Time is so fundamental to the everyday thinking of paleobiologists and geologists that it is seldom given close critical attention. Many of the currently debated issues in evolutionary history—catastrophic extinctions and punctuated vs. gradual morphological change, for instance—include assumptions about time and rate which are seldom made explicit. Methods now exist for calibrating evolutionary patterns through estimates of time components in the fossil and sedimentological records. There is a growing realization that increased precision in defining time frameworks can greatly clarify evolutionary problems. This has helped to stimulate renewed interest in the traditional topic of time and how it is represented in the biological and geological record.

The extent of morphologic innovation during the Ediacaran–Cambrian diversification of animals was unique in the history of metazoan life. This episode was also associated with extensive changes in the redox state of the oceans, in the structure of benthic and pelagic marine ecosystems, in the nature of marine sediments, and in the complexity of developmental interactions in Eumetazoa. But did the phylogenetic and morphologic breadth of this episode simply reflect the unusual outcome of recurrent evolutionary processes, or was it the unique result of circumstances, whether in the physical environment, in developmental mechanisms, or in ecological interactions? To better characterize the uniqueness of the events, I distinguish among these components on the basis of the extent of sensitivity to initial conditions and unpredictability, which generates a matrix of possibilities from fully contingent to fully deterministic. Discriminating between these differences is important for informing debates over determinism versus the contingency in the history of life, for understanding the nature of evolutionary theory, and for interpreting historically unique events.

The Plio-Pleistocene planktic foraminiferal sequence of the Globorotalia (Globoconella) puncticulata-inflata clade in Deep Sea Drilling Project Site 588, dated at 4.36 Ma to 0.05 Ma, records the branching history of the G. inflata lineage from the ancestral G. puncticulata lineage. The gradational nature of the divergence and the enormous morphological variability inherent in the G. inflata lineage have elicited different views on taxonomy and phylogeny of this clade. A pattern recognition technique, soft independent modeling of class analog (SIMCA), was used as an objective quantitative stratophenetic methodology to reconstruct the phylogenetic history.

Typical specimens of two species, G. puncticulata and G. inflata, were identified from a stratigraphic level dated at 2.76 Ma. Principal component models were built to characterize the morphometric patterns of the two morphotypes using SIMCA. The Globoconella specimens of the next lower and higher adjacent stratigraphic levels were evaluated against the models and classified into one of the two morphotypes. The newly classified specimens were then used to build new models for further tracing of lineages in lower and upper sections, respectively. Progression of such training and classification procedures through stratigraphic intervals resulted in a reconstruction of the evolutionary patterns of the two lineages. Cladogenesis gave rise to the descendant lineage, G. inflata, at about 3.5 Ma. The two co-existing species, G. inflata and G. puncticulata, differ only in size and show similarity in most characters at the beginning of their divergence. Other characters began to diverge later, at various rates. The gradients between planktic and benthic foraminiferal δ18O values show a continuous increase during the late Pliocene. The succession from G. puncticulata to G. inflata during the same time correlates with the progressively increased vertical stratification in temperature of surface waters. Globorotalia puncticulata became extinct at 2.35 Ma when the temperature gradient further increased, corresponding to the onset of extensive glaciation in the Northern Hemisphere.

The theoretical framework developed here delineates the biological and paleontological circumstances under which the observed durations and diversities of fossil taxa will be significantly affected by three time-dependent sampling biases. The pull of the Recent and the loss of fossiliferous sediments shorten the observed durations of ancient taxa, as compared to the observed durations of Recent taxa. By contrast, the duration truncation mechanism lengthens the observed durations of ancient, as compared to more recent, taxa. Both the pull of the Recent and loss of fossiliferous sediments decrease the observed diversity of ancient, as compared to contemporary, communities.

This theoretical framework is built on four parameters: the extinction and origination rates q and p; the fossilization rate, v; and the fossil loss rate, h. I derive formulas for the observed durations and diversities of fossil taxa in terms of these parameters and develop a method of estimating these parameters. The estimation procedure is applied to the bivalve families, and I provide statistical evidence for the conclusion that q – p < 0 for bivalve families, and hence that bivalve diversity has been increasing through time. The theory predicts that a certain fraction of an extinct fauna will be represented by only one fossil, and hence will have an observed duration of zero. As the record becomes poorer, this fraction increases. For bivalves I assume that families known from only one stage have an observed duration of zero, and exploit this assumption to estimate h and v/p.

