent laterally than a strong subvertical ridge on the exoccipital and basioccipital that lies posterior to the external foramen for the hypoglossal nerve, and the perilymphatic foramen faces away from the otic capsule in a posterior direction. Braincase morphology in the rauisuchians Saurosuchus galilei , Postosuchus kirkpatricki, and Tikisuchus romeri is reviewed. A matrix of 27 braincase characters for 12 archosaurian taxa is analysed. The most parsimonious hypothesis is consistent with the currently orthodox view of archosaurian phylogeny, except in that aetosaurians are more closely related to crocodylomorphs than is any rauisuchian. This phylogeny is used in a brief interpretation of the evolution of derived braincase features present in extant crocodilians. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 49–76.     

The fossil record of ichthyosaurs,completeness metrics and sampling biases

Ichthyosaurs were highly successful marine reptiles with an abundant and well‐studied fossil record. However, their occurrences through geological time and space are sporadic, and it is important to understand whether times of apparent species richness and rarity are real or the result of sampling bias. Here, we explore the skeletal completeness of 351 dated and identified ichthyosaur specimens, belonging to all 102 species, the first time that such a study has been carried out on vertebrates from the marine realm. No correlations were found between time series of different skeletal metrics and ichthyosaur diversity. There is a significant geographical variation in completeness, with the well‐studied northern hemisphere producing fossils of much higher quality than the southern hemisphere. Medium‐sized ichthyosaurs are significantly more complete than small or large taxa: the incompleteness of small specimens was expected, but it was a surprise that larger specimens were also relatively incomplete. Completeness varies greatly between facies, with fine‐grained, siliciclastic sediments preserving the most complete specimens. These findings may explain why the ichthyosaur diversity record is low at times, corresponding to facies of poor preservation potential, such as in the Early Cretaceous. Unexpectedly, we find a strong negative correlation between skeletal completeness and sea level, meaning the most complete specimens occurred at times of global low sea level, and vice versa. Completeness metrics, however, do not replicate the sampling signal and have limited use as a global‐scale sampling proxy.     

Quantifying the completeness of the bat fossil record

Bats (Chiroptera) are one of the most successful extant mammalian orders, uniquely capable of powered flight and laryngeal echolocation. The timing and evidence for evolution of their novel adaptations have been difficult to ascertain from the fossil record due to chronological gaps and the fragmentary nature of most fossil bat material. Here, we quantify the quality of the bat fossil record using skeletal and character completeness metrics, which respectively document for each taxon what proportion of a complete skeleton is preserved, and the proportion of phylogenetic characters that can be scored. Completeness scores were collected for 441 valid fossil bat species in 167 genera from the Eocene to the Pleistocene. All metrics record similar temporal patterns: peak completeness in the Lutetian stage reflects the presence of Lagerstätten, while subsequent stages have very low completeness, except an Aquitanian high and a Pleistocene peak in skeletal completeness. Bat completeness is not correlated with intensity of sampling through geological time but has a weak negative correlation with publication date. There is no correlation between taxonomic richness and completeness, as the bat record predominately consists of diagnostic but isolated teeth. Consequently, bat skeletal completeness is the lowest of any previously assessed tetrapod group, but character completeness is similar to parareptiles and birds. Bats have significantly higher character completeness in the northern hemisphere, probably due to heightened historical interest and presence of Lagerstätten. Taxa derived from caves are more complete than those from fluviolacustrine and marine deposits, but do not preserve highly complete specimens.     

Preservational bias controls the fossil record of pterosaurs

Pterosaurs, a Mesozoic group of flying archosaurs, have become a focal point for debates pertaining to the impact of sampling biases on our reading of the fossil record, as well as the utility of sampling proxies in palaeo‐diversity reconstructions. The completeness of the pterosaur fossil specimens themselves potentially provides additional information that is not captured in existing sampling proxies, and might shed new light on the group's evolutionary history. Here we assess the quality of the pterosaur fossil record via a character completeness metric based on the number of phylogenetic characters that can be scored for all known skeletons of 172 valid species, with averaged completeness values calculated for each geological stage. The fossil record of pterosaurs is observed to be strongly influenced by the occurrence and distribution of Lagerstätten. Peaks in completeness correlate with Lagerstätten deposits, and a recovered correlation between completeness and observed diversity is rendered non‐significant when Lagerstätten species are excluded. Intervals previously regarded as potential extinction events are shown to lack Lagerstätten and exhibit low completeness values: as such, the apparent low diversity in these intervals might be at least partly the result of poor fossil record quality. A positive correlation between temporal patterns in completeness of Cretaceous pterosaurs and birds further demonstrates the prominent role that Lagerstätten deposits have on the preservation of smaller bodied organisms, contrasting with a lack of correlation with the completeness of large‐bodied sauropodomorphs. However, we unexpectedly find a strong correlation between sauropodomorph and pterosaur completeness within the Triassic–Jurassic, but not the Cretaceous, potentially relating to a shared shift in environmental preference and thus preservation style through time. This study highlights the importance of understanding the relationship between various taphonomic controls when correcting for sampling bias, and provides additional evidence for the prominent role of sampling on observed patterns in pterosaur macroevolution.     

