Colony growth form in some Silurian heliolitid corals is analyzed by the measurement of their shape in profile. Data are presented for seven species, Stelliporella parvistella, Heliolites interstinctus, H. megastoma, H. daintreei, H. spongodes, Propora tubulata, and Plasmopora scita from three localities in Gotland, Sweden, and three localities in England. Intraspecific growth-form variation is presented on triangle diagrams. These plots allow variation to be compared between species present at each locality and between localities for each species. Results indicate that the overall potential for growth-form variation is genetically controlled and that levels of response to environmental stimuli may differ markedly between species. Stelliporella parvistella is a very plastic species, the only one developing branching growth in addition to other growth forms. Heliolites interstinctus is much less variable, dominantly tabular, domal, and low bulbous in form, but demonstrates a similar response. Propora tubulata has a tightly constrained bulbous growth form that shows little variation between localities. The other species are represented by few specimens, most of which parallel H. interstinctus. The likely moderating influences of light levels, substrate type, sedimentation rate, energy levels, and other variables on growth-form variation and species range are considered. The main environmental factor including ecophenotypic response is concluded to be sedimentation rate. A close correlation between this factor and growth form in S. parvistella indicates that form in this species is a particularly sensitive indicator of sedimentation rate and substrate conditions. No simple equations can be made between specific environments and one particular growth form in these corals.

For decades, theoretical morphological studies of different groups of organisms have been successfully pursued in biological, paleontological, and computational contexts, often with distinct modeling approaches and research questions. A regular influx of new perspectives and varied expertise has contributed to the emergence of a veritable multidisciplinary outlook for theoretical morphology. The broadening of this discipline is reflected in a substantial increase in the number of models, leading to a bewildering diversity that has yet to be scrutinized. In this work, we tackle this issue in a synthetic fashion, with a quantitative meta-analysis that allows an objective comparison of theoretical morphological models treated as entities. By analogy with empirical morphospace analyses of actual organisms, we performed a multivariate ordination of a representative sample of models, producing a metaspace of models in which patterns of similarity and difference are visualized. A phenetic tree was used to characterize the relationships between models. Four major groups have been identified, and their disparity analyzed. We suggest this typology as a useful starting point to identify a core set of fundamental principles and protocols for better interpretation of the plethora of current models and for more efficient construction of models in the future. This in turn can help in diversifying the scope of macroevolutionary, developmental, and bioenvironmental questions in theoretical morphology.

Persistent fossil taxa contravene paradigms of evolution: pervasive morphological change and taxic turnover. Comparative studies of taxic duration have often been approached from biogeographic, climatic, and ecological perspectives, with a focus on process. Here I use a morphological approach to study the pattern of longevity of a large family of marine living and fossil podocopid ostracodes, Trachyleberididae sensu lato. I test if geologically longer-lived genera are collectively morphologically more deviant from a group mean than their shorter-lived relatives by using both discrete morphological data and outline data. I discovered that long-lived genera are in general not significantly more or less morphologically deviant from the average morphology than their shorter-lived relatives. However, I found that contemporaneous subsets of long-lived trachyleberidids are often at least marginally significantly more deviant in discrete morphology than shorter-lived ones, especially in external morphology. No significant patterns of association between morphological deviation and durations in other subdivisions of the data emerged (i.e., whole data set, birth cohorts, groups of morphological characters, and outline data using both Fourier analysis and eigenshape analysis). This is in contrast to a previous finding that long-lived genera of crinoids within orders are often morphologically less deviant than their shorter-lived relatives than expected by chance.

Radiodonts (stem Euarthropoda) were ecologically diverse, but species generally displayed limited functional specialization of appendages along the body axis compared with crown group euarthropods. This is puzzling, because such functional specialization is considered to have been an important driver of euarthropod ecological diversification. One way to circumvent this constraint could have been the functional specialization of different parts of the frontal appendages, known to have been ecologically important in radiodonts. This hypothesis has yet to be tested explicitly. Here we redescribe the poorly known mid-Cambrian hurdiid radiodont Stanleycaris hirpex from the Burgess Shale (Stephen Formation) and quantitatively assess functional specialization of the frontal appendages of stem euarthropods. The appendages of Stanleycaris are composed of 14 podomeres, variously differentiated by their possession of pectinate endites, mono- to trifurcate medial gnathites, and outer spines. The oral cone is tetraradially organized and can be uniquely distinguished from those of other hurdiids by the presence of 28 rather than 32 smooth tridentate plates. Our phylogenetic analysis finds Stanleycaris in a grade of hurdiids retaining plesiomorphic raptorial appendicular functionality alongside derived adaptations for sweep feeding and large, bilaterally opposed gnathites. We conclude that the latter performed a masticatory function, convergent with gnathal structures like mandibles in various panarthropods. Taken together, Stanleycaris and similar hurdiids provide an extreme example of the evolution of division of labor within the appendage of a stem euarthropod and suggest that this innovation may have facilitated the functional transition, from raptorial to sweep feeding, at the origin of the hurdiid clade.

