Experiments show that diductor muscles of articulate brachiopods are normally relaxed, contracting only when the valves gape. Consequently, dead brachiopods rarely gape.

In quantitative paleobiology, one common format for image-derived information is the closed two-dimensional outline curve. Before the end of the last century a great variety of morphometric tools were on offer to deal with this data type, including diverse analyses of distance from a central point, “eigenshape analysis” of a carefully normalized tangent angle function, and Procrustes analysis of configurations of sliding semilandmarks. For the special case of outline forms that are nominally bilaterally symmetric, this paper offers a hybrid approach that fuses the Procrustes toolkit with a variant of a much older method, analysis of radial distance according to angles out of a center. The synthesis represents each outline as a regularly resampled data set of radially aligned shape coordinates analogous to bilateral Procrustes shape coordinates. When the center is iteratively located in one particular way, this representation has the same Procrustes distance statistics as a conventional radial representation, while the formalism of shape coordinates permits symmetrization, localizability, and thin-plate spline visualizations. We explain the method and sketch some of its advantages and limitations in the course of an example involving 99 Baculites inornatus outlines spanning 90 m of deposits from a high-precision measured section in the Campanian-age Rosario Formation cropping out at Punta San Jose, Baja California. This sequence shows substantial changes of pole curvature over time, changes that may involve a punctuation event regarding hydrostatics, hydrodynamics, or defense against predators. A concluding comment deals with implications of the developments here for Baculites microevolution and for morphometrics.

Interpreting the impact of climate change on vertebrates in the fossil record can be complicated by the effects of potential biotic drivers on morphological patterns observed in taxa. One promising area where this impact can be assessed is a high-resolution terrestrial record from the Bighorn Basin, Wyoming, that corresponds to the Paleocene–Eocene thermal maximum (PETM), a geologically rapid (~170 kyr) interval of sustained temperature and aridity shifts about 56 Ma. The PETM has been extensively studied, but different lines of research have not yet been brought together to compare the timing of shifts in abiotic drivers that include temperature and aridity proxies and those of biotic drivers, measured through changes in floral and faunal assemblages, to the timing of morphological change within mammalian species lineages. We used a suite of morphometric tools to document morphological changes in molar crown morphology of three lineages of stem erinaceid eulipotyphlans. We then compared the timing of morphological change to that of both abiotic and other biotic records through the PETM. In all three species lineages, we failed to recover any significant changes in tooth crown shape or size within the PETM. These results contrast with those documented previously for lineages of medium-sized mammals, which show significant dwarfing within the PETM. Our results suggest that biotic drivers such as shifts in community composition may have also played an important role in shaping species-level patterns during this dynamic interval in Earth history.

Accurate interpretation of origination and extinction of fossil species is crucial to answering a variety of questions in paleontology. Fossil datums, the observed age of first or last occurrences, are subject to sampling error as a result of preservation and low abundances near range endpoints. This sampling error can cause local range offset, an age difference between the observed first or last occurrence of a species and its true origination or extinction. Here, we develop and test a new technique, the Probable Datum Method (PDM), that can be used to assess the extent of local range offset for nannofossil species. The PDM estimates the original abundance of a taxon and its probable true age of first or last occurrence. The PDM uses a model in which original abundance is related to count abundance through preservation and the counting process. This model is empirically parameterized, including an experimental determination of false positive and error rates of a nannofossil count. The model is simulated then inverted to estimate likely original abundance and true datum age from count abundance data. We first test the PDM in a positive control experiment with known parameter values. This experiment shows that the PDM is robust and returns known values accurately. Next we apply the method to the origination of nannoplankton after the Cretaceous/Paleogene boundary to test whether first occurrences were synchronous between widely spaced locations. The PDM results suggest that observed diachrony of K/Pg originations cannot be explained by the effects of local range offset; rather, in some cases they indicate truly diachronous first occurrences between localities. Although the technique was developed to analyze nannoplankton ranges, the statistical nature of the PDM, its experimentally derived parameters, and its parsimonious nature should make it applicable to many micropaleontological studies that interpret patterns of origination and extinction.

Examination of the neanic apparatuses of known populations of Nephrolepidina praemarginata, N. morgani, and N. tournoueri reveals that the equatorial chamberlets are arranged in spirals, along the direction of connection of the oblique stolons, giving the optical effect of intersecting curves. In N. praemarginata commonly 34 left- and right-oriented primary spirals occur from the first annulus to the periphery, 21 secondary spirals from the third to fifth annulus, 13 ternary spirals from the fifth to eighth annulus, following the Fibonacci sequence.

