The origin of metazoan development: a palaeobiological perspective

The rapid diversification of early Metazoa remains one of the most puzzling events in the fossil record. Several models have been proposed to explain a critical aspect of this event: the origin of Metazoan development. These include the origin of the eukaryotic cell, environmental triggers, key innovations or selection among cell lineages. Here, the first three hypotheses are evaulated within a phylogenetic framework using fossil, molecular and developmental evidence. Many elements of metazoan development are widely distributed among unicellular eukaryotes, yet only 3 of the 23 multicellular eukaryotic lineages evolved complex development. Molecular evidence indicates the lineage leading to the eukaryotic cell is nearly as old as the eubacterial and archaebacterial lineages, although the symbiotic events established that the eukaryotic cell probably occurred about 1.5 billion years ago. Yet Metazoa did not appear until 1000 to 600 million years ago (Myr), suggesting the origin of metazoan development must be linked to either an environmental trigger, perhaps an increase in atmospheric oxygen, or key innovations such as the development of collagen. Yet the first model fails to explain the unique appearance of complex development in Metazoa, while the latter fails to explain the simultaneous diversification of several ‘protist’ groups along with the Metazoa. A more complete model of the origin of metazoan development combines environmental triggering of a series of innovations, with successive innovations generating radiations of metazoan clades as lineages breached functional thresholds. The elaboration of new cell classes and the appearance of such developmental innovations as cell sheets may have been of particular importance. Evolutionary biologists often implicitly assume that evolution is a uniformitarian, time-homogeneous process without strong temporal asymmetries in evolutionary mechanisms, rate or context. Yet evolutionary patterns do exhibit such asymmetries, raising the possibility that such innovations as metazoan development impose non-uniformities of evolutionary process.     

A molecular palaeobiological hypothesis for the origin of aplacophoran molluscs and their derivation from chiton-like ancestors

Aplacophorans have long been argued to be basal molluscs. We present a molecular phylogeny, including the aplacophorans Neomeniomorpha (Solenogastres) and Chaetodermomorpha (Caudofoveata), which recovered instead the clade Aculifera (Aplacophora + Polyplacophora). Our relaxed Bayesian molecular clock estimates an Early Ordovician appearance of the aculiferan crown group consistent with the presence of chiton-like molluscs with seven or eight dorsal shell plates by the Late Cambrian (approx. 501-490 Ma). Molecular, embryological and palaeontological data indicate that aplacophorans, as well as chitons, evolved from a paraphyletic assemblage of chiton-like ancestors. The recovery of cephalopods as a sister group to aculiferans suggests that the plesiomorphic condition in molluscs might be a morphology similar to that found in monoplacophorans.     

The concept of correlated progression as the basis of a model for the evolutionary origin of major new taxa

Understanding the evolutionary processes responsible for the long treks through morphospace associated with the origin of new higher taxa is hampered by the lack of a realistic and usable model that accounts for long-term phenotypic evolvability. The systems-related concept of correlated progression, in which all the traits are functionally linked and so constrained to evolve by small increments at a time in parallel with each other, provides the basis for such a model. Implications for the process of evolution at high taxonomic level are that: the evolving traits must be considered together as a system, and the exact sequence of incremental changes in characters is indeterminable; there are no identifiable key innovations; selection acts on the phenotype as a whole rather than on individual traits; and the selection force is therefore multidimensional. Application of the model to the pattern of evolution of traits and trait states as revealed by the fossil record of the stem groups of such taxa as mammals, turtles and tetrapods generates realistic testable hypotheses about how such groups evolved.     

The origin of higher taxa: macroevolutionary processes, and the case of the mammals

The origin of a new higher taxon is characterized by a long‐term phylogenetic trend, involving evolutionary changes in a large number of characters. At this phylogenetic level, the conflict between internal integration of the phenotype and its evolvability can be resolved by the correlated progression model, in which many disparate traits evolve by a sequence of small increments in loose correlation with one another, rather than by the modularity model. The trend leading to the new higher taxon implies the existence of a long ridge in an adaptive landscape. An evolutionary lineage tracking it requires adaptive changes in broad biological characteristics, involving many traits. Species selection is a possible additional driver of the trend. These conclusions are tested against the synapsid fossil record of the origin of mammals. The reconstructed sequence of acquisition of mammalian traits supports the correlated progression model. The adaptive ridge involved is postulated to have been a sequence of overlapping niches requiring increasing ability to remain active in daily and seasonally fluctuating environments by means of increasing internal regulation. An inferred speciation bias in favour of relatively small, relatively more progressive carnivores indicates that species selection was also involved in driving the trend. Palaeoenvironmental evidence indicates that ecological opportunity probably played a role at certain points along the lineage.     

