The aim of this analysis was to establish the basic mechanical principles of simple archosaur cranial form. In particular we estimated the influence of two key archosaur innovations, the secondary palate and the antorbital fenestra, on the optimal resistance of biting-induced loads. Although such simplified models cannot substitute for more complex cranial geometries, they can act as a clearly derived benchmark that can serve as a reference point for future studies incorporating more complex geometry. We created finite element (FE) models comprising either a tall, domed (oreinirostral) snout or a broad, flat (platyrostral) archosaur snout. Peak von Mises stress was recorded in models with and without a secondary palate and/or antorbital fenestra after the application of bite loads to the tooth row. We examined bilateral bending and unilateral torsion-inducing bites for a series of bite positions along the jaw, and conducted a sensitivity analysis of material properties. Pairwise comparison between different FE morphotypes revealed that oreinirostral models are stronger than their platyrostral counterparts. Oreinirostral models are also stronger in bending than in torsion, whereas platyrostral models are equally susceptible to either load type. As expected, we found that models with a fenestra always have greatest peak stresses and by inference are “weaker,” significantly so in oreinirostral forms and anterior biting platyrostral forms. Surprisingly, although adding a palate always lowers peak stress, this is rarely by large magnitudes and is not significant in bilateral bending bites. The palate is more important in unilateral torsion-inducing biting. Two basic principles of archosaur cranial construction can be derived from these simple models: (1) forms with a fenestra are suboptimally constructed with respect to biting, and (2) the presence or absence of a palate is significant to cranial integrity in unilaterally biting animals. Extrapolating these results to archosaur cranial evolution, it appears that if mechanical optimization were the only criterion on which skull form is based, then most archosaurs could in theory strengthen their skulls to increase resistance to biting forces. These strengthened morphotypes are generally not observed in the fossil record, however, and therefore archosaurs appear subject to various non-mechanical morphological constraints. Carnivorous theropod dinosaurs, for example, may retain large suboptimal fenestra despite generating large bite forces, owing to an interplay between craniofacial ossification and pneumatization. Furthermore, living crocodylians appear to strengthen their skull with a palate and filled fenestral opening in the most efficient way possible, despite being constrained perhaps by hydrodynamic factors to the weaker platyrostral morphotype. The future challenge is to ascertain whether these simple predictions are maintained when the biomechanics of complex cranial geometries are explored in more detail.

Analysis of two independent data sets with increased taxonomic resolution (genera rather than families) using the revised 2012 timescale reveals that an extinction periodicity first detected by Raup and Sepkoski (1984) for only the post-Paleozoic actually runs through the entire Phanerozoic. Although there is not a local peak of extinction every 27 Myr, an excess of the fraction of genus extinction by interval follows a 27-Myr timing interval and differs from a random distribution at the p ∼ 0.02 level. A 27-Myr periodicity in the spectrum of interval lengths no longer appears, removing the question of a possible artifact arising from it. Using a method originally developed in Bambach (2006) we identify 19 intervals of marked extinction intensity, including mass extinctions, spanning the last 470 Myr (and with another six present in the Cambrian) and find that ten of the 19 lie within ±3 Myr of the maxima in the spacing of the 27-Myr periodicity, which differs from a random distribution at the p = 0.004 level. These 19 intervals of marked extinction intensity also preferentially occur during decreasing diversity phases of a well-known 62-Myr periodicity in diversity (16 of 19, p = 0.002). Both periodicities appear to enhance the likelihood of increased severity of extinction, but the cause of neither periodicity is known. Variation in the strength of the many suggested causes of extinction coupled to the variation in combined effect of the two different periodicities as they shift in and out of phase is surely one of the reasons that definitive comparative study of the causes of major extinction events is so elusive.

Unfossiliferous marine sedimentary rocks of Phanerozoic age are known to all field-oriented paleontologists. These troublesome units are often encountered in the field, perhaps cursed roundly for a moment or two, and usually shrugged off in pursuit of the next fossiliferous interval. Paleontologists tend not to discuss barren units, and they rarely publish on the absence of a fauna from what appears to be unaltered marine rock. But aren't barren marine sediments revealing something important about their paleoenvironment and possibly about the paleoenvironments of conformably adjacent fossil-bearing units? Shouldn't paleontologists be just as interested in knowing the locations and ages of unfossiliferous sediments as they are fossiliferous strata?

