The Neoselachii are a monophyletic group including all of the extant sharks and rays. They underwent rapid diversification throughout the Jurassic and Cretaceous, going from low-diversity assemblages of members of extinct orders in the Late Triassic to diverse assemblages containing representatives of most extant clades by the end of the Cretaceous. The known fossil record of Mesozoic neoselachians is composed largely of isolated teeth, with articulated skeletal remains being known from a limited number of sites. The small tooth size of a large proportion of neoselachians, including almost all taxa in existence prior to the mid Cretaceous, led to very poor representation in older publications. Their state of knowledge has improved dramatically since 1970 with the increased use of bulk sampling for isolated dental remains. Despite this, the high proportion of Lazarus taxa from some stages suggests that the state of knowledge is still intermittent. Increase in assemblage diversity throughout the Jurassic and Cretaceous suggests that radiation events resulted in real and dramatic increases in diversity, and that the perceived diversification is not an artifact of poor knowledge. Cladogenesis inferred from the fossil record typically compares more favorably with divergence predicted from molecular analysis, where Batoidea form a discrete basal clade, than with divergence predicted from morphological analysis, where Batoidea are considered a derived crown group within the Squalea. The timing of diversification events is discussed in light of the known fossil record, cladistically generated divergence times, and the paleoenvironmental distribution of faunas.

Few paleontological studies of species distribution in time and space have adequately considered the effects of sample size. Most species occur very infrequently, and therefore sample size effects may be large relative to the faunal patterns reported. Examination of 10 carefully compiled large data sets (each more than 1,000 occurrences) reveals that the species-occurrence frequency distribution of each fits the log series distribution well and therefore sample size effects can be predicted. Results show that, if the materials used in assembling a large data set are resampled, as many as 25% of the species will not be found a second time even if both samples are of the same size. If the two samples are of unequal size, then the larger sample may have as many as 70% unique species and the smaller sample no unique species. The implications of these values are important to studies of species richness, origination, and extinction patterns, and biogeographic phenomena such as endemism or province boundaries. I provide graphs showing the predicted sample size effects for a range of data set size, species richness, and relative data size. For data sets that do not fit the log series distribution well, I provide example calculations and equations which are usable without a large computer. If these graphs or equations are not used, then I suggest that species which occur infrequently be eliminated from consideration. Studies in which sample size effects are not considered should include sample size information in sufficient detail that other workers might make their own evaluation of observed faunal patterns.

Morphologic diversification among rostroconch molluscs is explored using multiple phylogenies derived from different methods. In addition to strict parsimony, phylogenetic estimates were derived using four different methods that employ stratigraphic data. The resultant phylogenies are generally very similar to one another and to the phylogeny proposed by Pojeta and Runnegar (1976). All estimates (including the Pojeta and Runnegar estimate) imply much lower morphologic separations among post-Ordovician rostroconchs (measured here as the frequency of character state change per branch) than among Cambro-Ordovician rostroconchs. However, the data do not suggest that morphologic evolution became more constrained among rostroconchs as a whole, but instead suggest a reduced characteristic rate of morphologic change in the clade that happened to survive the end-Ordovician extinction. Likelihood ratio tests provide strongest support for the hypothesis that morphologic evolution was more constrained within a derived subclade (corresponding to previous definitions of the Conocardioidea, minus the Eopteriidae) than it was in a broad paraphylum (corresponding to the Ribeirioidea + Eopteriidae). Estimates from each of the phylogenetic methods lead to the same conclusions. Various metrics indicate that the pattern is not due to poor sampling of Cambro-Ordovician species and thus merits a biological explanation. Nonphylogenetic analyses of morphologic disparity suggest a similar history of morphologic evolution, including the apparent difference in characteristic rates of morphologic evolution between paraclade and subclade.

Morphological continuity in the fossil record is the principal evidence favoring evolution as a historical explanation for the diversity of life. Continuity is usually discussed on scales broader than the species level. Patterns of morphological variation characteristic of living species are useful in recognizing species on time planes in the fossil record, but the fossil record is rarely complete enough temporally or geographically to preserve more than a fraction of species living in a given interval. Transitions between known species are even rarer. Where transitions are preserved, new species appear to arise through anagenesis (transformation of an ancestral stock producing a modified descendant) and through cladogenesis (subdivision of an ancestral lineage where one or more descendants differ from the ancestral stock). Evolutionary species are often necessarily bounded arbitrarily in the dimension of time.