The origin of the vomiting response: a neuroanatomical hypothesis

The connections of area postrema include a set of nuclei with a common topographical location immediately deep to the ependyma or pia mater. These nuclei are all within two principal synapses of the area postrema and can be reached by more than one route from it. There is direct evidence that, like the area postrema, a number of these nuclei participate in vomiting. It is suggested that the paraventricular system may act as a distributed pattern generator for the several processes known to be integrated in the emetic response. It is also suggested that the other functions of the paraventricular nuclei, mainly homeostatic in mammals, have all evolved from a prototypical behaviour pattern of escape from, and subsequent avoidance of, noxious stimuli. Thus, through this system of nuclei, blood pressure, respiration, gastrointestinal motility and secretion, fluid and electrolyte balance, and the ingestion of food may all be influenced by noxious situations.     

New therapsid specimens and the origin of the secondary hard and soft palate of mammals

A newly prepared palate of the moschorhinid therocephalian Promoschorhynchus platyrhinus from the Upper Permian region of South Africa was used for reconsidering the initial evolutionary development of the secondary palate of mammals. The very distinctive choanal crests are considered to be indicative of strong choanal folds that were probably already fused anteriorly. This gingival bridge probably served as the basis for the outgrowing bony processes of the hard palate that, in therapsids, developed independently at least three times. The soft palate ( velum palatinum ) is considered to be a remnant of the choanal folds that were muscularized from behind. It has been assumed that, in therapsids, a ventral portion of the medial pterygoid musculature shifted its insertion onto the ventral side of the pterygoid; this portion is supposed to have differentiated into the mm. tensor tympani and tensor veli palatini . Investigation of serial sections of extant marsupials confirm the view that the levator of the velum is derived from the upper constrictor of the pharynx. The choanal folds and the secondary palate are discussed within the wider framework of the evolutionary biology of mammalian forerunners. It is suggested that the formation of a sealed mouth cavity is not only related to the improved passage of air, but also to sucking in neonate hatchlings. The importance of the secondary palate of mammals to a number of resulting basic adaptations (dentition, olfactory system, etc.) of mammals is discussed.     

Active centrum hypothesis: The origin of chiral homogeneity and the RNA-world

I propose a hypothesis on the origin of chiral homogeneity of bio-molecules based on chiral catalysis. The first chiral active centre may have formed on the surface of complexes comprising metal ions, amino acids, other coenzymes and oligomers (short RNAs). The complexes must have been dominated by short RNAs capable of self-reproduction with ligation. Most of the first complexes may have catalysed the production of nucleotides. A basic assumption is that such complexes can be assembled from their components almost freely, in a huge variety of combinations. This assumption implies that “a few” components can constitute “a huge” number of active centre types. Moreover, an experiment is proposed to test the performance of such complexes in vitro.If the complexes were built up freely from their elements, then Darwinian evolution would operate on the assembly mechanism of complexes. For the production of complexes, first their parts had to appear by forming a proper three-dimensional structure. Three possible re-building mechanisms of the proper geometric structure of complexes are proposed. First, the integration of RNA parts of complexes was assisted presumably by a pre-intron. Second, the binding of RNA parts of a complex may give rise to a “polluted” RNA world. Third, the pairing of short RNA parts and their geometric conformation may have been supported by a pre-genetic code.Finally, an evolutionary step-by-step scenario of the origin of homochirality and a “polluted” RNA world is also introduced based on the proposed combinatorial complex chemistry. Homochirality is evolved by Darwinian selection whenever the efficiency of the reflexive autocatalysis of a dynamical combinatorial library increases with the homochirality of the active centres of reactions cascades and the homochirality of the elements of the dynamical combinatorial library. Moreover, the potential importance of phospholipid membrane is also discussed.     