Factor analysis of morphometric data from fossil and recent Archaeogastropoda reveals that there has been a progressive reduction in the variety of archaeogastropod shell shapes through time. High and low spired genera have slowly disappeared while equidimensional forms, which have always been the most common, have become even more abundant. The taxonomic manifestation of this trend has long been appreciated and involves: 1) the extinction of most of the Pleurotomariacea, the Euomphalacea, Trochonematacea, and several other superfamilies, which collectively comprise most of the variation in shape among early archaeogastropods, and 2) the proliferation of the primarily equidimensional Trochacea and Neritacea. I suggest that this macroevolutionary trend may be explained by the general process I refer to as “stabilizing species selection.” More specifically, I suggest that the mechanism of stabilizing species selection in the case presented here may have been differential rates of origination.

I believe that the innovative concept of species selection has been burdened by the adherence of its advocates to the idea that differential extinction is its primary mechanism. This preference for differential extinction is rooted in our emphasis upon differential mortality in microevolution and is not based upon data of macroevolutionary change. I argue that differential origination may be as important as differential extinction in directing macroevolutionary change and that demographical and autecological properties of species may be its agents. Recognition of the importance of differential origination will have an important impact upon our concept of the relationship between adaptation and the evolutionary process.

Brinkmann (1929) argued that phyletic evolution in the Jurassic ammonite Kosmoceras is sufficiently linear that the amount of time missing at diastems can be estimated by the magnitude of morphologic jumps at those horizons. Brinkmann's unpublished raw data have been re-analyzed in order to evaluate the association between evolutionary jumps and physical breaks. All horizons in Brinkmann's 1400 centimeter stratigraphic section have been tested for morphological jumps. Up to 40 statistically significant morphological jumps exist in the section, the actual number depending on the sample interval used. A few of these correspond to the jumps found by Brinkmann and a few coincide with diastems. However, the correspondence between morphological jumps and physical breaks is not as consistent as was thought by Brinkmann. Furthermore, it is argued that the association between physical and biological discontinuities is not dependent on linear evolution but rather is the expected condition wherever phyletic evolution occurs. Due to the irregularity and unpredictability of evolutionary change in Kosmoceras, morphologic jumps cannot be used to assess the amount of time missing at diastems.

Although Phanerozoic increases in the global richness, local richness, and evenness of marine invertebrates are well documented, a common explanation for these patterns has been difficult to identify. Evidence is presented here from marine invertebrate communities that there is a Phanerozoic increase in the fundamental biodiversity number (θ), which describes diversity and relative abundance distributions in neutral ecological theory. If marine ecosystems behave according to the rules of Hubbell’s Neutral Theory of Biodiversity and Biogeography, the Phanerozoic increase in θ suggests three possible mechanisms for the parallel increases in global richness, local richness, and evenness: (1) an increase in the per-individual probability of speciation, (2) an increase in the area occupied by marine metacommunities, and (3) an increase in the density (per-area abundance) of marine organisms. Because speciation rates have declined over time and because there is no clear evidence for an increase in metacommunity area through the Phanerozoic, the most likely of these is an increase in the spatial density of marine invertebrates over the Phanerozoic, an interpretation supported by previous studies of fossil abundance. This, coupled with a Phanerozoic rise in body size, suggests that an increase in primary productivity through time is the primary cause of Phanerozoic increases in θ, global richness, local richness, local evenness, abundance, and body size.