Orthogenesis and punctuated equilibrium lie at opposite poles in a spectrum of speciation modes. Orthogenesis, highly constrained anagenesis, is probably rare. Cladogenesis appears to differ little from anagenesis once ancestral stocks are segregated. Limited evidence suggests that morphological differentiation during cladogenesis postdates genetic isolation. Hence punctuated equilibrium may be rare as well. Patterns of gradual change over time indicate that morphological evolution is reasonably viewed as continuous within and between species. Rates of evolution vary greatly (continuity does not require constancy). Rate distributions are truncated and biased by limits of stratigraphic completeness and time resolution: moderate to high rates of morphological evolution and species turnover are rarely recorded by fossils. Species durations are poorly characterized, but they appear to be so variable that there is no suggestion of periodicity. Species longevity is unpredictable. The episodic nature of faunal turnover suggests that extrinsic environmental factors rather than intrinsic homeostatic factors govern evolution at the species level.

We report on the bone microstructure of the Late Cretaceous birds Patagopteryx deferrariisi and members of the Enantiornithes. These birds represent the most primitive birds ever studied histologically. The occurrence of growth rings indicating alternating periods of slowed and fast growth suggests that these basal birds had slower growth rates, and differed physiologically from their modern relatives. Our findings also call into question previous ideas suggesting that nonavian theropods developed a full avian degree of homeothermic endothermy, which was later inherited by birds. On the contrary, our findings suggest that birds developed classic endothermy relatively late in their phylogenetic history.

One of the enduring puzzles to Stephen Jay Gould about life on Earth was the cause or causes of the fantastic diversity of animals that exploded in the fossil record starting around 530 Ma–the Cambrian explosion. In this contribution, we first review recent phylogenetic and molecular clock studies that estimate dates for high-level metazoan diversifications, in particular the origin of the major lineages of the bilaterally-symmetrical animals (Bilateria) including cnidarians. We next review possible “internal” triggers for the Cambrian explosion, and argue that pattern formation, those processes that delay the specification of cells and thereby allow for growth, was one major innovation that allowed for the evolution of distinct macroscopic body plans by the end of the Precambrian. Of potential “external” triggers there is no lack of candidates, including snowball earth episodes and a general increase in the oxygenation state of the world's oceans; the former could affect animal evolution by a mass extinction followed by ecological recovery, whereas the latter could affect the evolution of benthic animals through the transfer of reduced carbon from the pelagos to the benthos via fecal pellets. We argue that the most likely cause of the Cambrian explosion was the evolution of macrophagy, which resulted in the evolution of larger body sizes and eventually skeletons in response to increased benthic predation pressures. Benthic predation pressures also resulted in the evolution of mesozooplankton, which irrevocably linked the pelagos with the benthos, effectively establishing the Phanerozoic ocean. Hence, we suggest that the Cambrian explosion was the inevitable outcome of the evolution of macrophagy near the end of the Marinoan glacial interval.

Consideration of the ways in which ontogenetic development may be modified to give morphological novelty provides a conceptual framework that can greatly assist in formulating and testing hypotheses of patterns and constraints in evolution. Previous attempts to identify distinct modes of ontogenetic modification have been inconsistent or ambiguous in definition, and incomprehensive in description of interspecific morphological differences. This has resulted in a situation whereby almost all morphological evolution is attributed to heterochrony, and the remainder is commonly either assigned to vague or potentially overly inclusive alternative classes, or overlooked altogether.

The present paper recognizes six distinct modes of ontogenetic change, each a unique modification to morphological development: (1) rate modification, (2) timing modification, (3) heterotopy, (4) heterotypy, (5) heterometry, and (6) allometric repatterning. Heterochrony, modeled in terms of shape /time /size ontogenetic parameters, relates to parallelism between ontogenetic and phylogenetic shape change and results from a rate or timing modification to the ancestral trajectory of ontogenetic shape change. Loss of a particular morphological feature may be described in terms of timing modification (extreme postdisplacement) or heterometry, depending on the temporal development of the feature in the ancestor. Testing hypotheses of the operation of each mode entails examining the morphological development of the ancestor and descendant by using trajectory-based studies of ontogenetically dynamic features and non-trajectory-based studies of ontogenetically static features.