Ten more years of discovery: revisiting the quality of the sauropodomorph dinosaur fossil record

Spatiotemporal changes in fossil specimen completeness can bias our understanding of a group's evolutionary history. The quality of the sauropodomorph fossil record was assessed a decade ago, but the number of valid species has since increased by 60%, and 17% of the taxa from that study have since undergone taxonomic revision. Here, we assess how 10 years of additional research has changed our outlook on the group's fossil record. We quantified the completeness of all 307 sauropodomorph species currently considered valid using the skeletal completeness metric, which calculates the proportion of a complete skeleton preserved for each taxon. Taxonomic and stratigraphic age revisions, rather than new species, are the drivers of the most significant differences between the current results and those of the previous assessment. No statistical differences appeared when we use our new dataset to generate temporal completeness curves based solely on taxa known in 2009 or 1999. We now observe a severe drop in mean completeness values across the Jurassic–Cretaceous boundary that never recovers to pre-Cretaceous levels. Explaining this pattern is difficult, as we find no convincing evidence that it is related to environmental preferences or body size changes. Instead, it might result from: (1) reduction of terrestrial fossil preservation space due to sea level rise; (2) ecological specificities and relatively high diagnosability of Cretaceous species; and/or (3) increased sampling of newly explored sites with many previously unknown taxa. Revisiting patterns in this manner allows us to test the longevity of conclusions made in previous quantitative studies.     

Recognising angiosperm clades in the Early Cretaceous fossil record

Studies of the earliest Cretaceous angiosperms in the 1970s made only broad comparisons with living taxa, but discoveries of fossil flowers and increasingly robust molecular phylogenies of living angiosperms allow more secure recognition of extant clades. The middle to late Albian rise of tricolpate pollen and the first local dominance of angiosperm leaves mark the influx of near-basal lines of eudicots. Associated flowers indicate that palmately lobed ‘platanoids’ and Sapindopsis are both stem relatives of Platanus, while Nelumbites was related to Nelumbo (also Proteales) and Spanomera to Buxaceae. Monocots are attested by Aptian Liliacidites pollen and Acaciaephyllum leaves and Albian araceous inflorescences. Several Albian–Cenomanian fossils belong to Magnoliidae in the revised monophyletic sense, including Archaeanthus in Magnoliales and Virginianthus and Mauldinia in Laurales, while late Barremian pollen tetrads (Walkeripollis) are related to Winteraceae. In the basal ANITA grade, Nymphaeales are represented by Aptian and Albian flowers and whole plants (Monetianthus, Carpestella and Pluricarpellatia). Epidermal similarities of lower Potomac leaves to woody members of the ANITA grade are consistent with Albian flowers assignable to Austrobaileyales (Anacostia). Aptian to Cenomanian mesofossils represent both crown group Chloranthaceae (Asteropollis plant) and stem relatives of Chloranthaceae and/or Ceratophyllum (Canrightia, Zlatkocarpus, Pennipollis plant and possibly Appomattoxia).     

The fossil evidence of anthropoid brain evolution

The endocast of Aegyptopithecus, a 27 million year old ape, reveals that its brain was advanced over that of prosimians and comparable to that of modern anthropoids in relative size and in having expanded visual cortex, reduced olfactory bulbs, and a central sulcus separating primary somatic sensory and motor cortex. The early appearance of those features suggests that they may have been among the adaptations responsible for the evolution of anthropoids from prosimian ancestors. The frontal lobe was relatively smaller in Aegyptopithecus than in modern anthropoids. An endocast of Dolichocebus, one of the oldest known New World monkeys (25–30 million years old), reveals visual cortex expanded as in modern anthropoids. The 19 million year old Napak frontal bone displays a hominoid rather than cercopithecoid sulcal pattern. An 18 million year old endocast of the ape Dryopithecus (Proconsul) was neither monkey-like nor primitive, as originally described, but rather apelike and essentially modern in all observable features. The oldest undoubted Old World monkey endocast, from nine million year old Mesopithecus, reveals that the brain was modern in sulcal pattern and proportions. The sulcal pattern was like that of modern colobines, but that appears to be the more primitive condition, from which features characteristic of modern cercopithecine brains have evolved. The brain of six million year old Libypithecus was similar to that of Mesopithecus. A two million year old endocast of “Dolichopithecus” arvernensis displays a modern cercopithecine sulcal pattern.     

The early evolution of feathers: fossil evidence from Cretaceous amber of France

The developmental stages of feathers are of major importance in the evolution of body covering and the origin of avian flight. Until now, there were significant gaps in knowledge of early morphologies in theoretical stages of feathers as well as in palaeontological material. Here we report fossil evidence of an intermediate and critical stage in the incremental evolution of feathers which has been predicted by developmental theories but hitherto undocumented by evidence from both the recent and the fossil records. Seven feathers have been found in an Early Cretaceous (Late Albian, ca 100 Myr) amber of western France, which display a flattened shaft composed by the still distinct and incompletely fused bases of the barbs forming two irregular vanes. Considering their remarkably primitive features, and since recent discoveries have yielded feathers of modern type in some derived theropod dinosaurs, the Albian feathers from France might have been derived either from an early bird or from a non-avian dinosaur.     

Rates and patterns in the evolution of snake-like body form in squamate reptiles: evidence for repeated re-evolution of lost digits and long-term persistence of intermediate body forms

An important challenge in evolutionary biology is to understand how major changes in body form arise. The dramatic transition from a lizard-like to snake-like body form in squamate reptiles offers an exciting system for such research because this change is replicated dozens of times. Here, we use morphometric data for 258 species and a time-calibrated phylogeny to explore rates and patterns of body-form evolution across squamates. We also demonstrate how time-calibrated phylogenies may be used to make inferences about the time frame over which major morphological transitions occur. Using the morphometric data, we find that the transition from lizard-like to snake-like body form involves concerted evolution of limb reduction, digit loss, and body elongation. These correlations are similar across squamate clades, despite very different ecologies and >180 million years (My) of divergence. Using the time-calibrated phylogeny and ancestral reconstructions, we find that the dramatic transition between these body forms can occur in 20 My or less, but that seemingly intermediate morphologies can also persist for tens of millions of years. Finally, although loss of digits is common, we find statistically significant support for at least six examples of the re-evolution of lost digits in the forelimb and hind limb.     

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.