The Main Karoo Basin of South Africa contains a near-continuous sequence of continental deposition spanning ~80 Myr from the mid-Permian to the Early Jurassic. The terrestrial vertebrates of this sequence provide a high-resolution stratigraphic record of regional origination and extinction, especially for the mid–late Permian. Until now, data have only been surveyed at coarse stratigraphic resolution using methods that are biased by nonuniform sampling rates, limiting our understanding of the dynamics of diversification through this important time period. Here, we apply robust methods (gap-filler and modified gap-filler rates) for the inference of patterns of species richness, origination rates, and extinction rates to a subset of 1321 reliably-identified fossil occurrences resolved to approximately 50 m stratigraphic intervals. This data set provides an approximate time resolution of 0.3–0.6 Myr and shows that extinction rates increased considerably in the upper 100 m of the mid-Permian Abrahamskraal Formation, corresponding to the latest part of the Tapinocephalus Assemblage Zone (AZ). Origination rates were only weakly elevated in the same interval and were not sufficient to compensate for these extinctions. Subsampled species richness estimates for the lower part of the overlying Teekloof Formation (corresponding to the Pristerognathus and Tropidostoma AZs) are low, showing that species richness remained low for at least 1.5–3 million years after the main extinction pulse. A high unevenness of the taxon abundance–frequency distribution, which is classically associated with trophically unstable postextinction faunas, in fact developed shortly before the acme of elevated extinction rates due to the appearance and proliferation of the dicynodont Diictodon. Our findings provide strong support for a Capitanian (“end-Guadalupian”) extinction event among terrestrial vertebrates and suggest that further high-resolution quantitative studies may help resolve the lack of consensus among paleobiologists regarding this event.

The spinal quotient (S.Q.) is an osteologically defined estimate of the enlargement of the spinal cord at limb levels over that at interlimb levels. It is an efficient predictor of limb use in living reptiles and birds and may be used to predict limb function in fossil vertebrates. Among living reptiles, this ratio of limb to interlimb innervation is greatest in arboreal genera, followed by terrestrial sprawlers, aquatic forms, and undulatory forms. Birds show a wide range of brachial S.Q. values that are roughly commensurate with flight ability. S.Q. values for the manipulative forelimbs of some dinosaurs fall well above those of locomotory limbs. Dinosaur hind-limb values are either well within ranges predicted by living reptiles and birds (most taxa), or highly inflated (stegosaurs, sauropods). This inflation may be the result of presence of a glycogen body similar to that of birds. In no case does the lumbosacral S.Q. support the presence of a “sacral brain.”

The Mesozoic–Cenozoic transition is generally seen as a pivotal time in the evolution of benthic marine assemblages but the details of the timing and drivers of these changes are poorly known. The Atlantic and Gulf Coastal Plains of the United States contain assemblages preserved as original aragonitic and calcitic material in unconsolidated sediments. This makes coastal plain assemblages ideally suited to paleoecological analyses. Data derived from bulk samples of the Coffee Formation (lower/middle Campanian: Mississippi) as well as published faunal lists from comparable samples of the Severn (Maastrichtian: Maryland), Providence (Maastrichtian: Georgia and Alabama), Stone City (Eocene: Texas), and Gosport (Eocene: Alabama) Formations are used to assess changes in taxonomic diversity and ecomorphological group (life habit and trophic group) composition through this time interval.

These analyses find a significant decrease in rarefied-sample species richness from the Campanian through the Eocene, but no change in evenness. With the notable exception of the Stone City Formation, increases in carnivore (neogastropod) richness and abundance occur before the Campanian. Epifaunal suspension-feeding species are a smaller proportion of the sample richness in Eocene samples than in Cretaceous samples. Decreased relative epifaunal suspension-feeder biomass but unchanged relative numbers of epifaunal suspension-feeder individuals suggests a relative decrease in epifaunal suspension-feeder size. Infaunal suspension feeders increase in richness and abundance through the interval. The proportion of drilled bivalves and gastropods does not change through the interval. Changes found in the structure of local shallow-shelf benthic assemblages from the Campanian through the Eocene are generally small relative to the variability between samples. Formation-level variation between assemblages is high relative to the magnitude of the temporal signal, emphasizing the need for investigators to include multiple formations per interval in tests of temporal trends.

The rapid and accurate taxonomic identification of fossils is of great significance in paleontology, biostratigraphy, and other fields. However, taxonomic identification is often labor-intensive and tedious, and the requisition of extensive prior knowledge about a taxonomic group also requires long-term training. Moreover, identification results are often inconsistent across researchers and communities. Accordingly, in this study, we used deep learning to support taxonomic identification. We used web crawlers to collect the Fossil Image Dataset (FID) via the Internet, obtaining 415,339 images belonging to 50 fossil clades. Then we trained three powerful convolutional neural networks on a high-performance workstation. The Inception-ResNet-v2 architecture achieved an average accuracy of 0.90 in the test dataset when transfer learning was applied. The clades of microfossils and vertebrate fossils exhibited the highest identification accuracies of 0.95 and 0.90, respectively. In contrast, clades of sponges, bryozoans, and trace fossils with various morphologies or with few samples in the dataset exhibited a performance below 0.80. Visual explanation methods further highlighted the discrepancies among different fossil clades and suggested similarities between the identifications made by machine classifiers and taxonomists. Collecting large paleontological datasets from various sources, such as the literature, digitization of dark data, citizen-science data, and public data from the Internet may further enhance deep learning methods and their adoption. Such developments will also possibly lead to image-based systematic taxonomy to be replaced by machine-aided classification in the future. Pioneering studies can include microfossils and some invertebrate fossils. To contribute to this development, we deployed our model on a server for public access at www.ai-fossil.com.