To evaluate the errors made in applying the model to the bivalves, I estimate several parameters in two different ways and find the estimates to be consistent. Additionally, I argue that the errors caused by the simplifying assumptions of the model are less than the sampling errors inherent in estimating p, q, h, and v.

Predatory marine snails and their prey provide a unique look at many aspects of predation events, allowing behavioral inference and studies of coevolution. This study examines differential predation patterns, rates, and success of two co-occurring gastropod predator families which drill two co-occurring species of Turritella (Turritellidae: Gastropoda) in the Gulf of California. Both naticid and muricid predators, identified by the shapes of their respective boreholes, attacked the thinner-shelled Turritella leucostoma more frequently than the thicker-shelled Turritella gonostoma. Both species were drilled more frequently and more successfully by naticid, as compared to muricid, predators. Naticids drilled prey in the 40- to 70-mm size class most frequently. Prey over 100 mm in length were relatively safe from all drilling predators. Predator size (estimated by borehole diameter) in naticids was correlated with prey size in both species, but for a given-sized prey, predators on T. gonostoma were proportionally larger. There was no size correlation for muricid predators. Unsuccessful attempts (incomplete drilling) were started on the suture more often than were completed holes, for both predator families on both prey species. Naticids began drilling T. leucostoma on the suture significantly less than expected by chance. We looked for possible changes over evolutionary time by analyzing prey shells from Pleistocene and Recent storm deposits. We found no evidence of change in any aspect of implied predatory behavior over the past 100 k.y.

Reconstructing the tree of life requires deciphering major evolutionary transformations and the functional capacities of fossils with “transitional” morphologies. Some of the most iconic, well-studied fossils with transitional features are theropod dinosaurs, whose skeletons and feathered forelimbs record the origin and evolution of bird flight. However, in spite of over a century of discussion, the functions of forelimb feathers during the evolution of flight remain enigmatic. Both aerodynamic and non-aerodynamic roles have been proposed, but few of the form-function relationships assumed by these scenarios have been tested. Here, we use the developing wings of a typical extant ground bird (Chukar Partridge) as possible analogues/homologues of historical wing forms to provide the first empirical evaluation of aerodynamic potential in flapping theropod “protowings.” Immature ground birds with underdeveloped, rudimentary wings generate useful aerodynamic forces for a variety of locomotor tasks. Feather development in these birds resembles feather evolution in theropod dinosaurs, and reveals a predictable relationship between wing morphology and aerodynamic performance that can be used to infer performance in extinct theropods. By spinning an ontogenetic series of spread-wing preparations on a rotating propeller apparatus across a range of flow conditions and measuring aerodynamic force, we explored how changes in wing size, feather structure, and angular velocity might have affected aerodynamic performance in dinosaurs choosing to flap their incipient wings. At slow angular velocities, wings produced aerodynamic forces similar in magnitude to those produced by immature birds during behaviors like wing-assisted incline running. At fast angular velocities, wings produced forces sufficient to support body weight during flight. These findings provide a quantitative, biologically relevant bracket for theropod performance and suggest that protowings could have provided useful aerodynamic function early in maniraptoran history, with improvements in aerodynamic performance attending the evolution of larger wings, more effective feather morphologies, and faster angular velocities.

This paper is aimed at constraining the phylogenetic frame of the acquisition of endothermy by Archosauromorpha. We analyzed the bone histology of Azendohsaurus laaroussii. Stylopodial and zeugopodial bones show three tissue types: (1) avascular lamellar zonal bone formed at low growth rates; (2) a scaffold of parallel-fibered bone containing either small primary osteons or simple vascular canals; and (3) fibrolamellar bone formed at high growth rates. We used quantitative histology to infer the thermometabolic regime of this taxon. We define endothermy as the presence of any mechanism of nonshivering thermogenesis that increases both body temperature and resting metabolic rate. Thus, estimating the resting metabolic rate of an extinct organism may be a good proxy to infer its thermometabolic regime (endothermy vs. ectothermy). High resting metabolic rates have been shown to be primitive for the clade Prolacerta–Archosauriformes. Therefore, we inferred the resting metabolic rates of A. laaroussii, a sister group of this clade, and of 14 extinct related taxa, using phylogenetic eigenvector maps. All the inferences obtained are included in the range of variation of resting metabolic rates measured in mammals and birds, so we can reasonably assume that all these taxa (including Azendohsaurus) were endotherms. A parsimony optimization of the presence of endothermy on a phylogenetic tree of tetrapods shows that this derived character state was acquired by the last common ancestor of the clade Azendohsaurus–Archosauriformes and that there is a reversion in Crocodylia.

