We calculated water vapor conductance (a product of eggshell porosity) from the first definitively identified sauropod egg (Megaloolithus patagonicus) from the Auca Mahuevo locality in Argentina. We then compared the results with those from M. siruguei (an egg type long associated with sauropod dinosaurs) from the Pinyes locality in Spain. The 14-cm Auca Mahuevo egg has a thinner eggshell and 47 times fewer pores than the 22-cm M. siruguei specimen. The resulting water vapor conductance (GH2O) of the titanosaur and M. siruguei eggs is 341 and 3979 mg H2O day−1 Torr−1, respectively; these values are two and ten times greater than in avian eggs of comparable size, but lower than in eggs of most modern reptiles. Clutches from Auca Mahuevo typically contain 20–40 eggs; in contrast, M. siruguei clutches from the Pinyes site average nine eggs. The GH2O of M. siruguei exceeds that of the Argentine egg by an order of magnitude, supporting previous inferences of egg burial. The GH2O of the Argentine titanosaur egg closely approximates that of Troodon and some oviraptorid eggs, previously calculated as equal to or two times greater than, respectively, the GH2O of avian eggs of similar size. Higher embryonic growth rates (relative to modern reptiles), especially in some dinosaurs with large clutch mass, may have required incubation in a more open environment, where water conservation represented a more critical factor than in a buried clutch. The lower GH2O calculated for the two megaloolithid eggs is consistent with previous interpretations of nesting mode that are based on site taphonomy and nesting traces. This study indicates that at least some dinosaurs did not fully bury their eggs.

Medullosa stands apart from most Paleozoic seed plants in its combination of large leaf area, complex vascular structure, and extremely large water-conducting cells. To investigate the hydraulic consequences of these anatomical features and to compare them with other seed plants, we have adapted a model of water transport in xylem cells that accounts for resistance to flow from the lumen, pits, and pit membranes, and that can be used to compare extinct and extant plants in a quantitative way. Application of this model to Medullosa, the Paleozoic coniferophyte Cordaites, and the extant conifer Pinus shows that medullosan tracheids had the capacity to transport water at volume flow rates more comparable to those of angiosperm vessels than to those characteristic of ancient and modern coniferophyte tracheids. Tracheid structure in Medullosa, including the large pit membrane area per tracheid and the high ratio of tracheid diameter to wall thickness, suggests that its xylem cells operated at significant risk of embolism and implosion, making this plant unlikely to survive significant water stress These features further suggest that tracheids could not have furnished significant structural support, requiring either that other tissues supported these plants or that at least some medullosans were vines. In combination with high tracheid conductivity, distinctive anatomical characters of Medullosa such as the anomalous growth of vascular cambium and the large number of leaf traces that enter each petiole base suggest vascular adaptations to meet the evapotranspiration demands of its large leaves. The evolution of highly efficient conducting cells dictates a need to supply structural support via other tissues, both in tracheid-based stem seed plants and in vessel-bearing angiosperms.

The Paratethyan basins of eastern Europe and western Asia became isolated from marine influence in the Late Miocene, and were the sites of several remarkable endemic radiations of brackish and freshwater organisms. Here I describe the patterns of tempo and mode before and during the radiation of the gastropod Melanopsis in the Pannonian basin of eastern and central Europe, and I explore the underlying mechanisms of evolutionary change.

The most ancient melanopsid species in this area, M. impressa, was present in freshwater areas marginal to the basin well before the radiation. Widely spaced samples of M. impressa indicate that this species underwent a period of stasis lasting at least 7 m.y. The end of stasis corresponded with the extinction of the last of the normal marine fauna in the basin, suggesting that the lack of other fauna and/or reduced salinity in the basin permitted expansion of the melanopsids from the basin margins into the basin proper. Stasis ended with the onset of changes in size, shouldering, and ontogeny, which led eventually to M. fossilis. Change occurred over a 2-m.y. interval; a series of intermediates is present for all three characters. Within-sample correlations provide no evidence that the three characters are constructionally linked; instead they appear to be changing independently. The mode of change in the M. impressa–M. fossilis lineage appears to have been anagenetic. Alterations in the rate and direction of selection (and/or genetic links between characters) are probably required to explain the overall slowness of the change.