Principal components analysis (PCA) of 21 shell parameters (geometry, sculpture, aperture shape, and suture complexity) in 597 L. Devonian to L. Triassic ammonoid genera (spanning ~166 Myr) shows that eight basic morphotypes appeared within ~20 Myr of the first appearance of ammonoids. With one exception, these morphotypes persisted throughout the Paleozoic, occurring in ~75% of the ~5-Myr time bins used in this study. Morphotypes were not exclusive to particular lineages. Their persistence was not just a product of phylogenetic constraints or longevity, and multiple iterations of the same morphotypes occurred at different times and in different groups. Although mass extinction events severely condensed the range of morphologic variation and taxonomic diversity, the effects were short lived and most extinct morphotypes were usually iterated within 5 Myr. The most important effect of mass extinctions on ammonoid evolutionary history seems to have been their role in large scale taxonomic turnovers; they effectively eliminated previously dominant orders at the Frasnian/Famennian (F/F) (Agoniatitida), the Devonian/Mississippian (D/M) (Clymeniida), and the Permian/Triassic (P/T) (Goniatitida and Prolecanitida) extinctions. Survivors varied from two (P/T) to four (D/M) and five genera (F/F). These events generated sharp reductions in morphologic disparity at the D/M (58%) and at the P/T (59%), but there was a net increase at the F/F (38%). There was no obvious survival bias for particular morphotypes, but 64% are interpreted to have been Nautilus-like nektobenthic. The recurrence of particular combinations of morphology and their strong independence of phylogeny are strong arguments for functional constraint. Intervals between mass extinctions seem to have been relatively static in terms of morphotype numbers, in contrast to numbers of genera. Significant decreases in genus diversity (54%) and morphologic disparity (33%) commenced in the mid-Permian (Wordian/Capitanian boundary), well before the final P/T event.

Fourier analysis of percent extinction data for genera of fossil marine animals during the past 260 million years indicates the existence of a 26 m.y. extinction cycle. The first three harmonics, which account for 51.7 percent of the sum of squares, show the major extinction trends in the late Paleozoic, Mesozoic and Cenozoic. The first ten harmonics, which account for 89.5 percent of the sum of squares, are aligned with eight extinction peaks. The tenth harmonic, with a period of 26 million years, has an amplitude of 6.5 percent and accounts for 12.7 percent of the sum of squares.

Several different methods are used to test the validity of the apparent 26 m.y. extinction cycle. Analysis of variance indicated that the 26 m.y. extinction cycle is statistically significant at the 95 percent level. When the 260 m.y. sample interval is divided into equal 130 m.y. segments, the phases and amplitudes for the 26 m.y. cycle were almost equal within the two independent segments. When the 260 m.y. interval is divided into 10 shorter time segments of different lengths, the 26 m.y. cycle is represented in each time segment.

A series of experiments is run to test the influence of the Late Permian and Cretaceous peaks on the observed 26 m.y. cycle. When all peaks are smoothed out except for the two major peaks, a pseudo-cycle with a period of 26 m.y. is generated that accounts for 3.0 percent of the sum of squares. When the Permian and Cretaceous peaks are reduced to about half their observed height, the sum of squares accounted for by the 26 m.y. cycle is reduced from 12.7 to 11.0 percent. When the number of extinctions within each stage is used in the harmonic analysis in place of extinction percent, the 26 m.y. cycle accounts for 19.3 percent of the sum of squares.

Therefore, the evidence from harmonic analysis of fossil marine animals points toward a distinct and persistent 26 m.y. cycle in mass extinctions in the late Paleozoic, Mesozoic and Cenozoic.

