An investigation of shell morphometrics of the prosobranch genus Paramelania from Lake Tanganyika shows striking contrasts between some live and dead populations from identical localities. Living populations and surficial dead shells were collected from 15 sublittoral-profundal localities along the east side of the lake. Interpopulation variability in this endemic gastropod is clinal (N-S) for several shell characters. Intrapopulation variability of dead shell populations frequently exceeds that of live populations. Distinctive morphs may be present in a local dead shell population which are absent in the live population from the same locality. However, the phenodeviant dead shell morphology may occur among live snails elsewhere in the lake. Phenodeviant shells may be encrusted, but are unabraded and show no preferred orientation.

Radiocarbon ages of >2000 yr b.p. on dead phenodeviants suggest that surficial shell accumulations in Lake Tanganyika are strongly time-averaged. Although wave activity and biogenic concentration are important processes in Lake Tanganyika, neither is sufficient under modern lake conditions to account for these admixed assemblages. Winnowing and in situ stratigraphic condensation during periods of lowered lake levels is the most likely explanation for the occurrence of phenodeviant surficial shells. This hypothesis is consistent with the observation that extensive shell lags are exposed on the lake floor in areas currently inhabited by the same snails.

A model of biogeographic range fluctuations for particular Paramelania morphs during the Holocene, in concert with lake level fluctuations, can account for the shell assemblages seen on the lake floor today. The complex history of these accumulations suggests that taphonomic admixtures may obscure the interpretation of evolutionary sequences in the lacustrine stratigraphic record.

A new hypothesis concerning the evolutionary origin of pentameral symmetry in echinoderms is presented. The BA-A-BA pattern of Lovén's law in echinoids and the 2-1-2 symmetry of the edrioasteroid Stromatocystites appear to be morphogenetically related, and this pattern appears to be locked into development. This pattern may have originated from a unirayed 0-1-0, —A—, ancestor. I propose that a duplication of the uniray occurred and resulted in the addition of a pair of rays that followed Bateson's rules of symmetry to form a three-rayed 0-1-2, —A-BA, construction. This change occurred on the side that corresponds to the left side of the organism. This event made the individual asymmetric with respect to its anterior-posterior axis. Therefore I propose that morphogenetic regulation of bilaterality of the organism then led to homeotic expression of a mirror-image pair of rays on the opposite side. These two morphogenetic steps achieved the 2-1-2, BA-A-BA pattern. “Appendage status” of rays is assumed necessary to invoke the Batesonian mirror-image duplications of the model.

Three robust morphological characters emerge from the “rays-as-appendages” model: (1) 2-1-2, BAo-A-BA, organization; (2) “locked-in” pentamerism; and (3) a 2-3 pattern of right and left rays. Results based on ray homology research are presented for echinoids, asteroids, ophiuroids, edrioasteroids, ophiocistioids, and holothurians. I speculate that helicoplacoids may have 0-1-2 triradiate construction, and that solutes may have 0-1-0 uniray construction. The model has limits; it does not explain 1-1, or 2-1, or 2-2 or 1-1-1 organization of ambulacra.

By digitizing the enamel configuration and calculating the overall directional tendencies of the enamel edges in complex equid dental patterns, quantitative changes in functional attributes were traced through a phylogenetic series. Pronounced changes both in the directional emphasis of the enamel edges and in the chewing motion occurred and were interrelated. Two distinct trends were found. From early Eocene to Oligocene, edge alignment became increasingly confined to a single direction, but the teeth retained the phase I and phase II chewing angles characteristic of many primitive herbivorous and omnivorous mammals. The chewing direction in the horizontal plane also remained essentially unchanged. During the Oligocene and the early Miocene, both edge direction and chewing direction remained stable. However, the middle Miocene genus Merychippus represents a stage in which the enamel directional emphasis was lost in transition to a new anteroposterior emphasis, a trend that continued into the late Tertiary equine taxa. Concurrent with this transformation was a loss of all but vestiges of the phase I and II chewing angles in the vertical plane and a shift of the chewing direction in the horizontal plane to an almost transverse movement of the jaw.