The number of the spirals increases in larger specimens and in more embracing morphotypes, and especially in trybliolepidine specimens; the secondary and ternary spirals from the investigated N. praemarginata to N. tournoueri populations tend to start from more distal annuli. An interpretative model of the spiral growth of Nephrolepidina is attempted.

The angle formed by the basal annular stolon and distal oblique stolon in equatorial chamberlets ranges from 122° in N. praemarginata to mean values close to the golden angle (137.5°) in N. tournoueri.

The increase in the Fibonacci number of spirals during the evolution of the lineage, along with the disposition of the stolons between contiguous equatorial chamberlets, provides new evidence of evolutionary selection for specimens with optimally packed chamberlets.

Natural selection favors individuals with the most regular growth, which fills the equatorial space more efficiently, thus allowing these individuals to reach the adult stage faster. We refer to this new type of selection as “golden selection.”

To the student of evolution, Malmgren and Kennett's (1981) detailed study of the Globorotalia inflata lineage demonstrates the continuity of the stratigraphic record and the abundance of planktic microfossils in deep-sea sediments. Their work recalls the prediction (Prothero and Lazarus 1980) that planktic microfossils might serve as the laboratory animals of evolutionary studies and indicates the potential value of fossils in determining ancestry. However, for theorists to judge the evolutionary import of Malmgren and Kennett's high-resolution morphometric study, broader consideration of lineage relationships is needed. They portray the G. inflata lineage as a set of five successional taxa in simple ancestor-descendant relationship. Trends, established morphometrically and claimed to be “one of the clearest examples of gradualism in any group of fossils” (Malmgren and Kennett 1981, p. 236), are shown to progressively transform a compressed shell with peripheral keel (G. conoidea) into a globose shell (G. inflata) without a keel. While the succession of taxa given by Malmgren and Kennett has been supported in other studies of western South Pacific sequences (e.g., McInnes 1965; Walters 1965; Kennett 1966; 1973; Jenkins 1971; Scott 1979; Hornibrook 1982), it is not established that the order indicates simple phyletic transformation without branching, as Malmgren and Kennett imply. Crucial evidence concerns the history and status of keeled populations which resemble the ancestral G. conoidea but occur in the Pliocene, associated with advanced members of the G. inflata lineage. These populations are not mentioned by Malmgren and Kennett, yet, if they are also descendants of G. conoidea, their occurrence strongly suggests that lineage divergence occurred. I wish to review evidence in support of this view.

The typical marine animal has increased in biovolume by more than two orders of magnitude since the beginning of the Cambrian, but the causes of this trend remain unknown. We test the hypothesis that the efficiency of intra-organism oxygen delivery is a major constraint on body-size evolution in marine animals. To test this hypothesis, we compiled a dataset comprising 13,723 marine animal genera spanning the Phanerozoic. We coded each genus according to its respiratory medium, circulatory anatomy, and feeding mode. In extant genera, we find that respiratory medium and circulatory anatomy explain more of the difference in size than feeding modes. Likewise, we find that most of the Phanerozoic increase in mean biovolume is accounted for by size increase in taxa that accomplish oxygen delivery through closed circulatory systems. During the Cambrian, water-breathing animals with closed circulatory systems were smaller, on average, than contemporaries with open circulatory systems. However, genera with closed circulatory systems superseded in size genera with open circulatory systems by the Middle Ordovician, as part of their Phanerozoic-long trend of increasing size. In a regression analysis, respiratory and circulatory anatomy explain far more size variation in the living fauna than do feeding modes, even after accounting for taxonomic affinity at the class level. These findings suggest that ecological and environmental drivers of the Phanerozoic increase in the mean size of marine animals operated within strong, anatomically determined constraints.