Most Pannonian basin melanopsid species arose by rapid cladogenesis in the Middle Pannonian Stage. Physical factors in the basin probably influenced the timing of this diversification; contrasting patterns of variation and diversity between two melanopsid clades suggest that intrinsic factors influenced the extent of diversification.

We have known since Trueman's classic work of 1922 that the Lower Jurassic Gryphaea of Britain exhibit phyletic size increase and heterochronic change in shape. Since Hallam's revisionary work in the 1960s, we have recognized that pronounced and generalized juvenilization of form accompanied this increasing size. This extensive literature provides invertebrate paleontology's most famous example of a biometrically documented, continuous anagenetic trend within a discrete lineage. But Gryphaea has also provoked great frustration because a key datum, required for a full solution, had been theoretically recognized but practically unavailable. We could identify the evolution of shape as paedomorphic, but could not specify the mode of heterochrony for this paedomorphic result because we could not standardize samples by common age or developmental stage.

In this paper, we provide sclerochronological data on sizes and shapes at specified ages marked by annual growth bands in two Jurassic sequences of Gryphaea: the classic Lower Jurassic series showing phyletic size increase with paedomorphosis, and an independent Middle–Upper Jurassic series illustrating neither size increase nor heterochrony. We prove that size increase in the classic series occurs entirely by faster growth (larger descendant sizes at the same ages as ancestors), and not by extended age (for descendants lived no longer than ancestors). The well-marked paedomorphosis of form probably arose as a correlated consequence of growing larger by extending and maintaining rapid juvenile growth rates—thus marking the heterochronic mode as a case of neoteny. The independent upper sequence, acting as a different replicate in a natural experiment, shows neither size increase nor heterochrony but does exhibit (in contrast with the classic sequence) evolution toward greater longevity.

Hallam's flow tank experiments indicated a strong adaptive advantage in shell stability for both larger size and paedomorphic form. Neotenous development provides an evolutionary pathway to the simultaneous acquisition of both favored traits—thus showing that “constraints” due to “correlations of growth” (Darwin's own phrase for the phenomenon) may be positive in promoting joint evolutionary advantages, and not only neutral (in carrying spandrels along with primary adaptations), or negative (by imposing inadaptive “baggage” upon trends in form through developmental correlation with selected traits).

The most important topics discussed thus far by community paleoecologists are: community approach to paleoenvironmental reconstruction, community development in evolutionary time, and community constraints upon species evolution. The first step in community-paleoecological analysis is to reconstruct the paleocommunity or paleoecosystem. However there are severe methodological limitations to any inference drawn from the composition and structure of a fossil assemblage. These result from various taphonomic biases. These constraints upon reliability of a community-paleoecological analysis are the least severe in the case of Cenozoic (and possibly Cretaceous) tropical to subtropical, molluskdominated, subtidal, benthic communities.

The basic assumption of community paleoecology is that communities or biocoenoses represent a distinct, real level of biotic organization achieved through ecological integration of and coevolution among the species. This assumption seems to be invalid for two reasons. (1) The actual degree of community integration is in general insufficient to induce any driving forces for a structural development as predicted by the system theory. (2) The concept of biological reality and distinctness of the community level of biotic organization implies assignment of a significant role to group selection. The assumption that ecological communities achieve with time an equilibrium state representing an optimum habitat partitioning among the component species is invalid, at least as a generalization. This idea is largely falsified and refuted by a variety of ecological studies. Ecological communities are then merely an epiphenomenon of the overlap in distributional patterns of various organisms. There is no intrinsic, biotic mechanism inducing community dynamics in either ecological, or evolutionary time.

In spite of this conclusion, the so-called “community approach” under favorable taphonomic conditions is among the most reliable methods of paleoenvironmental reconstruction, and the data on so-called “community evolution” are relevant to the problem of a relationship between actually realized niche patterns and their environmental framework. The latter problem can also be approached through analysis of longevity-frequency-distributions of chronospecies found living together in various habitats. A preliminary investigation may indicate that the precondition to optimization of niche dimensions through coevolution among ecologically related species was met in subtidal benthic habitats but not in pelagic ones. To account for a niche subdivision among planktonic organisms, one has therefore to invoke a stochastic pattern of speciation rather than coevolutionary mechanisms.