In the Oligocene equids, the edges of the labial cusps did attain perpendicularity to the phase I chewing direction, which was oblique to the horizontal plane, but the chewing direction in the horizontal plane was far from perpendicular to the lingual enamel edge direction because the lingual cusps were configured for efficiency in compressive, not shearing, motion that dominated the phase II surfaces. In the later Tertiary the chewing direction and edges attained essentially a perpendicular relationship, which represents the theoretical optimum for a shearing system. The radical change in edge direction in the Miocene probably occurred as a result of an increase in translative movement (longer glide) between the surfaces, making the lingual edges subject to selection for perpendicularity and flattening of the occlusal surface, resulting in a condition where a single edge direction became optimal for all cusps. The extended chewing stroke translation was perhaps the key event initiating the other changes and would have resulted from increased chewing effort accompanying a shift to a diet of mainly grass. The loss of the dual (shearing and compressive) function of the occlusal surface suggests that the earlier diet may have contained significant proportions of other plant parts, such as fruits, seeds, or tender leaves that might be optimally chewed by compressive movements.

A new turnover rate metric is introduced that combines simplicity and precision. Like the related three-timer and gap-filler equations, it involves first identifying a cohort of taxa sampled in the time interval preceding the one of interest (call the intervals i0 and i1). Taxa sampled in i0 and i1 are two-timers (t2); those sampled in i0 and i2 but not i1 are part-timers (p); and taxa sampled only in either i1, i2, or i3 are newly notated here as either s1, s2, or s3. The gap-filler extinction proportion can be reformulated as (s1−s3)/(t2+p). The method proposed here is to substitute s3 with the second-highest of the three counts when the expected ordering s1≥s2≥s3 is violated. In simulation, this new estimator yields values that are highly correlated with those produced by the gap-filler equation but more precise. In particular, it rarely produces highly negative values even when sample sizes are quite small. It is mildly upwards biased when sampling is extremely poor and turnover rates are extremely low, but it is otherwise highly accurate. Examples of Phanerozoic extinction rates for four major marine invertebrate groups are given to illustrate the method’s improved precision. Based on the results, the procedure is recommended for general use.

Carbon (δ13C) and oxygen (δ18O) isotopic results are presented from 42 tooth enamel carbonate samples of rhinos (Family Rhinocerotidae) from a sequence of Florida Neogene localities between 9.5 and 4.5 Ma. These data are used to interpret ancient diets and test previous hypotheses of terrestrial/aquatic adaptations of two sympatric rhinos, Aphelops and Teleoceras. The long-limbed, shorter-crowned Aphelops traditionally has been reconstructed as an open-country browser (similar to the modern black rhino), whereas short-limbed, higher-crowned Teleoceras traditionally has been reconstructed as an amphibious grazer (similar to the modern hippo). Between about 9. 5 and 7 Ma the δ13C values (all <−11.0‰) from Florida Aphelops and Teleoceras indicate that both rhinos were feeding on C3 plants. This diet probably included a combination of browse and C3 grasses, although the exact proportions for each genus cannot be distinguished isotopically. In contrast, after the late Miocene global carbon shift as represented at 4.5 Ma in Florida, Aphelops was a browser (mean δ13C = −11.9‰), whereas Teleoceras was a mixed feeder/C4 grazer (mean δ13C = −7.0‰). Oxygen isotopic values indicate that neither Aphelops nor Teleoceras was principally aquatic. Given these new isotopic data, more plausible modern analogs for these two extinct rhinos are, respectively, the terrestrial browsing black rhino (Diceros bicornis) and the terrestrial grazing white rhino (Ceratotherium simum), which are sympatric today in Africa.

In the fossil record, biogeographic data at the species level are normally not adequately preserved to be utilized in the reconstruction of marine paleoprovinces; consequently, higher taxa are employed. Thus it is cogent to question whether modern marine provinces, which are generally recognized by their species content, are still distinctive at higher taxonomic levels. Similarity coefficients computed from the bivalve and gastropod faunas of six contiguous and five spatially separated modern provinces indicate that the present-day marine provinces are in fact recognizable at the generic and in most cases at the familial level. The deficiencies and biases of the fossil record obscure the distinctiveness and reduce the precision with which ancient provinces can be delimited. Nevertheless, paleoprovinces that are recognized from the distribution patterns of genera and families are clearly comparable to modern provinces that are determined by species distributions.

