Siwalik rocks of Pakistan are a virtually continuous, continental sedimentary sequence, extending in age from 18 to 1 ma b.p. This paper describes taphonomic features of late Miocene mammalian assemblages from a highly fossiliferous interval about 400 m thick, based on field documentation of sedimentary environments at 42 fossil localities and systematic fossil collection of 21 localities.

Within a broadly fluvial system, I recognize four sedimentary environments of bone accumulation, distinguished by lithology, unit-thickness, unit-geometry, contacts, sedimentary structures, and relationship to adjacent units. Each environment corresponds to an association of lithofacies. Facies Association I is interpreted as the persistent, major channel bodies of a meandering fluvial system; Facies Association II as coarse-grained flood deposits, such as crevasse splays, deposited beyond the main channels; Facies Association III as channel margins, including levees and swales; and Facies Association IV as predominantly subaerial floodplains.

Taphonomic features of bone assemblages from each facies association include skeletal-element composition, surface distribution of specimens, degree of articulation, hydraulic equivalence between organic and inorganic sedimentary particles, frequency of juvenile remains, size distribution of fauna, and an estimate of duration of accumulation of individual fossil localities. The distribution of these features among the four facies associations suggests that bone assemblages in Facies Associations I and II accumulated by the action of currents in river channels or floods, whereas bone assemblages in Facies Associations III and IV accumulated through concentration by biological agents and/or attrition at a repeatedly used site of predation.

Inclusion in fluvial accumulations depends on initial availability of skeletal remains and hydraulic characteristics of individual skeletal elements, but not taxonomic identity per se. For biological accumulations, however, taxonomic composition reflects the preferences of the individual agents of accumulation. The probability of preservation of taxa in fluvial accumulations is probably mainly a function of body size, as reflected in the sizes of isolated skeletal elements. Thus, in this Siwalik system, bone assemblages that experienced fluvial transport are better representations of original community composition than bone assemblages created by biological agents or passive accumulation.

Experiments were conducted to test the hypothesis that biomechanical constraints determine the morphology of regular echinoids. Hard-bottom-dwelling Tripneustes gratilla elatensis were transferred to an artifical sandy habitat to evaluate whether the change in substrate affects their height to diameter ratio (H/D). Within 1–2 months their H/D ratio increased significantly. This change was shown to be reversible to some extent. Surgical damage to the ambulacral system of one ray caused inactivation of tubefeet and atrophy of injured ambulacra. Test shape was also affected: the damaged ray was lower, and the nondamaged ambulacra deflected toward the treated one, producing bilateral symmetry as in recorded cases of teratology. Study of T. g. elatensis tetramers showed that while “perfect” tetramery was apparently associated with genetic aberration, “imperfect” tetramery results from mechanical injury at an early ontogenetic stage. Micromorphological study shows that in the longitudinal sutures, normally under tension, long and slender trabeculae develop, associated with long and well-aligned collagenous sutural fibers, while the latitudinal trabeculae and fibers are short and less organized. A mechanical effect is suggested by the oval cross-section of the fiber-anchoring trabeculae. Further, echinoid plates interact like soap bubbles, whereas the entire test behaves like a balloon, fastened to the substrate by the ambulacral tubefeet. All these observations support earlier hypotheses on the biomechanical control of echinoid test growth. A model is proposed in which the expansion of the inner mass, counteracted by the mechanical activity of the ambulacral tubefeet, mesenterial threads, and lantern muscles, affects sutural growth, thus controlling echinoid morphogenesis.

A morphometric survey among regular echinoids reveals an inverse relationship between ambulacral width and relative ambital height. Although both increase of ambulacral width and lowering of ambitus-line are evolutionary trends, it is suggested that they are a response to a mechanical effect. H/D ratio was not related to ambulacral width on the phylogenetic level. It is therefore suggested that the latter correlation is ontogenetically controlled. Aspects of irregular echonoid evolution, such as bilateral symmetry, flattening, and formation of the ambulacral petaloid, also are explained by this model.

The dominance of Paleozoic articulate brachiopods in once-muddy environments may be explained by an array of mechanisms and structures that reject nonfood particles, in some cases without interruption of feeding: (1) behavioral flexibility of the lophophore and its individual filaments; (2) persistent, variable-speed rejection currents on the mantle, which sometimes concentrate pseudofeces in topographically controlled vortices; (3) costae and alae (which have many other probable functions); (4) inhalant currents elevated above substrate; (5) marginal setae.

Some mantle currents parallel (and presumably augment) lophophore feeding currents; others diverge up to 90° to provide rejection while feeding continues. Contrary to previous reports, the lateral cilia seem to be involved in rejection and may reverse.