Shell form is not strictly linked to life habits in modern marine turritelliform gastropods. To test the usefulness of various morphological characters in determining life-mode, I present a set of predictions giving the expected distribution of characters occurring in turritelliform snails with three different life-modes. Burrowing species should lack sculpture, possess columellar folds and a flat whorl profile, and have an orthocline or prosocline aperture. Mobile epifaunal forms should have sculpture, a rounded whorl profile, a displaced tangential aperture and a smooth columella. Sedentary forms should resemble epifaunal forms but have non-tangential apertures. These predictions were tested with a sample of 105 Recent marine species. Each hypothesis was found to be a statistically valid generalization and in 92 of the species (88%) the life habits were correctly predicted. Accuracy may be further improved by considering additional features such as ratchet sculpture and disjunct or open coiling. These patterns of shell form can be used to interpret fossil species as burrowers, or as sedentary or active epifaunal forms. For example, the unusual Devonian murchisoniid gastropod Ptychocaulus verneuili is interpreted as an active burrower.

The relatively imperfect relationship between shell form and life-mode in turritelliform gastropods, as compared to the Bivalvia, apparently results in part from the behavioral complexity of the Gastropoda. Gastropods have a repertoire of activities which would place them in different life-modes at different times; snail morphology reflects this complexity.

The siphuncular tube is a key component of the buoyancy control and mechanical strength systems in both Nautilus and fossil cephalopods. We measured the rate of hydrostatically induced fluid flow across the tube wall and tube rupture strength of Nautilus pompilius at hydrostatic pressures in the range of 10–85 bars. We found that in fresh, undecayed tubes, rupture occurs at pressures of about 80–85 bars. This is equivalent to the strength of the shell proper and to the depth limit of the live animal. The siphuncular tube is neither markedly stronger, nor weaker, than the shell. Siphuncle rupture strength is constant in the last 20 chambers of the shell despite a strong decrease in the siphuncle strength index (ratio of tube thickness to radius). The notion that strength index gives an accurate indication of tube strength is therefore in error. This suggests that the geometry of the siphuncular tube can not be straightforwardly used as an index of living depth in fossil cephalopods. Rupture occurs at the siphuncle-septum contact. The junction of the tube to its mechanical supports is thus weaker than the tube itself. Measured flow rates are in the range of 1–20 ml/h/chamber. Flow rates increase linearly with applied pressure and in successively larger chambers as a result of size-related variation in surface area and thickness of the tube wall. Rates of osmotic pumping in live animals are up to three orders of magnitude lower than hydrostatically induced flow rates across the siphuncular tubes of empty shells. Pumping capacity of live animals is apparently limited by functional constraints of the osmotic pump rather than by the fluid conductance properties of the tube wall. Living depth in evolving cephalopod lineages may be limited ultimately by physiologic or chemical restrictions of the osmotic pumping mechanism rather than by mechanical strength of the shell or siphuncle.

Studies of taxonomic diversity over time commonly count and compare first- and last-appearance data (FADs and LADs) over a succession of temporal intervals, and interpret them with respect to taxon origination and extinction. Singleton taxa, which first appear and last appear in the same temporal interval, are often removed from analyses because they might result from preservational biases rather than evolutionary processes, or they might represent non-independent FADs and LADs. Should singleton taxa always be excluded? We argue that in the case of Paleozoic terebratulide brachiopods, although they may be sensitive to biases in sampling intensity, singleton genera should be included in diversity studies because they do not appear to result from more typical biases, such as Lagerstätten and temporal interval length, that arguably can result in artificially high numbers of singleton genera.