Taphonomic analysis of several late Pleistocene mastodon (Mammut americanum) skeletons excavated in southern Michigan provides compelling evidence of mastodon butchery by Paleo-Indians. The occurrence of butchery and details of butchering technique are inferred primarily from patterns of bone modification. An important aspect of butchering practice was production and use of tools fashioned from bones of the animal being butchered. Evidence for butchery and bone tool use includes matching marks on the conarticular surfaces of disarticulated pairs of bones; cutmarks on bones; green bone fracturing; use wear, secondary flaking, and impact features on bone fragments; and burned bone. Interpretation of these features is facilitated by information on patterns of bone distribution and disarticulation preserved in a primary depositional context. Preliminary comparisons among nine sites indicate that putative butchering sites differ consistently and in a variety of ways from sites that appear to record no human involvement. Although based on a small sample of sites, the apparent frequency of butchered individuals relative to those that were not butchered is unexpectedly high. These findings provide new evidence of a well-developed “bone technology” employed by the late Pleistocene human inhabitants of eastern North America. In addition, these data offer circumstantial support for the hypothesis that human hunting was an important factor in the late Pleistocene extinction of mastodons.

Bottom site remote camera photosequences at depths of 73–538 m on forereef slopes in Palau show that Nautilus belauensis is a highly mobile, chemosensitive, epibenthic scavenger and opportunistic predator. The overall depth range of this species is ca. 70–500 m, but photosequences indicate a preferred range of 150–300 m. Nautilus is active both nocturnally and diurnally, locating bait sites within 1–2 h. Associated macrofauna includes caridean shrimps, crabs, and eels; teleosts are rare below 100 m, but sharks are recorded in most photosequences below 250 m. Summarily, Nautilus exhibits a combination of characters that typify deep-sea strategy, including reproductive tactics, growth rate, and population dynamics. This and other evidence suggest that fossil Nautilidae may have been deep-water forms, in contrast to the typically shallower water ammonoids, and that Nautilus is a normal component of the deep forereef rather than a late Cretaceous refugee from shallow water.

It is shown that the abundant Jurassic oyster Gryphaea provides material adequate to test biogeographic and evolutionary hypotheses involving speciation. Phyletic size increase is widespread if not characteristic and tends to be marked by a sharp initial increase followed by a longer period of stability. The larger size of phyletically younger forms is apparently because of greater age of individuals rather than a higher growth rate. Species durations fall within the range of the majority of Jurassic bivalve species. The detailed study of European Gryphaea indicates a pattern of punctuated equilibria allied with morphological trends, some of which are paedomorphic. Gradualistic, species selection and cladistic models of change are not supported. World-wide analysis of Gryphaea species supports centres of origin and migration rather than vicariance, and a relationship exists between migrational and evolutionary events and changes in the physical environment.

Recent Nautilus pompilius from the Fiji Islands and N. macromaphalus from New Caledonia show decreasing cameral liquid volumes relative to total phragmocone volume during ontogeny. A maximal value of 32% of the phragmocone filled with cameral liquid was measured from a 190 g N. pompilius. No specimens of over 500 g total weight of either species exceeded 12%. These figures are in contrast to values derived for seven ammonoid species by Heptonstall (1970), who found values ranging between 19 and 52%.

The relationship between cameral liquid volume and salinity within single chambers engaged in the emptying process are examined in N. pompilius and N. macromphalus. Both species start with newly formed chambers filled with cameral liquid isotonic to seawater. Ionic removal by the siphuncular epithelium rapidly reduces the cameral liquid osmolarity, producing osmotic movement of the cameral liquid into the blood spaces of the siphuncle. In both species the lowest cameral liquid salinities occur when the chamber is slightly over half emptied. After this point, which coincides with decoupling of the cameral liquid from the siphuncle, cameral liquid volume continues to decrease, but cameral liquid salinity increases, indicating that the rate of ionic removal slows relative to liquid removal. In N. macromphalus decoupled cameral liquid salinity rises until it is nearly isotonic to seawater when the chamber is nearly emptied. In N. pompilius, however, the rate of ion removal in decoupled cameral liquid is not slowed as much as in N. macromphalus, since it rarely exceeds 40% seawater osmolarity even when the chamber is nearly emptied. The differences in emptying methods demonstrated in these two species are probably related to their different habitat depths: N. pompilius from Fiji is found in much deeper water and must employ more physiologic work to empty chambers at greater depth.