The modes identified here unite cases based on commonalities of observed modification to the process of morphological development at the structural scale. They may be heterogeneous or partially overlapping with regard to changes to genetic and cellular processes guiding development, which therefore require separate treatment and terminology. Consideration of the modes outlined here will provide a balanced framework within which questions of evolutionary change and constraint within phylogenetic lineages can be addressed more meaningfully.

Phylogenetic patterns of trilobite clades were used to deduce biogeographic patterns during the Middle Devonian, a time of active plate collision between North America (Laurentia) and other plates, coincident with several major episodes of sea-level rise and fall. The mapping of biogeographic states onto phylogenies for asteropyginid and proetid trilobites indicated that during their history these trilobite clades often shifted the areas they occupied, and also underwent vicariant differentiation, followed by range expansion, followed by subsequent vicariance. Biogeographic patterns in these individual phylogenies were evaluated and synthesized using a modified version of Brooks Parsimony Analysis, which is discussed. This method makes it possible using cladistic methods to distinguish between episodes of vicariance and episodes of dispersal. Two types of dispersal are recognized herein: (1) the individualistic responses of certain taxa in a single clade that cannot be generalized, i.e., traditional ad hoc dispersal, and (2) those patterns of congruent range expansion that are replicated across several clades. The latter are not treated as true dispersal, expansion of a taxon's range over a barrier accompanied by diversification, but rather as a result of the temporary removal of barriers to marine taxa, due either to relative sea-level rise or to the collision of formerly disjunct plates. These are interpreted as changes in the structure of areas, and this type of dispersal is referred to as geo-dispersal. Geo-dispersal was found to have occurred in the Middle Devonian trilobite fauna of Eastern North America.

Biogeographic analysis indicated that Eastern North America is a strongly supported area, with the Appalachian and Michigan Basins as sister areas. Armorica and the Canadian Arctic are also sister areas. Congruence was found between area cladograms produced by vicariance and dispersal analyses for Middle Devonian trilobites, suggesting that in some cases the geological processes governing vicariance, such as sea-level changes, were the same as those that caused dispersal.

Raup's (1976a) data on Phanerozoic species numbers are examined for species-area relationships, using published estimates of areas of continental seas. By means of multiple regression, species numbers are regressed on both estimated areas of seas and amounts of available rock for sampling, as measured by outcrop area and rock volume. Although the sampling effects apparently have the strongest influence on fossil species diversity, areas of seas substantially increase the total correlation, suggesting that Phanerozoic species numbers were in equilibrium with habitat area. This is further supported by the fact that estimated parameters in the regressions are fairly consistent with established island biogeographic theory. Much of the remaining residual variation can be explained by periods of disequilibrium.

Morphologic analysis of 281 species of ammonoids from Great Britain, the North American mid-continent, and the South Urals, at eight successive levels within the Namurian Series (ca. 18 Myr duration), using bivariate plots and principal-components analysis, permits definition of morphologic diversity and identification of morphotypic patterns in time and space. Namurian ammonoids exhibit the same general range of shell geometry that characterizes ammonoids as a whole; there were few post-Namurian innovations in the basic geometry of planispiral ammonoids. Within this overall range of geometry, there are eight preferred morphotypes: two were phylogenetically monopolized by long-ranging forms; three were generalized and reoccur in successive horizons; two others were homeomorphically utilized at different times by different lineages; and one represents morphologic innovation followed by radiation. Such patterns seem to represent combined effects of function, phylogeny, and ecology. Synchronous variations in isolated successions suggest global controls such as eustatic sea-level fluctuations, whereas provincial differences in diversity may be attributable to paleogeographic and ecologic factors. We predict that the Namurian record of ammonoid morphologic diversity and change will be found to be distinctive and differentiable from earlier and later intervals.

Most functional interpretations of ziphodont dentition are based on limited morphometric, behavioral, and taphonomic studies, but few are based on controlled observations of a modern ziphodont consumer. The purpose of this study is to determine through controlled feeding observations if the behaviors indicative of a ziphodont consumer are reflected by tooth marks left on bone surfaces by Varanus komodoensis, the Komodo monitor. We document feeding behavior, expand upon dental function, and correlate these aspects with tooth mark production. We also discuss the significance and limits of applying these data to fossil assemblages.

