Amphibian development in the fossil record

Ontogenetic series of extinct taxa are extremely rare and when preserved often incomplete and difficult to interpret. However, the fossil record of amphibians includes a number of well-preserved ontogenetic sequences for temnospondyl and lepospondyl taxa, which have provided valuable information about the development of these extinct groups. Here we summarize the current knowledge on fossil ontogenies of amphibians, their potential and limitations for relationship assessments, and discuss the insights they have provided for our understanding of the anatomy, life history, and ecology of extinct amphibians.     

Patterns of heterochrony in the fossil record

Recent analyses of the fossil record have revealed that heterochrony has played a significant role in the evolution of most marine invertebrate groups. Recognition of several different heterochronic processes has allowed their influence under different ecological regimes to be assessed. Furthermore, recent research has demonstrated significant differences in frequencies of these processes between groups and at their different stages of phylogenetic development. The fossil record shows that heterochron y has been an important component in the generation of evolutionary trends. Heterochrony is an important factor in macroevolution because it can result in relatively abrupt morphological change with only minimum alteration of the genome.     

Power spectra of extinction in the fossil record

Recent Fourier analyses of fossil extinction data have indicated that the power spectrum of extinction during the Phanerozoic may take the form of 1/f noise, a result which, it has been suggested, could be indicative of the presence of `critical dynamics'' in the processes giving rise to extinction. In this paper we examine extinction power spectra in some detail, using family-level data from two widely available compilations. We find that although the average form of the power spectrum roughly obeys the 1/f law, the spectrum can be represented more accurately by dividing it into two regimes: a low-frequency one which is well fit by an exponential, and a high-frequency one in which it follows a power law with a 1/f² form. We give explanations for the occurrence of each of these behaviours and for the position of the crossover between them.     

Cladistics and the hominid fossil record

Cladistic methodology has become common in phylogenetic analyses of the hominid fossil record. Even though it has correctly placed emphasis on morphology for the primary determination of affinities between groups and on explicit statements regarding traits and methods employed in making phylogenetic assessments, cladistics nonetheless has limitations when applied to the hominid fossil record. These include 1) the uncritical assumption of parsimony, 2) uncertainties in the identification of homoplasies, 3) difficulties in the appropriate delimitation of samples for analysis, 4) failure to account for normal patterns of variation, 5) methodological problems with the appropriate identification of morphological traits involving issues of biological relevance, intercorrelation, primary versus secondary characters, and the use of continuous variables, 6) issues of polarity identification, and 7) problems in hypothesis testing. While cladistics has focused attention on alternative phylogenetic reconstructions in hominid paleontology and on explicit statements regarding their morphological and methodological underpinnings, its biological limitations are too abundant for it to be more than a heuristic device for the preliminary ordering of complex human paleontological and neonatological data.     

Patterns of generic extinction in the fossil record

Analysis of the stratigraphic records of 19,897 fossil genera indicates that most classes and orders show largely congruent rises and falls in extinction intensity throughout the Phanerozoic. Even an ecologically homogeneous sample of reef genera shows the same basic extinction profile. The most likely explanation for the congruence is that extinction is physically rather than biologically driven and that it is dominated by the effects of geographically widespread environmental perturbations influencing most habitats. Significant departures from the congruence are uncommon but important because they indicate physiological or habitat selectivity. The similarity of the extinction records of reef organisms and the marine biota as a whole confirms that reefs and other faunas are responding to the same history of environmental stress.     

Atomism, transformism and the fossil record

The modern debate between exponents of classical evolutionary classification and of cladistic analysis of phylogenetic relationships mirrors to some extent the arguments that were put forward in the debate between Cuvier and Geoffroy Saint-Hilaire in 1830. Put into a historical perspective the problems of comparative biology centre around two complementary traditions, atomism and transformism.     