The origin and early evolution of metatherian mammals: the Cretaceous record

Metatherians, which comprise marsupials and their closest fossil relatives, were one of the most dominant clades of mammals during the Cretaceous and are the most diverse clade of living mammals after Placentalia. Our understanding of this group has increased greatly over the past 20 years, with the discovery of new specimens and the application of new analytical tools. Here we provide a review of the phylogenetic relationships of metatherians with respect to other mammals, discuss the taxonomic definition and diagnosis of Metatheria, outline the Cretaceous history of major metatherian clades, describe the paleobiology, biogeography, and macroevolution of Cretaceous metatherians, and provide a physical and climatic background of Cretaceous metatherian faunas. Metatherians are a clade of boreosphendian mammals that must have originated by the Late Jurassic, but the first unequivocal metatherian fossil is from the Early Cretaceous of Asia. Metatherians have the distinctive tightly interlocking occlusal molar pattern of tribosphenic mammals, but differ from Eutheria in their dental formula and tooth replacement pattern, which may be related to the metatherian reproductive process which includes an extended period of lactation followed by birth of extremely altricial young. Metatherians were widespread over Laurasia during the Cretaceous, with members present in Asia, Europe, and North America by the early Late Cretaceous. In particular, they were taxonomically and morphologically diverse and relatively abundant in the Late Cretaceous of western North America, where they have been used to examine patterns of biogeography, macroevolution, diversification, and extinction through the Late Cretaceous and across the Cretaceous-Paleogene (K-Pg) boundary. Metatherian diversification patterns suggest that they were not strongly affected by a Cretaceous Terrestrial Revolution, but they clearly underwent a severe extinction across the K-Pg boundary.     

The occurrence of thermophilous trees in the Scandes Mountains during the early Holocene: evidence for a diverse tree flora from macroscopic remains

1 Macroscopic remains of the fairly thermophilous tree species Alnus glutinosa , Tilia cordata and Betula pendula were recovered in subalpine and adjacent boreal environments far above and beyond their present-day distributional limits. This establishes that the early Holocene tree flora of the Scandes Mountains in northern Sweden was indeed richer than it is today.
2 Dates ranged between c . 8600 and 7000 radiocarbon years bp . These are much earlier than previous estimates by conventional pollen stratigraphical analyses of the arrival of these species at their maximum geographical limits. This highlights problems in using only pollen data for vegetation reconstruction, and suggests re-evaluation of earlier records.
3 The results, together with similar macrofossils for Picea abies and Larix sibirica in northern Sweden, suggest that many tree species spread rapidly and became established at their most extended range limits during the early Holocene. Abundances have subsequently varied in accordance with the ecology of individual species as well as with climatic change.
4 Palaeoclimatic inferences may suggest a strongly continental climate, i.e. warmer and drier summers and possibly fairly cold winters between 8600 and 7000 bp relative to the present. Some change towards a more oceanic climate regime with less pronounced seasonal contrasts may have occurred towards the end of the period.     

The early Triassic rhynchosaur Mesosuchus browni and the interrelationships of basal archosauromorph reptiles

Restudy of the unique diapsid reptile Mesosuchus browni Watson, from the Cynognathus Assemblage Zone (late Early Triassic to early Middle Triassic) of the Burgersdorp Formation (Tarkastad Subgroup; Beaufort Group) of South Africa, confirms that it is the most plesiomorphic known member of the Rhynchosauria. A new phylogenetic analysis of basal taxa of Archosauromorpha indicates that Choristodera falls outside of the Sauria, Prolacertiformes is a paraphyletic taxon with Prolacerta sharing a more recent common ancestor with Archosauriformes than with any other clade, Megalancosaurus and Drepanosaurus are sister taxa in the clade Drepanosauridae within Archosauromorpha, and are the sister group to the clade Tanystropheidae composed of Tanystropheus, Macrocnemus, and Langobardisaurus. Combination of the phylogenetic relationships of basal archosauromorphs and their known stratigraphic ranges reveals significant gaps in the fossil records of Late Permian and Triassic diapsids. Extensions of the temporal ranges of several lineages of diapsids into the Late Permian suggests that more groups of terrestrial reptiles survived the end-Permian mass extinction than thought previously.     

