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Many fossil assemblages are time-averaged, with multiple generations of organisms mixed into a single stratigraphic horizon. A time-averaged sample of a taxon should be more variable than a single-generation sample if enough morphologic change occurred during the interval of time-averaging. Time-averaging may also alter correlations between morphologic variables and obscure allometric relationships in an evolving population. To investigate these issues, we estimated the variability of six modern, single-generation samples of the bivalve Mercenaria campechiensis using Procrustes analysis and compared them with several time-averaged Pleistocene samples of M. campechiensis and M. permagna. Both the modern and the fossil samples ranged in variability, but these ranges were virtually identical. Morphology was quite stable over the hundreds to many thousands of years that passed as the assemblages accumulated, and the variabilities of the fossil samples could be used to estimate single-generation variability. At one fossil locality, the environment and paleocommunity changed partway through the collection interval; the morphology of Mercenaria appears stable above and below the transition but changes across it. This change is similar in magnitude to the differences between geographically separated modern populations, whereas temporal variation within single environmental settings is distinctly less than geographic variation. Analytical time-averaging (the mixing of fossils from different horizons) between paleocommunities increased variability slightly (but not significantly) above that found in living populations. While its constituent populations appear stable on millennial timescales, M. campechiensis has been evolutionarily static since at least the early to middle Pleistocene.
During the Ordovician Radiation, domination of benthic marine communities shifted away from trilobites, toward articulated brachiopods, and, to a lesser degree, toward bivalves and gastropods. Here, models are formulated that mathematically represent alternative hypothesized causes of this transition. These include models in which per-genus origination or extinction probabilities were constrained to be (1) constant, (2) diversity-dependent, (3) productivity-dependent, or (4) jointly dependent on productivity and diversity. Using a method adapted from capture-mark-recapture (CMR) population studies, we estimate origination, extinction, and sampling probabilities jointly in order to avoid confounding patterns in turnover probabilities with temporal variation in the quality of the fossil record. Using Akaike's Information Criterion (AIC), we assessed the fit of the alternative causal models relative to one another, and relative to a noncausal “phenomenological” alternative that placed no constraints on the pattern of temporal variation in origination or extinction. There were differences among taxa in the relative fit of these models, suggesting that the effects of productivity and diversity varied among higher taxa. In the aggregate, however, there was strong support for diversity-dependent origination. For extinction, poor fit of the alternative causal models suggests that we lack a good explanation for extinction patterns. These analyses support the hypothesis that diversity-dependent origination, particularly in trilobites, contributed to the Ordovician faunal transitions. By contrast, the effects of increased productivity, if indeed they were large enough to influence global diversification patterns, did not proceed in the hypothesized manner.
The characters and body parts of organisms are shaped by mechanical forces at two temporal scales. At the ontogenetic scale, the relevant forces are those of every day, exerted by muscles, other metabolism-powered processes, and normal interactions between the body and the external environment. At the phylogenetic scale, forces are strong enough to kill some individuals or to cause reproductive failure. These forces act more intermittently.
I explore these ideas by examining the characters of molluscan shells, which grow by the addition of skeletal material along the rim of the open end of a hollow, conical tube that is closed at its narrow (apical) end. In the idealized case of a null shell, the skeleton is a right circular cone, in which the magnitude and direction of growth are the same at each point along the rim. The rate of expansion of the cone is determined by the shell-builder's metabolism. Real shell-builders are exposed to, and themselves exert, forces that affect shell shape. These forces are generated by contact between the shell-secreting mantle margin and the substratum, by local or temporary deformations of the mantle margin imposed by other parts of the body and previously formed parts of the shell, and by contraction of muscles that connect the soft tissues to the inner shell surface. Early mollusks whose shells more or less resemble the null shell were slow-moving, epifaunal animals that clamped the shell against the substratum. Evolutionary increases in metabolic rate, associated with greater mobility and faster growth, made some ontogenetically important forces stronger and introduced new forces. As a result, the range of available phenotypes expanded. Refinements in genetic regulation of form, perhaps including an increase in the number of semiautonomous regulatory regions, further added to the specification and range of variation of characters that were subject either to evolutionary conservation or to natural selection. For example, the mantle margin in plesiomorphic gastropods appears to comprise one such region, which produces a growing shell margin in the form of a logarithmic spiral; in more-derived gastropods, the mantle margin may comprise two or more regions, which together produce a growing shell margin that departs strikingly from the logarithmic form of the outer shell lip.