Coupled evolutionary rates and the fossil record

A model for speciation is given in which a taxon’s phenotypic variation and concomitant variation in fitness are related to gradients within the environment. Phenotypic expressions within the population are shown to undergo abrupt transitions as a result of discontinuous fitness-functions. Evidence for rapid and abrupt phenotypic variation is explored by analyses of speciation (= origination) rates within the fossil record. In general, a high correlation exists among the area/volume changes of sedimentary rocks known for each geologic period and the apparent speciation rates seen in selected vascular/non-vascular plant groups. Regressions of speciation rates on rock area/volume plots indicate that the Devonian, Carboniferous, and Permian show significant divergences (= residual) rates from predicted origination rates. The Cretaceous shows the highest residual value as a consequence of the rapid appearance of angiosperm fossils. A similar pattern in diversity changes for the mandibulate terrestrial invertebrates is also apparent. The coupled evolution of the angiosperms with specific insect groups appears to be the most tenable explanation for the residual Cretaceous origination rate of the former group. It is postulated that the angiospems have evolved in part as the result of a phytochemical cost-function such that phytophagous insects are warded off, while potential pollinators are favored. Quantitative/qualitative differences observed in the distribution of secondary metabolites may be evidence for coupled evolution. A “predatorprey mediated co-existence” between phytophagous insects and angiosperms may have served as a factor in allowing the co-existence of less than optimal plant species providing an impetus for relatively rapid speciation turnover.     

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.     

Cyclicity in the fossil record mirrors rock outcrop area

In a recent article, Rohde & Muller (Rohde & Muller 2005 Nature 434, 208-210) identified a strong 62 Myr cyclicity in the history of marine diversity through the Phanerozoic. The data they presented were highly convincing, yet they were unable to explain what process might have generated this pattern. A significant correlation between observed genus-level diversity (after removal of long-term trends) and the amount of marine sedimentary rock measured at a surface outcrop in Western Europe is demonstrated. This suggests that cyclicity originates from long-term changes in sedimentary depositional and erosional regimes, and raises the strong possibility that the cyclicity apparent in the record of marine fossils is not a biological signal but a sampling signal.     

Molecular clocks and the incompleteness of the fossil record

Molecular clocks can be evaluated by comparing absolute rates of evolution and by performing relative-rate tests. Typically, calculations of absolute rates are based on earliest observed occurrences in the fossil record. Relative-rate tests, on the other hand, merely require an unambiguous outgroup. A major disadvantage of relative-rate tests is their insensitivity to concomitant and equal rate changes in all lineages. Apparent differences in absolute rates, in turn, may be artifacts that are attributable to an incomplete fossil record.Recently developed methods in quantitative biostratigraphy recognize the incompleteness of the fossil record and allow us to place confidence intervals on the endpoints of taxon ranges. These methods are applicable to taxa whose fossil records are of markedly different quality. When we extend these methods and integrate molecular and paleontologic data, we can test the null hypothesis that seemingly disparate rates of molecular evolution are in fact equal under the simplifying assumption that fossils are randomly and independently distributed over their temporal ranges and that fossils can be accurately placed in a phylogenetic context. We can also estimate the range of ticking rates, if any, that are compatible with known fossil data. Ultimately, more accurate rate estimates for widely divergent taxa should allow for more meaningful comparisons of evolutionary rates.DNA hybridization data for monotremes and marsupials suggest a 17-fold difference for 14 different rate calculations with a mean value of approximately 1% divergence per million years. Variation among marsupials is sevenfold. However, when we apply appropriate statistical tests and make additional allowances for fossils of uncertain taxonomic assignment, etc., all 14 rates are compatible with a molecular clock ticking at approximately 0.4% divergence per million years. In addition, this analysis brings relative- and absolute-rate tests into accord.     

Lemuriform origins as viewed from the fossil record

Fossils relevant to lemuriform origins are reviewed. Omanodon seems very close to the other early tooth-combed lemuriforms Karanisia, Wadilemur and Saharagalago, whereas Bugtilemur is rejected from the Lemuriformes. The Djebelemurinae, including Djebelemur and 'Anchomomys' milleri, are considered as stem lemuriforms preceding tooth comb differentiation; they are shown to be very distinct from European adapiforms. With tooth-combed lemuriforms present in Africa around 40 million years ago, and stem lemuriforms without tooth combs present on the same continent around 50-48 million years ago, a reasonable scenario can be proposed: tooth comb differentiation and lemuriform dispersal to Madagascar between 52-40 million years ago. The possible significance of Plesiopithecus for daubentoniid origins is raised. A critique of molecular dates is presented in the light of the fossil record. Azibiids are possibly early African prosimians. The timing of the dispersal of primates to Africa and the problem of strepsirhine origins are briefly examined.     