Repeated claims for the superiority of the gill of suspension-feeding bivalves over the “weak” individual filaments of the lophophore are probably false. In suspension-feeding bivalves, simultaneous feeding and rejection are likely to be hindered by fused gill elements and mucus-trapping of food. The energetically efficient articulates are predicted to have a competitive advantage over suspension-feeding bivalves when oxygen or food is limiting, as, for example, after a bolide impact.

Morphometric examination of cladogenesis and phyletic evolution in two late Neogene sister lineages of marine microfossils (Pterocanium prismatium and P. charybdeum, Radiolaria) from two equatorial Pacific sediment cores was undertaken to better understand the rate of cladogenesis and its relation to subsequent phyletic change. The origin of P. prismatium from P. charybdeum ∼4 ma ago has been estimated to take place over an interval of ∼500,000 yr. Results show that the speciation event consists of two distinct phases. The first phase, cladogenesis, occurred relatively rapidly (on the order of 50,000 yr). A second phase of relatively rapid divergent phyletic evolution away from the common ancestral state followed in both descendant branches and continued for at least 500,000 yr after completion of the cladogenetic event. Net evolutionary rates over the next 2 ma appear to be much lower. Individual characters change by as much as 2 population standard deviations during cladogenesis, and by a total of approximately 3 standard deviations over 2.5 ma of phyletic evolution. Up to 5 population standard deviations of change during ≦ 50,000 yr of cladogenesis, and 7 additional standard deviations of phyletic change over 500,000 yr are observed in multivariate (discriminant function) indices of morphologic difference. The measured pattern does not appear to be either strictly “punctuated” or strictly “gradual,” but instead shows features of both hypotheses.

Inadequacies in stratigraphic resolution or completeness can make true rates of morphologic change through geologic time impossible to estimate precisely. However, relative rates may be sufficient to test whether the tempo of change within species can account for morphologic differences across species boundaries, and hence to distinguish between gradual and punctuated patterns of evolution. The conditions under which these patterns can be distinguished statistically are explored by simulating varying degrees of within-species rate variability relative to across-species morphologic difference. The statistical methods are then applied to multiple-character morphologic data from closely spaced sequential populations of the Neogene bryozoan Metrarabdotos, using discriminant analysis to compare overall morphologies. In nine comparisons of ancestor-descendant species pairs, all show within-species rates of morphologic change that do not vary significantly from zero, hence accounting for none of the across-species difference. In all cases the ratio of within-species fluctuation to across-species difference is low enough to allow the punctuated pattern to be distinguished with virtual certainty. In at least seven of the cases, ancestor species persisted after giving rise to descendants, in conformity with the punctuated equilibrium mode of evolution.

The European Plio-Pleistocene Mimomys (Hintonia) occitanus-ostramosensis lineage of arvicolid rodents appears to present a genuine case of gradual evolution. Analysis of numerous dental samples recorded from 3 to 1.5 ma by several biometrical methods led to the following conclusions: samples from the whole geographical area order into a morphological continuum (chronomorphocline); the successive populations in the trend show important overlap of their variation; and the absence of stasis or any interruption in the trend, combined with its unidirectionality, suggests genuine phyletic gradualism. The chronomorphocline is characterized by an important increase in tooth hypsodonty, by the progressive and continuous development of lateral enamel tracks. Four phases of evolution can be recognized in the lineage, the rates of morphological changes being variable and related to climatic fluctuations (Reuverian, Pretiglian, Tiglian, Eburonian). Morphological changes are initiated by environmental fluctuations, although the polarity of climatic change is not a determining factor. Environmental changes act as stimulus and release drift processes. Phyletic gradualism occurring over such a wide area implies extensive gene flow, and suggests that only general, not local, environmental components act to produce the observed morphological changes.

In a nonhierarchical world, where selection on organisms regulated all nonrandom evolutionary change, the traditional equation of selection (a cause of sorting) with sorting itself (differential birth and death among varying organisms within a population) would rarely lead to error, even though the phenomena are logically distinct (for sorting is a simple description of differential “success,” and selection a causal process). But in a hierarchical world, with entities acting as evolutionary individuals (genes, organisms, and species among them) at several levels of ascending inclusion, sorting among entities at one level has a great range of potential causes. Direct selection upon entities themselves is but one possibility among many. This paper discusses why hierarchy demands that sorting and selection be disentangled. It then presents and illustrates an expanded taxonomy of sorting for a hierarchical world. For each of three levels (genes, organisms, and species), we show how sorting can arise from selection at the focal level itself, and as a consequence either of downward causation from processes acting on individuals at higher levels or upward causation from lower levels. We then discuss how hierarchy might illuminate a range of evolutionary questions based on both the logical structure of hierarchy and the historical pathways of its construction—for hierarchy is a property of nature, not only a conceptual scheme for organization.