Singleton genera can be critical and effective when used to test hypotheses regarding the existence and generation of latitudinal diversity gradients. Contrary to the anti-tropical diversity pattern of modern articulated brachiopods, Paleozoic terebratulides show a latitudinal diversity gradient that peaks in the Tropics. The hypothesis that the Tropics are either a diversity source or sink can be tested by comparing FAD and LAD latitudes. For singleton genera, FAD and LAD latitudes are taken from the same data points and must be removed for statistical comparisons to be valid. We suggest that taxon age distributions can accommodate singleton data, as the taxon age metric considers origination and extinction simultaneously. We generated taxon age distributions to test the hypothesis that the observed Paleozoic diversity gradient results from a latitudinal bias in generic turnover rate. We discovered that singletons are not randomly distributed over latitude, with proportionally more singleton genera occurring in the Tropics. In this case, singleton genera may reflect rapid evolutionary turnover of taxa, rather than simply preservational bias. Methods that can accommodate singleton taxa should be used to study the diversity of Paleozoic terebratulides and possibly other well-skeletonized marine metazoans.

Previous observations about the stable nature of coral-rich assemblages from the Middle Devonian Hamilton Group have led some researchers to invoke the primacy of ecological controls in maintaining biofacies structure through time. However, few analyses have examined the degree to which recurring biofacies vary quantitatively, and none have assessed lateral variability as a benchmark for testing the significance of temporal variability. Thus, the extent to which Hamilton biofacies persist and the mechanism(s) responsible for their hypothesized stability remain contentious. In this study, recurring coral-rich biofacies were evaluated from two stratigraphic horizons within the Middle Devonian Appalachian Basin to examine (1) the extent to which species assemblages persisted within the basin through space and time, and (2) whether ecological interactions may be a plausible mechanism for generating the degree of stasis observed in this case.

Variations in species composition and abundance were examined across multiple spatial scales within both sampled coral-rich horizons. This permitted the establishment of a baseline against which temporal differences in biofacies composition and structure could be evaluated. Although successive coral-rich horizons remained taxonomically stable, their dominance structures changed significantly through the 1.5 Myr study interval. Moreover, additional comparisons among older Hamilton coral-rich horizons corroborate our primary results. These findings support a model in which species respond individually to fluctuations in the physical environment, as indicated by the fluidity of their relative abundances geographically and temporally.

Vertebrates with terrestrial or freshwater ancestors colonized the sea from the Early Triassic onward and became competitively dominant members of many marine ecosystems throughout the Mesozoic and Cenozoic eras. The circumstances that led to initial marine colonization have, however, received little attention. One hypothesis is that mass extinction associated with ecosystem collapse provided opportunities for clades of amphibians, reptiles, birds, and mammals to enter marine environments. Another is that competitive pressures in donor ecosystems on land and in freshwater, coupled with abundant food in nearshore marine habitats, favored marine colonization. Here we test these hypotheses by compiling all known secondarily marine amniote clades and their times of colonization. Marine amniotes are defined as animals whose diet consists primarily of marine organisms and whose locomotion includes swimming, diving, or wading in salt water. We compared the number of clades entering during recovery phases from mass extinctions with the rate of entry of clades during nonrecovery intervals of the Mesozoic and Cenozoic. We conservatively identify 69 marine colonizations by amniotes. The only recovery interval for which prior mass extinction could have been a trigger for marine entry is the Early Triassic, when four clades colonized the sea over 7 Myr, significantly above the rates at which clades entered during other intervals. High nearshore productivity was a greater enticement to colonization than was a low diversity of potential marine competitors or predators in nearshore environments of a highly competitive terrestrial or freshwater donor biota. Rates of marine entry increased during the Cenozoic, in part because of rising productivity and in part thanks to the participation of warm-blooded birds and mammals, which broadened the range of thermal environments in which initial colonization of the sea became possible.

A method is provided to calculate the expected distribution of species' ranges for use as a basis of comparison for paleontological interpretation of species' range charts. If the method is used for such studies, the possibility of suggesting an environmental or biological cause for observed patterns, when in fact no such cause exists, will be diminished. The method is applied to a large, well-studied fossil data set to illustrate some possible species' range patterns which result only from sample size inequities. As examples, stepwise extinction and species richness patterns are predicted for situations where, in reality, none exist.