A fossil Halobates from the Mediterranean and the origin of sea skaters (Hemiptera, Gerridae)

Five species of sea skaters, genus Halobates Eschscholtz, are the only insects to have successfully colonized the open ocean. In addition, 38 species are found in sheltered coastal waters throughout tropical Indo-Pacific. The taxonomy of the genus is relatively well known, and the phylogeneuc relationships between extant species have recently been analysed (using cladistic methods). In the present paper, we describe the first fossil species of sea skaters, Halobates ruffoi sp. no v. from the Eocene deposit 'Pesciara di Bolca', in the province of Verona, northeastern Italy (geological age about 45 Myr). The significance of this fossil in setting the time scale for the reconstructed phylogeny and anagenesis of adaptive features of sea skaters, and in understanding the evolution and historical zoogeography of these marine insects is discussed.     

The origin and early evolution of birds

Birds evolved from and are phylogenetically recognized as members of the theropod dinosaurs; their first known member is the Late Jurassic Archaeopteryx, now represented by seven skeletons and a feather, and their closest known non-avian relatives are the dromaeosaurid theropods such as Deinonychus. Bird flight is widely thought to have evolved from the trees down, but Archaeopteryx and its outgroups show no obvious arboreal or tree-climbing characters, and its wing planform and wing loading do not resemble those of gliders. The ancestors of birds were bipedal, terrestrial, agile, cursorial and carnivorous or omnivorous. Apart from a perching foot and some skeletal fusions, a great many characters that are usually considered ‘avian’ (e.g. the furcula, the elongated forearm, the laterally flexing wrist and apparently feathers) evolved in non-avian theropods for reasons unrelated to birds or to flight. Soon after Archaeopteryx, avian features such as the pygostyle, fusion of the carpometacarpus, and elongated curved pedal claws with a reversed, fully descended and opposable hallux, indicate improved flying ability and arboreal habits. In the further evolution of birds, characters related to the flight apparatus phylogenetically preceded those related to the rest of the skeleton and skull. Mesozoic birds are more diverse and numerous than thought previously and the most diverse known group of Cretaceous birds, the Enantiornithes, was not even recognized until 1981. The vast majority of Mesozoic bird groups have no Tertiary records: Enantiornithes, Hesperornithiformes, Ichthyornithiformes and several other lineages disappeared by the end of the Cretaceous. By that time, a few Linnean ‘Orders’ of extant birds had appeared, but none of these taxa belongs to extant ‘families’, and it is not until the Paleocene or (in most cases) the Eocene that the majority of extant bird ‘Orders’ are known in the fossil record. There is no evidence for a major or mass extinction of birds at the end of the Cretaceous, nor for a sudden ‘bottleneck’ in diversity that fostered the early Tertiary origination of living bird ‘Orders’.     

The cranial anatomy and relationships of Secodontosaurus, an unusual mammal-like reptile (Synapsida: Sphenacodontidae) from the early Permian of Texas

The cranial anatomy of the Early Permian sphenacodontid synapsid Secodontosaurus is redescribed. There is no evidence for recognition of more than one species of Secodontosaurus, and S. willistoni is declared a junior subjective synonym of S. obtusidens. Numerous derived characters (autapomorphies), mainly related to the unusually slender and slightly elongated skull, distinguish Secodontosaurus. The cranial specializations are interpreted as an adaptation toward a feeding strategy which involved preying upon small tetrapods that attempted to avoid capture by hiding in crevices and burrows. A cladistic analysis supports the following hypotheses of relationships: (a) the Sphenacodontidae is the nearest sister taxon of Therapsida; (b) Secodontosaurus, Sphenacodon, Ctenospondylus and Dimetrodon share a more recent common ancestor with one another than any of them do with Haptodus, and (c) Secodontosaurus shares a more recent common ancestor with Dimetrodon than with either Sphenacodon or Ctenospondylus. The latter hypothesis suggests that the sphenacodont subfamily 'Sphenacodontinae' is paraphyletic and, therefore, invalid.     