The fossil record is notoriously imperfect and biased in representation, hindering our ability to place fossil specimens into an evolutionary context. For groups with fossil records mostly consisting of disarticulated parts (e.g., vertebrates, echinoderms, plants), the limited morphological information preserved sparks concerns about whether fossils retain reliable evidence of phylogenetic relationships and lends uncertainty to analyses of diversification, paleobiogeography, and biostratigraphy in Earth's history. To address whether a fragmentary past can be trusted, we need to assess whether incompleteness affects the quality of phylogenetic information contained in fossil data. Herein, we characterize skeletal incompleteness bias in a large dataset (6585 specimens; 14,417 skeletal elements) of fossil squamates (lizards, snakes, amphisbaenians, and mosasaurs). We show that jaws + palatal bones, vertebrae, and ribs appear more frequently in the fossil record than other parts of the skeleton. This incomplete anatomical representation in the fossil record is biased against regions of the skeleton that contain the majority of morphological phylogenetic characters used to assess squamate evolutionary relationships. Despite this bias, parsimony- and model-based comparative analyses indicate that the most frequently occurring parts of the skeleton in the fossil record retain similar levels of phylogenetic signal as parts of the skeleton that are rarer. These results demonstrate that the biased squamate fossil record contains reliable phylogenetic information and support our ability to place incomplete fossils in the tree of life.

Discriminant function analysis is used to examine the role of within-species differentiation in macroevolution of four Miocene bivalve genera. According to the punctuational model, macroevolution is decoupled from microevolution; within-species trends may be unrelated to between-species differences. The reductionist alternative suggests that intraspecific (primarily geographic) differentiation is the basis of macroevolution. If macroevolution depends on within-species differentiation, discriminant function analysis should separate interspecific and intraspecific populations of a genus using the same set of characters.

Chesapecten macroevolution apparently involved within-species differentiation. Shell length, height, and posterior auricle length differentiate geographic populations within species as well as interspecific populations. Results for Astarte, Anadara, and Eucrassatella are more consistent with the punctuational model. Characters which readily separate species of a genus are much less successful at differentiating intraspecific populations. Rank correlation techniques indicate that, for all three genera, the order of importance of characters in interspecific discrimination is not significantly related to their order in intraspecific discrimination. Decoupling of macroevolution from microevolution is implied for these taxa.

Insects in the fossil record are generally preserved in lacustrine shales or in amber. For those in lacustrine shales, preservation is usually via keroginization or mineralization. Given the extended period of microbial decay required to generate ions for mineralization, there is a predicted inherent bias toward lower preservation quality for this pathway by most taphonomic indices compared with keroginization. This study tests this hypothesis by comparing multiple measures of preservation quality between sites with similar sedimentology in the Eocene Green River Formation of Colorado. Here, insects are either mineralized in iron oxides (likely after pyrite) at the Paleoburn site or keroginized at the Anvil Points site.

Generally, the prediction that keroginization preserves soft-bodied fossils with higher preservational quality than mineralization is affirmed, but with some caveats. Beetles, known for their robust cuticles, are proportionately more abundant at the Paleoburn site, but eight of the nine orders recorded are shared between sites. As predicted, insects show lower preservation fidelity at the Paleoburn site, but they also show higher degrees of disarticulation. This second bias should be acquired primarily during the biostratinomy stage, and not early diagenesis. Nonetheless, higher-energy biostratinomic conditions may be compatible with taphonomic conditions that promote mineralization over keroginization.

Comparing the inherent taphonomic bias of different preservation pathways is often difficult, since fossil deposits may be preserved millions of years or thousands of kilometers apart. By studying two different preservation pathways of insects within the same formation, we can affirm that keroginization does indeed preserve recalcitrant organic matter with higher quality than pyritization or iron-oxide mineralization. Additionally, some guidelines can be proposed concerning the body parts and taxa that can be compared, and for what purpose, when contrasting mineralized and keroginized soft-bodied deposits.

The paleoecologist has been hampered in his studies of ancient communities by an inability to obtain reasonable estimates of standing crops and diversities. Finding the diversity of an ancient community is no simple matter, because a fossil assemblage is a mixture of living and recently dead individuals from that community. Considerable numbers of each group may be missing. The relationship of the elements in a fossil population for an individual species can be expressed as follows: F = Fl + Fd, where F is the average number of fossil individuals collected per unit area, Fl is the part of the sample produced by the burial of live individuals, and Fd is the part produced by the individuals already dead at the time of burial. Fl is a product of the standing crop S (number of individuals per unit area), multiplied by the probability of failure P (proportion of the standing crop which can be expected to fail to escape anastrophic burial), or Fl = (S) (P). Fd is the product of the death rate D (number of individuals which die per age-class time unit) multiplied by the residence time R (average length of time expressed in age-class time units that the remains of an individual will persist after death), or Fd = (D) (R). The probability of failure, residence time, and death rate can all be estimated from modern communities; the death rate can also be estimated from the fossils. Thus, the standing crop of an individual species can be expressed as follows:

This equation can be modified to eliminate the need to estimate D, as follows where i is the subscript indicating the number of the specific age class where the youngest is 1 and oldest is n. Once the standing crop for each species in the community has been estimated, a meaningful diversity can be calculated. Study of the model reveals that in populations where high-early mortality is the rule the likelihood of error in estimating population parameters is lower than for constant and low-early mortality populations.