The age (mortality) profiles of ungulate species in a fossil fauna are useful for reconstructing modes of death and of bone accumulation. Crown height measurements probably provide the most practical means of obtaining age profiles in most fossil samples. Reliable, interpretable profiles will result when individual age is estimated from crown height using a mathematical model first suggested by C. A. Spinage and when the individual ages are grouped into relatively broad age classes. One potential objection to the model is that it requires tedious calculation and there is a strong possibility of calculation error when many crown heights are involved. This problem may be circumvented by the use of an interactive BASIC program appended to this paper. The user supplies the raw crown heights in a fossil sample, together with estimates of initial unworn crown height, ages of dental shedding and eruption, and maximum possible individual age. The program then tabulates the number of individuals in successive 10%-of-potential-lifespan intervals.

Field and laboratory data indicate that all four species of Thais (Gastropoda, Muricacea) from rocky shores of the northeastern Pacific selectively attack barnacles at the margins of parietal (lateral) and opercular plates. Attacks are also more likely to be successful at plate margins. Such preferential attack and differential attack success may account for the evolutionary reduction in the number of parietal plates exhibited within seven of the eight families of balanomorph barnacles. Additional evidence suggests that predation by drilling gastropods may have also favored the evolution of strong external ribbing on these plates within some balanomorph lineages.

Radiation and plate reduction in the Balanomorpha occurred in concert with extensive radiation in muricacean gastropods, starting in the Late Cretaceous. Of the three skeletal conformations exhibited by open-surface dwelling barnacles (8, 6 and 4 parietal plates), only genera of the four-plated form have become proportionally more common in the Recent. Further, in a rather striking evolutionary experiment, a lineage of barnacles evolving free from gastropod predation retained primitive skeletal features. This supports the interpretation that predation by drilling gastropods has been an important selective force behind the widespread, parallel evolutionary tendency towards plate reduction in the Balanomorpha; a trend recognized by Darwin (1854), but for which he offered no explanation.

Ecologists attempt to explain species diversity within Recent seabird communities in terms of Recent oceanographic and ecological phenomena. However, many of the principal oceanographic processes that are thought to structure Recent seabird systems are functions of geological processes operating at many temporal and spatial scales. For example, major oceanic currents, such as the North Pacific Gyre, are functions of the relative positions of continents and Antarctic glaciation, whereas regional air masses, submarine topography, and coastline shape affect local processes such as upwelling. I hypothesize that the long-term development of these abiotic processes has influenced the relative diversity and community composition of North Pacific seabirds. To explore this hypothesis, I divided the history of North Pacific seabirds into seven intervals of time. Using published descriptions, I summarized the tectonic and oceanographic events that occurred during each of these time intervals, and related changes in species diversity to changes in the physical environment. Over the past 95 years, at least 94 species of fossil seabirds have been described from marine deposits of the North Pacific. Most of these species are from Middle Miocene through Pliocene (16.0-1.6 Ma) sediments of southern California, although species from Eocene to Early Miocene (52.0-22.0 Ma) deposits are from Japan, British Columbia, Washington, and Oregon. During the history of the North Pacific seabirds, thee were many widespread (global) and local (California Current) oceanographic events, but the underlying physical processes that affected the diversity of seabirds throughout the North Pacific were tectonic changes that led to sequential stages of thermal isolation and refrigeration of Antarctica. Besides these broad-scale phenomena, the uplift of the Isthmus of Panama and the initiation and intensification of coastal upwelling in the California Current directly affected the diversity of seabirds in southern California. For the most part, Middle to Late Miocene and Pliocene seabird faunas from California resembled Recent communities, but with several exceptions: (1) the extreme diversity and abundance, but subsequent extinction of gannets (Morus) and flightless alcids (mancallids); (2) the comparatively low abundance and diversity of cormorants and shags (Phalacrocoracidae) until the Late Pliocene; and (3) the absence of marine Laridae until the Late Pliocene. By affecting both the physical and biological environments, geological factors such as Antarctic glaciation, eustatic changes in sea levels, and local tectonic activities influenced and will continue to influence the structure of seabird systems in the North Pacific.