Goat carcasses were fed to 11 captive individuals. V. komodoensis modifies bone surfaces extensively. Individuals exhibit a “medial-caudal arc'7 when defleshing, followed by inertial swallowing. Bone crushing was not observed. The vast majority of tooth marks are scores, with pits being significantly less common. Tooth furrows and punctures are rare. “Edge marks” are produced on flat elements. Marks are elongate and narrow, with variable lengths and curvature. Over one-third of the marks occur within parallel clusters. Striations are evident on 5% of all marks.

Both feeding behavior and tooth marks indicate that ziphodont crowns are ideal for defleshing by being drawn distally through a carcass. Crowns are poorly built for crushing, and within-bone nutrients are acquired through swallowing. Mark production is a by-product of the distal crown movement during the flesh removal process. Scores are the consequence of apical dragging. Edge marks and striated scores result respectively from distal and mesial carinae contact. Mark curvature is the consequence of arcing motions. Parallel clusters may result from repetitive defleshing strokes and/or from multiple crown contacts during a stroke.

These observations can be used to draw functional, behavioral, and taphonomic interpretations from fossil assemblages. When they are provisionally applied to theropod tooth marks, similar crown function and defleshing behavior with little bone crushing is apparent. Differences occur concerning mark frequency and curvature, relating potentially to taphonomic biases and rostral motion, respectively.

The mid-late Ediacaran Period (~579–541 Ma) is characterized by globally distributed marine soft-bodied organisms of unclear phylogenetic affinities colloquially called the “Ediacara biota.” Despite an absence of systematic agreement, previous workers have tested for underlying factors that may control the occurrence of Ediacaran macrofossils in space and time. Three taxonomically distinct “assemblages,” termed the Avalon, White Sea, and Nama, were identified and informally incorporated into Ediacaran biostratigraphy. After ~15 years of new fossil discoveries and taxonomic revision, we retest the validity of these assemblages using a comprehensive database of Ediacaran macrofossil occurrences. Using multivariate analysis, we also test the degree to which taphonomy, time, and paleoenvironment explain the taxonomic composition of these assemblages. We find that: (1) the three assemblages remain distinct taxonomic groupings; (2) there is little support for a large-scale litho-taphonomic bias present in the Ediacaran; and (3) there is significant chronostratigraphic overlap between the taxonomically and geographically distinct Avalonian and White Sea assemblages ca. 560–557 Ma. Furthermore, both assemblages show narrow bathymetric ranges, reinforcing that they were paleoenvironmental–ecological biotopes and spatially restricted in marine settings. Meanwhile, the Nama assemblage appears to be a unique faunal stage, defined by a global loss of diversity, coincident with a noted expansion of bathymetrically unrestricted, long-ranging Ediacara taxa. These data reinforce that Ediacaran biodiversity and stratigraphic ranges of its representative taxa must first statistically account for varying likelihood of preservation at a local scale to ultimately aggregate the Ediacaran macrofossil record into a global biostratigraphic context.

Many areas of paleobiological research require reliable extinction metrics. Branching-and-extinction simulations and data on Phanerozoic marine families and genera are used to investigate the relationship between interval length and commonly used extinction metrics. Normalization of extinction metrics for interval length is problematic, even when interval length is known without error, because normalization implicitly assumes some model of variation in extinction risk within an interval. If extinction risk within an interval were constant, or if it varied but played no role in the definition of stratigraphic intervals, then Van Valen's time-normalized extinction metric would provide a measure of average extinction risk that is effectively unbiased by interval length. When extinction risk varies greatly within an interval and interval boundaries are drawn at times of heavy extinction, extinction metrics that normalize for interval length are negatively correlated with interval length. Despite its intuitive appeal, the per-taxon extinction rate (proportional extinction per million years) is biased by interval length under a wide range of extinction models.

Empirically, time-normalized extinction metrics for Phanerozoic families and genera are negatively correlated with interval length. This is consistent with an extinction model in which many times of very low risk are punctuated by a few times of very high risk which in turn determine stage boundaries. Origination and extinction patterns are similar, but origination intensity varies less among stages than extinction intensity. This observation has at least two plausible explanations: that origination episodes are more protracted than extinction episodes, and that biologic groups do not respond in unison to origination opportunities the way they seem to respond during extinction events. For families and genera, there is enough variation in extinction intensity among stages that stage length can be ignored when studying certain extinction patterns over the entire Phanerozoic.