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.

The carbon stable isotope (δ13C) composition of the calcitic tests of planktonic foraminifera has an important role as a geochemical tracer of ocean carbon system changes associated with the Cretaceous/Paleogene (K/Pg) mass extinction event and its aftermath. Questions remain, however, about the extent of δ13C isotopic disequilibrium effects and the impact of depth habitat evolution on test calcite δ13C among rapidly evolving Paleocene species, and the influence this has on reconstructed surface-to-deep ocean dissolved inorganic carbon (DIC) gradients. A synthesis of new and existing multispecies data, on the relationship between δ13C and δ18O and test size, sheds light on these issues. Results suggest that early Paleocene species quickly radiated into a range of depths habitats in a thermally stratified water column. Negative δ18O gradients with increasing test size in some species of Praemurica suggest either ontogenetic or ecotypic dependence on calcification temperature that may reflect depth/light controlled variability in symbiont photosynthetic activity. The pattern of positive δ13C test-size correlations allows us to (1) identify metabolic disequilibrium δ13C effects in small foraminifera tests, as occur in the immediate aftermath of the K/Pg event, (2) constrain the timing of evolution of foraminiferal photosymbiosis to 63.5 Ma, ∼0.9 Myr earlier than previously suggested, and (3) identify the apparent loss of symbiosis in a late-ranging morphotype of Praemurica. These findings have implications for interpreting δ13C DIC gradients at a resolution appropriate for incoming highly resolved K/Pg core records.

Attempts at understanding evolutionary relationships among Paleozoic Dipnoi (lungfish) using cladistic methodology have proved totally unsatisfactory (Miles 1977; Marshall 1987). We attempt to reconstruct the relationships between the better known genera using a method that involves the recognition of lineages based on evolving functional complexes, particularly those involved with food reduction and respiration. Within these broadly defined lineages, we have defined sub-lineages based on evolutionary patterns shown by structures that have been stratigraphically dated; such patterns are found inter alia in the roofing bones and the external dermal bones of the mandible. A number of new suborders and families are recognised; genera for which further morphological data are required before they can be assigned to a higher taxon are indicated; two generic synonyms are recognised.

In appendices, short descriptions are given of two new genera—Pillararhynchus from the Gogo Formation (Upper Devonian) of Western Australia, and Sorbitorhynchus from the Emsian of Guangxi, China.

Analysis of an exhaustive data base of Namurian ammonoid shell characters indicates that the morphology of the Goniatitida can be explained in terms of functional constraints, resulting particularly from hydrostatic and hydrodynamic properties. Modes of life ranging from benthic to pelagic are inferred on this basis for various goniatitid morphotypes; all morphologic features and facies associations are normally compatible with these inferences. Neutral buoyancy is shown to have been likely for all goniatitids. By contrast, the prolecanitids (Order Agoniatitida) exhibit a number of hydrostatic and morphologic anomalies; these anomalies are not explicable using the same principles and remain problematic. This is noteworthy, in that prolecanitids survived the Permian/Triassic extinctions and gave rise to the diverse ceratitic radiation in the Triassic.

The applicability of these results to ammonoids outside the Namurian is assessed, and, in particular, morphologic parallels with Mesozoic ammonites are discussed.

Scanning electron microscope (SEM) examination of bone surfaces from the Pleasant Lake mastodon, excavated in southern Michigan, documents features indicative of butchery. These features are identified by comparison with modern bones modified by human and natural processes. We report new studies of (1) marks made by bone tools during removal of meat from and disarticulation of carcasses and (2) use wear developed on bone tools. We also apply previously developed criteria for recognizing stone tool cutmarks and stages in the burning of bone. The Pleasant Lake site, dated to between 10,395 ± 100 and 12,845 ± 165 b.p., provides compelling evidence of mastodon butchery and bone tool use. Another site, near New Hudson, Michigan, provides replication of much of this evidence. Together these sites offer new examples of patterns of bone modification and extend the geographic and temporal representation of the much discussed, but still controversial, late Pleistocene bone technology.