Constraining patterns of growth using directly observable and quantifiable characteristics can reveal a wealth of information regarding the biology of the Ediacara biota—the oldest macroscopic, complex community-forming organisms in the fossil record. However, these rely on individuals captured at an instant in time at various growth stages, and so different interpretations can be derived from the same material. Here we leverage newly discovered and well-preserved Dickinsonia costata Sprigg, 1947 from South Australia, combined with hundreds of previously described specimens, to test competing hypotheses for the location of module addition. We find considerable variation in the relationship between the total number of modules and body size that cannot be explained solely by expansion and contraction of individuals. Patterns derived assuming new modules differentiated at the anterior result in numerous examples in which the oldest module(s) must decrease in size with overall growth, potentially falsifying this hypothesis. Observed polarity as well as the consistent posterior location of defects and indentations support module formation at this end in D. costata. Regardless, changes in repeated units with growth share similarities with those regulated by morphogen gradients in metazoans today, suggesting that these genetic pathways were operating in Ediacaran animals.

The mean taxon duration, , and the 50 percent point of the Lyellian curve, T50, are directly proportional to one another: . However, there is a disagreement in the literature over the correct value of the proportionality constant, C. Various authors have claimed that C ≈ 1.44, C = 2.00 and C ≈ 2.89. I critically examine the assumptions underlying these three choices of C and argue that C ≈ 1.44, which is based on an exponential model of survivorship, is more widely applicable than the other values of C.

Diversity data from stochastic phylogenies and from uniformly spaced Gaussian curves were subjected to Q-mode factor analysis in order to determine whether a few factors would account for a large percentage of the original variance. In both analyses, a small number of factors show systematic variations in time and account for more than 90% of original data variance. To further study the question of evolutionary pulsations, turnover rates were calculated between successive samples. These turnover rates indicate that stochastic phylogenies have pulses similar to those recorded in the fossil record. Large scale environmental changes are not required to explain such pulses. Therefore the observed existence in the real world of biologic diversity associations and evolutionary pulsations can as equally well be accounted for in a stochastic world (in which each species is an independent variable) as in a deterministic world. This supports the notion that there may be stochastic laws in paleontology akin to the gas laws of chemistry.

A cladistic analysis of the three recognized patterns of central nervous visual organization among teleosts reveals that there is a pattern of intermediate complexity representing the plesiomorphic condition for teleosts, and that there is a simple visual pattern in two unrelated teleost groups which can be concluded to be a secondarily reduced derived condition, as well as an elaborate pattern which is present only in acanthomorph teleosts, thus likely representing a synapomorphy for this taxon. The elaborate central nervous visual pattern, therefore, is one of many functional-anatomical advanced features characterizing the acanthomorphs. Furthermore, when neontological and paleontological data is compared with the paleoecological record of early acanthomorph history during the Late Cretaceous, it is consistent with a hypothesis that this acanthomorph synapomorphic functional-anatomical complex arose likely in ctenothrissiforms as an adaptation to the life in the reorganizing reefs of that geologic period.

The pelvic structure in non-avian archosaurs plays a key role in understanding the evolution of terrestrial locomotor patterns because the pelvis contains major attachment sites for proximal hind limb musculature. In order to investigate patterns of pelvic evolution in archosaurs, this study compiled three pelvic indices, as well as femoral head orientation, for 92 archosaur taxa. With the metrics and a reconstructed supertree, we examined the correlated evolution of the pelvis and femur, the correlation among pelvic components, and temporal trends in the evolution of the pelvis. The result shows that archosaurs with medially directed femoral heads have more cranially shifted iliac centroids and more posteriorly rotated pubes than taxa with anteromedially directed femoral heads. The craniad shift of the iliac centroid might be correlated to the posterior rotation of pubis. The pelvic structures of pterosaurs, ornithischians, sauropods, and avetheropods occupy a different morphospace from basal archosaurs, pseudosuchians, basal dinosauromorphs, basal theropods, and basal sauropodomorphs in having more cranially expanded ilia, more posteriorly rotated pubes, and medially deflected femoral heads. This may imply that pterosaurs and those derived dinosaurs independently underwent similar shifts in thigh muscles and locomotion. The evolutionary model fitting supports the early-burst model for iliac and pubic metrics in more inclusive archosaur clades, indicating that larger changes of archosaur pelves occurred in early times of the clade's history.