Pterosaurs were the first flying vertebrates and formed important components of terrestrial and marginal marine ecosystems during the Mesozoic. They became extinct during the latest Cretaceous (latest Maastrichtian), at, or near, the Cretaceous/Paleogene boundary, following an apparent decline in diversity in the Late Cretaceous. This reduction in species richness has been linked to the ecological radiation of birds in the Early Cretaceous and the proposal that birds competitively excluded pterosaurs from many key niches. However, although competition is often posited as a causal mechanism for many of the clade-clade replacements observed in the fossil record, these hypotheses are rarely tested. Here we present a detailed examination of pterosaur diversity through time, including both taxic and phylogenetically corrected diversity estimates and comparison of these estimates with a model describing temporal variation in the number of pterosaur-bearing formations (a proxy for rock availability). Both taxic and phylogenetic diversity curves are strongly correlated with numbers of pterosaur-bearing formations, suggesting that a significant part of the signal contained within pterosaur diversity patterns may be controlled by geological and taphonomic megabiases rather than macroevolutionary processes. There is no evidence for a long-term decline in pterosaur diversity through the Cretaceous, although a reduction in morphological, ecological, and phylogenetic diversity does appear to have occurred in the latest Cretaceous. Competitive replacement of pterosaurs by birds is difficult to support on the basis of diversity patterns.

For some higher taxa, species can be identified in the fossil record with a high degree of reliability. The great geological durations of species indicate that phyletic evolution is normally so slow that little change occurs within a lineage during 105–107 generations. Failure to recognize sibling species in the fossil record has no bearing on this conclusion because they embody virtually no morphological change. Although slowness is the rule, we have no more precise assessment of morphological rates of phyletic evolution for any major taxon. Morphological data that have been assembled to assess rates of phyletic evolution have been meager, unrepresentative, and predominantly reflective of nothing more than body size. Net selection pressures within long segments of phylogeny—even ones that embrace large amounts of evolution—are infinitesimal and seemingly unsustainable against random fluctuations. This suggests that natural selection operates in a highly episodic fashion.

Rates of adaptive radiation and extinction at the species level can be estimated for many taxa and, from them, rates of speciation in adaptive radiation. Species selection should universally tend to increase rate of speciation and decrease rate of extinction, yet these rates are positively correlated in the animal world, apparently because they are linked by common controls: both rate of speciation and rate of extinction seem to increase with level of stereotypical behavior and to decrease with dispersal ability. Only a few “supertaxa” have been able to combine high rates of speciation with moderate rates of extinction.

The sequences of skeletal disarticulation in a broad range of African mammals in a tropical savanna environment suggest that in general the process is very consistent. There is some variation among species, but this cannot yet be convincingly related to higher taxonomic categories or to body size. Disarticulation begins shortly after death, proceeds more or less continuously, and is almost complete after 5 yr. Because of the overall predictability of disarticulation, the stage of disarticulation reached by comparable fossil skeletons provides useful information about their taphonomic history prior to burial.

High-resolution palynological data sets from shallow marine Triassic-Jurassic (Tr/J) boundary beds of two principal sections in Europe (Hochalplgraben in Austria and St. Audrie's Bay in the United Kingdom) were analyzed to reconstruct changes in vegetation, biodiversity, and climate. In Hochalplgraben, a hardwood gymnosperm forest with conifers and seed ferns is replaced by vegetation with dominant ferns, club mosses and liverworts, which concurs with an increased diversification of spore types during the latest Rhaetian. Multivariate statistical analysis reveals a trend to warmer and wetter conditions across the Tr/J boundary in Hochalplgraben. The vegetation changes in St. Audrie's Bay are markedly different. Here, a mixed gymnosperm forest is replaced by monotonous vegetation consisting mainly of Cheirolepidiaceae (80–100%). This change is caused by a transition to a warmer and more arid climate. The observed diversity decrease in St. Audrie's Bay affirms this interpretation. Although both sections show major vegetation changes, neither of them demonstrates a distinctive floral mass extinction. A compilation of Tr/J boundary sections across the world demonstrates the presence of Cheirolepidiaceae-dominated forests in the Pangaean interior and increases in abundance of spore-producing plants adjacent to the Tethys Ocean. We propose that the non-uniform vegetation changes reflected in the Tr/J palynological records are the result of environmental changes caused by Central Atlantic Magmatic Province volcanism. The increase in greenhouse gases caused a warmer climate and an enhanced thermal contrast between the continent and the seas. Consequently, the monsoon system got stronger and induced a drier continental interior and more intensive rainfall near the margins of the Tethys Ocean.