Using computer simulations and a quantitative method for describing bifurcating structures, the morphology of branching patterns seen in early land plants is analyzed. Four types or models of random branching (regular, geometric, binomial, and poisson) are shown to adequately describe the range of observed branching in most early land plants. Approximately 57% of all randomly generated computer patterns show reiterative branching events (=three successive identical modes of branching). Artificial canalization of reiterative events results in branching patterns structurally analogous with that of ancient fossil plants. Simulated phylogenetic changes among early land plant lineages, based on parsimonious transitions in branching patterns, indicate that most observed trends can be related directly to those seen in randomly generating branching patterns in which “size” is increased. The trimerophyte to progymnosperm trend in changing branching patterns is an exception, since the binomial model describing the progymnosperms has not been simulated by random processes.

While the apparent phylogenetic changes among early land plant groups do not require deterministic explanations, the transition from regular to geometric branching and the “canalization” of reiterative branching patterns may represent a grade level response to selective pressures related to mechanical design and vegetative reproduction.

The Brontotheriidae (Perissodactyla, Mammalia) are often used as an illustration of vertebrate macroevolutionary trends because their morphological evolution includes significant size increases accompanied by the disproportionate lengthening of bony frontonasal horns. The positive phylogenetic allometry for horn length vs. skull length is among the strongest known of such relationships in vertebrate phylogeny. Hypotheses explaining the change from small, incipient horns in Eocene ancestors to longer horns in Oligocene descendants have included two heterochronic mechanisms, hypermorphosis (extrapolation) and predisplacement (earlier onset time of horn growth). These proposed peramorphic mechanisms derive from interpretation of adult intergeneric allometries in logarithmic data spaces. Analysis of the raw (unlogged) data shows that the simple allometric model previously used is not an appropriate model for this specific problem. The heterochronic interpretations derived from them are therefore unsupported (but not disproven) by the allometries. A more appropriate allometric model for the data (full model) does not support any heterochronic interpretation. Previously unaccounted for in the heterochronic hypotheses is a complication due to body-size scaling effects on life history stage lengths. Neontological scaling patterns suggest that brontothere size increases were probably accompanied by increasing life spans and longer developmental stages. This effect broadens the types of heterochronies that may be postulated. Semiquantitative analyses comparing brontotheres with similarly sized extant ungulates show the hypothesized effect of larger size on brontothere life history stages. A scaled descendant ontogeny introduces the problem of relative vs. absolute time frames within which to view ontogenetic onset times. Thus, predisplacements, postdisplacements, or nondisplacements may be viewed as relative or absolute with respect to ancestral ontogenies. This raises a fundamental question about how development scales, which in turn affects how heterochronies are interpreted. A scaling effect suggests that brontothere horns are more likely postdisplaced in the traditional absolute time sense. Paradoxically then, while the descendant adult horn is peramorphic, its onset time may have shifted in a paedomorphic direction. Data for two Oligocene juvenile brontotheres suggest that most horn growth occurred late in their longer (i.e., descendant) ontogenies (hypermorphosis), and that the horns probably grew at faster rates (acceleration) than in Eocene taxa.

One of the major evolutionary transitions of the mammaliaform lineage was the origin of a typically mammalian pattern of growth. This is characterized by rapid juvenile growth followed by abrupt cessation of growth at adult size and may be linked with other important mammaliaform apomorphies of dental replacement and morphology. Investigation of growth patterns in the tritylodontid cynodont Oligokyphus and the basal mammaliaform Morganucodon provides insight into this crucial transition. We collected mandibular depth measurements from large samples of Morganucodon and Oligokyphus and constructed distributions of mandibular depth versus frequency for each species. These were compared with distributions from species from three different growth classes of extant amniote: testudines + crocodilians, mammals + birds, and lepidosaurs. Discriminant function analysis was used to differentiate between known growth classes by using different combinations of three measures of mandibular depth distribution shape (skew, kurtosis, and coefficient of variation) as proxies for different juvenile and adult growth patterns. Classification of the fossil species showed that Morganucodon closely resembled extant placental mammals in having rapid juvenile growth followed by truncated, determinate adult growth. Oligokyphus showed intermediate growth patterns, with more extended adult growth patterns than Morganucodon and slightly slower juvenile growth. This suggests a gradual evolution of mammalian growth patterns across the cynodont to mammaliaform transition, possibly with the origin of rapid juvenile growth preceding that of truncated, determinate adult growth. In turn, acquisition of both these aspects of mammalian growth was likely necessary for the evolution of diphyodont tooth replacement in the mammaliaform lineage.