Archaeocyaths are the most abundant sponges from the Cambrian period, having formed the first animal reef communities more than 500 million years ago. The Archaeocyatha are index fossils for correlating rocks of similar ages globally because of their abundance, extensive geographic distribution, detailed anatomy, and well-established taxonomy. Their ecological significance remains incompletely explored, yet they are known to have strongly competitively interacted, unlike modern sponges. This study examines the feeding ecology of the fossil remains of Siberian archaeocyath assemblages. As suspension feeders, archaeocyaths filtered plankton from the water column through pores in their outer wall. Here we outline a new method to estimate the limit on the upper size of plankton that could be consumed by an archaeocyath during life. The archaeocyaths examined were predominantly feeding on nanoplankton and microplankton such as phytoplankton and protozooplankton. Size-frequency distributions of pore sizes from six different Siberian archaeocyath assemblages, ranging from Tommotian to Botoman in age, reveal significantly different upper limits to the prey consumed at each locality. Some of the assemblages contain specimens that could have fed on larger organisms extending into the mesoplankton, including micro-invertebrates as a possible food resource. These results show that during the establishment of the first animal reef systems, prey partitioning was established as a way of reducing competition. This method has applicability for understanding the construction and the functioning of the first reef systems, as well as helping to understand modern reef systems and their development though time and space.

It was once thought that the endemic carnivorous mammals of South America, the metatherian sparassodonts, were driven extinct by North American carnivorans through competitive exclusion. However, sparassodonts went extinct before most groups of carnivorans entered South America; only the endemic Cyonasua-group procyonids (Cyonasua and Chapalmalania), which immigrated to South America nearly 4 million years earlier than other carnivorans, significantly overlapped with sparassodonts in time. In this study, we examine the functional morphology of the dentition of Cyonasua and Chapalmalania through quantitative analysis to determine the dietary habits of these taxa and the degree to which they may have ecologically overlapped sparassodonts and large predatory Neogene didelphimorphians. We find Cyonasua and Chapalmalania to be more carnivorous than extant procyonids, other than Bassariscus, in agreement with previous studies, but more omnivorous than most other carnivorans and all meat-eating South American metatherians, including sparassodonts. The extreme ecological dissimilarity between Cyonasua-group procyonids and members of the endemic South American predator guild may explain why procyonids were able to successfully establish themselves in South America several million years earlier than most other northern mammals (including all other carnivorans): they moved into a previously unoccupied ecological niche (large omnivore) and avoided direct competition with incumbent native species, a situation similar to that documented in historical cases of biological invasion. The omnivorous diets and climbing/swimming abilities of procyonids may have increased their chances for a successful over-water dispersal relative to other carnivorans, further favoring their successful establishment in South America.

The inner ear of the two higher clades of modern cetaceans (Neoceti) is highly adapted for hearing infrasonic (mysticetes) or ultrasonic (odontocetes) frequencies. Within odontocetes, Platanistoidea comprises a single extant riverine representative, Platanista gangetica, and a diversity of mainly extinct marine species from the late Oligocene onward. Recent studies drawing on features including the disparate tympanoperiotic have not yet provided a consensus phylogenetic hypothesis for platanistoids. Further, cochlear morphology and evolutionary patterns have never been reported. Here, we describe for the first time the inner ear morphology of late Oligocene–early Miocene extinct marine platanistoids and their evolutionary patterns. We initially hypothesized that extinct marine platanistoids lacked a specialized inner ear like P. gangetica and thus, their morphology and inferred hearing abilities were more similar to those of pelagic odontocetes. Our results reveal there is no “typical” platanistoid cochlear type, as the group displays a disparate range of cochlear anatomies, but all are consistent with high-frequency hearing. Stem odontocete Prosqualodon australis and platanistoid Otekaikea huata present a tympanal recess in their cochlea, of yet uncertain function in the hearing mechanism in cetaceans. The more basal morphology of Aondelphis talen indicates it had lower high-frequency hearing than other platanistoids. Finally, Platanista has the most derived cochlear morphology, adding to evidence that it is an outlier within the group and consistent with a >9-Myr-long separation from its sister genus Zarhachis. The evolution of a singular sound production morphology within Platanistidae may have facilitated the survival of Platanista to the present day.