Late Paleocene planktic foraminifera exhibit strong positive correlations between carbon isotopic values and test mass, but negative correlations between oxygen isotopic values and test mass. Based on analogy with modern taxa, these trends are probably ecotypic and may or may not apply to an ontogenetic series. Among Acarinina and Morozovella species, the magnitude and direction of these trends resemble those of modern planktic foraminifera with dinoflagellate photosymbionts. This is consistent with current models of photosymbiosis and calcification in planktic foraminifera and suggests that Acarinina and Morozovella relied heavily on photosymbionts for nutrition.

Acarinina and Morozovella species resemble modern, strongly photosymbiotic taxa in their association with low and mid latitude nearsurface water masses. However, their test morphologies differ greatly from those of extant taxa that bear obligate photosymbionts. Earliest Paleocene taxa that exhibit a similar paleohabitat association and similar size-related isotopic trends are characterized by still different test morphologies. These comparisons suggest that (1) throughout geologic time, strong reliance on photosymbiont activity has been closely linked to habitat, but not to test morphology; (2) photosymbiosis has been a common and convergently evolved strategy of planktic foraminifera over geologic time, and (3) modern relationships between planktic foraminiferal test morphology and photosymbiont dependence are largely an artifact of geologically recent phylogenetic relationships and shared ecologic strategies.

Intersite comparison suggests that the stable isotopic signals of narrowly constrained size fractions of a late Paleocene Acarinina or Morozovella species can be used to reconstruct the magnitude and direction of relative variation in equilibrium stable isotopic values throughout its geographic and temporal range. This is supported by analogy with extant photosymbiotic taxa. However, since photosynthetic depletion of 12C leaves 13C-enriched HCO3-for calcification, the carbon isotopic values of Acarinina and Morozovella tests may have been consistently greater than paleoseawater values. Failure to account for this effect could lead to overestimation of late Paleocene marine productivity.

The well-known decline of global background extinction intensity was caused by the sorting of higher taxonomic groups. Two factors were responsible. First, probabilities of familial origination and extinction in these groups (taxonomic orders) were highly correlated. Groups whose families had high probabilities of origination and extinction tended to have highly volatile diversity paths and, consequently, short life spans. Second, orders with high probabilities of familial origination and extinction were rarely replaced by new high-turnover orders. Thus, because high-turnover orders tended to become extinct without replacement, the global background extinction intensity declined. Since familial origination and extinction probabilities are correlated, global background origination intensity inevitably declined as well. As a consequence of these processes, virtually all groups of organisms now living have low probabilities of familial origination and extinction.

Simulations of branching evolution were used to obtain the expected relationships among probabilities (of origination and extinction), volatilities, and longevities for the entire range of possible probabilities, and these relationships were compared to those obtained from the empirical record. In the simulations, only the probabilities of origination and extinction were specified, so volatilities and clade longevities were determined entirely by the probabilities. The similarity between results obtained by simulation and those obtained by analysis of the empirical record further supports the inference that the observed decline of background extinction (and origination) intensity can be explained largely by the loss of high-probability groups to induced volatility.

Five generalizations derived from the shell form of prosobranch gastropods are developed. (1) A univalve of more than one volution whose aperture lies in a plane that passes through the axis of coiling does not live with the aperture parallel to the substrate. (2) A univalve of more than one volution whose aperture lies in a plane that is tangential to the body whorl does live with the plane of the aperture parallel to the substrate. (3) Gastropods with tangential apertures, when extended, support the shell so that the center of mass of the shell and its contents is over the midline of the cephalopedal mass; this balancing of the shell may be accomplished either by regulatory detorsion, by inclination or by a combination thereof, to keep the center of gravity of the shell as low as possible. (4) Angulations or re-entrants in the gastropod aperture are usually indicative of inhalent or exhalent areas; inhalent areas are directed as far anteriorly as possible. (5) Gastropods having elongated apertures possess only a single gill and develop a water current through the mantle cavity from anterior to posterior along the long axis of the aperture; this axis is subparallel to the anterior-posterior axis of the foot.

These generalizations are then used as the basis for some deductive interpretations of behavioral modes of Paleozoic Gastropoda.