Carnivore-rich fossil sites are uncommon in the fossil record and, accordingly, provide valuable opportunities to study predators from vantages that are rarely applied to ancient faunas. Through stable isotopes of carbon and a Bayesian mixing model, we analyze time-successive (nearly contemporaneous), late Miocene carnivoran populations from two fossil sites (Batallones-1 and Batallones-3) from central Spain. Stable isotopes of carbon in tooth enamel provide a reliable and direct methodology to track ancient diets. These two carnivoran-dominated fossil sites display differences in the composition and abundance of the carnivoran species, with some species present at both sites and some present only at one site. This disparity has been interpreted as the consequence of habitat differences between Batallones-1, the older site, and Batallones-3, the younger site. However, carbon isotope values of carnivore and herbivore tooth enamel suggest a common habitat of C3 woodland originally present at both sites. The differences in the carnivoran faunas rather may be the consequence of the dynamics of species entrance and exit from the Madrid Basin during the time elapsed between Batallones-1 and Batallones-3 and changes in population densities due to biotic factors. We infer higher levels of interspecific competition in Batallones-3 than in Batallones-1 because of the larger number of similar-sized, sympatric predators; the clear overlap in their δ13C values (except for the amphicyonid Magericyon anceps); and similarity of their preferred prey: the hipparionine horses. Finally, carbon stable isotopic composition of Indarctos arctoides teeth implies that this ursid was a carnivorous omnivore rather than a herbivorous omnivore. This work demonstrates the insights that stable isotopes can provide in characterizing the feeding ecology and trophic interactions of ancient carnivoran taxa.

Several important questions in evolutionary biology and paleobiology involve sources of variation in extinction rates. In all cases of which we are aware, extinction rates have been estimated from data in which the probability that an observation (e.g., a fossil taxon) will occur is related both to extinction rates and to what we term encounter probabilities. Any statistical method for analyzing fossil data should at a minimum permit separate inferences on these two components. We develop a method for estimating taxonomic extinction rates from stratigraphic range data and for testing hypotheses about variability in these rates. We use this method to estimate extinction rates and to test the hypothesis of constant extinction rates for several sets of stratigraphic range data. The results of our tests support the hypothesis that extinction rates varied over the geologic time periods examined. We also present a test that can be used to identify periods of high or low extinction probabilities and provide an example using Phanerozoic invertebrate data. Extinction rates should be analyzed using stochastic models, in which it is recognized that stratigraphic samples are random variates and that sampling is imperfect.

In marsupial mammals and their extinct relatives—collectively, metatherians—only the last premolar is replaced, but the timing of dental eruption is variable within the group. Our knowledge of fossils metatherians is limited, but is critical to understanding several aspects of the evolution and morphological diversification of this clade. We analyzed the sequence of eruption of 76 specimens of metatherians, including Sparassodonta, an extinct clade of specialized carnivores from South America. In Sparassodonta (1) the P3/p3 erupt simultaneously, in common with some didelphids (in other didelphids, p3 erupts before P3, whereas in the remaining didelphids, some peramelids, one caenolestid, and Pucadelphys this order is reversed); (2) the upper and lower molars at the same locus erupt more in synchrony than in other carnivorous metatherians in which the lower molars clearly precede the upper equivalents; (3) the upper canine in thylacosmilids and proborhyaenids is hypselodont; (4) species with similar molar morphologies have different morphologies of the deciduous premolars, suggesting diverse diets among the juveniles of different taxa; (5) deciduous teeth are functional for a long period of time, with thylacosmilids even retaining a functional DP3 in the permanent dentition. The retention of the DP3 and the hypertrophied and hypselodont upper canine of thylacosmilids represent clear heterochronic shifts. Specializations in the timing of dental eruption and in the deciduous tooth shape of sparassodonts are evolutionary mechanisms that circumvent constraints imposed by the metatherian replacement pattern and increase morphological disparity during ontogeny.

Analysis of relative changes in plate position during ontogeny was made on a number of Cenozoic spatangoid echinoids: species of Breynia, Lovenia, Protenaster and Echinocardium. Contrary to the generally held view that adjacent ambulacral and interambulacral columns in echinoids always remain in a fixed relative position during ontogeny, many spatangoids show great fluidity in plate growth, with adjacent columns ‘sliding past’ one another during ontogeny. Furthermore, in many genera plates from one column may undergo strong lateral growth and bisect pairs of plates in adjacent rows. This phenomenon of relative plate movement, both meridional and equatorial, is herein termed ‘plate translocation’. It is considered to occur by localized resorption and redeposition in a narrow zone along adjacent plate boundaries. Plate translocation has been of considerable phylogenetic significance to spatangoids and, combined with an increase in differential allometries between plates, has been one of the most important factors in the evolution of the Spatangoida. Furthermore, the initiation of plate translocation in the apical system in certain Jurassic echinoids may have been the trigger for the migration of the periproct out of the apical system, and a major factor in the evolution of irregular echinoids. Heterochrony in plate growth has been critical in controlling the course of evolution in many lineages of spatangoid echinoids.