The morphospace occupied by accretionary shells can be described by (1) the number of semiautonomous developmental modules, (2) selective regimes observable as phenotypic adaptive evolution, and (3) metabolic rate. The perspective outlined here implies that shells initially occupied a limited morphospace encompassing one or two modules, adaptation as an epifaunal clamping animal, and slow growth (low expansion rates) and metabolism. Further compartmentalization, together with increased metabolic rates in ecologically dominant taxa, caused the morphospace to expand both in the number of independent descriptors and in the range of values that each parameter spans. These trends in morphospace may characterize all major multicellular clades.
The precise knowledge of the number and nature of the species belonging to a fossil assemblage as well as of the structure of each species (e.g., age, sex) is of great importance in paleontology. Mixture analysis based on the method of maximum likelihood is a modern statistical technique that concerns the problem of samples consisting of several components, the composition of which is not known. Nonparametric bootstrap and jackknife techniques are used to calculate a confidence interval for each estimated parameter (prior probability, mean, standard deviation) of each group. The bootstrap method is also used to evaluate mathematically how many groups are present in a sample. Experimental density smoothing using the kernel method appears to be a better solution than the use of histograms for the estimation of a distribution. This paper presents some basic concepts and procedures and discusses some preliminary results concerning sex ratios and mortality profile assessments using bones and tooth metric data of small (Ovis antiqua) and large (Bos primigenius) bovines from European Pleistocene sites.
Four vascular plant lineages, the ferns, sphenopsids, progymnosperms, and seed plants, evolved laminated leaves in the Paleozoic. A principal coordinate analysis of 641 leaf species from North American and European floras ranging in age from Middle Devonian through the end of the Permian shows that the clades followed parallel trajectories of evolution: each clade exhibits rapid radiation of leaf morphologies from simple (and similar) forms in the Late Devonian/Early Carboniferous to diverse, differentiated leaf forms, with strong constraint on further diversification beginning in the mid Carboniferous. Similar morphospace trajectories have been documented in studies of morphological evolution in animals; however, plant fossils present unique opportunities for understanding the developmental processes that underlie such patterns. Detailed comparison of venation in Paleozoic leaves with that of modern leaves for which developmental mechanisms are known suggests developmental interpretations for the origination and early evolution of leaves. The parallel evolution of a marginal meristem by the modification of developmental mechanisms available in the common ancestor of all groups resulted in the pattern of leaf evolution repeated by each clade. Early steps of leaf evolution were followed by constraint on further diversification as the possible elaborations of marginal growth were exhausted. Hypotheses of development in Paleozoic leaves can be tested by the study of living plants with analogous leaf morphologies.
The dichotomy between short-necked, large-headed “pliosaurs” and long-necked, small-headed “plesiosaurs” has formed the basis of plesiosaur taxonomy for over one hundred years. Recent work has cast doubt on the taxonomic validity of this dichotomy, suggesting that the pliosaur morphotype may have evolved independently in more than one clade. This paper quantifies the variation in body proportion in the clade Plesiosauria using principal component analysis and demonstrates that the traditional plesiosaur/pliosaur dichotomy is an oversimplified view of the range of morphologies present in the group. The topology of the clade is mapped into the morphospace, demonstrating that the pliosaur morphotype evolved three times from two different regions of morphospace. Both the range of body morphologies displayed by plesiosaurs and the evolutionary history of those morphologies, are more complex than previously supposed.
Finite-element analysis of circular septum models indicates that (1) anticlastic fluting weakened the last septa of the same radius of curvature by a factor of about 2.5 relative to the tensile stresses in a sphere of nacre, (2) septa with ammonitic sutures were stronger than those with goniatitic sutures of the same thickness, and (3) septa with more “complex” ammonitic sutures were stronger at the edge between lobes and saddles than “simple” ones. These results contradict recent claims that ammonoid septa became weaker as sutural complexity increased from goniatitic through ammonitic, so that the most complex sutures were limited to the shallowest habitats. The smaller marginal flutes of complex septa were relatively strong, allowing them to be thinner than the central septum and still act as elastic wall supports. Many Mesozoic ammonoids with highly sinuous sutures occurred in deep epeiric and open-ocean habitats, whereas it is those with secondarily reduced, ceratitic sutures that were typically associated with restricted shallow basins.