The depositional history of Upper Miocene through Recent sediments from DSDP Site 214 (Ninetyeast Ridge, Indian Ocean) is reexamined. Samples of the Globorotalia tumida planktic foraminiferal lineage, originally obtained from these sediments by Malmgren et al. (1983), serve as the empirical basis for the recognition of punctuated anagenesis as a distinct mode of phenotypic evolution and have been the subject of numerous additional investigations. However, conclusions reached by previous authors depend strictly on the validity of the original chronostratigraphic interpretation of these sediments. Graphic correlation analysis of first- and last-appearance datum levels for a total of 41 planktic foraminiferal, radiolarian, and calcareous nannoplankton taxa provides evidence for a more complex depositional history at this deep-sea site than originally believed. Based on a conservative model of variation in the pattern of sediment accumulation rates, the lowermost portion of the studied section (6.5-4.3 Ma) represents an interval of temporally condensed sediment accumulation (1.88 cm/1,000 yr) followed by an interval (4.3-2.8 Ma) of temporally expanded sediment accumulation (3.97 cm/1,000 yr). This interval, in turn, is followed by a depositional hiatus or an extremely condensed interval, at least 800,000 yr in duration, which is followed by another relatively condensed (1.36 cm/1,000 yr) interval from 2.0 Ma-Recent. Although this chronostratigraphic reinterpretation deviates substantially from the original, which recognized Site 214 as being both temporally continuous and exhibiting a constant sediment accumulation rate from the Upper Miocene through the Upper Pliocene, it is more consistent with expectations based on Neogene eustatic sea-level fluctuations and global surveys of Neogene hiatus distributions. Age assignments for samples of the Gr. tumida lineage based on the revised chronostratigraphic model reverse some findings of previous investigators with respect to the distinctiveness of phenotypic evolutionary rates characterizing the transition from Gr. plesiotumida to Gr. tumida. Finally, a brief survey of similar marine invertebrate lineage studies shows that changes in the rate of phenotypic evolution often appear to coincide with major physical changes in the paleoceanographic environment. Such correspondences may be due, at least in part, to the effect of these environmental changes on sediment accumulation rates. Paleobiologists who seek to understand patterns of phenotypic change over time must remove the effects of variations in sediment accumulation rates from their data before evolutionary hypothesis testing and remain aware of the limitations imposed on their interpretations by the uncertain nature of chronostratigraphic inference.

The Lower Tertiary Gulf Coast was subjected to repeated transgressive-regressive cycles of varying rate and extent which had a marked effect on species longevities. In some neogastropods dispersal and geographic distribution played important roles in species longevities. Species with planktic larval stages that inhabited broad areas of the Gulf Coast could maintain their distributions even during regressive phases when delta building was strong. Wide distributions, free gene flow, and resistance to isolation gave planktic species an evolutionary stability that resulted in greater species longevity. Species with nonplanktic development could attain wide geographic ranges when barriers were minimized, but they were unable to maintain their distributions in the face of regressive progradation. Thus their populations were isolated, and extinction and speciation accelerated, giving them lesser evolutionary longevity. Although planktic volutids tend to be widespread, planktic fasciolariids and mitrids were often restricted in distribution and therefore probably stenotopic. If dispersal and environmental tolerance are considered as separate forces, they seem to act in combination to produce very long, very short or intermediate species longevities.

Lengths of crinoid stalk segments consisting of multiple columnals (pluricolumnals) from the Fort Payne Formation of south-central Kentucky (Mississippian) were tested for uniformity using simulations. The results indicate that stalk segment lengths are too uniform to have been produced by a random process operating on homogeneous stalks.

The most parsimonious explanation for the uniformity of pluricolumnal lengths is based on analogy with stalks of living isocrinids. The stalks of these crinoids are organized into multicolumnal segments of approximately uniform length: columnals within each segment are connected by “through-going” ligament and “intercolumnal” ligament. The articulation between segments, however, consists of only intercolumnal ligament. Preliminary results from experiments on Recent carcasses indicate that the latter is the primary plane of disarticulation following death.

The preservational pattern of the Mississippian stalks suggests that their ligamentary organization was similar to that of the isocrinids. Since the plane of primary disarticulation in isocrinids corresponds to the autotomy plane, Mississippian crinoids also may have been capable of autotomizing their stalks.