Even though Neogene outcrops along the temperate Pacific coast of South America harbor a rich marine vertebrate fossil record, no studies have examined the diversification patterns of these taxa. Here, we analyze diversification trends based on the stratigraphic ranges of 86 genera of marine vertebrates, including sharks, rays, chimaeras, marine mammals, and seabirds. The richness of genera shows a hump-shaped trend, with maximum values around the late Miocene, driven by a large pulse of origination during mid-Miocene and higher extinction rates during the Pliocene. Trends varied markedly among taxa and departed largely from expectations based on global diversification patterns. Moreover, these trends cannot be explained solely as a sampling artifact derived from sampling intensity (i.e., number of occurrences) or sedimentary rock availably (i.e., number of geologic maps). A large fraction of genera (42%) went globally extinct by the late Pliocene–Pleistocene, and the extinction was highly selective according to different ecological and life-history traits. An analysis using “randomForest” showed that taxonomic structure and the geographic midpoint of distribution could explain up to 83% of extinction of genera. The extinction was taxonomically clumped (i.e., disproportionally high in Cetacea and very low in Carcharhiniformes) and concentrated in the northern area of the temperate Pacific coast of South America. Our results suggest that the particular paleogeographic, paleoclimatic, and paleoceanographic events that took place during the Neogene along the temperate Pacific coast of South America had a significant effect on the structure of marine biodiversity.

We have investigated whether changes in marine faunal diversity during the Pleistocene would have been affected by changes in sea level. Our estimate of changes in area of shallow marine seas between low stand and high stand of sea level (approximately 200 m) leads to a prediction of approximately a 25% change in species diversity, 10% change in generic diversity and less than a 5% change in familial diversity. This agrees with the observation that large scale generic or familial changes have not been widely noted for the Pleistocene. As in other intervals of geologic time, changes in the sizes of faunal provinces owing to changes in sea level are less likely to be the significant factor in changing marine diversity than are changes in the number of faunal provinces.

The class Echinoidea apparently originated during the Ordovician Period and diversified slowly through the Paleozoic Era. The clade then mushroomed in diversity beginning in Late Triassic time and continued expanding into the present. Although this evolutionary history is generally accepted, the taphonomic overprint affecting it has not been explored. To gain a more accurate perception of the evolutionary history of the group, I have compared the diversity history of the family Cidaridae (Echinodermata: Echinoidea) with the preservational style of fossil type species using literature-derived data. The Cidaridae apparently originated in Middle Triassic time and diversified slowly through the Neocomian (Early Cretaceous). Diversity was maintained through the remainder of the Cretaceous and Tertiary Periods, reflecting the diversity history of the subclass. Characterization of the preservational style of type fossil material for the family revealed the following breakdown of preservational states: 60% of species were described on the basis of disarticulated skeletal material, primarily spines; 20% based on intact coronas denuded of spines, apical system, Aristotle's lantern and peristomial plates; 10% based on large coronal fragments; and 10% based on other skeletal elements. This distribution may represent the effect of a disarticulation threshold on the condition of echinoid carcasses before final burial and suggests that preservation of intact specimens may be very unlikely. For cidaroids, previous work has suggested that this threshold is likely to be reached after 7 days of decay.

Comparison of the diversity history of the Cidaridae with the preservation data reveals that characteristic patterns of taphonomic overprint have affected the group since its origination in Middle Triassic time, and the nature of that overprint has changed over time: the early diversity history of the group is characterized by occurrences of fragmented fossil material, with spines predominant; further radiation of the group in mid-Jurassic time coincided with an increase in modes of preservation, ranging between exceptionally well-preserved material and disarticulated skeletal elements. Finally, type material is more rarely described from younger stratigraphic intervals (Miocene–Pleistocene) and consists predominantly of disarticulated skeletal elements and coronal fragments larger than an interambulacrum in size. Intact, denuded coronas are noticeably lacking.

The number of type species of Cidaridae described in each stratigraphic interval has not been consistent during post-Paleozoic time. Middle Triassic, Malm (Upper Jurassic), Senonian (Upper Cretaceous) and Eocene series yielded significantly (α = .05) higher numbers of type specimens per million years, while the Lias (Lower Jurassic), Dogger (Mid-Jurassic), Lower Cretaceous and Paleocene yielded significantly (α = .05) lower numbers of type specimens per million years. This may be the result of a combination of taxonomic, sampling, and geographical biases.