The punctuated equilibrium hypothesis, largely derivable from Mayr's peripheral isolate model of speciation, is but one of many possible paleontologic patterns of speciation which may be tested against the deep-sea fossil record of planktonic microfossils. The nature of reproduction and variation within a species has a profound effect on its expected tempo and mode of evolutionary change. Biologic data on pelagic plankton and on protistan genetics and reproduction suggest that speciation in pelagic holoplanktonic protists may also be parapatric, “equal” allopatric, or the result of hybridization. Each of these models is shown to make testable predictions of paleontologic pattern. Published records of speciation in planktonic deep-sea microfossil data are compatible with these alternative models. Existing data sets are not yet sufficiently complete to provide strong tests. Future work needs to integrate taxonomic, paleoecologic, biogeographic, and biometric data in studies of individual speciation events. Micropaleontologic studies of morphologic change through time may thereby contribute to our understanding of speciation mechanisms in the pelagic protistan plankton.

Taxonomic diversity and morphological disparity are different measures of biodiversity that together can describe large-scale evolutionary patterns. Diversity measures are often corrected by extending lineages back in time or adding additional taxa necessitated by a phylogeny, but disparity analyses focus on observed taxa only. This is problematic because some morphologies required by phylogeny are not included, some of which may help fill poorly sampled time bins. Moreover the taxic nature of disparity analyses makes it difficult to compare disparity measures with phylogenetically corrected diversity or morphological evolutionary rate curves. We present a general method for using phylogeny to correct measures of disparity, by including reconstructed ancestors in the disparity analysis. We apply this method to discrete character data sets focusing on Triassic archosaurs, Cenozoic carnivoramorph mammals, and Cretaceous-Cenozoic euarchontogliran mammals. Phylogenetic corrections do not simply mirror the taxic disparity patterns, but affect the three analyses in heterogeneous ways. Adding reconstructed ancestors can inflate morphospace, and the amount and direction of expansion differs depending on the taxonomic group in question. In some cases phylogenetic corrections give a temporal disparity curve indistinguishable from the taxic trend, but in other cases disparity is elevated in earlier time intervals relative to later bins, due to the extension of unsampled morphologies further back in time. The phylogenetic disparity curve for archosaurs differs little from the taxic curve, supporting a previously documented pattern of decoupled disparity and rates of morphological change in dinosaurs and their early contemporaries. Although phylogenetic corrections should not be used blindly, they are helpful when studying clades with major unsampled gaps in their fossil records.

Living echinoids provide a structural analogue to the rigid thecae of many extinct echinoderms and are here used to examine some consequences of the increased rigidity and decreased number of plates which occurred in several lineages of echinoderms. A special feature of the echinoids is the large spines which protect the echinoid tests from failure under impact. The spines absorb energy or spread loads because the spines break or the tissue connecting the spine to the test is stretched or torn. Spines that absorb impacts might be superfluous in echinoderms encased in a rigid test or theca if they have flexible stems or are in low energy habitats, but spines should be useful to stemless thecate echinoderms on hard substrata. In the absence of spines, a flexible theca of overlapping plates might be superior to a rigid theca. Greater rigidity does not necessarily provide greater protection.

Tracts of podial pores are sites of weakness in all tests examined [Eucidaris tribuloides (Lamarck), Echinometra viridis Agassiz, Diadema antillarum Philippi, Lytechinus variegatus (Lamarck), Tripneustes ventricosus (Lamarck)]. Therefore arrangement of pores affects strength of rigid thecae. Sutures are stronger than plates in most species examined, but cracks cross sutures, and tests of E. tribuloides frequently break along sutures. It is not yet proven that sutures increase toughness of rigid tests by relieving stress on plates or stopping cracks. If resisting failure under impact is important, then preconditions for acquiring rigidity are life in a low energy habitat or accessory structures which absorb or spread impacts.