Exceptional examples of planispiral ammonites that were infested by epizoans during life display alterations of their normal coiling. Most commonly, the epizoan(s) settled on the venter of the ammonite and constituted an obstacle for the whorl tube grown one whorl later; this caused lateral deviation of the whorl tube and tilting of the ammonoid because of changes in the hydrostatic condition; from here on, the whorl tube periodically crossed the venter of the preceding whorl, thereby producing a zigzag coiling pattern. Some epizoans, which were particularly centered on the midventer, provoked detachment between whorls. In a few cases, lateral placement of the epizoan did not directly obstruct the normal growth path of the ammonite but induced a trochospiral coiling pattern. Both the zigzag and the trochospiral pattern were created when the ammonite tried to maintain the growth direction within the vertical plane at the same time as whorls remained in contact along a differentiated dorsal epithelium. The aperture reacted to changes in growth direction, to maintain also a permanent angle with the vertical direction. Growth direction, then, was a major morphogenetic parameter in ammonites, because it contained the necessary instructions for correct shell coiling. The model based on the observation of fabricational defects has been tested by a theoretical model by which the different situations so far observed are simulated and in which the parameters are the morphogenetic instructions inferred to have been present in the biological system.
We investigated the petrography and biochemistry of varved sediments from a Pleistocene mass occurrence of fossil vertebrates in the lake basin of Neumark-Nord (Sachsen-Anhalt, Germany). The carbonate portions of the varves appear to be cyanobacterial layers that have been decomposed by benthic bacteria. The biochemical results obtained by absorption spectroscopy and RP-HPLC with UV-detection show that pigments, and probably toxins, characteristic for cyanobacteria are preserved in the sediment. The results of this study indicate the presence in the lake of large amounts of toxic cyanobacteria that probably occurred in seasonal blooms. It seems likely that these toxic cyanobacterial blooms caused the mass death of the large mammals preserved at the Neumark-Nord locality. This model may explain comparable lithologies and vertebrate occurrences in other Tertiary lake sediments such as the Messel oil shale.
The Burgess Shale arthropod Leanchoilia superlataWalcott 1912, commonly preserves a three-dimensional axial structure generally interpreted as gut contents. Thin-section examination shows this instead to be phosphatized biserially repeated midgut glands, including exceptional preservation of subcellular features. The preferential mineralization of these structures is related to their unusually high chemical reactivity and probably to an internal source of phosphate. Sub-millimetric lineations previously interpreted as annular musculature are in fact planar, sometimes radially arranged, subdivisions of these glands. Ventral rows of isolated phosphate patches appear to represent the same tissue.
In extant arthropods, extensively developed midgut glands are related to a rich but infrequent diet with a primary function in storage. Their conspicuous occurrence in unambiguous fossil predators such as Sidneyia and Laggania (Anomalocaris) suggests they served a similar role in the Cambrian; by extension, their conspicuous occurrence in Leanchoilia suggests it was a predator or scavenger.
Phosphatized midguts with a structure essentially indistinguishable from that of Leanchoilia are also found in Burgess Shale Odaraia, Canadaspis, Perspicaris, Sidneyia, Anomalocaris, and Opabinia. All are characterized by a distinctive sub-millimetric arrangement of planar elements that is not found in extant arthropods or trilobites, suggesting they diverged before the last common ancestor of extant forms; i.e., they represent stem-group arthropods.
Three-dimensionally preserved guts are widely preserved in the Lower Cambrian Chengjiang biota but, unlike those in the Burgess Shale, appear to be filled with sediment. Although generally interpreted as evidence of deposit feeding, the form of these structures points to early permineralization of (sediment-free) midgut glands that were subsequently altered to clay minerals. There is no evidence of deposit feeding in the Chengjiang; indeed, there is a case to be made for deposit feeding not being generally exploited generally until after the Cambrian.
Fossils with three-dimensionally preserved axes from the Lower Cambrian Sirius Passet biota have been interpreted as lobopodians; however, most of the putative lobopodian features find alternative interpretations as aspects of Leanchoilia-type midgut glands. Although Kerygmachela is reliably identified as a stem-group arthropod, its phylogenetic position remains unresolved owing to the non-preservation of critical external features and to the plesiomorphic nature of its Leanchoilia-type midgut.