The quality of the fossil record of Mesozoic birds

The Mesozoic fossil record has proved critical for understanding the early evolution and subsequent radiation of birds. Little is known, however, about its relative completeness: just how 'good' is the fossil record of birds from the Mesozoic? This question has come to prominence recently in the debate over differences in estimated dates of origin of major clades of birds from molecular and palaeontological data. Using a dataset comprising all known fossil taxa, we present analyses that go some way towards answering this question. Whereas avian diversity remains poorly represented in the Mesozoic, many relatively complete bird specimens have been discovered. New taxa have been added to the phylogenetic tree of basal birds, but its overall shape remains constant, suggesting that the broad outlines of early avian evolution are consistently represented: no stage in the Mesozoic is characterized by an overabundance of scrappy fossils compared with more complete specimens. Examples of Neornithes (modern orders) are known from later stages in the Cretaceous, but their fossils are rarer and scrappier than those of basal bird groups, which we suggest is a biological, rather than a geological, signal.     

Mass extinction events and the plant fossil record

Five mass extinction events have punctuated the geological record of marine invertebrate life. They are characterized by faunal extinction rates and magnitudes that far exceed those observed elsewhere in the geological record. Despite compelling evidence that these extinction events were probably driven by dramatic global environmental change, they were originally thought to have little macroecological or evolutionary consequence for terrestrial plants. New high-resolution regional palaeoecological studies are beginning to challenge this orthodoxy, providing evidence for extensive ecological upheaval, high species-level turnover and recovery intervals lasting millions of years. The challenge ahead is to establish the geographical extent of the ecological upheaval, because reconstructing the vegetation dynamics associated with these events will elucidate the role of floral change in faunal mass extinction and provide a better understanding of how plants have historically responded to global environmental change similar to that anticipated for our future.     

Geographical range and speciation in fossil and living molluscs

The notion of a positive relation between geographical range and speciation rate or speciation probability may go back to Darwin, but a negative relation between these parameters is equally plausible. Here, we test these alternatives in fossil and living molluscan taxa. Late Cretaceous gastropod genera exhibit a strong negative relation between the geographical ranges of constituent species and speciation rate per species per million years; this result is robust to sampling biases against small-bodied taxa and is not attributable to phylogenetic effects. They also exhibit weak inverse or non-significant relations between geographical range and (i) the total number of species produced over the 18 million year timeframe, and (ii) the number of species in a single timeplane. Sister-group comparisons using extant molluscan species also show a non-significant relation between median geographical range and species richness of genera. These results support the view that the factors promoting broad geographical ranges also tend to damp speciation rates. They also demonstrate that a strong inverse relation between per-species speciation rate and geographical range need not be reflected in analyses conducted within a single timeplane, underscoring the inadequacy of treating net speciation as a proxy for raw per-taxon rates.     

The fossil record of the Peronosporomycetes (Oomycota)

Evidence of fossil Peronosporomycetes has been slow to accumulate. In this review various fossils historically assigmed to the Peronosporomycets are dicussed briefly and an explanation is provided as to why the fossil record of this grouop has remained inconsistent. In recent year there has been several new reports of fossil peronosporomycetes based on structurally preserved oogonium-antheridium complexes from Derovonian and Carboniferous rocks that demonstrate the existence of these organisms as fossils and refute the long-standing assumption that they are too delicate to be preserved. Among these are serral tyoes characterized by oogonial surface members of the group. To date at last three groups of fossil vascular plants (i.e. lycophytes, ferns and seed ferns) are known to host peronosporomycetes aas endophytes; however only one form has been identified as a parasite.