The extreme morphological simplicity of lingulide brachiopod shells makes them particularly useful for investigating the species-level taxonomic resolution of the fossil record as well as the relationships between taxonomy, morphological complexity, and evolutionary rates. Lingulides have undergone little change in shell morphology and have had low taxonomic diversity since the Paleozoic. Is this pattern an evolutionary phenomenon or an artifact of the shell's simplicity? Multivariate methods were used to establish morphogroups among seven populations of four extant species of Glottidia. Six characters (three shell dimensions and three internal septa) were measured for 162 specimens from field and museum collections. All populations follow similar allometric trajectories: internal septa display positive allometry and shell dimensions display negative allometry. The allometric pattern may reflect D'Arcy Thompson's Principle of Similitude. Principal component analysis does not reveal any distinct clusters in Glottidia morphospace but suggests that some differences independent from ontogeny exist among the populations. Size-free canonical variate analysis indicates the presence of five size-invariant groups that are statistically distinct. Bootstrap-corrected error rates indicate that four specimens are enough to classify a sample correctly at α = 0.05 and eight specimens at α = 0.01. The groups are consistent with neontological classification with the exception of two populations of G. pyramidata identified by discriminant analysis as two distinct groups. The size-free morphogroups reflect geographic separation rather than ontogenetic or substrate differences among the populations.

Despite the morphological simplicity of the shell, size-free multivariate analysis of Glottidia delineates groups that offer taxonomic resolution comparable with the neontological classification. The method offers a promising tool for identifying natural morphogroups on the basis of few morphological characters. Moreover, the agreement between neontological taxonomy and the morphogroups suggests that the size-free approach can be applicable for evaluating the reality of the low diversity and turnover rates observed in the fossil record of lingulide brachiopods (= Family Lingulidae). Assuming that the neontological species of Glottidia are biologically meaningful, this study shows that morphological simplicity of lingulides does not necessarily result in taxonomic underresolution. Our analysis, as well as several previous case studies, suggests that taxonomic diversity and turnover rates do not have to be dependent on the morphological complexity of preservable parts. In many cases, when rigorous quantitative methods are employed, the differences in the rates of morphological evolution may be a real evolutionary phenomenon and not artifacts of morphological complexity.

Both fossil preservation and sampling methods affect perceived patterns of biotic diversity. Artificial range truncations, for example, may lead to incongruences between apparent- and actual-diversity curves. Thus, a catastrophic extinction event may appear gradual. Recent advances in biostratigraphic-gap analysis provide models for the distribution of gap lengths between fossil occurrence horizons and provide methods to place confidence intervals on local taxon ranges and remove the biases caused by artificial range truncations. Confidence intervals for a set of local taxon ranges may then be evaluated collectively to test a hypothesis of co-extinction/co-emigration or co-origination/co-immigration. In the case of terminal Cretaceous ammonites from Seymour Island, range-chart data are compatible with an abrupt extinction event, although the test statistic is not minimized at the stratigraphic horizon that was suggested by Macellari (1986).

The hyrax, a living primitive subungulate, makes an ideal model in terms of its molar morphology for early lophodont ungulates. Cine fluoroscopic studies of mastication show that the proposed Phase II in jaw movement, experimentally demonstrated in primates, but surmised in ungulates only from molar wear facets, does in fact exist in this animal, but does not appear to be an important component of the masticatory cycle. Comparison of tooth wear in early Tertiary ungulates with contemporaneous primates shows that, whereas primates enlarge the Phase II wear facets on the molars as they move into more herbivorous niches, the same is not true for ungulates, and in fact Phase II may be lost altogether in grazing forms. It is concluded that “grinding” in herbivorous ungulates is a process fundamentally different from that described for herbivorous primates.

Discovery of several new specimens of the gigantic Eocene ground bird Diatryma gigantea from the Willwood Formation of northwestern Wyoming, has prompted an analysis of its feeding apparatus and an assessment of the mode of life of this unusual bird. Diatryma exhibits many of the features predicted by biomechanical models to occur in animals delivering large dorsoventral bite forces. Similarly, the mandible of Diatryma, which was modeled as a curved beam, appears well equipped to withstand such forces, especially if they were applied asymmetrically. Interpretation of these size-independent biomechanical properties in light of the large absolute skull size of Diatryma suggests a formidable feeding apparatus. The absence of modern analogues complicates the determination of just how this unique skull morphology correlates with diet. Suggestions that Diatryma was an herbivore seem improbable in that they require the postulation of excessively high safety factors in the construction of the skull. The traditional hypothesis of Diatryma as a carnivorous bird accords as well or better with the data at hand. Carnivory raises the probability of “accidental” encounter with bones, thus explaining the high safety factors. In fact, the skull and mandible of Diatryma are so massive that bone crushing may have been an important behavior. Diatryma could have been a scavenger. However, limb allometry and phylogenetic interpretation of limb proportions call into question the picture of Diatryma as a slow, plodding graviportal animal, suggesting that active predation was within its behavioral repertoire.