Fossil evidence for a herbaceous diversification of early eudicot angiosperms during the Early Cretaceous

Eudicot flowering plants comprise roughly 70% of land plant species diversity today, but their early evolution is not well understood. Fossil evidence has been largely restricted to their distinctive tricolpate pollen grains and this has limited our understanding of the ecological strategies that characterized their primary radiation. I describe megafossils of an Early Cretaceous eudicot from the Potomac Group in Maryland and Virginia, USA that are complete enough to allow reconstruction of important life-history traits. I draw on quantitative and qualitative analysis of functional traits, phylogenetic analysis and sedimentological evidence to reconstruct the biology of this extinct species. These plants were small and locally rare but widespread, fast-growing herbs. They had complex leaves and they were colonizers of bright, wet, disturbance-prone habitats. Other early eudicot megafossils appear to be herbaceous rather than woody, suggesting that this habit was characteristic of their primary radiation. A mostly herbaceous initial diversification of eudicots could simultaneously explain the heretofore sparse megafossil record as well as their rapid diversification during the Early Cretaceous because the angiosperm capacity for fast reproduction and fast evolution is best expressed in herbs.     

The origin of homeostasis in the early earth

Summary An equation is developed that describes the condition of homeostasis in a general molecular system containing catalysts. In a prebiotic environment, this condition first results from a critical level of catalytic feedback in feedback loops containing differing organic molecular species. This critical level results in temporary exponential growth in concentrations of those catalyst species participating in the feedback loops, leading to homeostasis as the steady-state endpoint. None of the molecules in any feedback loop need be self-replicating for this autocatalysis to occur. Homeostasis is regarded as a definition of life at the lowest possible hierarchical level. A general mathematical boundary condition is derived for the critical level of catalytic feedback mentioned above-in effect, an origin of life condition. The paper argues that any natural prebiotic system of organic molecules in an H₂O medium will automatically form many catalytic feedback loops, even if of very low catalytic efficiency. The analysis in this paper indicates that high temperatures strongly increase the efficiency of such catalytic feedback. If the temperature and total concentration of carbon in the system (e.g., in CO₂, CH₄, etc.) are sufficiently high, the critical condition for initial exponential growth will be attained. High initial temperatures for the earth are predicted by the planetesimal accretion model.     

Ancestral xerophobia: a hypothesis on the whole plant ecophysiology of early angiosperms

Today, angiosperms are fundamental players in the diversity and biogeochemical functioning of the planet. Yet despite the omnipresence of angiosperms in today's ecosystems, the basic evolutionary understanding of how the earliest angiosperms functioned remains unknown. Here we synthesize ecophysiological, paleobotanical, paleoecological, and phylogenetic lines of evidence about early angiosperms and their environments. In doing so, we arrive at a hypothesis that early angiosperms evolved in evermoist tropical terrestrial habitats, where three of their emblematic innovations – including net‐veined leaves, xylem vessels, and flowers – found ecophysiological advantages. However, the adaptation of early angiosperm ecophysiology to wet habitats did not initially promote massive diversification and ecological dominance. Instead, wet habitats were permissive for the ecological roothold of the clade, a critical phase of early diversification that entailed experimentation with a range of functional innovations in the leaves, wood, and flowers. Later, our results suggest that some of these innovations were co‐opted gradually for new roles in the evolution of greater productivity and drought tolerance, which are characteristics seen across the vast majority of derived and ecologically dominant angiosperms today.     

Records of a lake's life in time: the sediments

The evidence from analysis of cores from lake sediments is used to identify the influences which, acting through time, have brought about changes in the lakes of the English Lake District. These are: i. climatic changes, recorded mainly in microfossil assemblages ii. soil dynamics on catchments — the natural soil development of an interglacial cycle and its effect on water composition, and iii. perturbations of input resulting from activities of man.The present position of each of 14 lakes in a series of increasing eutrophication is shown to be the result of two processes of modification by man. The first was a significant reduction in volume of the hypolimnion of the shallow lakes, consequent on the accelerated rate of sediment accumulation which followed deforestation and cultivation of catchments in all the lakes — this did not affect the biology of the deep lakes. The second has been the introduction of human and animal wastes into some of the lakes during the last 120 years, which imposed on affected lakes a process of more rapid change which was more intense in the shallow lakes.