The faunas of three fossil sites in Africa that contain time successive groups of fossil Hominoidea have been analyzed to determine their paleoenvironments. The basis for the analysis is an assessment of the ecological diversity of the fauna, which is expressed in terms of four categories: taxonomic composition, body size, feeding habits and locomotor zonal adaptation. This method has shown that the community structures of the three fossil faunas are significantly different, and comparisons with the community structure of modern habitats suggest that the environment of the early Miocene fauna of Songhor, and the primitive apes associated with it, was probably a type of lowland forest; the habitat of Ramapithecus and the Fort Ternan middle Miocene fauna compares best with modern woodland-bushland habitats; and the habitat of Homo habilis at Olduvai appears to have been intermediate between grassland and woodland-bushland. If man evolved from one of the early forest living apes, as seems likely on present evidence, an adaptive shift from forest to non-forest habitats must have occurred at some stage in his evolution. The evidence from Fort Ternan shows that in Africa Ramapithecus made this adaptive shift, and it is also now becoming clear that several genera of Eurasian apes, including Ramapithecus, made a similar environmental change at the same time.

Specific conductance was calculated for secondary xylem in seven Carboniferous stem taxa utilizing an equation derived from the Hagen-Poiseuille relation. Arborescent and lianoid representatives of major pteridophytic (Calamitaceae, Lepidodenraceae, Sphenophyllaceae) and gymnospermous (Cordaitaceae, Medullosaceae) groups were examined. In the calamite Arthropitys communis and the seed plant Cordaites (Cordaixylon sp. and Mesoxylon sp.), conductance corresponded approximately to the low end of the range for both extant conifers and angiosperms. A substantially higher conductance was determined for the wood of Arthropitys deltoides, conforming to the high end of the range for conifers and the low-middle part of the range for angiosperms. The highest conductance values were found in Sphenophyllum plurifoliatum, Medullosa noei, and Paralycopodites brevifolius and corresponded to the middle-high portion of the range for vessel-containing angiosperms. This outcome is particularly significant in light of the fact that tracheary elements in the fossils are imperforate. The results indicate that conductance in secondary xylem of some of the most ancient, woody groups was comparable to that in extant plants and that highly effective conducting tissue developed relatively early in plant evolution. Moreover, it is suggested that the general relationship between wood anatomy, growth habit, and ecology demonstrated for living plants can also be extended back in time to include fossil plants.

A physical, model-based approach to body temperatures in dinosaurs allows us to predict what ranges of body temperatures and what thermoregulatory strategies were available to those dinosaurs. We argue that

1. 1. The huge range of body sizes in the dinosaurs likely resulted in very different thermal problems and strategies for animals at either end of this size continuum.

2. 2. Body temperatures of the smallest adult dinosaurs and of hatchlings and small juveniles would have been largely insensitive to metabolic rates in the absence of insulation. The smallest animals in which metabolic heating resulted in predicted body temperatures ≥ 2°C above operative temperatures (Te) weigh 10 kg. Body temperature would respond rapidly enough to changes in Te to make behavioral thermoregulation possible.

3. 3. Body temperatures of large dinosaurs (>1000 kg) likely were sensitive to both metabolic rate and the delivery of heat to the body surface by blood flow. Our model suggests that they could adjust body temperature by adjusting metabolic rate and blood flow. Behavioral thermoregulation by changing microhabitat selection would likely have been of limited utility because body temperatures would have responded only slowly to changes in Te.

4. 4. Endothermic metabolic rates may have put large dinosaurs at risk for overheating unless they had adaptations to shed the heat as necessary. This would have been particularly true for dinosaurs with masses > 10,000 kg, but simulations suggest that for animals as small as 1000 kg in the Tropics and in temperate latitudes during the summer, steady-state body temperatures would have exceeded 40°C. Slow response of body temperatures to changes in Te suggests that use of day-night thermal differences would have buffered dinosaurs from diel warming but would not have lowered body temperatures sufficiently for animals experiencing high mean daily Te.

5. 5. Endothermic metabolism and metabolic heating might have been useful for intermediate and large-sized (100–3000 kg) dinosaurs but often in situations that demanded marked seasonal adjustment of metabolic rates and/or precise control of metabolism (and heat-loss mechanisms) as typically seen in endotherms.