It is commonly believed that the brains of the ancestors of modern carnivores (miacids) were superior to (e.g., larger than) those of other early carnivores (creodonts and mesonychids). Examination of the fossil record of brains of early carnivores reveals no evidence to support that belief. Moreover, evolutionary trends towards increasing relative brain size and an expansion of neocortex are seen in both miacids and creodonts. The neocortex expanded in a different way in miacids than in creodonts and mesonychids (evidenced by different sulcal patterns), but the biological significance of the observed differences is unknown.

The megafauna and associated behavioral traces of two deep-sea benthic environments, the central Arctic and Antarctic, with a surface primary productivity differential of 104 were compared to assess the role of food availability in foraging strategy and community structure. Bottom photographs, analyzed for megafauna and trace density and diversity at comparable depths in the Arctic Canada Basin and the Antarctic Bellingshausen Basin, indicated that trace frequency was inversely proportional to organism density but that trace diversity directly reflected organism diversity. Those traces identified in the fossil record to represent efficient foraging strategies, i.e., the Nereites facies, were conspicuously absent at all depths in the Arctic and present at all depths in the Antarctic, in contradiction of the paradigm of increasing behavioral complexity and sediment exploitation as food availability decreases. Presence or absence of surface-grazing organisms seems to exert a greater influence on trace diversity than depth or nutrient supply. Trace density, however, may reflect episodic sedimentation events which intermittently influence the deep-sea trophic regime.

We investigated the hydrodynamic properties of cephalopod shell sculpture in two ways: 1) flow visualization experiments with sculptured shells; and 2) application of drag coefficient data for simple geometric bodies to cephalopod shells. Results of this work suggest:

1) the hydrodynamic effect of shell sculpture depends primarily on the size of the sculptural elements relative to the size of the shell and on the positions of sculpture elements on the shell and relative to each other.

2) sculpture is detrimental to swimming (reduces hydrodynamic efficiency) if it exceeds the height of the lower part of the shell's boundary layer.

3) sculpture is advantageous to swimming (increases efficiency) if it remains immersed in the boundary layer and induces premature conversion to turbulent boundary layer flow. To be hydrodynamically optimal, small shells (diam ≈ 10 cm) must have rough (sculptured) surfaces, whereas large shells (diam ≈ 100 cm) require smooth surfaces. Thus, in order to maintain maximum efficiency throughout life, the ontogeny of small individuals, or species, should be characterized by progressive roughening of the shell, while large forms should become increasingly smooth. Such allometries are observed among many ammonoids.

4) sculpture always has an effect on the flow around a cephalopod shell. In some species this effect was probably negligible, while in others, those with compressed shells especially, it was probably of major importance. In these species, sculpture appears to have functioned primarily to increase swimming ability.

Diversity dynamics of the Permian–Triassic land plants in South China are studied by analyzing paleobotanical data. Our results indicate that the total diversity of land-plant megafossil genera and species across the Permian/Triassic boundary (PTB) of South China underwent a progressive decline from the early Late Permian (Wuchiapingian) to the Early-Middle Triassic. In contrast, the diversity of land-plant microfossil genera exhibited only a small fluctuation across the PTB of South China, showing an increase at the PTB. Overall, land plants across the PTB of South China show a greater stability in diversity dynamics than marine faunas. The highest extinction rate (90.91%) and the lowest origination rate (18.18%) of land-plant megafossil genera occurred at the early Early Triassic (Induan), but the temporal duration of the higher genus extinction rates (>60%) in land plants was about 23.4 Myr, from the Wuchiapingian to the early Middle Triassic (Anisian), which is longer than that of the coeval marine faunas (3–11 Myr). Moreover, the change of genus turnover rates in land-plant megafossils steadily fluctuated from the late Early Permian to the Late Triassic. More stable diversity and turnover rate as well as longer extinction duration suggest that land plants near the PTB of South China may have been involved in a gradual floral reorganization and evolutionary replacement rather than a mass extinction like those in the coeval marine faunas.