Digitate shell morphologies have evolved repeatedly in planktonic foraminifera throughout the Cretaceous and Cenozoic. Digitate species are usually rare in fossil and modern assemblages but show increased abundance and diversity at times during the Cretaceous and middle Eocene. In this paper we discuss the morphology and stratigraphic distribution of digitate planktonic foraminifera and establish the isotopic depth ecology of fossil ones to draw parallels with modern counterparts. δ18O and δ13C values of six extinct and two modern digitate species, from six time slices (Cenomanian, Turonian, Eocene, Miocene, Pleistocene and Holocene) have similar isotopic depth ecologies, consistently registering the most negative δ13C and usually the most positive δ18O compared to coexisting species. These results indicate a similar deep, subthermocline (é150 m) habitat, characterized by lower temperatures, reduced oxygen, and enrichment of dissolved inorganic carbon. This is consistent with water-column plankton studies that provide insight into the depth preferences of the three modern digitate species; in over 70% of observations digitates occurred in nets below 150 m, and down to 2000 m. The correlation between digitate species and subsurface habitats across multiple epochs suggests that elongated chambers were advantageous for survival in a deep mesopelagic habitat, where food is usually scarce. Increased abundance and diversity of digitates in association with some early and mid-Cretaceous oceanic anoxic events, in middle Eocene regions of coastal and equatorial upwelling, and occasionally in some modern upwelling regions, suggests an additional link with episodes of enhanced ocean productivity associated with expansion of the oxygen minimum zone (OMZ). We suggest that the primary function of digitate chambers was as a feeding specialization that increased effective shell size and food gathering efficiency, for survival in a usually food-poor environment, close to the OMZ. Episodes of increased digitate abundance and diversity indicate expansion of the deep-water ecologic opportunity under conditions that were unfavorable to other planktonic species. Our results provide evidence of iterative evolution reflecting common functional constraints on planktonic foraminifera shell morphology within similar subsurface habitats. They also highlight the potential of digitate species to act as indicators of deep watermasses, especially where there was expansion of the OMZ.

Because of the fragmentary preservation of the earliest Cooksonia-like terrestrial plant macrofossils, younger Devonian fossils with complete anatomical preservation and documented gametophytes often have received greater attention concerning the early evolution of vascular plants and the alternation of generations. Despite preservational deficits, however, possible physiologies of Cooksonia-like fossils can be constrained by considering the overall axis size in conjunction with the potential range of cell types and sizes, because their lack of organ differentiation requires that all plant functions be performed by the same axis. Once desiccation resistance, support, and transport functions are taken into account, smaller fossils do not have volume enough left over for an extensive aerated photosynthetic tissue, thus arguing for physiological dependence on an unpreserved gametophyte. As in many mosses, axial anatomy is more likely to have ensured continued spore dispersal despite desiccation of the sporophyte than to have provided photosynthetic independence. Suppositions concerning size constraints on physiology are supported by size comparisons with fossils of demonstrable physiological independence, by preserved anatomical detail, and by size correlations between axis, sporangia, and sporangial stalk in Silurian and Early Devonian taxa. Several Cooksonia-like taxa lump fossils with axial widths spanning over an order of magnitude—from necessary physiological dependence to potential photosynthetic competence—informing understanding of the evolution of an independent sporophyte and the phylogenetic relationships of early vascular plants.

The extinction of endemic brachiopods in North America at the end of the Ordovician and recolonization by European species has been related to glacio-eustatic lowering of sea level which disrupted conditions in epicontinental seas. North American species may have been narrowly adapted to relatively stable conditions of broad, tropical shallow seas. European invaders may have been less specialized because they were adapted to conditions in both the open ocean and in narrow European epicontinental seas. Being less narrowly adapted, European species probably were better able to cope with changing environmental conditions than were North American species.

During the Lower and Middle Llandovery, shallow water, low diversity communities of Pentamerus Community depth were unstable and characterized by repeated extinctions and invasions. Following the crisis at the Ordovician-Silurian boundary 3 to 5 million years were needed to reestablish communities that were persistent in geologic time.