The photosynthetic gas exchange capacities of early angiosperms remain enigmatic. Nevertheless, many hypotheses about the causes of early angiosperm success and how angiosperms influenced Mesozoic ecosystem function hinge on understanding the maximum capacity for early angiosperm metabolism. We applied structure-functional analyses of leaf veins and stomatal pore geometry to determine the hydraulic and diffusive gas exchange capacities of Early Cretaceous fossil leaves. All of the late Aptian—early Albian angiosperms measured possessed low vein density and low maximal stomatal pore area, indicating low leaf gas exchange capacities in comparison to modern ecologically dominant angiosperms. Gas exchange capacities for Early Cretaceous angiosperms were equivalent or lower than ferns and gymnosperms. Fossil leaf taxa from Aptian to Paleocene sediments previously identified as putative stem-lineages to Austrobaileyales and Chloranthales had the same gas exchange capacities and possibly leaf water relations of their living relatives. Our results provide fossil evidence for the hypothesis that high leaf gas exchange capacity is a derived feature of later angiosperm evolution. In addition, the leaf gas exchange functions of austrobaileyoid and chloranthoid fossils support the hypothesis that comparative research on the biology of living basal angiosperm lineages reveals genuine signals of Early Cretaceous angiosperm ecophysiology.

Mammalian carnivores are rarely incorporated in paleoenvironmental reconstructions, largely because of their rarity within the fossil record. However, multivariate statistical modeling can be successfully used to quantify specific anatomical features as environmental predictors. Here we explore morphological variability of the humerus in a closely related group of predators (Felidae) to investigate the relationship between morphometric descriptors and habitat categories. We analyze linear measurements of the humerus in three different morphometric combinations (log-transformed, size-free, and ratio), and explore four distinct ways of categorizing habitat adaptations. Open, Mixed, and Closed categories are defined according to criteria based on traditional descriptions of species, distributions, and biome occupancy. Extensive exploratory work is presented using linear discriminant analyses and several fossils are included to provide paleoecological reconstructions.

We found no significant differences in the predictive power of distinct morphometric descriptors or habitat criteria, although sample splitting into small and large cat guilds greatly improves the stability of the models. Significant insights emerge for three long-canine cats: Smilodon populator, Paramachairodus orientalis, and Dinofelis sp. from Olduvai Gorge (East Africa). S. populator and P. orientalis are both predicted to have been closed-habitat adapted taxa. The false “sabertooth” Dinofelis sp. from Olduvai Gorge is predicted to be adapted to mixed habitat. The application of felid humerus ecomorphology to the carnivoran record of Olduvai Gorge shows that the older stratigraphic levels (Bed I, 1.99–1.79 Ma) included a broader range of environments than Beds II or V, where there is an abundance of cats adapted to open environments.

The extinct shark Carcharocles megalodon is one of the largest marine apex predators ever to exist. Nonetheless, little is known about its body-size variations through time and space. Here, we studied the body-size trends of C. megalodon through its temporal and geographic range to better understand its ecology and evolution. Given that this species was the last of the megatooth lineage, a group of species that shows a purported size increase through time, we hypothesized that C. megalodon also displayed this trend, increasing in size over time and reaching its largest size prior to extinction. We found that C. megalodon body-size distribution was left-skewed (suggesting a long-term selective pressure favoring larger individuals), and presented significant geographic variation (possibly as a result of the heterogeneous ecological constraints of this cosmopolitan species) over geologic time. Finally, we found that stasis was the general mode of size evolution of C. megalodon (i.e., no net changes over time), contrasting with the trends of the megatooth lineage and our hypothesis. Given that C. megalodon is a relatively long-lived species with a widely distributed fossil record, we further used this study system to provide a deep-time perspective to the understanding of the body-size trends of marine apex predators. For instance, our results suggest that (1) a selective pressure in predatory sharks for consuming a broader range of prey may favor larger individuals and produce left-skewed distributions on a geologic time scale; (2) body-size variations in cosmopolitan apex marine predators may depend on their interactions with geographically discrete communities; and (3) the inherent characteristics of shark species can produce stable sizes over geologic time, regardless of the size trends of their lineages.

Inferring the causes for change in the fossil record has been a persistent problem in evolutionary biology. Three independent lines of evidence indicate that a lineage of the fossil stickleback fish Gasterosteus doryssus experienced directional natural selection for reduction of armor. Nonetheless, application to this lineage of three methods to infer natural selection in the fossil record could not exclude random process as the cause for armor change. Excluding stabilizing selection and genetic drift as the mechanisms for biostratigraphic patterns in the fossil record when directional natural selection was the actual cause is very difficult. Biostratigraphic sequences with extremely fine temporal resolution among samples and other favorable properties must be used to infer directional selection in the fossil record.