Cantius is an early Eocene adapid primate with an exceptionally well known fossil record. Measurements were recorded to describe the size and shape of upper and lower first molars collected from a measured stratigraphic section in the Clarks Fork Basin of Wyoming. Rates of change of size and shape are quantified by calculating evolutionary rates in standard deviation units per generation (haldanes). Temporal scaling of rates shows that change in size was generated by a significantly nonrandom directional process, while change in shape was generated by a significantly nonrandom stabilizing process. Size change in Cantius is interpreted to be the result of weak directional selection, and shape change is interpreted to be the result of strong stabilizing selection.

A simple principle predicts a tendency, or vector, toward increasing organismal complexity in the history of life: As the parts of an organism accumulate variations in evolution, they should tend to become more different from each other. In other words, the variance among the parts, or what I call the “internal variance” of the organism, will tend to increase spontaneously. Internal variance is complexity, I argue, albeit complexity in a purely structural sense, divorced from any notion of function. If the principle is correct, this tendency should exist in all lineages, and the resulting trend (if there is one) will be driven, or more precisely, driven by constraint (as opposed to selection). The existence of a trend is uncertain, because the internal-variance principle predicts only that the range of options offered up to selection will be increasingly complex, on average. And it is unclear whether selection will enhance this vector, act neutrally, or oppose it, perhaps negating it. The vector might also be negated if variations producing certain kinds of developmental truncations are especially common in evolution.

Constraint-driven trends—or what I call large-scale trends of the fourth kind—have been in bad odor in evolutionary studies since the Modern Synthesis. Indeed, one such trend, orthogenesis, is famous for having been discredited. In Stephen Jay Gould's last book, The Structure of Evolutionary Thought, he tried to rehabilitate this category (although not orthogenesis), showing how constraint-driven trends could be produced by processes well within the mainstream of contemporary evolutionary theory. The internal-variance principle contributes to Gould's project by adding another candidate trend to this category.

We investigate whether musculoskeletal anatomy and three-dimensional (3-D) body proportions were modified during the evolution of large (>6000 kg) body size in Allosauroidea (Dinosauria: Theropoda). Three adaptations for maintaining locomotor performance at large body size, related to muscle leverage, mass, and body proportions, are tested and all are unsupported in this analysis. Predictions from 3-D musculoskeletal models of medium-sized (Allosaurus) and large-bodied (Acrocanthosaurus) allosauroids suggest that muscle leverage scaled close to isometry, well below the positive allometry required to compensate for declining muscle cross-sectional area with increasing body size. Regression analyses on a larger allosauroid data set finds slight positive allometry in the moment arms of major hip extensors, but isometry is included within confidence limits. Contrary to other recent studies of large-bodied theropod clades, we found no compelling evidence for significant positive allometry in muscle mass between exemplar medium- and large-bodied allosauroids. Indeed, despite the uncertainty in quantitative soft tissue reconstruction, we find strong evidence for negative allometry in the caudofemoralis longus muscle, the single largest hip extensor in non-avian theropods. Finally, we found significant inter-study variability in center-of-mass predictions for allosauroids, but overall observe that consistently proportioned soft tissue reconstructions produced similar predictions across the group, providing no support for a caudal shift in the center of mass in larger taxa that might otherwise reduce demands on hip extensor muscles during stance. Our data set provides further quantitative support to studies that argue for a significant decline in locomotor performance with increasing body size in non-avian theropods. However, although key pelvic limb synapomorphies of derived allosauroids (e.g., dorsomedially inclined femoral head) evolved at intermediate body sizes, they may nonetheless have improved mass support.

An intertidal sample of 118 to 155 Terebratalia transversa on Saltspring Island, B.C., was censused in 1974, 1975, and 1976. Growth in length was a maximum of 7.8 mm per year and declined with size (age). As in fossil articulates, mortality rate was independent of age (size). Recruitment was patchy in time and space, was multi-annual or continuous, and was concentrated near conspecific adults. There were as many as 800 individuals/m2. None moved or changed orientation. Morphologic variation of Terebratalia valves is not directly controlled by the intensity of waves or currents.

Paleontological survivorship curves based on size-frequency data are subject to ambiguous interpretation because two critical assumptions (direct relationship of size and age, constant population structure) are difficult to justify.