To understand how well fossil assemblages represent original communities, paleoecologists seek comparisons between death assemblages and their source communities. These comparisons have traditionally used nearshore, marine invertebrate assemblages for their logistical ease, high abundance, and comparable census data from living communities. For large marine vertebrates, like cetaceans, measuring their diversity in ocean ecosystems is difficult and expensive. Cetaceans, however, often beach or strand themselves along the coast, and archived data on stranded cetaceans have been recorded, in some areas, over several decades. If the stranding record is interpreted as a death assemblage, then the stranding record may represent a viable alternative for measuring diversity in living communities on directly adjacent coastlines. This study assessed the fidelity of the cetacean stranding record in the eastern North Pacific Ocean. The living community in this region has been studied for over 100 years and, recently, extensive and systematic live transect surveys using ship-based observing platforms have produced a valuable source of live diversity data. Over this same period, the U.S. Marine Mammal Stranding Program has collected and archived a record of cetacean strandings along the U.S. Pacific coastline, providing an ideal death assemblage for comparison. Using fidelity metrics commonly used in marine invertebrate taphonomy, I determined that the stranding record samples the living cetacean community with high fidelity, across fine and coarse taxonomic ranks, and at large geographic scales (>1000 km of coastline). The stranding record is also richer than the live surveys, with live-dead ratios between 1.1 and 1.3. The stranding record recovers similar rank-order relative abundances as live surveys, with statistical significance. Also, I applied sample-based rarefaction methods to generate collector's curves for strandings along the U.S. Pacific Coast to better evaluate the spatiotemporal characteristics of the stranding record. Results indicate that saturation (i.e., sampling >95% assemblage) at species, genus, and family levels occurs in less than five years of sampling, with families accumulating faster than species, and larger geographic regions (i.e., longer coastlines) accumulating taxa the most rapidly. The high fidelity of the stranding record, measured both in richness and by ranked relative abundance, implies that ecological structure from living cetacean communities is recorded in the death assemblage, a finding that parallels marine invertebrate assemblages, though at far larger spatial scales. These results have implications for studying cetacean ecology in both modern and ancient environments: first, these results imply that the stranding record, over sufficiently long time intervals, yields a richer assemblage than using line-transect methods, and faithfully records aspects of community structure; and second, these results imply that geochronologically well-constrained fossil cetacean assemblages might preserve ecologically relevant features of community structure, depending on depositional and taphonomic conditions.

Late Eocene to middle Oligocene floras from the area of the present Rio Grande depression represent a time sequence from the upper part of mixed subtropical forest (Bernalillo flora), to an ecotone between broadleaved sclerophyll and mixed conifer forest (Red Rock Ranch flora), to subalpine conifer forest (Hillsboro and Hermosa floras). The implied difference in mean annual temperature of ~ 11°C suggests that altitude increased 2,000 m in 6–8 my. Construction of the Datil-Mogollon volcanic pile, averaging about 1,200 to 1,500 m thick, is thought to be largely responsible for the forest zonation, but regional doming accompanying volcanism may also be involved. The Oligocene subalpine conifer forests now occur in the piñon-juniper belt 900 and 1,200 m below the present subalpine zone, consistent with the subsidence that formed the Rio Grande rift beginning in the late Oligocene/early Miocene and continuing to the present. Later epeirogenic uplift of ~ 1200 m is implied by fossil plants in the Galisteo (~ 40 my bp), Creede (~ 27 my bp) and Tesuque (~ 14 my bp) Formations that border, or are in the rift.

Brachiopods generally have not been considered to be typical or significant faunal components of modern or ancient hydrothermal vent and cold-seep settings. The Early Cretaceous (Neocomian) rhynchonellide brachiopod Peregrinella has long been viewed as a paleontological curiosity because of its distinctive morphology, status as the largest Mesozoic brachiopod, anomalous stratigraphic associations, and widespread, yet discontinuous paleogeographic distribution. Examination of all worldwide Peregrinella occurrences (14) indicates restriction of this brachiopod to ancient cold-seeps. It is probable that Peregrinella grew to large sizes in such great abundances at fossil cold-seep sites because of a richly organic food supply generated by localized fluid seepage and bacterial chemosynthetic activity. Living brachiopods are not known to harbor chemosymbiotic bacteria in their tissues; however, direct chemoautotrophic utilization of reduced fluids by Peregrinella cannot be rejected or demonstrated at present. Peregrinella occurs at widely separated cold-seeps of Neocomian age (e.g., California, Mexico, Tibet, Europe), yet its mode of dispersal and larval development is unknown. In modern hydrothermal vents of the deep-sea, organism dispersal occurs along oceanic ridges, where benthic faunas display both planktotrophic and nonplanktotrophic larval-mode types. Peregrinella may represent a Mesozoic relic of a long-lived “lineage” of vent-seep associated rhynchonellides from the Paleozoic (e.g., ?Eoperegrinella, Dzieduszyckia), but major gaps in the stratigraphic record between these rhynchonellide occurrences, and the lack of rigorous phylogenetic analysis for these groups preclude a clear resolution of the origin(s) of vent-seep brachiopods at present.