We compare refined data sets for Atlantic benthic foraminiferal oxygen isotope ratios and for North American mammalian diversity, faunal turnover, and body mass distributions. Each data set spans the late Paleocene through Pleistocene and has temporal resolution of 1.0 m.y.; the mammal data are restricted to western North America. We use the isotope data to compute five separate time series: oxygen isotope ratios at the midpoint of each 1.0-m.y. bin; changes in these ratios across bins; absolute values of these changes (= isotopic volatility); standard deviations of multiple isotope measurements within each bin; and standard deviations that have been detrended and corrected for serial correlation. For the mammals, we compute 12 different variables: standing diversity at the start of each bin; per-lineage origination and extinction rates; total turnover; net diversification; the absolute value of net diversification (= diversification volatility); change in proportional representation of major orders, as measured by a simple index and by a G-statistic; and the mean, standard deviation, skewness, and kurtosis of body mass. Simple and liberal statistical analyses fail to show any consistent relationship between any two isotope and mammalian time series, other than some unavoidable correlations between a few untransformed, highly autocorrelated time series like the raw isotope and mean body mass curves. Standard methods of detrending and differencing remove these correlations. Some of the major climate shifts indicated by oxygen isotope records do correspond to major ecological and evolutionary transitions in the mammalian biota, but the nature of these correspondences is unpredictable, and several other such transitions occur at times of relatively little global climate change. We conclude that given currently available climate records, we cannot show that the impact of climate change on the broad patterns of mammalian evolution involves linear forcings; instead, we see only the relatively unpredictable effects of a few major events. Over the scale of the whole Cenozoic, intrinsic, biotic factors like logistic diversity dynamics and within-lineage evolutionary trends seem to be far more important.

The history of diversity, origination and extinction of Cambro-Ordovician nautiloid cephalopods is explored to determine if differences in evolutionary rates between nautiloid orders are sufficient to document significantly high or low rates of evolutionary turnover (taxotely of Raup and Marshall 1980). The stratigraphic ranges of 425 nautiloid genera are analyzed for this purpose.

Evolutionary rates for five of the seven time intervals analyzed fall within frequency distributions of rates which are thought to be characteristic for a given time interval (horotelic distribution of Simpson 1944). Sufficient heterogeneity is present among extinction rates of Arenigian orders and origination rates of Caradocian orders to reject the null hypotheses of horotely in favor of taxotely. The orders Ellesmerocerida and Tarphycerida, each with a significantly high rate of extinction (P ≥ 0.99), and the Actinocerida, with a significantly low rate of extinction (P ≥ 0.99), were responsible for taxotely during the Arenigian. The Oncocerida and Discosorida, each with a significantly high rate of origination (P ≥ 0.99), were responsible for taxotely during the Caradocian. In each case, taxotely is attributable to the influence of North American endemics. This effect is believed to be more the result of real biological aspects of nautiloid evolution than an artifact of the fossil record.

In this paper we reconstruct the phylogeny of a clade of pachypleurosaurs (Reptilia: Sauropterygia) occurring in Triassic-age deposits in the Monte San Giorgio region, Switzerland. We also present the phylogeny of this clade as a case study for two paradigms of phylogeny reconstruction: cladistics and phenetic/stratigraphic methods. While this dichotomy is not held rigidly by all workers, its advancement by cladists leads us to retain it initially for rhetorical purposes. We review the philosophical bases of species, species concepts, and speciation, as well as cladograms and phylogenies, before introducing the experimental system.

Data are presented from cladistic analyses, phenetic analyses, and stratigraphic information. Phylogeny of the clade is interpreted from both paradigms, and the interpretations are found to be inconsistent. Resolution of the phylogeny rests on the emphasis of one type of data over another. An interpretation of cladogenesis within the genus Neusticosaurus entails rejection of suggestive phenetic and stratigraphic data, whereas an anagenetic interpretation entails reversal of autapomorphies in ancestral taxa. Anagenesis is deemed to be the more probable interpretation, based on the strength of the stratigraphic and phenetic data relative to the character data. Implications of the test case results for phylogeny reconstruction in general are discussed, ending with a call for pluralism in approach.