This paper presents scaling equations relating suspension-feeding rates to body size for articulate brachiopods and bivalve molluscs, two classes which represent a significant component of the fossil record of marine benthic communities. Clearance (feeding) rates of five species of living articulate brachiopods and three species of epifaunal suspension-feeding bivalve molluscs collected from mid-latitude fjords of Newfoundland and New Zealand were measured in similar experimental conditions. In comparisons within and between the two classes, we found that both plectolophous and spirolophous brachiopods had significantly lower feeding rates than mytilids, which are filibranchs, but that a sympatric primitive eulamellibranch veneroid bivalve had rates comparable to the brachiopods. Articulate brachiopods do not appear to feed effectively at the high algal concentrations which bivalves can exploit. The data on comparative suspension-feeding rates support the hypothesis that past changes in diversity and distribution of bivalves and brachiopods may be related to an overall increase in energy flux and escalation of metabolic rates during the Phanerozoic.

The significant impact of extant carnivores, particularly spotted hyenas, on the depositional history and physical characteristics of archaeofaunal and paleontological assemblages is well recognized. We focus on the behavioral ecology of extant spotted hyenas (Crocuta crocuta) in relation to bone accumulations produced by one East African clan at communal dens. Limbs and skulls of prey animals more frequently appear at dens than do other carcass portions. These items reflect the relative abundance of prey species near dens; carnivore remains are poorly represented. Comparative analysis reveals that bones are deposited far more slowly (<7 carcass portions per month) and accumulations tend to be smaller at Crocuta dens than at dens of either brown (Parahyaena brunnea) or striped (Hyaena hyaena) hyenas. We propose that extant Crocuta bone accumulation rates and sizes are likely affected by prey species abundance, clan size, social interactions within the clan, and the type and availability of den sites. We also suggest that the potential for intraspecific behavioral variability in bone accumulation patterns is important when comparisons are made among spotted hyena populations and across hyena species. For example, accumulation patterns may be dramatically influenced by the temporal span, potentially ranging from days to hundreds or thousands of years, in which bones are collected, depending on the species-specific history of occupation at a given site. Understanding the behavioral and ecological variability likely to influence bone accumulation patterns at dens used by different hyaenids will allow taphonomists and zooarchaeologists to refine their knowledge of mechanisms underlying site formation processes and potential causes of variability in deeper-time den assemblages.

Morphologic discrimination of species of scleractinian reef corals has long been plagued by a shortage of independent characters and by high ecophenotypic plasticity. Because of these two factors, many species appear to intergrade morphologically. We outline a newly developed protocol for the morphometric recognition of species, which uses size and shape coordinates derived from landmark data. The landmarks consist of spatially homologous points digitized in three dimensions on upper calical surfaces. The approach is more powerful than linear measurements at detecting subtle distinctions among species; and the distinctions are easy to visualize and interpret biologically, which increases the accuracy and resolution of subsequent phylogenetic and large-scale faunal analyses.

As an example, we distinguish morphospecies in collections of Porites made at three Caribbean locations. Size and shape coordinates are analyzed using principal component analysis, average linkage cluster analysis, and a series of iterative discriminant analyses. Positions of different corallites from the same colonies are examined on cluster dendrograms to determine cutoffs for group recognition, and discriminant classifications for different corallites from the same colonies are compared to maximize group assignments. The results yield seven morphospecies, which are generally in 90% agreement with classification of the same animals using allozyme electrophoresis. Measures of corallite size and the relative heights and locations of the pali and septal denticles all reveal unique patterns of variation among morphospecies.